Desert Puma - Evolutionary Ecology and Conservation

About Island Press
Island Press is the only nonprofit organization in the United States whose prin-
cipal purpose is the publication of books on environmental issues and natural
resource management. We provide solutions-oriented information to profes-
sionals, public officials, business and community leaders, and concerned citizens
who are shaping responses to environmental problems.
In 2001, Island Press celebrates its seventeenth anniversary as the leading
provider of timely and practical books that take a multidisciplinary approach to
critical environmental concerns. Our growing list of titles reflects our commit-
ment to bringing the best of an expanding body of literature to the environ-
mental community throughout North America and the world.
Support for Island Press is provided by The Bullitt Foundation, The Mary
Flagler Cary Charitable Trust, The Nathan Cummings Foundation, Geraldine
R. Dodge Foundation, Doris Duke Charitable Foundation, The Charles Engel-
hard Foundation, The Ford Foundation, The George Gund Foundation, The
Vira I. Heinz Endowment, The William and Flora Hewlett Foundation, W.
Alton Jones Foundation, The John D. and Catherine T. MacArthur Foundation,
The Andrew W. Mellon Foundation, The Charles Stewart Mott Foundation,
The Curtis and Edith Munson Foundation, National Fish and Wildlife Foun-
dation, The New-Land Foundation, Oak Foundation, The Overbrook Founda-
tion, The David and Lucile Packard Foundation, The Pew Charitable Trusts,
Rockefeller Brothers Fund, The Winslow Foundation, and other generous
donors.
About the Hornocker Wildlife Institute

The Hornocker Wildlife Institute was founded by Maurice Hornocker in 1985
as an independent, nonprofit organization with the mission to conduct long-
term ecological research, support graduate and post-graduate students, and dis-
seminate information to the public and to the agencies responsible for conser-
vation and management of wildlife and wildlands. In the year 2000, the institute
became part of the Wildlife Conservation Society, known for its field science
and conservation projects around the world for more than one hundred years.
The institute now forms the core of the WCS Global Carnivore Program, which
directs and advises carnivore research and conservation efforts worldwide from
its offices in Bozeman, Montana. Desert Puma is the first in a series of publica-
tions on long-term scientific projects under the HWI/WCS banner.
Desert Puma
To our son Oren,
and our parents
Rita and James Logan,
Joan and George Sweanor
Desert Puma
E v o l u t i o n a ry E co l o g y
a n d Co n s e r v a t i o n o f a n
E n d u r i n g Ca r n i v o re
Kenneth A. Logan
and Linda L. Sweanor
HORNOCKER WILDLIFE INSTITUTE,

A Program of the Wildlife Conservation So c i e t y
Washington • Covelo • London

Copyright © 2001 Kenneth A. Logan and Linda L. Sweanor
All rights reserved under International and Pan-American Copyright Conventions. No

part of this book may be reproduced in any form or by any means without permission
in writing from the publisher: Island Press, 1718 Connecticut Avenue, N.W., Suite
300, Washington, DC 20009.
ISLAND PRESS is a trademark of The Center for Resource Economics.
Library of Congress Cataloging-in-Publication Data
Logan, Kenneth A.
Desert Puma : evolutionary ecology and conservation of an enduring
carnivore / Kenneth A. Logan and Linda L. Sweanor.
p. cm.
Includes bibliographical references (p. ).
ISBN 1-55963-866-4 (cloth : alk. paper) — ISBN 1-55963-867-2 (paper :
alk. paper)
1. Puma—New Mexico. 2. Desert animals—New Mexico. I. Sweanor,
Linda L. II. Title.
QL737.C23 L64 2001
599.75’24’097896—dc21
2001003356
British Cataloguing-in-Publication Data available.
Printed on recycled, acid-free paper
Manufactured in the United States of America

10 9 8 7 6 5 4 3 2 1
Contents
List of Figures, Tables, and Photos xv

Foreword xxiii
Acknowledgments xxvii
Part 1. Setting the Stage

Chapter 1. Introduction 3
Rationale and Approach 5
Synopsis of Previous Studies 6
Chapter 2. Pumas Past and Present 9
Puma Phylogeny 9
Puma Distribution and Status 15
Pumas in the Southwest 16
Taxonomy 16
Description 18
Statistic 21
Chapter 3. Our Outdoor Laboratory 23
Physiography and Geology 26
Climate 27
Flora 29
Fauna 30
History of Human Use 33
History of Puma Exploitation 36
vii
viii Contents
Chapter 4. Studying Wild Pumas 39

Life Afield 39
To Catch a Puma 42
Monitoring Desert Mule Deer and Desert Bighorn Sheep 51
Radiotelemetry 54
Investigating the Dead 58
Overview of Analytical and Statistical Methods 59
Part 2. Puma Life History Strategies and

Population Dynamics
Chapter 5. A Puma Population in the Desert 63
Research Hypotheses and Predictions 63
Terms for Pumas 64
Counting Pumas 65
Chapter 6. Puma Population Structure 69
Sex Structure 69
Cubs 69 • Subadults 73 • Adults 73
Age Structure 75
The Population 75 • Adult Pumas 78
Summary 80
Statistics 81
Chapter 7. Reproduction 83
Natality 83
Timing of Births 88
Mating, Gestation, and Birth Intervals 91
Puberty and First Litters 93
Parental Investment 96
Reproductive Success 96
Females 98 • Males 103
Summary 110
Statistics 111
Contents ix
Chapter 8. Mortality and Survival 115

Human-Caused Mortality 115
Natural Mortality 117
Cubs 117 • Subadults 122 • Adults 127
Why Do Pumas Kill Other Pumas? 139
Summary 142
Statistics 143
Chapter 9. Independence of Puma Progeny, and Philopatry, Emi-
gration, and Immigration 145
Independence of Progeny 146
Philopatry and Dispersal 148
Emigration 153
Recruitment of Progeny and Immigrants 154
Summary 154
Chapter 10. Puma Population Density, Growth, and
Metapopulation Structure 157
Experimentally Removing Pumas 157
Density 160
Rates of Population Increase 169
Metapopulation Dynamics 175
Summary 179
Part 3. Puma Behavior and Social Organization

Chapter 11. How Should Desert Pumas Behave? 183
Two-Strategies Hypothesis 183
Self-Limiting Hypothesis 184
Chapter 12. Adult Home Range Characteristics 189
Delineating the Home Range 189
Seasonal and Annual Home Range 191
Birth-Interval Home Range 192
x Contents
Lifetime Home Range 195

What Factors Influence Adult Home Range Size? 195
Home Range Size, Prey Abundance, and Puma Density 201
Does Prey Abundance Affect Home Range Size? 202 • Does Puma
Density Affect Home Range Size? 204 • Does Home Range Size
Reflect an Attempt at Population Self-Limitation? 210
Adult Home Range Fidelity 211
Method 1—Fidelity Index 213 • Method 2—Distances between
Mean Locations 214 • Home Range Shifts in Pumas 216 •
Homing by Translocated Pumas 222 • Benefits of Fidelity and the
Two-Strategies Hypothesis 223 • Fidelity in Desert Pumas and the
Self-Limiting Hypothesis 225
Summary 225
Statistic 227
Chapter 13. Subadult Ranging Behavior 231
Philopatry in Females 232
Female Dispersal 233
Male Dispersal 236
Frustrated Dispersal 239
Why Do Pumas Disperse? 240
Transient Behavior in Pumas 244
Summary 245
Statistic 246
Chapter 14. Interactions between Pumas 247
Spatial Relationships 247
Home Range Overlap Indices 249 • Nearest-Neighbor Analysis
254
Spatiotemporal Relationships 255
Direct Interactions 257
Associations between Independent Pumas 260 • Associations
between Family Members 267
Communication among Pumas 269
Vocalizations 269 • Chemical Communication 272
Contents xi
Summary 276
Statistics 277
Chapter 15. Adaptive Significance of Puma Social
Organization 281
The Social Structure of Desert Pumas 281
Female Structure 282 • Male Structure 283
The Self-Limiting Hypothesis 284
Land Tenure and Territoriality 285 • Do Desert Pumas Exhibit Land
Tenure or Territoriality? 286
The Two-Strategies Hypothesis 288
Female Strategy 288 • Male Strategy 290
Pumas and Other Big Cats—Similar Strategies? 294
Summary 297
Part 4. Puma–Prey Relationships

Chapter 16. Puma Diet 301
Patterns of Pumas and Prey 301
Puma Diet on the San Andres Mountains 302
Summary 308
Chapter 17. Pumas and Desert Mule Deer 311
Hypotheses, Predictions, and Terms 311
Characteristics of Dead Deer 312
Fates of Radio-Collared Deer 314
Mule Deer Population Dynamics 321
Puma Predation and Mule Deer Population Growth 322
Some Behavioral Interactions between Deer and Pumas 331
Did Puma Predation Limit the Deer Population? 332
Did Pumas Limit Their Own Density and Not Harm Their Food
Supply? 333
Cases of Pumas and Other Carnivores Limiting Prey
Populations 335
What Limits the Puma Population? 336
xii Contents
How Would a Puma Population Respond to a Prey Crash? 337

Summary 338
Statistics 339
Chapter 18. Pumas and Desert Bighorn Sheep 341
Hypothesis and Predictions 341
Pre-history, History, and Threats 342
Sheep Population Characteristics during Our Research 346
Fates of Radio-Collared Sheep 347
Survival Rates and Agent-Specific Mortality 350
Did Puma Predation Limit the Sheep Population? 354
Finale of the Sheep Population 354
Pumas and Other Sheep Populations 356
Summary 357
Statistic 357
Chapter 19. Synthesis: Pumas and Weather Modulate
Large-Mammal Population Dynamics on the San Andres
Mountains 359
Part 5. Pumas and People

Chapter 20. Conservation and Management of Wild
Pumas 365
Threats to Pumas 367
Habitat Loss 367 • Puma Overkill 371
Alleviating Threats 377
Habitat Conserv a t i on 378 • Preventing Unnecessary Ove rk i ll 383 •
Ad a p t i veManagement—Inv o lving People 384
Summary 395
Statistic 395
Chapter 21. Epilogue 397
Appendix 1. Morphological Measurements of Pumas At Least 3.5 Months Old on

the San Andres Mountains, New Mexico, 1985–1995 399
Contents xiii
Appendix 2. Reproductive Chronology of Mated Pairs of Pumas on the San

Andres Mountains, New Mexico, 1987–1994 413
Appendix 3. Methods and Estimates of Annual Home Range Size for Pumas on
the San Andres Mountains, New Mexico 415
Appendix 4. A Deterministic, Discrete Time Model That Simulated Mule Deer
Population Dynamics in the Treatment Area, San Andres Mountains, New
Mexico, 1987–1995. See Chapter 17 for its application. 419
References 423
Index 451
List of Figures, Tables, and Photos
Figures
3-1 The puma study area, San Andres Mountains, New Mexico. 24
3-2 Annual and growing season precipitation, San Andres Mountains,
New Mexico, 1985–1994. 28
4-1 Locations of resident adult pumas on the San Andres Mountains,
New Mexico, 1985–1995. 55
6-1 Masses of puma cubs, San Andres Mountains, New Mexico,
1986–1994. 71
6-2 Age structure of the puma population in the Treatment Area and
Reference Area each January, 1989–1995, San Andres Mountains,
New Mexico. 76
6-3 Adult puma age structure in the Treatment Area and Reference Area in
three broad age classes, San Andres Mountains, New Mexico. 80
7-1 Distribution of births of seventy-eight puma litters by month,
1986–1994, San Andres Mountains, New Mexico. 88
7-2 Scatter plot of the number of litters produced by adult female and
male pumas monitored for at least twelve months by radiotelemetry,
San Andres Mountains, New Mexico. 100
8-1 Puma cub survival rates from birth to thirteen months, San Andres
Mountains, New Mexico. 119
8-2 Causes of death in puma cubs, San Andres Mountains, New
Mexico. 121
8-3 Relationship of adult puma density to frequency of independent puma
deaths from intraspecies strife in the Treatment Area and Reference
Area, San Andres Mountains, New Mexico. 135
9-1 Dispersal moves of pumas from the San Andres Mountains to other
habitats in New Mexico. 151
xv
xvi List of Figures, Tables, and Photos
10-1 Observed exponential rates of increase for adult pumas in the

Treatment Area and Reference Area using January population
estimates, San Andres Mountains, New Mexico. 171
12-1 The means and standard deviations of annual, birth interval, and
lifetime home range sizes, based on the 90 percent adaptive kernel
home range estimator, for adult pumas on the San Andres Mountains,
New Mexico, 1986–1994. 192
12-2 Home range size for female pumas during three consecutive
cub-rearing periods on the San Andres Mountains, New Mexico,
1986–1994. 194
12-3 The home range of puma F15 during three consecutive cub-rearing
periods, San Andres Mountains, New Mexico. 194
12-4 Adult puma home range size correlated with adult puma density in five
study areas (Alberta, British Columbia, Idaho, New Mexico, and Utah)
in North America. 204
12-5 Changes in annual home range size and density of adult pumas in the
Treatment Area, San Andres Mountains, New Mexico. 207
12-6 Puma M3’s annual home range, 1988–1990 (a), and 1991–1993 (b), in
the Treatment Area, San Andres Mountains, New Mexico. 209
12-7 A simplified example of a fidelity index. 212
12-8 Frequency distribution of the fidelity index (FI1) for adult male and
female pumas, based on the 90 percent adaptive kernel home range
estimator, San Andres Mountains, New Mexico, 1986–1994. 214
12-9 Fidelity index (mean and standard deviation) for adult pumas on the
12-10 Examples of strong annual home range fidelity in adult pumas on the
12-11 Puma M36’s annual home ranges during six of seven consecutive years,
12-12 Examples of home range shifts exhibited by adult pumas over
consecutive years, San Andres Mountains, New Mexico. 217
14-1 Mean and standard deviation of the annual spatial overlap and
spatiotemporal overlap between pairs of same-sex adult pumas on the
14-2 Number of associations between independent pumas that were related
to breeding, non-breeding, or unknown activities on the San Andres
Mountains, New Mexico, 1985–1995. 263
14-3 Home ranges for adult pumas M19 and M29 in 1991, San Andres
17-1. Rates of survival, puma predation, and death from other causes for
radio-collared mule deer, San Andres Mountains, New Mexico. 318
List of Figures, Tables, and Photos xvii
17-2 Desert mule deer population trend on the San Andres Mountains, New
Mexico. 325
17-3 Relationship of deer population growth rates to puma predation rates
on radio-collared deer, San Andres Mountains, New Mexico. 326
17-4 Relationship of deer population growth rates to fawn production, San
Andres Mountains, New Mexico. 327
17-5 Relationship of adult puma numbers to predation rates on radio-
collared deer, San Andres Mountains, New Mexico. 328
17-6 Mule deer and puma population trends on the San Andres Mountains,
New Mexico. 334
18-1 Number of adult and yearling desert bighorn sheep observed in annual
surveys and radio-collared during each year, San Andres Mountains,
New Mexico. 348
18-2 Rates of survival, puma predation, and deaths from other causes for
radio-collared desert bighorn sheep, San Andres Mountains, New
Mexico. 354
20-1 Trends in number of puma-hunting permits issued and number of
pumas killed by hunters in New Mexico. 373
20-2 A hypothetical zone-management approach for pumas in New
Mexico. 385
A3-1 Home range sizes of pumas M5 and F45 plotted against the number of
locations used to determine their size. 416
Tables
4-1 Summary of puma snare capture efforts on the San Andres Mountains,
New Mexico, 1985–1995. 46
5-1 Percent accuracy for the estimated number of adult pumas per year,
1986–1994, in the Treatment Area and Reference Area, San Andres
6-1 Sex ratios of adult pumas in the Treatment Area and Reference Area
each year, 1988–1994, San Andres Mountains, New Mexico. 74
6-2 Sex ratios of adult pumas in the Treatment Area and Reference Area
each January, 1989–1995, San Andres Mountains, New Mexico. 75
6-3 Proportion of pumas in three broad age classes in the Treatment Area
population each January, 1989–1995, San Andres Mountains, New
Mexico. 77
6-4 Proportion of pumas in three broad age classes in the Reference Area
population each January, 1989–1995, San Andres Mountains, New
Mexico. 77
xviii List of Figures, Tables, and Photos
6-5 Mean ages of adult pumas in the Treatment Area and Reference Area
each January, 1989–1995, San Andres Mountains, New Mexico. 78
8-1 Annual survival rates for adult pumas, 1987–1994, San Andres
8-2 Number, percentage, and ages of adult pumas that died of natural causes
on the San Andres Mountains, New Mexico, 1985–1995. 130
9-1 Dispersal distances for independent pumas born on the San Andres
Mountains, New Mexico, and for pumas with origins outside the San
Andres Mountains, 1986–1994. 150
10-1 Puma population, number of pumas removed, and post-removal
population in the Treatment Area, December 1990–July 1991, San
10-2 Estimated puma population each January in the Treatment Area
(1988–1995) and Reference Area (1989–1995), San Andres Mountains,
New Mexico. 161
10-3 Estimated density of pumas each January in the Treatment Area
New Mexico. 162
10-4 Estimated puma population each January, 1989–1995, San Andres
10-5 Yearly adult puma density estimates for the Treatment Area
New Mexico. 164
10-6 Biological year adult puma density estimates for the Treatment Area
(1987–1988 to 1993–1994) and Reference Area (1988–1989 to
1993–1994), San Andres Mountains, New Mexico. 165
10-7 Estimates of puma population composition and density in North
American studies that employed intensive capture-mark-recapture and
radiotelemetry techniques. 167
10-8 Observed exponential rates of increase for the adult puma population
in the Treatment Area and Reference Area, San Andres Mountains,
New Mexico. 170
12-1 Female puma home range size during an entire birth interval, San
12-2 Cumulative home ranges for ten adult male and six adult female pumas
that were radio-monitored for forty-eight consecutive months on the
12-3 Home range estimates for adult pumas in North America. 199
12-4 Annual home range size for adult pumas during years of relatively low
and high deer densities, San Andres Mountains, New Mexico. 203
List of Figures, Tables, and Photos xix
12-5 Mean annual home range size and density for adult male and female
pumas on the San Andres Mountains, New Mexico, 1988–1994. 205
12-6 Distance between a female puma’s mean locations during consecutive
six-month periods as she raised young cubs, older cubs, regained
solitary status, and raised her subsequent litter of young cubs, San
14-1 Spatial relationships between adult same-sex pumas in other North
American study areas. 248
14-2 Number of adult pumas of the same and opposite sex with which each
puma shared parts of its home range, San Andres Mountains, New
Mexico. 251
14-3 Percent home range overlap between pairs of adult pumas in the San
14-4 Percentage of a puma’s home range shared by all other adult pumas of
the same sex, San Andres Mountains, New Mexico. 253
14-5 Percent home range overlap between adult pumas on the San Andres
Mountains, New Mexico, during 1992. 254
14-6 Clark-Evans ratios for distances between arithmetic centers of nearest
neighbors on the San Andres Mountains, New Mexico. 256
14-7 Spatiotemporal overlap indices for pumas on the San Andres
14-8 Associations between independent pumas on the San Andres
14-9 Comparisons of adult M-M, F-F, and M-F dyad associations in the San
Andres Mountains puma population, New Mexico. 262
14-10 Scrape sites used by four resident adult male pumas on the San Andres
Mountains from September 1985 to September 1988. 274
16-1 Prey killed by pumas on the San Andres Mountains, New Mexico,
1985–1995. 304
16-2 Prey items identified by hair and bone remains in 832 puma fecal
samples and four stomachs collected from the San Andres Mountains,
New Mexico, during 1985–1995. 306
16-3 Puma prey utilization based on 832 puma fecal samples and four
stomachs collected from the San Andres Mountains, New Mexico,
during 1985–1995. 307
17-1 Age and sex of desert mule deer that died on the San Andres
17-2 Survival rates of radio-collared desert mule deer, San Andres
17-3 Rates of puma predation and deaths from other causes in
xx List of Figures, Tables, and Photos
radio-collared desert mule deer, San Andres Mountains, New Mexico,

1987–1995. 317
17-4 Survival rates and agent-specific mortality rates of radio-collared deer,
San Andres Mountains, New Mexico, 1987–1994. 320
17-5 Mule deer population composition, San Andres Mountains, New
Mexico, 1988–1994. 323
17-6 Model parameter estimates used to simulate trends in the deer population
on the San Andres Mountains, New Mexico, 1987–1995. 324
17-7 Finite rates of increase estimated by modeling the desert mule
deer population on the San Andres Mountains, New Mexico,
1987–1995. 325
18-1 Desert bighorn sheep observed in surveys conducted on the San Andres
18-2 Survival rates of radio-collared desert bighorn sheep, San Andres
18-3 Rates of puma predation and other causes of death in radio-collared
desert bighorn sheep, 1987–1994, San Andres Mountains, New
Mexico. 353
A3-1 Annual home range size for twenty-four adult male and thirty adult
female pumas on the San Andres Mountains, New Mexico,
1986–1994. 416
Photos
1 Adult male puma M3 was resident on the San Andres Mountains until his
death at twelve years of age. 18
2 Puma F47’s four-week-old cubs at the birth nursery. 20
3 San Andres Peak after sunup. 25
4 Upper Sulphur Canyon. 25
5 Puma paw petroglyph near Three Rivers, New Mexico. 33
6 Hembrillo Basin Camp. 40
7 San Andrecito Camp. 41
8 Puma tracks in the sand. 41
9 Puma M7 caught in a snare by his right forefoot. 43
10 Linda Sweanor using a pole syringe to inject puma F37 with immobilizing
drugs. 44
11 Ken Logan hauling puma F37 for translocation to northern New Mexico.
45
12 Puma F27 at her birth nursery. 46
13 Adult female puma F47 with eartag and radio collar. 48
List of Figures, Tables, and Photos xxi
14 Six-week-old cub F162 with eartag, tattoo, and radio collar. 49

15 New Mexico Department of Game and Fish biologist Amy Fisher with a
radio-collared mule deer. 49
16 Puma F147’s three cubs at five weeks of age. 84
17 Puma F54’s birth nursery, where she had four cubs. 85
18 Orphaned puma F10 at ten months of age and nine days after her mother,
F2, died. 122
19 Puma F15’s skull showing fatal bite wounds to the braincase and old
damage to the zygomatic arch. 131
20 Puma M22’s skull showing mended canine puncture wounds. 133
21 Three eleven-month-old siblings F107, M108, and F109 with Ken Logan
and Frank Smith. 149
22 Aerial telemetry was critical to estimating puma home ranges and
determining puma interrelationships. 190
23 Brian Spreadbury with four-month-old puma F54. 234
24 Puma F40 was killed and eaten by territorial puma M19. 266
25 Male pumas were responsible for all of the killing of other pumas on the
San Andres Mountains. 291
26 Desert mule deer were the most important prey to pumas on the San
Andres Mountains. 303
27 A mule deer buck killed and eaten by a puma. 314
28 A desert bighorn ewe killed and eaten by a puma. 348
29 A yearling desert bighorn ram with ear canals occluded with scabs. 349
30 The puma is a keystone species in wild desert ecosystems, and it can serve
as an umbrella species in conservation efforts. 361
Foreword
My involvement in New Mexico began in May 1984. Dr. Wain Evans of the
New Mexico Department of Game and Fish asked if I would meet with the New
Mexico wildlife governing body, the State Game Commission, to discuss puma
research and management. The outcome of the meeting was an invitation to
develop a research project that would provide knowledge to guide the state’s
puma management program.
I welcomed this opportunity for two reasons: First, I was eager to study this
adaptable carnivore in an entirely different environment from that of my expe-
rience, spanning some twenty years, in the northern Rockies. Would there be
similarities and would there be differences in the way pumas live in a relatively
simplified desert environment? And what could we learn to help establish uni-
fying principles for this species to aid in its conservation throughout its range?
Secondly, this opportunity coincided with my founding of the Hornocker
Wildlife Institute and the proposed puma research fit the nonprofit Institute’s
mission statement:
The Hornocker Wildlife Institute, Inc., was founded to stimulate
curiosity about the natural world and its complexity; to provide
a direct, straight-forward framework in which to satisfy that
curiosity through observation and learning; to create an under-
standing of man’s place in the world’s fragile ecological structure
and his responsibility to it.
The Institute is designed to conduct intensive long-term
research with special emphasis on threatened and endangered
species and their wild environments; to train and develop supe-
rior post-graduate and graduate scientists; to make new knowl-
edge available to the scientific community, to the agencies
charged with managing wilderness and wild lands, and to the
xxiii
xxiv Foreword
public. Research focuses on scholarly, creative efforts designed to

make lasting contributions to our knowledge of the natural
world.
The project in New Mexico that we envisioned would become a flagship

effort of the fledgling Institute.
At the urging of Commission Chairman James Koch and members Thomas
Arvas and David Salman, I worked with Dr. Wain Evans, Department Director
Harold Olson, and others in developing the research. Over a period of several
months, we traveled to different potential study sites in New Mexico, settling on
White Sands Missile Range as the best area in which successful research could
be conducted. Major General Niles Fulwyler, Commander at White Sands,
agreed, and a Memorandum of Understanding was drafted. The result of these
deliberations was a ten-year contract between the State of New Mexico and the
Hornocker Wildlife Institute to conduct a comprehensive study of pumas in a
desert environment.
I already knew who I would approach to do the work. Ken Logan had
invited me to visit his puma project in northern Wyoming in 1982. He was
studying an exploited puma population for his master’s degree project at the
University of Wyoming. I was impressed with his approach and dedication and
had maintained contact with him over the years. He and Linda Sweanor,
another dedicated wildlife researcher and student at the University, had worked
together on bighorn sheep projects in northwestern Wyoming, and later mar-
ried. They eagerly accepted the offer to work as a team on the New Mexico
puma project.
At the same time, I invited Frank Smith to join our team. Frank had thirty
years of experience tracking and controlling pumas for the New Mexico Depart-
ment of Game and Fish and was considered the most knowledgeable person in
the region. I met this exceptional individual on my first trip to the field in New
Mexico and was convinced he would be a major player in a successful effort. He
agreed, and for seven years Frank contributed immeasurably to the project.
This book is a result of the ten-year puma project and is testimony to the
wisdom of long-term field study and observation. It fulfills, above and beyond,
the original mission statement of the Hornocker Wildlife Institute and rightfully
takes its place as a signature project. It is a tribute to the commitment and pas-
sionate dedication of its authors; it is a tribute to the vision of those New Mex-
ico game commissioners who committed resources to long-term field research in
a day and age when this is a rarity.
Several years ago I wrote the following in the foreword of Ted Bailey’s book,
The African Leopard, about our leopard research in South Africa:
Foreword xxv
Because they are consummate predators, cats sit at the apex of the
food chain. As such, they act as bellwethers of the condition of
the environment—healthy populations of cats mean healthy
populations of prey and a healthy environment. Yet one-third of
the world’s species of cats are currently threatened, if not endan-
gered. As the world’s human population grows and irreversibly
alters many environments, we may lose some species of cats
before we learn anything about them. Dedicated research like Dr.
Bailey’s is essential for gaining the new knowledge that conserva-
tion efforts require. His work will become a landmark in guiding
the conservation of leopards throughout much of their range.
Those comments are appropriate here. Ken’s and Linda’s work, in the finest
tradition of scholarly and rigorous scientific inquiry, has advanced our knowl-
edge of pumas far beyond any previous work. The fact that this new knowledge
can be utilized throughout this species range goes without saying. Their work
truly will become a landmark in puma conservation.
MAURICE HORNOCKER
Senior Scientist
Hornocker Wildlife Institute/
Wildlife Conservation Society
Acknowledgments
A dream came true for field ecologists like us, to study pumas for ten years in
one of the most beautiful desert settings in North America. For that, we thank
Dr. Maurice Hornocker. He was invited to New Mexico by the state Game and
Fish Department and the State Game Commission to design the project and
assume responsibility for its conduct. He hired us to do the work. Maurice was
unflagging in his support for us, both in the triumphs and in the struggles of the
research. We benefited immensely from his sage advice. At critical times when
funding was unavailable, Maurice and his Hornocker Wildlife Institute (HWI)
came through to continue our financial support. This project simply would not
have happened without him. In addition, he served as the major advisor for both
of our graduate programs at the University of Idaho.
Ecological studies of this magnitude, intensity, and difficulty require a dedi-
cated corps of researchers. We are forever grateful to the ones that worked with
us. J. Frank Smith, former Conservation Officer and Depredation Animal Con-
trol Officer for the New Mexico Department of Game and Fish and self-made
naturalist, was our cohort for seven years. Even with his thirty-plus years’ expe-
rience in tracking pumas, he was always yearning to learn more. His extensive
experience was especially important in giving the project a jump start in the
beginning. Brian Spreadbury hailed from British Columbia, Canada, where he
studied pumas for his master’s degree project; he worked with us three years.
Brian’s considerable experience in the bush enabled him to accomplish all that
we asked of him with ease and to the highest standards. His training in search
and rescue made us considerably more comfortable when we went on remote
forays to find pumas. If disaster struck, we knew Brian would do everything pos-
sible to get us out alive. After his work on our project, he returned to British
Columbia, found employment as a wildlife biologist with a coal mining com-
pany for a while, then became a park warden for Parks Canada. Toni Ruth was
finishing up her master’s degree project on puma–human interactions in Big
xxvii
xxviii Acknowledgments
Bend National Park, Texas, when we hired her to study the pumas we translo-
cated to northern New Mexico. Later, she came to our southern New Mexico
study area to assist us. In all, Toni spent three years on our project, then trans-
ferred to another HWI study of puma-wolf interactions in and around Glacier
National Park, Montana, for her Ph.D. studies. Later, she went on to study
puma-wolf relationships in Yellowstone National Park. Jennifer Cashman came
from Arizona, where she had studied desert mule deer water requirements and
puma diet in the Sonora Desert. She worked with us the last three years of the
project and later worked on HWI’s black bear research in New Mexico. Jeff
Augustine, a Wisconsinan, had studied beavers and coyotes but endeavored to
study the big cats. On our project, Jeff always came through with his enthusi-
asm and careful field work. At our camps, we could always count on him to
strike up enjoyable, probing discussions on evolution, ecology, and conserva-
tion. Jeff worked the last two years of our project, then went to Siberia to study
Amur leopards for HWI. Patricia Sweanor worked with us only a few months,
yet we benefited from the perspectives on large-mammal ecology she developed
while studying moose in Sweden. She left for Colorado and assisted the National
Park Service in the restoration of bighorn sheep in western U.S. parks. These
people never shirked the primitive living conditions in the desert; in fact, they
thrived there, performing yeoman’s services to accomplish our research objec-
tives. Many of the observations upon which this book is based are due to their
efforts. Experiences we shared with these people will forever be etched in our
memories.
Back at Hornocker Wildlife Institute headquarters at the University of
Idaho, people were toiling away, tending to the fusillade of paperwork that
attended the project’s multi-agency agreement. We thank Dr. Howard Quigley,
Molly Parrish, and Linda Harris for their attentiveness to our needs. Howard
became the Institute’s president during the last two years of the field research
and thereafter made sure that we were supported financially so we could write
this book. We will always be grateful for his faith in us and his constant patience.
We also thank Sandra Martin for securing the funds that sustained us.
We thank our pilots Bob Pavelka, Ed Pavelka, and Carl Hendrickson of Ed’s
Flying Service (Alamogordo, New Mexico). They were the ultimate profession-
als, always providing us meticulously maintained aircraft and safe flying as we
tracked radio-collared animals.
The New Mexico Department of Game and Fish financed the field research,
initial data analysis, and development of New Mexico’s puma management plan
mostly with funds provided by the Federal Aid in Wildlife Restoration Act and
New Mexico’s Share With Wildlife tax checkoff program. Dr. Wain Evans and
Wally Haussamen were particularly helpful in coordinating our field work with
Acknowledgments xxix
Department operations. Amy Fisher, Darrel Weybright, and Doug Humphreys

planned and executed the capture operations and aerial surveys of desert mule
deer and desert bighorn sheep, making it possible for us to gather detailed infor-
mation on the relationships of pumas to deer and sheep. Larry Temple facilitated
the translocation of pumas from our southern study area to northern New Mex-
ico and helped radio-track those cats. These long-term studies are dependent on
leadership and foresight of individuals within agencies who understand the need
to keep them going even in the face of political opposition. For that crucial sup-
port, we are grateful to Wain and directors Harold Olson, William Montoya,
Jerry Maracchini, and state game commissioners James Koch, Dr. Thomas
Arvas, and David Salman. We especially appreciate the leadership of Jimmy
Gonzalez, who personally saw to it that our research findings were taken to the
public of New Mexico, Department employees, and other wildlife professionals.
In addition, he had us directly involved in the development of New Mexico’s
puma management plan.
The U.S. Army, White Sands Missile Range, hosted our project. Major Gen-
eral Niles Fulwyler made it possible for us to conduct the research on the mis-
sile range. His support was a natural outgrowth of his personal interests in biol-
ogy, paleontology, archeology, and geology. Daisan Taylor, Patrick Morrow, and
Dave Holderman were missile range wildlife biologists who dutifully assisted us
by ushering the obligate reams of paperwork through the daunting army
bureaucracy, thus “breaking trail” for us to work in the field. Dave’s keen inter-
est in applying scientific information to wildlife management was especially
encouraging; later he was key to the incorporation of our information in wildlife
management on the missile range. Military police game wardens—and Sergeant
Artemis Hogan in particular—were tireless in keeping poachers off White Sands
Missile Range. We are especially grateful to John Collins, Edwin Erickson, and
Daniel Ansbach III for scheduling our research activities as missile-range mis-
sions, thus eliminating conflicts and keeping us out of harm’s way. We were
proud of the mission code “Alpha-Alpha” that they assigned to us for being “out
there all of the time.”
The U.S. Fish and Wildlife Service aided our project on two fronts. San
Andres National Wildlife Refuge personnel Patty Hoban, Mara Weisenberger,
Gary Montoya, and Steve Berendzen provided us with critical information on
desert bighorn sheep and allowed us to use camps on the refuge. Patty, Mara,
and Steve helped us capture some puma cubs. And Mara provided us with some
detailed observations of pumas. Federal aid coordinators Dave Parsons and Lau-
rel Wiley administered this project in accordance with the Federal Aid in
Wildlife Restoration Act.
We rarely got to see some vital people with whom we interacted daily dur-
xxx Acknowledgments
ing the field work. They were the two-way radio station operators in Santa Fe,
Las Cruces, and Raton who relayed our messages back and forth. Thank you for
helping us to keep people and animals safe and to efficiently coordinate field
activities.
We also thank the Jornada Experimental Range and the National Aeronau-
tics and Space Administration, White Sands Test Facility for access to their areas
adjacent to White Sands Missile Range.
We are extremely grateful to veterinarians Maury Brown, Frank Coons,
Michael Richards, and Brett Snyder for their unselfish professionalism. They
operated on puma F147’s fractured leg and rehabilitated her, making it possible
for her to return to the wild and successfully raise cubs. We also thank the Rio
Grande Zoological Park in Albuquerque for providing the place to rehabilitate
F147.
A special thanks goes to all those carnivore researchers and conservationists
in North America who wrote the commander of White Sands Missile Range in
support of our research when it was reviewed at the halfway point to decide on
continuation or termination. Your letters were helpful. In addition, several
wildlife managers and puma researchers shared their valuable opinions with us
about what was needed to further puma conservation. We especially benefited
from the feedback of Nick Smith, Jimmy Gonzalez, Charles Hayes, and Bill
Dunn of the New Mexico Department of Game and Fish; Tom Beck of the Col-
orado Division of Wildlife; John Beecham of Idaho Department of Fish and
Game; Rich DeSimone of the Montana Department of Fish, Wildlife and Parks;
Steve Torres and Vern Bleich of the California Department of Fish and Game;
Jimmy Rutledge of the Texas Parks and Wildlife Department; and Kerry Mur-
phy of the National Park Service, Yellowstone National Park.
Many people in northern New Mexico were essential to the success of the
puma translocation portion of this project. We acknowledge the support from
the communities and surrounding areas of Raton, Cimarron, Ute Park, Eagle
Nest, Angel Fire, and Folsom. We are particularly thankful for the cooperation
from Vermejo Park Ranch, the C.S. Cattle Company, and the National Rifle
Association Whittington Center. Appreciation for access to private land is
extended to the Hennigan Ranch, Chase Ranch, Philmont Scout Ranch, and
Sauble Circle Dot Ranch. We are grateful to the late Rick Poe, our pilot, who
flew us all around northern New Mexico to find the wayward radio-collared
pumas.
Scientific reviewers of draft manuscripts provided invaluable recommenda-
tions that improved this book. We are grateful to Doctors Maurice Hornocker,
Howard Quigley, Earnest Ables, Edward Garton, James Peek, Arthur Rourke
(all from the University of Idaho), Walter Boyce (University of California,
Acknowledgments xxxi
Davis), Paul Krausman (University of Arizona), Brian Miller (Denver Zoologi-

cal Foundation), and Wain Evans (New Mexico Department of Game and Fish,
retired). Some of the University of Idaho professors were also members of our
graduate programs and contributed immensely to our education.
Special thanks are extended to Tim and Karen Hixon, Summerlee Founda-
tion, McCune Charitable Trust, Richard King Mellon Foundation, Pond Foun-
dation, Thaw Foundation, Harry Bettis and Laura Moore Cunningham Foun-
dation, and Larry Westbrook for providing funding for the writing of this book.
We are very grateful to our editors at Island Press, particularly Barbara Dean,
Barbara Youngblood, Cecilia González, and Erin Johnson for improving the
manuscript and for guiding the book to completion.
For us, this project was an expedition, not only into a region of the Chi-
huahua Desert that was little known, but also into our own physical, intellec-
tual, and emotional limits. Ten years is a long time to maintain peak intensity
in laborious field research. But the place and the life kept us always looking and
stepping forward. Without a doubt, our deepest satisfaction was in trying to fig-
ure out why life works the way it does on a piece of Earth, the San Andres
Mountains. Every canyon, ridge, and mountain peak became familiar to us. It
was inevitable. Daily, we searched for or followed pumas wherever they might
go, which was everywhere in those desert mountains. A clandestine helicopter
crew could have blindfolded us and plunked us down alone in some secluded
spot in the middle of night, but by daybreak, or even before, we would have
known where we were. The rigors and pleasures of the land, its seasons, and the
animals defined our way of life. We will always regard the San Andres Moun-
tains as “our home.”
The San Andres Mountain range is unique in that it is the largest single
block of ecologically intact Chihuahua Desert mountains remaining in North
America. This vibrant ecosystem deserves concerted attention by the agencies
responsible for its land and resources, and a caring public, to ensure that its nat-
ural values are preserved. It is our hope that scientific information contained in
this book will promote that effort.
Finally, to the pumas, thank you for the wonder. We hope our work will help
people to understand you and to contribute to your endurance.
—KENNETH A. LOGAN and LINDA L. SWEANOR

P a rt I
Setting the Stage

Chapter 1
Introduction
C a r n i vo res stir within us a primitive curiosity. We have been honing our senses
to these flesh-eaters ever since our human ancestors re a l i zed that they could be
killed and eaten by such beasts. This curiosity has gradually developed into a
highly sophisticated form we call science. Now, we delve into evolution, biol-
ogy, and ecology to better understand carnivo res and our relationships to
them.
C a r n i vo res profoundly impact animals and ecosystems. T h rough predation,
they transfer matter and its associated energy into their own trophic level and
d i ve rt some of what they do not consume to scavengers, detritivo res, and
m i c roorganisms (Ricklefs 1990). W h e re carnivo res are the highest-level con-
sumers and closely linked with the dominant herbivo res, they can alter pro-
duction in plant communities (Mc L a ren and Peterson 1994). Carnivo res can
impact the dynamics of prey populations by limiting (Be r g e rud et al. 1983,
Messier 1991, Gasaway et al. 1992) or regulating (Messier 1995) their num-
bers, and they are a major factor contributing to cyclic pre d a t o r - p rey systems
( K rebs et al. 1995). Carnivo res partially shaped the evolution of prey by the
selection for morphological, physiological, and behavioral variations that
enhance surv i val. For example, cervids have acute senses for predator detection
and slender elongate limbs powe red by large proximal muscle groups for rapid
escape (Putman 1988). In addition, prey evo l ved fitness-enhancing antipre d a-
tor strategies, such as social grouping (Berger 1978), vigilance (Hunter and
Skinner 1998), crypsis (Caro and Fitzgibbon 1992), specialized locomotion
( C a ro 1986, Geist 1998), avoidance (Mech 1977b), migration (Be r g e rud and
Page 1987), and birth synchrony (Rutberg 1987). Interspecific competition
3
4 PART I. SETTING THE STAGE
among carnivo res can affect behavioral patterns, influence re s o u rce part i t i o n-
ing, and may have been an evo l u t i o n a ry force contributing to assemblages of
c a r n i vo res in rich environments (Schaller 1972, Mills 1990, Caro 1994,
Durant 1998).
Carnivores influenced human evolution. Carnivory by humans may have
had its humble beginnings when ancestral hominids scavenged the kills of car-
nivores (Blumenschine 1991). This new, rich food source enabled hominids to
stride out from the African tropics into temperate habitats and beyond. Living
among big carnivores surely induced strong selective forces for beings that could
outwit such dangerous competitors and predators. And the high-quality, easy-
to-digest carnivorous diet was essential for the development of a much more
complex neural system (Aiello and Wheeler 1995). Humans may partly owe the
evolution of our large encephalized brain and extreme abilities for reason and
cunning to large carnivores (Wilson 1980:133–137).
Modern humans are still influenced by carnivores. Carnivores kill and eat
our livestock, pets, and wild prey, and they sometimes physically threaten, or
even kill and eat, humans. Yet their form, intelligence, and beauty are awe-
inspiring and are perhaps even the nascence for domestication of their gentler
forms. They are our companions, and, in special cases, we depend on them to
aid us in disability, search and rescue, and law enforcement. Felids, in particu-
lar, may provide us healthier lives. Because felid genome organization is similar
to that of humans (Nash and O’Brien 1982, Rettenberger et al. 1995, Wienberg
et al. 1997), cats, wild and domestic, are being used in coordination with the
Human Genome Project to study genetic links to hereditary defects and diseases
that afflict both cats and humans (O’Brien 1997).
Scientific knowledge and personal experience have cultivated human appre-
ciation for the beauty and ecological role of carnivores. Hence, we have deemed
it necessary to conserve them and even to make them the focus of complex,
landscape-scale conservation strategies (Hummel et al. 1991, Noss et al. 1996,
Weber and Rabinowitz 1996). For some of the most endangered carnivores,
conservation efforts have extended beyond the people that live on the land with
the animals to include national and international governments and nature con-
servation organizations (see Schaller 1993, Seidensticker 1997). As the human
population burgeons and uses up more space and resources, we need to know
more about carnivores and how we influence their populations and habitats if
we want them as members of Earth’s biota. This book on the puma (Puma con -
color) is our attempt to convey a scientific understanding of this magnificent car-
nivore and to present ways in which enlightened humans can continue to coex-
ist with the puma.
CHAPTER 1. INTRODUCTION 5
Rationale and Approach

This puma research originated from a contract secured by Dr. Maurice
Hornocker with the New Mexico Department of Game and Fish in 1985, fol-
lowing several months of consultation and project development between Dr.
Hornocker and Department personnel. At that time, the Department and its
governing body, the State Game Commission, requested basic biological and
ecological information that wildlife managers could use to develop management
strategies for pumas and their prey. Hence, our research was mission-oriented,
taking the form of applied ecology. This was the first intensive, long-term inves-
tigation of pumas living in a desert; therefore, much of our work describes in
detail the natural history of pumas in that environment. Moreover, we tested
hypotheses on pumas and other mammals previously posed in the scientific lit-
erature. And as patterns emerged from our data, we developed other hypothe-
ses. In some instances, we offer measured speculation for natural phenomena we
observed but for which we could not gather quantitative data; our hope is that
scientists may be able to build on these ideas.
The goal of New Mexico wildlife managers was to use scientific information
to develop management and conservation strategies on pumas to address con-
cerns of their most politically influential constituents. Their operational purpose
was to manage for a self-perpetuating puma population while dealing with inter-
related issues, including maximizing opportunities for recreational puma hunt-
ing; minimizing competition between hunters and pumas for big game animals,
especially mule deer; minimizing puma predation on desert bighorn sheep, a
state-listed endangered species; and minimizing puma predation on livestock.
In consideration of the wildlife managers’ needs, we developed three main
research objectives: (1) to describe and quantify puma population dynamics, (2)
to describe and quantify puma social organization, and (3) to describe and
quantify the relationships of pumas to their prey, specifically desert mule deer
(Odocoileus hemionus crooki) and desert bighorn sheep (Ovis canadensis mexi -
cana). Understanding puma population dynamics and life history strategies is
essential to developing scientific approaches to puma management and conser-
vation. It is also the first step in testing hypotheses about puma social organiza-
tion and effects of puma predation on prey. Social organization is important to
investigate because it may limit the population (Seidensticker et al. 1973), and
it influences sex ratios, reproduction, mortality, dispersal, recruitment, genetic
makeup, and gene flow. Studying relationships of pumas to their prey is critical
to understanding how puma predation affects prey populations and how prey
affect puma populations.
When we started this project, we made a significant long-term commitment.

Our experience with pumas taught us that their low population densities, cryp-
tic lifestyle, and use of complex habitats would make them extremely difficult
and challenging to study. We anticipated that it would take at least two years of
the project simply to develop an intensive population-monitoring routine to
reliably quantify puma population characteristics and behavior in our large
desert study area. In addition, evaluating effects of puma predation and other
limiting factors on mule deer and bighorn sheep population fluctuations would
require several years. Also, we wanted to experimentally reduce puma density on
the study area to test hypotheses relevant to puma population dynamics, social
organization, and puma-prey interactions. This required an intensive study of
puma and prey population dynamics before puma removal, a subsequent
removal of pumas at a rate high enough to cause a substantial population reduc-
tion (i.e., more than 50 percent of adult pumas), and, finally, a comprehensive
study of the dynamics after the removal. In our estimation, we could accomplish
the research goal and objectives in a minimum of ten years. Our work became
the first long-term research of pumas in a desert environment. In the process, we
studied the largest number of individual pumas ever in an ecological project.
As we studied the pumas, we constantly asked ourselves, “Why do pumas
live and behave the way they do?” This simple question characterized our sec-
ond and not-so-simple goal, which was to explain the evolutionary ecology of
pumas. Evolutionary ecology integrates all that we understand about puma pale-
ontology, evolution, genetics, biology, and ecology. It is a synthesis that helps us
to comprehend where pumas came from, how they adapted to environments,
how they affect the environment, and what impacts humans may have on them.
Evolution—with natural selection its key mechanism for change—is so far the
best scientific explanation for life on Earth (Trivers 1985, Stearns 1992, Mayr
1996, Freeman and Herron 1998). Thus, we made it the central theme of this
book. We hope to render a more complete concept of this beast, something akin
to a photographer metering the light, composing the scene, and turning the
camera lens to snap into sharper focus an image that grasps the essence of the
subject.
Synopsis of Previous Studies

Although pumas have been subjects of numerous short-term studies, there have
been few intensive, continuous investigations of eight or more years, which
would approach the natural life span of this carnivore in the wild. The first of
these was the classic puma research by Maurice Hornocker and colleagues in the
River of No Return Wilderness in central Idaho from 1964 to 1972 (Hornocker
CHAPTER 1. INTRODUCTION 7
1969, Hornocker 1970, Seidensticker et al. 1973). The first five years of research
involved intensive capture, marking, and recapture techniques to quantify the
puma population, and ground and aerial surveys of deer and elk to assess the
impact of puma predation on those species. Not until the subsequent three-year
companion study by Seidensticker et al. (1973) were the biologists able to
employ the new technology of radiotelemetry to describe puma social organiza-
tion. They also developed the basic techniques that would thereafter be used in
all subsequent field studies of puma population dynamics, behavior, and habi-
tat use. More importantly, they developed original biological concepts about
pumas. Hornocker (1969:462, 464) described a type of nonaggressive territori-
ality in pumas maintained through a “mutual avoidance reaction” and con-
cluded that the primary function of such behavior was to “limit population
size.” The biologists developed that idea further into the only published hypoth-
esis on the evolution of puma social organization. They postulated that the
social organization evolved to limit the density of breeding pumas to a level
below that set by their prey (Seidensticker et al. 1973). This was one of the
hypotheses we tested relative to the way pumas lived in our desert study area.
In Nevada, Dave Ashman and colleagues with the Nevada Department of
Wildlife studied pumas from 1972 to 1982 in eleven mountain ranges in the
northeast and central parts of the state. The bulk of their work was done in the
Ruby Mountains and the Monitor Range. They focused on puma population
dynamics, movements, and predation (Ashman et al. 1983). From 1978 to
1989, a group of graduate students assembled by Fred Lindzey studied a puma
population in the Boulder-Escalante area of south-central Utah. They investi-
gated a diverse range of topics, including puma population dynamics, energet-
ics, predation, habitat selection, effects of sport-hunting, and survey methods
(Lindzey et al. 1989). Allen Anderson studied pumas on the Uncompahgre
Plateau in southwestern Colorado from 1981 to 1988. Although his main focus
was puma population biology, he also provided information on puma move-
ment patterns, intraspecific interactions, morphological and physiological char-
acteristics, and predation (Anderson et al. 1992). Martin Jalkotzy and Ian Ross
studied puma population dynamics from 1981 to 1989 along the eastern slope
of the Rocky Mountains in southwestern Alberta (Ross and Jalkotzy 1992).
They later provided information on puma predation (Ross and Jalkotzy 1996,
Ross et al. 1997). Kerry Murphy studied the ecology of pumas in the Northern
Yellowstone Ecosystem from 1987 to 1996. He collaborated with a host of other
biologists and geneticists to investigate puma-prey relationships, competition
between pumas and bears, and puma reproductive success (Murphy 1998). The
longest study is that of the endangered Florida panther, which has been investi-
gated since 1981. Interagency and interdisciplinary teams of researchers have
looked into practically every aspect of the remnant population of this feline,
including population and social biology, predation, habitat selection, diseases,
and genetics (Belden 1982, Belden et al. 1988, Maehr et al. 1989, O’Brien et al.
1990, Roelke et al. 1993a, Roelke et al. 1993b, Maehr and Cox 1995). The
number of individual pumas studied in these investigations ranged from forty-
seven in Colorado (Anderson et al. 1992) to about a hundred in Nevada (Ash-
man et al. 1983) and Utah (Lindzey et al. 1989). By comparison, we studied
294–295 individual pumas in ten years.
Along with our own work, we treated all these studies, and many of the
short-term studies, as independent sources of data on pumas because they were
done in different conditions, including climate (e.g., temperate, desert, tropi-
cal), habitat (e.g., continuous, fragmented, isolated), prey diversity and density
(e.g., high, low), and selective pressures (e.g., hunted, nonhunted, other carniv-
orous competitors, lack of competitors). This allowed us to contrast our find-
ings—on puma population characteristics, life history strategies, behavior, social
organization, and relationships of pumas to their prey—with pumas in other
environments.
This book tells our story of puma research in twenty-one chapters appor-
tioned among five parts. Part 1 sets the stage for the research and includes back-
ground information on development of the project, puma phylogeny, taxon-
omy, and description. We also describe our study area and how we studied
pumas. Part 2 covers puma life history strategies and population dynamics. Part
3 describes puma behavior and social organization. Part 4 analyzes relationships
of pumas to their prey, particularly desert mule deer and desert bighorn sheep.
Part 5 presents our ideas for conserving and managing pumas in the wild. Four
appendices cover puma morphometrics, reproductive chronology, home range
estimation, and mule deer population dynamics.
Chapter 2
Pumas Past and Present
Puma Phylogeny
Paleontologists use fossils to construct a phylogeny (i.e., evolutionary history) of
carnivores, albeit, an incomplete one. The present record is that the Carnivora
evolved from the basal family Miacidae (small, mostly arboreal carnivores) dur-
ing the Eocene epoch (Vaughan 1997), about fifty million years ago. The most
specialized carnivores of all, the true cats (family Felidae) apparently evolved in
the Miocene about twenty million years ago. Their ancestors probably were of
the genus Pseudaelurus, the most common felid in the early and middle
Miocene, and were lynx-sized predators with arboreal and terrestrial stalk-and-
ambush hunting modes (Werdelin 1996). In the middle Miocene, felids began
their radiation into the great sabertooths (Machairodus, Homotherium, Megan -
tereon, Smilodon) and other cats with conical canines (Felis, Puma, Panthera,
Metailurus, Miracinonyx, Acinonyx) that came to occupy the Pliocene and Pleis-
tocene epochs about 5.3 million to twelve thousand years ago (Hunt 1996).
Phylogenetic relationships for thirty-four of the thirty-seven species of extant
felids, based on maternally inherited mitochondrial genes, have been arranged
into eight major clades and four unaligned species (Johnson and O’Brien 1997,
Pecon-Slattery and O’Brien 1998). Puma is the namesake for the clade that also
includes the cheetah (Acinonyx jubatus) and jaguarundi (He r p a i l u rus
yaguarondi); all members were thought to have originated in North America.
Other groups are ocelot lineage (ocelot Leopardus pardalis, tigrina L. tigrinus,
margay L. wiedii, pampas cat Oncifelis colocolo, Geoffroy’s cat O. geoffroyi),
9
domestic cat lineage (domestic cat Felis catus, jungle cat F. chaus, African wild
cat F. libyca, sand cat F. margarita, black-footed cat F. nigripes, Chinese desert cat
F. bieti, European wild cat F. silvestris), panthera group (lion Panthera leo, leop-
ard P. pardus, tiger P. tigris, snow leopard P. uncia, jaguar P. onca, clouded leop-
ard Neofelis nebulosa), lynx genus (bobcat Lynx rufus, Canadian lynx L. canaden -
sis, Siberian lynx L. lynx), Asian leopard cat group (Asian leopard cat
Prionailurus bengalensis, flat-headed cat P. planiceps, fishing cat P. viverrinus),
caracal group (caracal Caracal caracal, African golden cat Catopuma aurata), and
bay cat group (Asian golden cat C. temminckii). Unaligned species include the
serval (Leptailurus serval), rusty-spotted cat (Prionailurus rubiginosus), Pallas’ cat
(Otocolobus manul), and marbled cat (Pardofelis marmorata).
The puma group is considered to be the most ancestral, having diverged
about 8.5 million years ago. By comparison, the other large cats in panthera
group last shared a common ancestor about six million years ago (Johnson and
O’Brien 1997, Pecon-Slattery and O’Brien 1998). Within the puma group, the
cheetah diverged first from a common ancestor about five to eight million years
ago; puma diverged from a common ancestor with the jaguarundi about four to
five million years ago (Johnson and O’Brien 1997, Culver 1999). Extant puma
lineages last shared a common ancestor about 0.39 million years ago (Culver et
al. 2000). About the same time, Homo sapiens was emerging in Africa (Freeman
and Herron 1998).
An obscure puma fossil record has lead to a paleontology that is oblique to
the phylogenetics of Puma concolor. Puma-like felids have been relatively com-
mon in North America since about the late Blancan land mammal age about 2.5
million years ago. Miracinonyx inexpectatus was a form intermediate to modern
pumas and cheetahs, but larger, and may have been ancestral to Puma concolor
(Kurtén 1976). A transition from one form to the other may have occurred dur-
ing the Rancholabrean land mammal age when P. concolor became widespread
in North America; the earliest fossil finds are from the Sangamonian interglacial
about 0.2–0.3 million years ago (Kurtén and Anderson 1980). The puma was
thought to be indigenous to North America (Adams 1979). In South America,
P. concolor lacked probable ancestors and was thought to have immigrated from
the north during the Ensenadan land mammal age about 0.5–0.8 million years
ago (Webb 1976, Savage and Russell 1983). According to this scenario, the pre-
sumed Ensenadan appearance of P. concolor in South America predates the ear-
liest fossils of the species found in North America. Van Valkenburgh et al.
(1990) suggested that P. concolor may have evolved from an unknown species in
the Neotropics because Puma has shorter limbs and larger head and teeth than
M. inexpectatus, which are characteristics of a more primitive condition.
Phylogeographic research (i.e., study of the phylogenetic relationships and
CHAPTER 2. PUMAS PAST AND PRESENT 11
geographic distributions of individuals in a species) on pumas across the West-

ern Hemisphere by Culver and her colleagues (Culver 1999, Culver et al. 2000)
suggest that pumas emerged in eastern South America and then dispersed to
other parts of South America and eventually to North America. Culver (1999)
found ancestral mitochondrial DNA haplotypes (mtDNA is maternally inher-
ited; a haplotype is a set of alleles from closely linked loci carried by an individ-
ual and usually inherited as a unit) exclusively in extant pumas from Brazil and
Paraguay south of the Amazon River. Patterns of genetic variation in mitochon-
drial and microsatellite markers (microsatellites are nuclear DNA sequences
inherited from both parents and comprising variable numbers of tandemly
repeated bases, usually five to forty, for example, CACACACACA) supported
their contention that eastern South America was the site of origin. Pumas in
eastern South America (i.e., the central population) had the highest genetic vari-
ation, while pumas in North America and southern South America (i.e., periph-
eral populations) had the lowest variation. This sequence of evolution and
migration and Culver’s estimated coalescence time of 0.39 million years for
puma mitochondrial DNA haplotypes resolves the estimated dates for the
appearance of P. concolor in South America and the earliest puma fossils in North
America. Culver (1999) points out that a puma ancestor still may have
descended from the North American M. inexpectatus, dispersed to South Amer-
ica, and then speciated into the modern puma, or that the puma descended from
a yet-unknown South American ancestor (also see Van Valkenburgh et al. 1990).
Puma ancestors might have entered South America from North America in the
midst of the “Great American Interchange” during the Uquian land mammal
age about 1.5–3 million years ago (Marshall et al. 1982). A small founder pop-
ulation crossing an isthmus of variable integrity, the presence of other geo-
graphic barriers in South America (e.g., broad river systems, high rugged moun-
tain ranges), behavioral shifts in response to novel environments and
competitors, and concomitant extinctions of indigenous species would have
provided a constellation of conditions ripe for evolution.
For the North American puma, Culver’s research tells a story of mass extinc-
tion, then recolonization. Based on the low variance in the number of
microsatellite repeats, the presence of only one mitochondrial haplotype (rela-
tive to other phylogeographic puma groups), and a molecular clock based on
microsatellite allele variance, the North American puma population probably
experienced a founder event about ten thousand to twelve thousand years ago.
Culver and colleagues (2000) believe that the puma that inhabited North Amer-
ica (up to about 0.3 million years ago) went extinct along with the many other
large mammals during the late Pleistocene (see below). The geneticists conclude,
“Most likely modern North American pumas descended from a founder event
involving a small number of individuals who migrated ‘out of South America’

approximately 10,000–12,000 years before the present and subsequent to the
abrupt Pleistocene extinction of large North American mammal species” (Cul-
ver et al. 2000:196).
While pumas were living in Pleistocene North America, the large mammal
fauna was far richer than in the Holocene epoch. Mammals were huge compared
to their modern relatives. This was likely because the environments they lived in
were highly productive, and large-bodied animals had survival, and hence selec-
tive, advantages in a perilous world of large carnivores. Diversity of the herbi-
vore fauna was similar to that of Africa’s Pleistocene and modern fauna (Stuart
1991, Burney 1993, Van Valkenburgh and Hertel 1993). Four glaciations, three
intervening interglacials (Kurtén and Anderson 1980), and attendant habitat
and animal dynamics provided selective forces for evolution of this assemblage.
In addition, Eurasian mammals immigrated across the Bering Bridge (Kurtén
and Anderson 1980). In late Pleistocene North America, there were fifty-six her-
bivore species larger than 30 kg, including twenty-nine that exceeded 300 kg.
Moreover, a rich carnivore assemblage, including fifteen species coyote-sized or
larger, preyed or scavenged on the herbivores. Today by comparison, there are
only eleven herbivores larger than 30 kg, three herbivores larger than 300 kg,
and seven carnivore species equal to or larger than the coyote (Van Valkenburg
and Hertel 1993).
Stratigraphic ranges for Pleistocene mammals in North America constructed
by Kurtén and Anderson (1980) display the array of large herbivores that pumas
may have preyed upon and carnivores with which they may have competed.
There were two species of capybaras (Hydrochoerus, Neochoerus), perhaps as
many as thirteen species of horses, onagers, and asses (Equus), three species of
tapirs (Tapirus), two species of peccaries (Mylohyus, Platygonus), seven species of
camels (Camelops, Hemiauchenia), seven species of deer (Odocoileus, Navaho -
c e ros, Sangamona, Alces, Cervalces, Cerv u s), nine species of pronghorns
(Capromeryx, Tetrameryx, Stockoceros, Antilocapra), and three species of moun-
tain goats and mountain sheep (Oreamnos, Ovis). There were other mammals
that were simply too enormous for pumas to kill, including giant ground sloths
(Eremotherium, Nothrotheriops, Glossotherium), two species of bison (Bison), a
mastodont (Mammut), and mammoths (Mammuthus).
A diverse carnivore fauna, acting as predator and scavenger, was supported
by the herbivores. There were five species of bears, including the cave bear
(Tremarctos floridanus), lesser short-faced bear (Arctodus pristinus), giant short-
faced bear (A. simus), grizzly (Ursus arctos), and black bear (U. americanus).
Besides the puma there were five other species of large cats, including the saber-
tooth (Smilodon fatalis), scimitar cat (Homotherium serum), lion (Panthera leo
atrox), jaguar (P. onca), and cheetah (M. trumani). Among the canids, there were
the timber wolf (Canis lupus), red wolf (C. rufus), dire wolf (C. dirus), coyote (C.
latrans), and dhole (Cuon alpinus). The selective pressures imposed by this wide
range of prey and competitors no doubt helped to shape the evolution of the
puma.
Excavations at four sites in southern New Mexico (i.e., Burnet Cave and Dry
Cave in Eddy County, Blackwater Draw in Lea County, U-Bar Cave in Hidalgo
County) indicate that the region supported some of these beasts in the late Pleis-
tocene (Harris 1987). Large herbivores and carnivores identified in those
deposits included Shasta ground sloth (Nothrotheriops shastensis), tapir, five
species of horses, peccary (Platygonus), three species of camels (Camelops, Hemi -
auchenia), two species of deer (Odocoileus, Navahoceros), three species of prong-
horns (Antilocapra, Stockoceros, Tetrameryx), bison, and possibly Harrington’s
mountain goat (Oreamnos harringtoni). Carnivores found at the caves were the
coyote, gray wolf, dire wolf, giant short-faced bear, sabertooth, jaguar, and
puma.
This diversity of large mammals in North America came to a relatively
abrupt end in a wave of extinctions during the late Pleistocene about
11,500–10,500 years ago (Stuart 1991). Mass extinctions were concurrently
occurring in South America; in fact, no other continent lost as many animals
(Burney 1993). Although the exact causes of the extinctions are not clear, some
hypotheses have been forwarded to explain them. Severe environmental changes
brought by a shifting climate at the end of the last ice age with attendant trans-
formations of vegetation, redistribution of fauna, and new competitive relation-
ships may have triggered extinctions among nonadaptive species. Another
hypothesis suggests paleolithic hunters who came to North America from Eura-
sia about twelve thousand years ago via the Bering Bridge and were south of the
ice sheets by about 11,500 years ago overkilled the large and conspicuous slow-
breeding mammals not adapted to this novel predator. These technologically
advanced humans effectively used sharp, stone-tipped spears to kill large mam-
mals, even mammoths. It may be more likely that combined effects of environ-
mental shifts and overkill by humans caused extinctions. Large herbivore distri-
butions and population numbers may have been reduced initially by disruptive
environmental transition and habitat loss, making the herbivores vulnerable to
overkill by an increasing population of paleolithic hunters. As the large herbi-
vores disappeared, so did many of their specialist predators (Kurtén and Ander-
son 1980, Stuart 1991). Still another, the “keystone herbivore hypothesis”
(Owen-Smith 1987, 1989), suggests that paleolithic hunters eliminated the
megaherbivores (species with adult mass greater than 1,000 kg) that were essen-
tial to maintaining a diverse habitat mosaic, thus promoting a diverse, large-her-
bivore fauna. In the absence of megaherbivores, grasslands turned to woodlands,

triggering a decline in open-country herbivores, and, we would suggest, making
others more vulnerable to predation. Time lags between the decline of herbi-
vores and the decline of their predators may have triggered inversely density-
dependent predation and further hastened the decline of herbivores. As large
herbivores disappeared, so did their predators. All of the sabertooth cats went
extinct, and most conical-toothed cats suffered restrictions in their ranges (Mar-
tin 1980). Pumas, it seems, were victims too.
Regardless of their point of origin, pumas thrived after catastrophic late
Pleistocene extinctions, an expected result for such a eurytopic species (Erwin
1998). Because of their moderate size and ability to sustain themselves on a wide
range of prey in various habitats, they had advantages over specialist carnivores.
As other large carnivores went extinct, interspecific competition would have
declined. Moreover, pumas were adapted to dense woodlands and rugged terrain
where other post-Pleistocene herbivores also thrived, perhaps because they were
less conspicuous to human hunters. In addition, because of their smaller size,
they would be expected to mature faster, have greater fecundity, and, hence,
have greater intrinsic rates of increase than the doomed megaherbivores (Eisen-
berg 1981, Read and Harvey 1989).
Post-Pleistocene North America that lay before the puma had a relatively
impoverished large-mammal fauna. Ecosystems in the Southwest, in particular,
would become much simpler. In the Chihuahua Desert, the puma’s large-mam-
mal prey would be composed only of desert mule deer, white-tailed deer
(Odocoileus virginianus), collared peccaries (Pecari tajacu), and desert bighorn
sheep. Competitors with which pumas would have to contend were still formi-
dable but reduced in diversity to gray wolves, coyotes, jaguars, and on occasion
grizzly bears and black bears.
Mule deer and desert bighorn sheep benefited from the low diversity in the
mammal fauna following the extinctions. Mule deer emerged as a new form
about nine thousand to eleven thousand years ago, presumably the result of
i n t e r b reeding of white-tailed does and black-tailed bucks (Geist 1998).
Bighorn sheep dispersing from the north became established in what is now the
northern Chihuahua De s e rt about thirteen thousand to fifteen thousand ye a r s
ago; they began showing characteristics of the modern desert bighorn sheep
b e t ween eight thousand and eleven thousand years ago (Geist 1985). Com-
p a red to the relationships that must have evo l ved between the Pleistocene
puma and its prey during hundreds of thousands, if not millions, of years of
c o e volution, the Holocene relationships of pumas with their prey in the So u t h-
west are re l a t i vely young. Neve rtheless, for the past eight thousand to eleve n
thousand years, the finely tuned mountain pre d a t o r, the puma, would provide
a strong selective force to which mule deer and desert bighorn sheep would
need to adapt to surv i ve .
Puma Distribution and Status

Historically, pumas have had the broadest geographic distribution of any terres-
trial mammal in the Western Hemisphere, except for humans. Pumas ranged
from northern British Columbia to Patagonia and from the Atlantic to the
Pacific coasts (Young 1946). The huge geographic range of the puma is testi-
mony to its extraordinary adaptability. Pumas have occupied practically every
type of biogeographic zone, including boreal foothills, temperate mountains and
forests, tropical rainforests, grasslands, and deserts (Young 1946; Ian Ross, per-
sonal communication) along an elevation gradient ranging from sea level to
3,350 m in North America, and to 4,500 m in South America (Nowak 1991).
However, European immigrants to the Americas since the 1500s brought
predator control and habitat loss. Settlements, agriculture, and industry trans-
formed habitats, making them unsuitable for pumas. Many prey species were
extirpated. Pumas were killed outright mainly to protect livestock and game ani-
mals and to allay human fears. Basically, pumas were considered to be “bad ani-
mals.” By the late 1800s in North America, eastern puma populations were
extinct or severely reduced, and by the early 1900s, western populations were
diminished (Nowak 1976). For pumas in North America, this meant a contrac-
tion to about one-half of their modern geographic range (Logan and Sweanor
2000).
Legal protection for pumas in North America has been quite recent. Since
1965, regulations on killing pumas by all of the western United States (except
for Texas) and provinces of Canada have enabled populations to recover from
historical low levels. In a survey we conducted in 1997 on puma population sta-
tus in fourteen western states and two western Canadian provinces (Logan and
Sweanor 2000), 62 percent reported increasing or stable trends, 25 percent
reported declining or stable trends, and 13 percent did not know the trend in
their puma populations. Passage of the Endangered Species Act in 1973 gave
pumas in eastern United States full protection and may have prevented the
extinction of the Florida panther subspecies (P. concolor coryi). Even in recent
years, pumas have been sighted in several other eastern states and provinces, but
very rarely have those reports been authenticated. In addition, it is not known
whether those individuals were escaped captive animals or wild.
Status of the puma across its range in Central and South America is sketchy.
It appears that pumas still occupy almost all of their historical range except for
the densely human-inhabited areas and landscapes altered for agriculture and
livestock. Puma abundance has been reduced in eastern Brazil and northeastern
Argentina (Nowell and Jackson 1996). Lopez-Gonzalez (1999) reported that
puma hunting was prohibited throughout South America, except in Peru, and
that the species had legal protection in Central America, except in El Salvador,
where the puma was nearly extinct.
Pumas in the Southwest

As European humans immigrated to the Southwest beginning in the late 1500s,
and with an accelerating rate of immigration during the late 1800s and early
1900s, the large carnivore fauna in the desert was depleted. Humans organized
extermination campaigns sanctioned by state and federal governments with the
intent to protect livestock and big-game animals. Carnivores were killed by any
means possible, including shooting, trapping, and poisoning (Brown 1984,
1985). Wolves were extirpated from the Southwest by about 1925 (Brown
1984), followed by jaguars in about 1930 (Findley et al. 1975, Hoffmeister
1986), and grizzly bears in about 1935 (Brown 1985). Pumas survived this
onslaught probably because of their solitary, highly cryptic nature and their
propensity to inhabit the most rugged terrain, but even so, puma populations
declined.
In New Mexico, puma numbers were severely reduced. Puma killing was
encouraged statewide with a bounty that lasted from 1867 until 1923 (New
Mexico Department of Game and Fish Operational Plan, 1987–1995, Nowak
1976). Young (1946:28) reported that the puma in New Mexico “of late years,
due to intensified hunting, is not as common as it was at the beginning of the
present century. The animal may now be said to be confined mainly to the
rougher mountainous sections west of the Rio Grande.” Not until 1971, when
the legal status of the puma was changed to effectively curb human-caused mor-
tality, did the species have the opportunity to increase and recolonize historical
habitat. Today, there are pumas inhabiting most habitats in New Mexico.
Taxonomy
The puma belongs to the order Carnivora, suborder Feliformia (i.e., the felids,
herpestids, hyaenids, viverrids), family Felidae, subfamily Felinae (which also
includes small cats of the genera Caracal, Catopuma, Felis, Herpailurus, Leopar -
dus, Leptailurus, Lynx, Oncifelis, Oreailurus, Otocolobus, Prionailurus, Profelis)
(Wozencraft 1993). Linneaus originally named the puma Felis concolor in 1771,
but Jardine renamed the genus Puma in 1834 (Wozencraft 1993). Nowak
(1991) presented three taxonomic arrangements for the thirty-seven species in
the Felidae family from three authors (Ewer 1973, Hemmer 1978, Leyhausen
1979) and also from one of his own, which was a composite from a variety of
sources. Each arrangement had a varying number of genera. The puma was
placed in the genera Profelis and Felis once each, and in Puma twice. Nowak’s
arrangement included twenty-nine species in the genus Felis of which the puma
was the largest cat. More recently, Wozencraft (1993) upgraded many of the
subgenera within Felis to full generic status and placed the puma back in the
monotypic genus Puma in support of Jardine’s 1834 recognition of the unique
evolution of the species (Nowell and Jackson 1996).
Goldman, in 1946, compiled the first intensive study of the phylogeography
of the puma. He re c o g n i zed thirty puma subspecies in the Western Hemisphere
based on morphological and pelage characteristics from a total of 764 specimens
composed of skulls and skins collected from throughout the puma’s range. How-
e ve r, the validity of some of these subspecies was questionable due to small sam-
ple sizes and an unknown age distribution of the specimens. Some of the skulls
examined we re without skins, and some of the skins we re without skulls. Ni n e-
teen of the subspecies were described from fewer than nine specimens each,
including six subspecies that we re described from only one specimen each. Go l d-
man (1946) described thirteen subspecies in No rth America (including Canada,
the United States, and Mexico). The subspecies Felis [Pu m a] concolor azteca,
which he described from 228 specimens (the largest number in the collection),
had a geographic range that included almost all of New Mexico, most of Arizo n a ,
and the western half of Mexico. This would be the subspecies that we studied.
Modern molecular genetic techniques tell a different story about puma phy-
logeography. Culver (1999) examined subspecies of puma by using three mito-
chondrial genes and ten microsatellite loci in biological samples collected from
a total of 315 pumas from throughout the historical range. Twelve of her speci-
mens were from pumas that we studied on the San Andres Mountains in New
Mexico. Culver (1999) identified six phylogeographic groups: (1) North Amer-
ica (Canada south to Guatemala and Belize), (2) Central America (Nicaragua,
Costa Rica, and Panama), (3) eastern South America (Brazil south of the Ama-
zon River and Paraguay), (4) northern South America (Columbia, Venezuela,
Guyana, French Guyana, Ecuador, Peru, and Bolivia), (5) central South Amer-
ica (northeastern Argentina and Uruguay), and (6) southern South America
(Chile, southwestern Argentina). Hence, Culver (1999, 2000) recommended six
subspecies for puma and named them based on the oldest Latin name among
the previous subspecies that were combined. They included P. c. couguar for
North America, P. c. costaricensis for Central America, P. c. concolor for northern
South America, P. c. capricornensis for eastern South America, P. c. cabrerae for
central South America, and P. c. puma for southern South America.
Description
The puma is the fourth-largest cat in world. It is comparable in mass to the
African leopard and snow leopard. The jaguar is the largest cat in the Western
Hemisphere, but its body size overlaps with that of the puma. At the extremes,
jaguars can be 80 percent larger than pumas. The African lion and tiger are two
to three times larger. In the Western Hemisphere, the puma and the jaguar are
the only extant large wild felids. Puma body size, in general, increases with lati-
tude (Kurtén 1973). Puma masses compiled by Anderson (1983) clearly indi-
cate sexual dimorphism. Males weighed significantly more than females (aver-
aging 40 percent heavier) in five out of six North American subspecies for which
data were available; insufficient sample size may have precluded the sixth. Mean
masses for adult specimens (twenty-four months or older) ranged from 52.8 to
68.0 kg for males and from 34.4 to 48.0 kg for females. The heaviest puma on
record was a 125.2 kg eviscerated male killed in Arizona (Musgrave 1926). Adult
male pumas reached total lengths (body plus tail) of 202.2–230.8 cm and adult
females measured 183.6–201.7 cm. Shoulder height measurements for adults
ranged from 56.0 to 78.7 cm for males and from 53.4 to 76.2 cm for females.
Adult desert pumas (twenty-four months or older) that we studied on the
San Andres Mountains exhibited similar morphometric characteristics (see
PHOTO 1. Adult male puma M3 was resident on the San Andres Mountains until his death
at twelve years of age.
Appendix 1). On average, males weighed 70 percent more than females. Male
masses ranged from 45.4 to 71.7 kg and female masses ranged from 24.5 to 40.4
kg [1]. Total lengths for males ranged from 200 to 227 cm, and females ranged
from 172 to 205 cm. The body and tail comprised about 63 percent and 37 per-
cent of the total body length, respectively. Height at shoulder for males ranged
from 54 to 67.5 cm, and for females ranged from 43 to 61 cm.
Pumas have a relatively compact head and a slender, elongate, muscular, lithe
body on powerful limbs. Forelimbs are somewhat shorter and heavier than the
aft and have larger paws on supine wrists, adaptations for handling prey and for
climbing. The hind limbs, proportionately the longest in large felids (Gonyea
1976), propel the puma during quick bursts and help it to catapult cliffs, rock
outcrops, and trees. The long, muscular tail is cranked to and fro apparently for
maintaining balance while attacking prey, hustling over broken terrain, and
climbing trees.
Retractile claws on the feet are used for self-defense, and for seizing prey,
gaining traction on slippery media, and climbing trees. Forefeet have five toes,
four of which support the puma on the ground. Although the pollex is nonsup-
porting, it has the largest claw and is used for combat, climbing, and grasping
prey. Hind feet have four supporting toes. Claws on the hind feet are not as
robust, recurved, or sharp as the foreclaws, but they are used for traction and
climbing. Toe pads are oval and the plantar pad has three distinct rear lobes, a
common feature in cats. Width measurements of plantar pads can be used to
help distinguish tracks of adult male and female pumas within the same popu-
lation. We used this as a guide in our study to quickly classify tracks that we
encountered. Tracks with forefoot plantar pads measuring 55 mm wide or more
were usually of adult males, and those measuring 50 mm wide or less were usu-
ally of females (see Appendix 1). However, sometimes there was confusion when
subadult males had foot sizes similar to those of females.
The outward appearance of an adult puma’s pelage is principally of one color
(hence the epithet concolor), with varying hues of tawny, reddish brown, and
grayish-brown. We observed all these variations within our study population. A
black “mustache” accents the sides of the muzzle at the base of the vibrissae and
contrasts sharply against the white hair around the mouth. Long, stout, white,
black, or brown vibrissae are highly innervated and are probably important for
directing a puma’s killing bite or a mother’s affections to her cubs. The nose pad
can be bright pink, neutral, or brown, and is sometimes flecked with black.
Black patches of hair are on the backs of the pinnas and may have a sparse
median gray patch. The front of the pinnas have white hair, thicker at the mar-
gins. Whitish hair covers the underparts of the neck, chest, abdomen, legs, and
tail. The dorsal side of the tail resembles the color of the puma’s back. The tip
of the tail is black. Dark bars transverse the inner sides of the forelimbs at and
below the angle of the elbows; they are black in younger animals and fade to
brown with age. Sometimes white flecks of hair are scattered on the top of the
head, nape, back, and shoulders. We believe the white hairs grow from skin cells
traumatized from claw and teeth punctures during fights between pumas, or
from tick bites. Although the adult puma lacks the stripes or spots present on
many of the other felids, its pelage camouflages it among like-colored objects
and light patterns in the rocks, grass, and shrub cover of its habitats. We can
attest to this during literally hundreds of approaches to observe pumas up close.
Their pelage colors and textures appear to meld with their surroundings.
Cubs are born fully furred and with black spots on reddish-brown to gray-
brown coats and with black rings on their tails. These coat patterns effectively
hide cubs among the rocks, holes, and low vegetation at nurseries. Birth weights
average 508.3 grams (Anderson 1983). Within two weeks of birth, a cub’s eyes
are fully open (Young 1946) and are blue. We noted that the eyes of cubs as
young as five months had turned to the brown or amber color of adult pumas.
Pelage spots and rings fade into light brown dapples by the time cubs reach nine
months old. Dapples disappear usually before twenty-four months of age, about
the time of sexual maturity. But we saw faint dapples persist on forelimbs and
hind limbs of known-age pumas that were thirty months old.
PHOTO 2. Puma F47’s four-week-old cubs at the birth nursery.

Genders of adult pumas can be distinguished by their external sex organs.

Males have a spot of black hair about 2.5 cm in diameter that encircles the open-
ing of the penis sheath and is about 12 cm anterior-ventral to the anus. The
scrotum, situated between the anus and the black spot, is mostly covered with
whitish hair but with flecks of silver and brown. The female’s vulva is directly
below the anus, and it may be encircled by a line of black hairs that is sometimes
faint and broken. We used these characteristics to accurately determine the sex
of pumas that we treed in Wyoming during a previous study (Logan et al. 1986)
either with the naked eye or with binoculars. The male’s sex organs are visually
evident, but those of the female are usually hidden beneath the base of the tail.
The puma has thirty teeth, with an upper/lower dental formula of incisors,
3/3; canines, 1/1; premolars, 3/2; molars, 1/1. The relatively small number of
teeth and vestigial teeth (P2, M1) may be the result of selection for shorter jaws
that deliver more effective biting action—a functional adaptation for solitary
hunters of large, potentially dangerous prey. Predatory behavior and a strict car-
nivorous diet are reflected in the teeth morphology. Large conical canines grab
prey; puncture skin, muscle, ligaments, and sinew; and separate skeletal joints.
Robust carnassials shear apart tissues and break bones. Small, closely spaced inci-
sors pluck hair or feathers from the skin of prey, strip tissues from bone, and are
used for self-grooming.
The form and function of the puma is that of a highly adaptive asocial pred-
ator suited for environments with rugged terrain and closed vegetative cover
where prey is dispersed and clumped. The puma preys on mammals ranging in
size from hares (Iriarte et al. 1990) to adult elk (Cervus elaphus) (Hornocker
1970). Of the large felids, from leopards to tigers, pumas routinely kill the
largest prey relative to their own mass (Packer 1986). Selective pressures
designed the puma as an ambush and stalking predator that attacks its prey with
a quick powerful rush, overwhelms the victim with staggering strength, and
delivers a killing bite in seconds.
LN N5LN5)
1. Summary morphometric statistics on adult pumas (twenty-four months
and older) captured on the San Andres Mountains, New Mexico (refer to
Appendix 1): Male masses (kg): n = 27, x– = 56.3 ± 6.1, median = 55.3.
Female masses: n = 33, x– = 33.1 ± 3.9, median = 33.1. Test for difference in
mass between males and females: two-sample t-test: t = 17.497, 58 d.f., P <
0.0001.
Chapter 3
Our Outdoor Laboratory
The San Andres Mountains comprised our 2,059 km2 study area in south-cen-
tral New Mexico (Fig. 3-1). The range extends north from San Augustin Pass on
U.S. Highway 70 in Doña Ana County to Mockingbird Gap in Socorro
County. The area was almost completely within White Sands Missile Range,
which was under the jurisdiction of the U.S. Army. About 93 km2 of the study
area was outside the western boundary of White Sands Missile Range and was
primarily public domain administered by the U.S. Department of Interior’s
Bureau of Land Management, with small private inholdings owned by cattle
ranchers who leased Bureau and state trust lands for grazing. Only about thir-
teen people, including an average of four puma research team members, inhab-
ited the San Andres Mountains year-round, for a human density of about 0.65
per 100 km2.
We chose the San Andres Mountains as our study area for a number of rea-
sons: (1) We preferred a desert area because intensive, long-term research on
pumas in a desert environment had not been done. This gave us the greatest
opportunity to contribute new knowledge about the species. (2) The mountains
comprised a large area of puma habitat not contiguous with other large blocks
of puma-inhabited areas; thus, we could isolate a puma population for intensive
study. (3) The San Andres Mountains constitute a self-contained ecosystem,
where none of the large mammals migrate. (4) The area was relatively undis-
turbed by humans because access and activities were highly restricted by the
U.S. Army. (5) Through agreements with cooperating land and natural
resources management agencies, we could minimize human-caused confound-
ing variables by prohibiting puma hunting and furbearer trapping and restrict-
23
FIGURE 3-1. The puma study area (2,059 km2), San Andres Mountains, New Mexico. The
sixteen camps are depicted as triangles and include San Nicholas (SN), East San Andres
(ES), San Andrecito (SA), Dead Man (DM), Lost Man (LM), Hembrillo Basin (HB),
Hembrillo Narrows (HN), Sulphur (S), Buckhorn (BH), Rhode’s Pass (RP), Hardin (H),
Rosebud (RB), Good Fortune (GF), Puma Basin (PB), Edifice (E), and Mockingbird Gap
(MG).
PHOTO 3. San Andres Peak after sunup.
Slide @318%
PHOTO 4. Upper Sulphur Canyon.

ing mule deer hunting. In addition, because livestock was prohibited on all but
5 percent of the area, livestock were not an alternate source of food for pumas,
nor could they compete with native ungulates. The only human manipulations
of the study population would be our own experimental removal of pumas. (6)
A network of dirt roads and primitive trails accessible by four-wheel-drive vehi-
cle enabled us to thoroughly explore the mountains for pumas.
We partitioned the study area into two subunits to provide us with a Treat-
ment Area (TA) and a Reference Area (RA). The 703 km2 Treatment Area in the
southern one-third of the study area was delineated because it held all of the
radio-collared mule deer and almost all of the radio-collared desert bighorn
sheep in our study. We removed pumas from that area to experimentally study
puma population dynamics and how puma predation influenced survival rates
and agent-specific mortality in radio-collared deer and sheep. The Reference
Area was 1,356 km2 in size; no manipulations of the puma population occurred
there, allowing us to observe puma population dynamics under natural condi-
tions (i.e., in the absence of human exploitation).
Physiography and Geology

The San Andres Mountains are within the easternmost part of the Mexican
Highland section of the Basin and Range physiographic province (Hawley et al.
1976). The mountain range is long and narrow, measuring over 127 km long
and 9–30 km wide. It is part of a broken chain of mountains extending to the
Oscura Mountains to the north and terminating in the Organ and Franklin
Mountains to the south (Fig. 3-1). Two large desert basins flank the range: the
Jornada del Muerto to the west and the Tularosa to the east. The Tularosa Basin
spans some 97 km east to west at its widest point; the Jornada del Muerto, about
55 km.
The San Andres Mountains are composed of a westward-tilted fault block of
Precambrian granite and Paleozoic-age limestone, dolomite, sandstone, and
shale (Eidenback 1983, Kottlowski et al. 1956). A major fault zone borders the
range to the east and separates the great collapsed crest of the Tularosa Basin
from the steep east face of the mountains. The precipitous east escarpment rises
up to 1,500 m above the basin floor and is characterized by cliff-forming lime-
stones (Van Devender and Toolin 1983). The east face has been eroded back
from the original fault boundary. Alluvial fans formed from the weathering
granite and limestone bedrock are broken by a network of arroyos and outcrops
of Precambrian granite and schist. Fault activity during the last ten million years
has disturbed some of the alluvial deposits and underlying bedrock, and fault
scarps formed during the past two million years are present along most of the
CHAPTER 3. OUR OUTDOOR LABORATORY 27
basin-range margin (Hawley 1983). The western slope is relatively gentle. There,
tilted sedimentary rocks dip 10–20 degrees westward into the younger rocks and
valley fill of the Jornada del Muerto syncline (Kottlowski et al. 1956). Within
the central portion of the mountain range there are fourteen major east-west,
convoluted canyons that drain numerous other north-south branches. Internal
basins are interrupted by subsidiary tilted blocks, and folds.
The southern end of the San Andres Mountains is more geologically similar
to the adjacent Organ Mountains to the south and consists of a course-grained
phase of the Organ Mountains monzonite batholith. Mockingbird Gap, at the
north end of the range, is a broad pass in a down-faulted anticlinal axis. The
general north-south trend of the range continues northward in a series of low
hills called the Little Burros and is separated by another broad gap from the
Oscura Mountains. But the Oscura fault blocks dip eastward and are bounded
on the west by a fault-line scarp (Kottlowski et al. 1956). Elevations within the
our study area range from about 1,280 m along the east piedmont to 2,730 m
at Salinas Peak.
The San Andres Mountains are relatively well watered. There are approxi-
mately one hundred natural springs distributed throughout the study area that
produce perennial surface water. Another thirty springs are intermittent. Per-
haps as many as one hundred other sites supply ephemeral water, charged only
during the monsoon season. Flows at springs are variable, ranging from a few
centimeters deep in a reach of up to 1 m, to shallow, narrow streams that reach
about 0.5–1 km. However, the latter only occur at three springs. The average
distance between springs is about 1 km, with a range of about 0.06 to 6.8 km
(Boykin et al. 1996).
Climate
The San Andres Mountains have a semi-arid climate classified as w a rm temperate
in Brown et al. (1998), A Classification of No rth American Biotic Communities.
Precipitation falls mostly as rain during the monsoon season of July through Se p-
tember. This results from large-scale systems that push tropical and subtropical
moist air from the Gulf of California and the Gulf of Mexico nort h w a rd into
New Mexico, bringing intense, but brief, scattered rain showers. Winter moisture
fluctuates depending on the re l a t i velatitude of the jet stream, and storms that do
occur generally produce slow, drizzling rain or light snow. Weather is seasonal,
with hot summers, warm springs and falls, and cold winters.
We summarized precipitation data for the Ash Canyon gauge because it
occurred at mid-elevation (1,731 m) in the central portion of the Treatment
Area and complete data have been collected there since 1937 by the U.S.
FIGURE 3-2. Annual and growing season (July to September) precipitation, San Andres
Mountains, New Mexico, 1985–1994.
Department of Agriculture’s Jornada Experimental Range (Fig. 3-2). During our

research period, 1985–1994, mean annual precipitation was 43.6 ± 11.4 cm (±
1 SD), which was 10.8 cm greater than the mean annual precipitation during
the previous forty-eight years (1937–1984; ( x– = 32.8 ± 10.2 cm). Moreover, the
first, third, and fifth highest annual precipitation occurred during our study
(1991, 1992, 1986, respectively). The monsoons bring almost half of the annual
precipitation. Thus, the growing season in the San Andres Mountains is marked
from July to September. During our research period, an average of 19.8 ± 5.9
cm of precipitation fell during those months. Below-average precipitation dur-
ing the growing seasons of 1992–1994 contributed to severe drought that
resulted in the complete drying of some of the perennial springs. These drought
conditions would prove to be pivotal in the way pumas affected the desert mule
deer and desert bighorn sheep populations on the San Andres Mountains.
Snow fell infrequently during December through February. Continuous
snow cover did not persist for more than a few days except along the north-fac-
ing slopes at higher elevations. Individual storms generally brought less than 2.5
cm of snow to the lower elevations, whereas higher elevations occasionally
received 12 cm or more. The most severe storm occurred in December 1987 and
brought 46 cm of snow to elevations above 1,372 m.
Temperature records from the Jornada Experimental Station (elevation
1,349 m, 15 km west of our study area) for 1985–1994 gave a mean annual
temperature of 14.5 degrees Celsius. July was the hottest month, with tempera-
tures averaging 25.5 degrees Celsius. January was the coldest month; tempera-
tures averaged 3.4 degrees Celsius. Extreme temperatures in summer reached

42.8 degrees Celsius and in winter dipped to –24 degrees Celsius.
The “windy season” typically occurred in March and April. Winds often
exceeding 50 km per hour blew across the desert from the west and southwest,
parching the herbaceous vegetation and whatever winter moisture was stored in
the thin topsoil. Wind velocities as high as 187 km per hour were clocked at the
summit of Salinas Peak (White Sands Missile Range climate records).
Flora
Woodrat (Neotoma spp.) middens examined by botanists on the San Andres
Mountains indicate when the extant regional biota emerged. A mixed-conifer
forest dominated by Douglas-fir (Pseudotsuga menziesii), blue spruce (Picea pun -
gens), and ponderosa pine (Pinus ponderosa) was present at the higher elevations
during the late Wisconsin glacial age, about fifteen thousand years ago. Lower
elevations supported an open juniper (Juniperus spp.) –oak (Quercus spp.) wood-
land (Van Devender and Toolin 1983). During the early Holocene (about nine
thousand years ago) the climate became warmer and dryer and desert commu-
nities began to develop. The vegetation and climate have been virtually modern
for about the last 4,400 years.
Today, the San Andres Mountains lie at the northern edge of the Chihuahua
Desert, which extends mostly southward into northern Mexico and westward
into southwestern New Mexico and southeastern Arizona. Flora on the San
Andres Mountains mostly represents the Chihuahuan Interior Chaparral, Chi-
huahuan Desert Scrub, and the Chihuahuan (Semidesert) Grassland biotic com-
munities described in Brown et al. (1998). Localized cooler and moister condi-
tions support small communities of Ponderosa pine and Gambel oak (Quercus
gambelli) on Salinas Peak, and Gambel oak with an understory of snowberry
(Symphoricarpos rotundifolius) on San Andres Peak.
Because plant communities on the San Andres Mountains had not been pre-
viously described and mapped, we developed our own scheme to describe the
vegetative settings for our observations. We recognized seven vegetative cover
types, including tall mixed desert shrub, low mixed desert shrub, grass, mixed
mountain shrub, dry channel, moist riparian, and piñon (Pinus edulis) –juniper
(Juniperus spp.). Vegetation was highly influenced by elevation, aspect, parent
material, and soil type. Sometimes two or more vegetation types occurred as co-
dominants, and shrubs and grass were frequent sub-dominants in the piñon-
juniper type.
Dominant plants in the tall mixed desert shrub type were generally taller
than 1 m. Creosote (Larrea divaricata), mesquite (Prosopis glandulosa), mimosa
(Mimosa biuncifera), and southwestern condalia (Condalia lycioides) typify this

community and were usually found in the desert basins, on alluvial fans, and on
south- and west-facing slopes. Stony uplands supported ocotillo (Fouquieria
splendens) and sotol (Dasylirion wheeleri). Drier slopes supported low mixed
desert shrub vegetation such as mariola (Parthenium incanum), feathered dalea
(Dalea formosa), prickly pear (Opuntia spp.), yucca (Yucca baccata), and agave
(Agave parryi).
Grass communities were composed of black, blue, and sideoats grama
(Bouteloua eriopoda, B. gracilis, B. curtipendula), three-awn (Aristida spp.), and
muhly (Muhlenbergia spp.). Black grama grew in the limestone hills in associa-
tion with ocotillo and hairy mountain mahogany (Cercocarpus breviflorus).
Muhly typically occurred on alluvial slopes. Localized stands of invader species
such as burro grass (Schleropogon brevifolius) and snakeweed (Gutierrezia spp.)
were indicative of past overgrazing; however, native grasses have recovered dra-
matically in most areas (Van Devender and Toolin 1983).
Protected slopes and higher elevations supported mixed mountain shrubs
generally taller than 1 m, including mountain mahogany, skunkbush sumac
(Rhus trilobata), Wright’s silktassel (Garryea wrightii), oak, and pale hop tree
(Ptelea trifoliata). Mountain shrubs usually integrated with desert shrubs where
the soil became shallow and stony.
Piñon and juniper grew in the coolest areas—generally north slopes and the
highest elevations. Coverage varied from a few scattered trees to 100 percent.
Trees were typically short and usually did not exceed heights of 6 m. Of the
junipers, one-seed (J. monosperma) was the most represented. Alligator-bark
junipers (J. depeana) were only found in the most protected areas.
Dry channel vegetation grew in the major drainage channels. Wide canyon
bottoms were gravelly, sandy, and well drained, and they supported vegetation
such as Apache plume (Fallugia paradoxa), brickellbush (Brickellia laciniata),
algerita (Berberis trifoliata), and desert willow (Chilopsis linearis).
Moist riparian vegetation grew where springs, seeps, and a high water table
occurred. Cattails (Typha angustifolia) grew in the water, and on the banks were
hackberry (Celtis reticulata), Velvet ash (Fraxinus velutina), and cottonwood
(Populus deltoides weslizeni) trees, the latter two species reaching heights of up to
9 m. At some perennial springs, and where the water table was high, the exotic
salt cedar (Tamarix ramosissima) had invaded.
Fauna
Four native ungulate species occurred on the study area: desert mule deer, desert
bighorn sheep, pronghorn (Antilocapra americana), and javelina (Pecari tajacu).
None of these animals made seasonal or elevational migrations. Mule deer were
the most abundant ungulate throughout the mountain range, numbering in the
thousands. They even inhabited the most rugged terrain preferred by the desert
bighorn sheep. They were sparse or nonexistent in the broad desert basins flank-
ing the mountains. Sport-hunting of the deer was prohibited on the San Andres
Mountains between 1985 and 1989. However, as a result of pressure from sport-
hunters in southern New Mexico, limited hunting for buck deer was allowed by
the New Mexico Department of Game and Fish in the northern one-quarter of
our study area (i.e., north end of the Reference Area) in alternate years includ-
ing 1990, 1992, and 1994. In those years, forty-four, eighty-two, and sixty-
seven bucks, respectively, were killed.
The San Andres Mountains supported the only extant, naturally occurring
population of desert bighorn sheep in New Mexico. This herd was a peripheral,
isolated population at the eastern edge of the geographic range for the species.
During this study, only a remnant population of thirty to forty desert bighorns
remained. They ranged primarily in the steep, rugged terrain of the east escarp-
ment, and almost all of them lived in the Treatment Area. The desert bighorn
sheep is a New Mexico state-listed endangered species, with fewer than 280
sheep statewide (Pederson 1996).
Pronghorns infrequently ranged into the foothills in the northern portion of
the San Andres Mountains. Typically they traveled in small herds of fewer than
twenty animals. The open desert flats were the areas they usually inhabited.
Sport-hunting of pronghorns was limited; about thirty to forty buck permits
were issued to hunters each year (White Sands Missile Range 1994).
Javelina probably numbered fewer than fifty animals on the San Andres
Mountains. Small, scattered bands of fewer than ten animals each occurred in
the denser piñon-juniper communities in the northern and southern portions of
our study area.
Oryx (Or yx gazella) were introduced onto the San Andres Mountains in
1969 (Saiz 1975). The fifty-one animals in the original herd were from trans-
planted stock from Africa’s Kalahari Desert. Oryx preferred the broad desert
basins east and west of the mountains, and the population increased and
expanded its range during the ten years of our study. Consequently, oryx began
to use the large canyons that dissected our study area on a more frequent basis.
By the close of the study in 1995, the oryx population numbered approximately
one to two thousand animals (White Sands Missile Range 1994). At the begin-
ning of our study, there were probably no more than twenty-five oryx in the
mountains at any time, but by 1995 there may have been fifty to one hundred.
Because oryx were at relatively low numbers, ate a diet consisting of more than
80 percent grasses (Smith et al. 1998), and had no need for surface water (Saiz
1975), they probably competed little, if at all, with mule deer and desert
bighorn sheep. The New Mexico Department of Game and Fish attempted to
limit the oryx population on White Sands Missile Range through sport-hunt-
ing.
Feral horses, remnants of the recently past ranching culture, ranged in the
Tularosa Basin on White Sands Missile Range. However, we never observed
horses on our study area.
Cattle grazed in valley bottoms and foothills along the western boundary of
the study area. Their numbers were small, probably never exceeding more than
one hundred head at any one time. Herds were composed of cow-calf groups
with few bulls.
A variety of small mammals inhabiting the San Andres Mountains were
potential puma prey. The most abundant were the black-tailed jackrabbit (Lepus
californicus), desert cottontail (Sylvilagus audubonii), New Mexico ground squir-
rel (Spermophilus mexicanus), rock squirrel (S. variegatus), white-tailed antelope
squirrel (Ammospermophilus leucurus), and woodrat (Neotoma mexicana). We
rarely observed porcupines (Erethizon dorsatum).
About 183 species of birds and thirty-five species of reptiles and amphibians
have been documented on the San Andres Mountains (White Sands Missile
Range 1994). Ground-dwelling birds included three species of quail. Gambel’s
(Callipepla gambelii) and scaled (C. squamata) quail were found in the dry chan-
nel and riparian communities, and were most abundant during years of high
precipitation. Montezuma quail (Cytonx montezumae) apparently occurred in
small numbers with a patchy distribution. Wild turkeys (Meleagris gallopavo)
were represented by probably fewer than ten animals. Avian scavengers included
turkey vultures (Cathartes aura), golden eagles (Aquila chrysaetos), red-tailed
hawks (Buteo jamaicensis), and ravens (Corvus spp.), but these occurred in such
low numbers that they rarely clued us to locations of dead animals.
Other carnivores on the study area included the coyote (Canis latrans), gray
fox (Urocyon cinereoargenteus), badger (Taxidea taxus), bobcat (Lynx rufus), ring-
tail (Bassariscus astutus), and the striped (Mephitis mephitis) and hog-nosed
skunks (Conepatus mesoleucus). Coyotes usually inhabited the upper basins of
drainages. We frequently saw gray foxes and their tracks throughout the moun-
tains, but we rarely saw bobcats or their tracks.
Historical accounts of jaguars, wolves (Canis lupus baileyi), and bears (Ursus
americanus, U. arctos) on the San Andres Mountains are either extremely rare or
nonexistent. We found one anecdotal account of a Biological Survey hunter
whose dogs “jumped,” but did not catch, a jaguar in the San Andres Mountains
in 1937 (Halloran 1946). Black bears apparently occurred in low numbers as
late as the 1890s (Halloran 1946) but were extirpated about 1908 (J. P. Wood,
Jr. personal communication). No records indicated the presence of grizzly bears
or wolves on the mountains. However, because wolves once occupied other
ranges in southern New Mexico—including the Black Range and Mogollon
Mountains to the west and the Sacramento Mountains to the east—all of which
were within dispersal range for wolves, it is likely that the San Andres Moun-
tains supported at least a small population of wolves from time to time. Such a
population could have been easily extirpated by the earliest non-native human
settlers.
History of Human Use

Human presence on the San Andres Mountains has waxed and waned since pre-
historic times. Nomadic bands of paleolithic hunters pursued now-extinct large
herbivores in the northern San Andres Mountains and adjacent basins about
eleven thousand years ago. Those people also used the area to quarry fine-
grained chert from which they manufactured their hunting and processing tools.
They left some of their fluted projectile points and scrapers at campsites in
Mockingbird Gap and at the mouth of Rhodes Canyon (Beckett 1983, Breter-
nitz and Doyel 1983). By the time modern climatic conditions prevailed some
Slide @318%
PHOTO 5. Puma paw petroglyph near Three Rivers, New Mexico.

four to five thousand years ago, paleolithic adaptations had been replaced by
subsistence hunting-and-gathering traditions known as Archaic or Desert Cul-
ture. People hunted modern animal species and relied heavily on wild plant
foods such as juniper berries, piñon nuts, mesquite beans, agave, yucca, and
prickly pear. They left behind baskets, grinding tools, and projectile points in
the San Andres Mountains (Beckett 1983, Eidenbach 1983). During about
1,700–1,200 years ago people in the mountains had a more sedentary lifestyle.
Some of their pithouses and shards of their pottery are still there. They also prac-
ticed horticulture, growing corn and beans. We found some of their discarded
corncobs in dry undercut ledges and shallow caves. These were probably the
people that created a red ochre pictograph of a puma that we found on a lime-
stone cliff face only 30 m from our Hembrillo Narrows Camp, near the center
of our study area. About the same time, relatives of those people, living about
70 km to the northeast at the western foot of the Sacramento Mountains, etched
an oversized likeness of a puma’s paw into basalt. These are the first known
observations of pumas to be recorded by humans in the region. For some
unknown reason, perhaps due to an extended drought in combination with
social and economic conflicts, people essentially abandoned the San Andres
Mountains about six hundred years ago. Thereafter, aboriginal activity was spo-
radic (Breternitz and Doyel 1983).
When the Spanish arrived in southern New Mexico in 1540, most of the
native peoples they encountered were settled in pueblos along the Rio Grande.
Nomadic Apache, who had arrived about 1500, raided the pueblos and villages
of settlers (Sale and Laumbach 1989). Apache groups used the San Andres
Mountains for seasonal hunting and gathering and as a base from which to con-
duct raids. They also used the mountains as a refuge from punitive pursuits of
the Spanish, Mexican, and United States military. Raids continued into the
1880s when the last Apache bands were confined to reservations. In 1880, one
of the last battles of the Apache-American war took place in Hembrillo Canyon,
where the famous Apache Chief Victorio fought the U.S. 9th Cavalry (Thrapp
1980, Breternitz and Doyel 1983, Sale and Laumbach 1989). In the canyon,
breastworks and pictographs of horse-mounted men testify to those parlous
times. Our Hembrillo Basin Camp lay in the middle of the battlefield. Cavalry-
issue brass littered the ground; here and there on the limestone were splashes of
lead from bullets. The battle marked the end of aboriginal peoples’ use of the
San Andres Mountains.
Anglo and Mexican homesteaders established the first permanent ranches
on the San Andres Mountains in the late 1860s. The 1880s brought an influx
of Texas cattlemen to the area. Drought and overgrazing in the 1890s forced
many ranchers to shift to raising goats and sheep. Settlement in the mountains
continued through the late 1930s. Ranches we re generally small family outfits
that raised angora goats and sheep for the mohair and wool industries. They
also kept cattle, horses, and mules for their own uses. Family goat herds num-
b e red from a few hundred up to two thousand head (Eidenbach 1989). Go a t
operations we re active throughout the San Andres Mountains. We counted at
least nineteen ruins with corrals and chutes designed for handling goats and
s h e e p. It is likely that during the period when goats and sheep we re raised, mule
deer and desert bighorn sheep we re subjected to the greatest competition for
food and water. Overgrazing may have helped to transform mountain grass-
lands to communities dominated by shrubs and trees. Na t i ve sheep, in part i c-
ular, we re also at the greatest risk of contracting diseases and parasites fro m
their domestic kin.
Old-time ranchers who lived in Bear Den Canyon, upper Cottonwood
Canyon, and Deadman Canyon, all within the heart of the San Andres Moun-
tains, recollected to us that pumas were fairly common in the mountains, always
presenting a threat especially to goats, sheep, and horse and mule foals. They
consistently described mule deer as abundant or common, except during
droughts. Mule deer meat was a staple food. Desert bighorn sheep, they said,
were always in low numbers.
Hard-rock mining activity was widespread in the mountains during the
period of settlement. Miners were principally searching for galena, copper,
barite, and quartz (Sandoval 1979b, Breternitz and Doyel 1983).
The U.S. Government began acquiring land in the area in 1912. That year
the Jornada Experimental Range, under the jurisdiction of the U.S. Forest Ser-
vice, was formed along the west flank of the San Andres Mountains and
extended westward onto the Jornada del Muerto. In 1933, White Sands
National Monument was established in the center of the Tularosa Basin. Eight
years later, the San Andres National Wildlife Refuge, encompassing 230 km2,
was created near the southern end of the San Andres Mountains, primarily for
the protection of desert bighorn sheep and other desert wildlife (Hoban 1986).
White Sands Proving Ground (later named White Sands Missile Range) was
established in 1945, principally to aid efforts to win World War II. This had a
great impact not only on the land in the region but also on the world. On July
16, 1945, the first atomic bomb was exploded in the desert basin at Trinity Site,
12 km north of the northern tip of the San Andres Mountains. Inhabitants of
the region had to evacuate to make room for the missile range; hence, the his-
toric era of ranching and mining came to an end, and tight restrictions were
placed on other public uses. The San Andres Mountains functioned as a tremen-
dous natural buffer zone for military research and development that took place
mainly in the desert basins. Once again, the mountains were left practically
undisturbed by humans. During the forty years leading up to our study, the
mountains recovered from some effects of previous human use.
History of Puma Exploitation

Pumas were killed by ranchers and government hunters on the San Andres
Mountains during the late 1800s and early 1900s mainly to protect livestock
(Eidenbach and Morgan 1994). J. P. Wood Jr. (personal communication), who
lived in Bear Den Canyon, perceived that local ranchers kept pumas from
becoming a significant problem by keeping them in “check” through trapping
and hunting with hounds. U.S. Fish and Wildlife personnel, while carrying out
predator control to protect wildlife during 1940 to 1979, killed fourteen pumas
on or near the San Andres National Wildlife Refuge (Muñoz 1983). Another
five pumas were killed by hunters on other parts of the San Andres Mountains
from 1966 to 1971 for the same reason (Anderson and Taylor 1983). The toxi-
cant compound 1,080 (sodium monofluoroacetate) was used to control carni-
vores on the refuge during 1949 and 1951 (Muñoz 1983). We found three of
the former 1080 bait stations; they were marked by weathered juniper posts, the
top one-quarter painted red. One still had rusting wire wrapped around the base
that once held deadly bait. There were no records of how many animals were
poisoned in this way.
Sport-hunters with hounds killed eight pumas in the mountains from
December 1979 to February 1985, including one male (age not recorded), four
females (ages not recorded), and one adult female and her two male cubs. Yet
another predator-control action during September 1980 to May 1984 was car-
ried out to protect the remnant population of desert bighorn sheep on the San
Andres Mountains (Sandoval 1979a, Evans 1983). Trappers for the New Mex-
ico Department of Game and Fish, U.S. Fish and Wildlife Service Animal Dam-
age Control, and the San Andres National Wildlife Refuge killed forty-two
pumas, including thirty-four adults. This last control action was terminated
after New Mexico Department of Game and Fish biologists evaluated its effec-
tiveness and concluded that the removal of the pumas did not cause an increase
in desert bighorn sheep survival (Evans 1983). Assuming that a total of forty
adult pumas were killed on the San Andres Mountains during the forty-three
months of exploitation (December 1979–February 1985), then the kill rate
averaged eleven adult pumas per year on the San Andres Mountains. The kill
rate for all fifty pumas (i.e., adults, subadults, and cubs) averaged fourteen
pumas per year.
The Oscura Mountains constitute a 673 km2 patch of puma habitat 9 km
northeast of the north end of the study area. Puma control was implemented
there too from October 1980 to October 1981 for the purpose of protecting the
desert bighorn sheep. The rationale was that if the puma population was deci-
mated there, then movement of pumas onto the San Andres Mountains would
be minimized, further reducing puma predation risk to the sheep. An Animal
Damage Control agent killed thirteen pumas during the one-year period. Dur-
ing our ten-year study, however, no pumas were killed for sport-hunting or con-
trol on the Oscura Mountains.
On the 272 km2 Organ Mountains, connected to the south end of our study
area, puma-hunting pressure was relatively light. A total of five pumas (three
males, two females) were killed from 1986 to early 1995 (New Mexico Depart-
ment of Game and Fish kill records).
Original—and rather crude—records kept for the forty-two pumas killed on
the San Andres Mountains for predator control during the period 1980 to 1984
indicated thirty-four were adults (twenty males, fourteen females), two were
yearlings (one male, one female), and six were cubs (i.e., dependent offspring;
one male, five females). Similar records kept for pumas killed in the Oscura
Mountains indicated eleven were adults (six males, five females) and two were
yearling females. We inspected the preserved skulls of twenty-nine of the pumas
(sixteen from the San Andres Mountains, thirteen from the Oscura Mountains)
that were deposited at the Museum of Southwestern Biology (University of New
Mexico, Albuquerque) to compare dental and suture characteristics with skulls
of known-age and approximately known-age pumas we studied on the San
Andres Mountains. The estimated mean age of nine adult males was 61.2 ± 32.3
months (± 1 SD), and the mean age of eleven adult females was 80.4 ± 36
months. One male and two females were thirteen to eighteen months old. One
male, three females, and two pumas of unknown sex were less than twelve
months old.
Beginning in March 1985, and throughout the time of our study, the San
Andres Mountains were closed to puma hunting, except for about 5 percent of
the study area that lay outside of the west White Sands Missile Range boundary.
The New Mexico Department of Game and Fish did not want to offend ranch-
ers operating in that area by completely eliminating puma hunting. However,
even there, all pumas tagged by us were protected by state hunting regulations.
Fur-bearer trapping was also eliminated on the White Sands Missile Range por-
tion of our study area in order to minimize risks to pumas. This regulation was
particularly important to protect cubs.
Chapter 4
Studying Wild Pumas
Pumas are cryptic, crepuscular, and nocturnal, and normally shun humans.
They are scarce in their complex habitats. These traits make pumas one of the
most difficult large-terrestrial mammals to study in the world. Field ecologists
normally “observe” pumas by their tracks, beds, nurseries, playgrounds, drop-
pings, and scraps of prey. Because chances to observe wild pumas directly are
extremely rare, direct, quantitative data must come from pumas wearing collars
containing radio transmitters. To attain our research objectives, we employed
those means of data gathering and diligent field ecology. To do that, we lived
among the pumas.
Life Afield
We used sixteen base camps located at strategic locations to efficiently cover
large parts of the San Andres Mountains (Fig. 3-1). The camps, along with a net-
work of primitive roads accessible to four-wheel-drive vehicles, facilitated a thor-
ough exploration of the entire study area. We trucked in all our staples and
brought some basic comforts to the camps by outfitting them with stoves,
lanterns, and cots. Most of our camps were sheltered by large tents. Four of the
camps were in abandoned or rarely used structures, including a 1940s-vintage
trailer, a hunter checkstation, and a former military police station. Our favorite
was San Andrecito Camp, which consisted of a 3 m ¥ 3 m shack built out of
missile-engine crates. We retrofitted bunk beds, shelves, and windows with
material salvaged from a dilapidated abandoned barn about 10 km away. We
covered the shack with tar paper to keep out blowing sand. The camp, perched
39
on a promontory, overlooked extraordinary puma habitat at the confluence of

two north and south drainages that fed into a rugged, winding, east-draining
canyon. Two springs within 100 m of camp provided vital water for a variety of
wildlife. Pumas that came there to drink, hunt, and socialize passed us in the
night, leaving their footprints in the sand. On a few occasions we listened and
even watched pumas caterwauling.
Our team normally consisted of four persons. During our research from 6
August 1985 to 31 March 1995, our team spent on average 254 days a year in
the field (range = 230–289). We accumulated a total of 2,445 field days and
about 9,780 total person-days of field research. We were on the San Andres
Mountains year-round, normally in ten-day stints, then back in civilization for
four days for rest, baths, supplies for the next stint, and to tend to necessary
paperwork and needs of agency administrators. However, puma activities some-
times dictated extended stays in the field. Before each ten-day stint, our research
team met to discuss observations and progress in the previous stint and the
short-term objectives we aimed to accomplish in the field.
Each biologist was usually stationed at a different camp. We kept in contact
with each other with two-way radios. This allowed us to cover different portions
of the study area. From the camps and roads, we ranged out on foot in system-
atic searches for evidence of pumas. The usual dry desert conditions and sandy
Slide @318%
PHOTO 6. Hembrillo Basin Camp.

Slide @318%
PHOTO 7. San Andrecito Camp.
Slide @171%
PHOTO 8. Puma tracks in the sand.

or dusty soil surfaces often preserved puma tracks for several days; puma scrapes
would persist for weeks at a time. Observations of puma sign gave us a rudi-
mentary understanding of how the cats were using the landscape. All observa-
tions were recorded in field journals and plotted on 7.5-minute (1:24,000-scale)
U.S. Geological Survey topographic maps. Using that information, we set out
to capture the pumas. Once satisfied that we had captured and identified all of
the pumas in an area served by a camp, we moved to the next camp in another
part of the study area and repeated the process. We continued this process year-
round and year after year for the duration of the study.
To Catch a Puma
The only reliable way to study puma population dynamics and social organiza-
tion is to capture as many pumas in the population as is possible, individually
tag them, release them, and radio-track their movements. To capture pumas,
most other studies used packs of trained dogs to trail pumas by scent, then bay
them in trees or on rock ledges. Once bayed, pumas were chemically immobi-
lized, then lowered to the ground for examination (Hornocker 1970, Logan et
al. 1986). Success and safety of this method are largely dependent upon the pres-
ence of trees, which pumas use to escape the pursuing dogs. This mode of escape
is probably the result of a long evolutionary history with other competitors and
predators such as wolves and bears. Still, even in treed areas, pumas are some-
times bayed on the ground and mauled by dogs, resulting in severe injuries or
death to the pumas or dogs (Shaw 1977, Barnhurst 1986, Anderson et al. 1992).
The San Andres Mountains presented us serious problems because trees that
could be used by pumas for escape were sparse. Initially, we tried using Frank
Smith’s dogs to capture pumas, but without success. Either the dogs could not
catch up to the pumas in the dry, rugged terrain or they bayed pumas momen-
tarily on ledges from which the cats later escaped using routes the dogs could
not follow. In one instance, the puma fought the dogs, knocking one of them
off a cliff, breaking its back. We realized early in the project that dogs simply
were not going to yield consistent, safe captures of pumas. We estimated that
80–90 percent of catches would probably be on the ground or on rimrocks and
ledges—dangerous places for dogs to hold pumas at bay and for biologists to
handle the animals. Cubs would especially be vulnerable to mauling by dogs.
Knowing these high stakes, we decided to try a different method for capturing
pumas.
We built on Frank’s extensive knowledge of catching nuisance and depre d a t-
ing pumas and black bears with foothold traps and snares. We tried the No. 41/2
CHAPTER 4. STUDYING WILD PUMAS 43
Slide @318%
PHOTO 9. Puma M7 caught in a snare by his right forefoot.
steel traps (Woodstream Corp., Animal Trap Div., Lititz, Pennsylvania 17543)
sparingly early in the study (6 August 1985–11 January 1986) and had limited
success. In fact, we only caught one adult female puma with rubber-padded traps
after 114 trap-days (one trap-day is defined as one trap set for one day). Be c a u s e
of the great risk of injury that those heavy, powe rfully jawed traps brought to
small pumas and especially nontarget animals (e.g., deer, sheep, javelina, coyotes,
foxes, bobcats, rabbits, hares), we abandoned the technique. Instead, we focused
on foothold snares to capture pumas that weighed 10 kg or more.
Snare assemblies (Schimetz/Aldrich Spring Activated Animal Snare, Sekiu,
Washington 93831) were modified to minimize injuries to pumas and to avoid
capturing nontarget animals. We attached two to four rubber bungee cords into
the snare assembly to absorb shock as the captured animal struggled. In addi-
tion, we attached a slide stop to the foot loop, which minimized its closure to
18–19 cm circumference. This assisted blood circulation to the puma’s foot and
allowed smaller-footed, nontarget animals to pull free from the loop (Logan et
al. 1999).
Moreover, we carefully chose snare sites to minimize injury to captured
pumas and risk to nontarget animals. Preferred sites had limber bushes with
multiple basal stems to securely anchor the snare drag, and a safety area with a
Slide @171%
PHOTO 10. Linda Sweanor using a

pole syringe to inject puma F37 with
immobilizing drugs.
circumference 5 m or more around the anchor point. The safety area was cleared
of trees, old fence wire, and potentially injurious vegetation (e.g., cacti, yuccas,
agaves) to minimize the probability that a struggling puma might be injured.
Snares were not set near cliffs or water. We avoided setting snares on trails inten-
sively used by deer and oryx. But when paths indicated infrequent ungulate use,
we positioned stick hurdles horizontally about 60–70 cm above the snare.
Ungulates generally went around or jumped over the sticks; naive pumas walked
under (Logan et al. 1999).
We set one to six snares on paths used by pumas and at puma kill caches and
scrape sites. We set them at carcasses of ungulates that died from other causes in
case pumas came to scavenge. In addition, we set snares along likely travel routes
even if we were not certain that pumas used them. Where a travel way was too
wide to constrict a puma’s movements through a snare set, we augmented the set
with a sight lure consisting of a shiny piece of tin, and occasionally feathers, dan-
gled from a string or wire over the snare. In total, we established 1,211 snare
sites throughout the San Andres Mountains.
Our intent was to capture and tag all of the adult and independent subadult
Slide @171%
PHOTO 11. Ken Logan hauling puma

F37 for translocation to northern New
Mexico.
pumas on the study area. We targeted all of them initially; then as the marked
population of pumas grew, we targeted the unmarked or unidentified pumas in
the population. Later, we recaptured pumas that had radio collars that were non-
functional or had waning battery power. Pumas with well-functioning radio col-
lars were reliably identified in the field by radiotelemetry; therefore, we persist-
ently tried to avoid recapturing those cats. To prevent excessive stress and trauma
to captured animals, we checked all snares each day by 10 a.m. during spring
and summer and by 12 p.m. during fall and winter. In the hottest part of sum-
mer we sometimes checked snares twice per day (again at about 4 p.m.). But
during short periods of exceptionally hot, cold, or snowy weather, we deacti-
vated all snares to eliminate risk altogether.
From 6 August 1985 to 23 February 1995, we accumulated a total of 40,419
snare-days (Table 4-1), and captured 107 individual pumas (forty-eight males,
fifty-nine females) a total of 209 times. Snare captures of marked pumas (n =
132) outnumbered pumas captured for the first time (n = 78) by snares or trap.
Our average capture success with snares was one puma per 193 snare-days.
Although we tried to avoid unnecessary recaptures, we still made thirty-one
Slide @318%
PHOTO 12. Puma F27 at her birth nursery.
Table 4-1. Summary of puma snare capture efforts on the San Andres
Mountains, New Mexico, 1985–1995.
No. pumas
first captured Total pumas No. snare No. snare No. snare- Snare-days
Year and markeda recapturedb captures sites daysc per capture
1985d 3 1 4 16 426 107
1986 7 0 7 129 1,446 207
1987 6 9 15 211 2,149 143
1988 14 13 27 227 5,072 188
1989 7 15 22 215 3,987 181
1990 7 18 25 244 4,675 187
1991 5 34 39 287 7,689 197
1992 15 16 31 258 4,066 131
1993 6 16 22 282 5,793 263
1994 7 10 17 268 4,778 281
1995d 1 0 1 69 452 452
aPumas first captured with snares and one trap included thirty-six males and forty-two females.
bMarked pumas recaptured included thirty-five males recaptured seventy-seven times and thirty-four
females recaptured fifty-five times.
cSteel leg-hold traps were used in 114 trap-days. Only one female was caught. We ceased using traps
22 January 1986.
dCapture efforts were abbreviated in 1985 and 1995, extending from August to December in 1985
and from January to February in 1995.

recaptures of nontarget pumas (radio-collared pumas that did not need collar
replacement). Each male and female was captured an average of 2.4 (range =
1–6) and 1.6 times (range = 1–5), respectively (Logan et al. 1999).
We immobilized pumas captured in foothold devices with ketamine
hydrochloride (100 mg/ml, Vetalar®, Parke-Davis, Div. of Warner-Lambert
Co., Morris Plains, New Jersey 07959) dosed at about 12 mg per kg body mass.
The drug was injected remotely, usually into the caudal thigh muscles, by pole
syringe (length 3.05 m) or by 3–5 ml aluminum darts fired from a CO2-pow-
ered pistol. After induction, we quickly removed pumas from snares, covered
their heads, and tethered their legs. If pumas required further calming, we
injected xylazine hydrochloride (20 mg/ml, Rompun®, Haver-Lockhart, Bay
Vet Div. Cutter Laboratories, Inc., Shawnee, Kansas 66201) intramuscularly
with a hand syringe; the dose was about 0.5 mg per kg body mass. We protected
immobilized pumas from hyperthermia in hot ambient temperatures by placing
them in shade and by cooling them with water that we doused over their bod-
ies, especially the head, chest, abdominal, and inguinal regions. We protected
them from hypothermia in cold temperatures by wrapping them in a thermal
blanket and placing them in sunlight. We monitored heart rate, respiration rate,
and circulation (via capillary refill time) during the period of immobilization.
When we removed pumas from the Treatment Area, we used the same tech-
niques for their capture and immobilization. Pumas were removed for two rea-
sons. First, we honored a request from the cooperating agencies to remove indi-
vidual pumas that killed two or more of the state-endangered desert bighorn
sheep in a relatively short time span. Secondly, we removed pumas to experi-
mentally reduce population density. In all, we removed fourteen pumas, includ-
ing six adults of each sex and two subadult females. One adult male was removed
6 April 1989 because he killed three sheep in a three-month period (Janu-
ary–March 1989). The other thirteen pumas were removed for the experimen-
tal reduction in the Treatment Area puma population from 7 December 1990
to 21 June 1991. The pumas were captured and translocated, without injury, to
northern New Mexico. We hauled them by truck in individual wooden crates
(dimensions: 122 cm long, 24 cm high, 24 cm wide) less than forty-eight hours
after capture. They were given food and water during transport but were released
without supplemental food at predetermined release sites. All fourteen pumas
became subjects of our evaluation of the survival and behavior of translocated
pumas (Ruth et al. 1998).
Ours was the first study to routinely capture nursling cubs so we could accu-
rately quantify neonatal sex ratio and litter size and individually tag newborns
in the population. These data would furnish us information on sex ratio shifts,
cub survival rates, and recruitment rates for progeny born in the San Andres
Slide @318%
PHOTO 13. Adult female puma F47 with eartag and radio collar.
Mountains population and immigrant recruits from other puma populations.

Nursling cubs were small (less than 10 kg) and vulnerable, and, hence, required
greater care. We usually caught them by hand and sometimes with the aid of a
catchpole. In a few instances we had our small non-contentious hound “Spotty”
assist us by baying fleet cubs in trees, on the ground, or in rock piles. During the
study, we hand captured 163 cubs a total of 186 times. We restrained them
without immobilizing drugs; instead, we used a catchpole and our gloved hands.
We placed littermates in new burlap sacks to await our examination and tagging
procedures and removed them some distance from the nursery to minimize our
scent there. The sacks had open mesh for ventilation. Littermates on standby
received minimal direct human contact and were relatively calm. After we exam-
ined and tagged all the cubs, we returned them to their nursery, then vacated the
area posthaste.
We sexed and thoroughly examined all the pumas we captured. Drug-
sedated pumas were weighed with a spring scale and measured with a steel met-
ric tape. But because small cubs we handled were usually very active, accurate
morphological measurements, other than mass, were impossible. Early in the
study, we estimated ages of pumas based on dental characteristics described in
Ashman et al. (1983:23–26). We kept photographs, measurements, and written
Slide @171%
PHOTO 14. Six-week-old cub F162

with eartag, tattoo, and radio collar.
Slide @318%
PHOTO 15. New Mexico Department of Game and Fish biologist Amy Fisher with a radio-
collared mule deer.
descriptions of dental characteristics. As the study progressed, we noted dental

characteristics of the known-age pumas in the population, beginning with cubs
for which we knew practically the dates of their birth. In addition, we also noted
other changes in pelage and eye color. We used those characteristics to age
pumas with previously unknown histories, and, when necessary, to adjust esti-
mated ages of pumas that we captured earlier in the study.
All captured pumas were marked for life by tattooing an identification num-
ber on the inner surface of one pinna. In the other pinna, we inserted a 25
mm–diameter, colored, numbered plastic tag (Duflex TagsTM, Fearing Manu-
facturing Co., South St. Paul, Minnesota 55075). The eartags gave us a marker
that we could see from a distance and were particularly important for identify-
ing tagged progeny that were not radio-collared. Each adult and subadult puma
was fitted with a collar containing a 150 or 151 MHz transmitter with a mor-
tality mode set to turn on after six to thirteen hours of immobility (configura-
tion MOD-500 transmitter with X-1 shock crystal and S6A mortality sensor,
CLM collar, transmitter cast in polymer. Telonics Inc., Mesa, Arizona 85204).
Radio collars weighed at most about 450 grams. Some of the cubs that were
more than thirty-one days old also were radio-collared (configuration MOD-
225 transmitter with S6A mortality sensor, also by Telonics Inc.). We built their
collars out of expandable material that would break away within twelve months.
Cub collars weighed about 113 grams. The tattoos, eartags, and radio collars
made it possible for us to accurately estimate the number of pumas in the pop-
ulation and to determine relationships between them, such as relatedness and
breeding pairs.
There were occasions when we could not tag all progeny in a litter; conse-
quently, we later had questions regarding the relatedness of untagged independ-
ent individuals that we captured. Were these new immigrants? Or were they
unmarked progeny? To answer the questions, we collected tissues from most of
the pumas whenever we had them in hand. Usually, the plugs of pinna skin and
cartilage that we displaced with the eartags were collected because these were the
easiest to preserve in the desert field conditions. We also collected muscle tissues
from deceased individuals. We sent tissue samples to the Laboratory of Genetic
Diversity at the National Cancer Institute in Frederick, Maryland, where Dr.
Melanie Culver analyzed alleles at highly polymorphic loci (i.e., microsatellites)
and compared them with those of candidate mothers, siblings, and fathers to
estimate the likelihood of relatedness.
Our snaring technique proved to be relatively safe for capturing pumas
weighing 10 kg or more (Logan et al. 1999); life-threatening injuries occurred
in only 2.4 percent of total puma snare captures. Four pumas died from capture
injuries. One adult female suffered a fatal spinal injury; the other three cats (two
adult females and one adult male) suffered severe fractures to the ulna and radius
of the leg caught in the snare. A fourteen-month-old female cub also suffered a
broken ulna and radius. Fortunately, puma F147, otherwise known as “Lefty”
(named for her undamaged foreleg), was treated, rehabilitated, and released onto
her natal area where she established a home range and raised cubs. Deaths of
nine cubs in three litters also resulted from our research activities. One of the
adult females that we euthanized because of capture injuries was apparently rais-
ing four newborn cubs. We determined this from her placental scars and her
milk-producing mammary glands. In another incident, a first-time mother
apparently abandoned all four of her forty-two-day-old cubs after we marked
two of them. In the final case, we discovered too late that a six-month-old
female cub had slipped her expandable radio collar over her front leg; the collar
rubbed her skin, causing an open wound. Although we captured the cub and
placed her in veterinary hospital, she succumbed to septicemia. This was the
only incidence of a radio-collar injury to pumas we studied.
During our research we captured, tagged, and released 241 individual
pumas; live pumas were handled a total of 396 times. Seventy-eight pumas
(thirty-six males, forty-two females) were initially captured with foothold snares.
Another 163 pumas (eighty-one males, eighty-two females) were cubs when we
initially captured them by hand. We radio-collared 126 pumas (forty-nine
males, seventy-seven females). Of pumas radio-collared for the first time, sixty-
two were adults, twenty-six were subadults, and thirty-eight were cubs. We also
accounted for another fifty-three to fifty-four pumas in the population that we
could not tag. Forty-six were cubs; of these, we knew thirteen died while still
dependent on their mothers and nine were still dependent on mothers at the
close of the study. Of the remaining eight unmarked pumas, four were found
dead at ages ranging from twelve to thirty months, two were detected as new
immigrants at the close of the study, and one to two females (one of which may
have been tagged and wearing a nonfunctional radio collar) successfully pro-
duced cubs (we visually observed one litter and found tracks of another toward
the end of the study). In total, we observed 294–295 pumas on the San Andres
Mountains; this constituted the largest number of pumas ever studied in a sin-
gle population.
Monitoring Desert Mule Deer and Desert Bighorn Sheep

To study effects of puma predation on the deer and sheep populations we had
to accomplish two basic tasks: (1) monitor the dynamics of the deer and sheep
populations, and (2) determine the extent of puma predation on those animals.
Because our team’s efforts were mainly focused on puma population dynamics
and predation, we relied largely on biologists of the New Mexico Department of

Game and Fish and the U.S. Fish and Wildlife Service to furnish us information
via annual surveys on the ungulate populations.
Mule deer population composition counts gave us an estimate of the sexes
and broad age classes available as puma prey, as well as an index to fawn pro-
duction. Population composition was estimated by helicopter surveys carried
out by Game and Fish biologists during six years and by ground-based surveys
by our team during four years. Aerial and ground survey sampling techniques
met or exceeded recommendations established by Humphreys and Elenowitz
(1988:Appendix A) to stabilize herd composition ratios and to achieve similar
(i.e., no difference in) precision in ground and aerial surveys. A minimum of
twenty-five groups and one hundred does were counted in all surveys, regardless
of the method. Counts were made in winter in nine years and fall in one year.
Deer were classified as bucks and does older than one year and fawns of the
year (born July–October). When possible, we distinguished between yearling
(i.e., one year old) and adult (two years or older) deer. Helicopter-based obser-
vations were made by two experienced Game and Fish personnel in a Bell 206
Ranger or Hughes 500D. Counts were made during low-level flights between 7
a.m. and 10:45 a.m. in a one-day period. Ground-based observations were made
with 7–10-power binoculars and 15–45-power spotting scopes. Ground surveys
were conducted during successive days until minimal requirements were
exceeded. Recounts of deer were eliminated by not counting deer whenever
there was a possibility of re-observation.
We estimated deer survival rates and assayed causes of mortality for a sample
of 175 radio-collared deer (ninety-one bucks, eighty-four does) that were one
year or older and lived in the Treatment Area during 1987–1995. We tried to
maintain a pool of at least thirty radio-collared deer of each sex in the Treatment
Area per year. Deer were captured throughout the Treatment Area during nine
capture operations from October 1986 to October 1993. In two operations,
deer were caught by using the drive net technique (Beasom et al. 1980). In the
remaining seven operations, deer were caught in a net fired from a helicopter
(Bell 206 Ranger or Hughes 500D). Even though radio-collared deer were dis-
persed throughout the Treatment Area, captured individuals and their capture
locations were not intended to be random representations. Instead, capture
operations were more practical—meant to capture as many deer as were needed
to meet sample-size objectives with the most efficient use of personnel and
equipment (i.e., money). Hence, capture efforts normally concentrated on deer
that were the most conspicuous to observers in the helicopter. Deer were some-
times physically restrained. But if they required immobilization, they were
injected intravenously with xylazine hydrochloride (20 mg/ml) dosed at about 1
mg per kg estimated body mass. The drug was reversed with an intravenous
injection of yohimbine hydrochloride (AntagonilTM, 5 mg/ml, Wildlife Labora-
tories, Fort Collins, Colorado 80524) dosed at 0.2 mg per kg body mass. Deer
were examined to record sex and physical condition; a rough estimate of age was
derived from dental characteristics (Robinette et al. 1957). Deer one year or
older were each fitted with a collar containing a 148 or 149 MHz transmitter
with a mortality mode set to turn on after two hours of constant immobility
(configuration MOD-500 transmitter with S6A mortality sensor by Telonics
Inc.). We included in our research only radio-collared deer that survived more
than fourteen days after capture to minimize the effect that capture trauma
might have on deer survival.
We relied on a deterministic, discrete time model to simulate the dynamics
of the deer population (see Chapter 17 and Appendix 4). Parameters of the
model used the estimates of deer population composition and survival along
with estimates of other characteristics gathered on the San Andres Mountains
deer population and existing information from other desert mule deer popula-
tions in southern New Mexico.
Desert bighorn sheep population characteristics were monitored annually by
ground- and helicopter-based surveys conducted by U.S. Fish and Wildlife and
New Mexico Game and Fish biologists. Observed sheep were classified as adult
rams and ewes, yearling rams and ewes, and lambs (distinguished by sex when
possible). However, due to funding and personnel constraints, the surveys were
not conducted in a standardized fashion. Biologists made ground-based surveys
in nine years using binoculars and spotting scopes to search habitats used by
radio-collared sheep. During 1986 through 1989 and in 1994, ground-based
counts were made by a biologist who searched for the sheep throughout each
year. In 1990, 1991, and 1992, the ground-based counts were done during the
month of December, 31 August to 10 September, and from May through
December, respectively. A helicopter survey was conducted in 1993 by two
observers in a Hughes 500D. Biologists in all surveys primarily homed on radio-
collared sheep and counted and classified all sheep they observed.
We monitored a total of forty-three radio-collared sheep (sixteen rams,
twenty-seven ewes) from August 1985 through March 1995. The sheep were
captured by Game and Fish biologists using a net-gun fired from a helicopter
(Hughes 500D or Bell 206 Ranger) in ten capture operations conducted from
October 1980 to October 1993. All sheep were physically restrained, then
examined to record sex, age, and general physical condition. Ages of lambs and
yearlings were determined from their estimated birth dates. As those individuals
advanced in years, their ages were known. Otherwise, ages of adults were esti-
mated by counting horn growth annuli (Geist 1966, Hemming 1969). Radio
collars placed on sheep were similar to those on the collared deer. Radio-collared
sheep were used to find other sheep in the population; therefore, early in the
study, Fish and Wildlife and Game and Fish biologists attempted to maintain a
relatively high percentage of radio-collared individuals in the population (more
than 50 percent of the observed) by capturing sheep annually from 1980
through 1985, and again in 1988. However, from November 1989 to October
1993 the percentage of radio-collared sheep waned to less than 50 percent
because of a temporary hands-off policy adopted by the cooperating agencies
(i.e., U.S. Fish and Wildlife Service, New Mexico Department of Game and
Fish, White Sands Missile Range). As with the radio-collared deer, we only used
radio-collared sheep that survived more than fourteen days after capture for
analysis of annual survival rates and agent-specific mortality rates.
Radiotelemetry
Radiotelemetry was the channel that linked our observations of sign on the
ground to real-time observations of pumas. Without it, our task of quantifying
puma population dynamics would have been far more difficult. It enabled us to
identify tagged individuals that made sign we found and to deduce if untagged
pumas might be present. We were also able to forecast births of litters from pre-
viously recorded associations of potentially breeding pairs. Radiotelemetry made
possible our detailed study of puma social organization. In addition, the tech-
nology facilitated our study of interactions between pumas and prey by helping
us to find puma kills and enabling us to quantify survival rates and puma pre-
dation rates on deer and sheep.
We located radio-collared pumas 13,947 times by fixing their positions from
the ground (31 percent of locations) and from airplanes (69 percent of loca-
tions). Of the total, 8,490 locations were of adult pumas that lived on the San
Andres Mountains study area (Fig. 4-1). The remainder were of subadults and
cubs on the study area and of adults and subadults that ranged partially or exclu-
sively on the Organ and Oscura Mountains. Ground locations were acquired
opportunistically by using portable radio receivers attached to handheld direc-
tional antennas (TR-2 receiver with TS-1 scanner/programmer, RA-2A antenna
by Telonics Inc.) to take line-of-sight bearings on the strongest aural signal from
the transmitter. Three or more bearings taken from different locations were
mapped on 7.5-minute-series topographic maps. Plotted bearings produced a
triangle or polygon. The approximate geometric center of the triangle or poly-
gon was plotted to represent the location of the radio-collared puma and was
recorded as Universal Transverse Mercator (UTM) grid coordinates to the near-
est 0.01 km. Error around each location was estimated by using the distance (m)
FIGURE 4-1. Locations of resident adult pumas on the San Andres Mountains, New Mexico,
1985–1995. We obtained 8,490 locations between the north and south study area bound-
aries; of those, 99 percent were also within the east and west boundaries. Another 304 loca-
tions were obtained in the Organ and Oscura Mountains.
from the center of the triangle or polygon to the most distant vertex. This dis-
tance was used as the radius of a circle that contained the location of the puma
(Saltz and Alkon 1985).
We flew in light fixed-wing aircraft (Cessna 182, 172 XP, or T337) 370 times
and logged 1,852 hours to locate radio-collared pumas. Normally we flew once
each weekend, but sometimes flights were canceled because of poor weather or
White Sands Missile Range missions. In each flight, we attempted to locate all
of the radio-collared pumas on the study area, as well as dispersing individuals;
their numbers varied from ten to forty pumas. Aircraft were rigged with direc-
tional antenna mounted on each wing strut, which were connected via coaxial
cables to an antenna switch box, a receiver with scanner, and the intercom sys-
tem. Aerial locations of radio-collared pumas were fixed at sites where we made
at least three passes over peak aural signals. The puma’s location was recorded as
a UTM coordinate to the nearest 0.01 km. Variations in terrain at puma loca-
tions and differences in our flight altitude as influenced by terrain and air tur-
bulence created error in our fixes, which we estimated by flying concentric cir-
cles around the locations to mark where the signal waned. We expressed this
error as a radius (m) of a circle around the fixed location.
Accuracy of radiotelemetry locations was tested during the first three years at
173 of 2,170 locations by finding test collars, puma sign at the locations,
dropped collars, dead pumas, and by subsequently walking in on pumas to see
them. In all instances the radio collars were within the error radii of the loca-
tions (Sweanor 1990). Thereafter, we confirmed radiotelemetry locations on an
opportunistic basis by using the same techniques. All of our recorded
radiotelemetry locations had estimated error radii of 500 m or less, and 91.5
percent of the locations had error radii of 300 m or less. At all actual puma loca-
tions and radiotelemetry locations with error that included single or co-domi-
nant vegetation cover types, we recorded the vegetation and terrain features. Ele-
vation was recorded as indicated on the 7.5-minute-series topographic maps
(converted to meters); we also noted the aspect and percent slope (measured
with a USGS Topo Map Land Area and Slope Indicator, Reproduction Special-
ties, Inc., Denver, Colorado 80222).
By walking in on free-ranging radio-collared pumas (i.e., they we re not
caught in snares), we we re able to visually observe 579 individuals in 287 obser-
vations, the most that wild pumas have been viewed during a study. The desert
mountains, with their re l a t i vely low, sparse vegetative cove r, made it easier for
us to safely approach and visually observe pumas. Normally, we attempted
observations of mothers to locate their nurseries and to get initial counts of
n ewborns. Subsequently, we approached families to count cubs for surv i va l
estimates. In addition, we would sneak in on pumas to see if they made a kill,
to check on their physical state after being captured, or to see if consort pairs
we re actually together. We also checked the well-being of some newly inde-
pendent subadults.
We usually approached pumas from upslope and when possible from a
promontory, such as a cliff, rock outcrop, or boulder pile. Our aim was to gain
a visual advantage from a safe distance to observe the cats without causing them
to move from their chosen locations. We hoped our upslope position also would
enable us to appear ominous to pumas that detected us and leave them an easy
escape route downslope if they chose to flee. We usually approached in groups
of two or more people, but sometimes we were alone. We looked for the slight-
est movements and kept our hearing trained on two dimensions—listening for
the minutest variation in the signal from the radio receiver and listening for any
sound from the pumas.
Our closest approaches unintentionally brought us to within 5–10 m of
pumas. Sometimes, because boulders or rock outcrops shielded transmitter sig-
nals at close range, we accidentally got too close before we saw the cats. Some-
times pumas would slink silently away, using any terrain, vegetation, or large
rocks for cover. Other times they simply froze except for moving their eyes, ears,
or heads to track our every move. Some, in ultra-slow-motion, shifted their bod-
ies into position to watch us. Others slowly positioned their bodies and gathered
their feet beneath them, then bolted away. Steadfast ones seemed secure in cover,
but watched us intently; a few individuals seemed totally aloof and fell asleep.
Only during six close encounters were we threatened by pumas. In every
case, females were probably trying to protect their three- to twelve-week-old
cubs. Three mothers charged us but aborted their rushes as we stood our
ground, yelled at them, and, twice, defended ourselves with a wooden stick or a
metal catchpole. One mother stalked Frank head on, approached to within 5 m,
and seemed unfazed even when Frank discharged his .22 magnum pistol twice
in the air. The puma turned away only after Frank stepped onto a ledge that ele-
vated him about 0.5 m higher than the level of the puma. Two females growled
at us as they lay prone at their nursery, feet set beneath them, head and ears
perked toward us, eyes locked onto ours. In the latter two instances, we sensed
attacks were probably imminent, so we retreated by slowly backing away. Usu-
ally we observed families that were unaware of our presence or were apparently
indifferent to it. This allowed us unique opportunities to watch cubs interact
with their mothers and siblings.
We employed similar radiotelemetry techniques on radio-collared deer and
sheep. This enabled us to find puma kills and to directly estimate annual sur-
vival rates and rates of death attributed to puma predation or other causes. We
monitored the status (alive or dead) of radio-collared deer and sheep from the
ground on a weekly basis. In addition, we flew in airplanes monthly to check the
status of all the radio-collared animals and to specifically search for any animals
we may have missed from the ground. When transmitters indicated death by
emitting a twofold increase in beat frequency, we investigated. If animals were
dead (sometimes the transmitters temporarily failed), we performed necropsies
to determine probable proximate causes of death.
Investigating the Dead

To determine the most influential mortality factors to the puma, deer, and sheep
populations in our study, we tried to ascertain why animals died. We also
recorded other pertinent information, including sex, age, and, when possible,
the physical state of the animal prior to death. Extremely high ambient temper-
atures, especially during May through September, caused such rapid decompo-
sition of carcasses that trying to ascertain condition prior to death was practi-
cally impossible unless we found the animals within hours of death. For pumas
that were dead less than twenty-four hours and died for reasons that were
obscure to us, we collected organ tissues and sent them to the New Mexico
Department of Agriculture Veterinary Diagnostic Services in Albuquerque to
assay cause of death.
Specific criteria were used to categorize dead prey animals as puma prey,
probable puma prey, and animals that died of other causes. In our analysis of the
effects of puma predation on deer and sheep, we combined the categories of
puma prey and probable puma prey. Animals that died of other causes were not
associated with any evidence that implicated pumas as the proximate cause of
death. By using these categories, we could assess the strength of puma predation
as a limiting factor of deer and sheep relative to other nonpredator causes of
mortality.
We classified animals that were undoubtedly killed by a puma as puma prey.
Evidence for puma attack and consumption included puma canine punctures to
the back of the neck, the throat, or the head; puma feeding patterns (i.e., pluck-
ing or cropping of hair with the incisors, opening of the carcass first at the
abdomen or thorax and consumption of vital organs, expulsion of the stomach,
consumption of bones, breakage of large bones); puma tracks in an attack
sequence or at the cache; puma feces, scrapes, or prey carcass drag line; and cov-
erage of the prey with ground debris (e.g., soil, leaves, sticks).
Some animals we found dead were in such advanced states of decomposition
or consumption that we could not find puma canine punctures. But because
other evidence pointed to puma predation, we called them probable puma prey.
Evidence included puma feeding patterns; puma tracks, feces, or scrapes; prey
drag lines; and coverage of the prey with ground debris.
We also collected puma feces and contents of puma stomachs opportunisti-
cally, noted the exact circumstances of their deposition (e.g., random find, asso-
ciation with a specific kill or puma, location and habitat features), and air dried
them. They were analyzed by Mike Elmer at the University of Idaho. He pro-
vided us a detailed analysis of the diet of pumas on the San Andres Mountains
(Elmer 1997).
CHAPTER 4. STUDYING WLD PUMAS 59
In the course of our field research, we kept detailed journals of our day-to-
day activities as well as data forms on each of our observations. We had specific
data forms for captured animals, puma locations, and status of radio-collared
deer and sheep, and for pumas and prey animals that died. We transferred data
on forms to electronic databases that we used to facilitate our analyses by com-
puter.
Overview of Analytical and Statistical Methods

Exact analytical and statistical techniques that we used are included in each of
the following chapters addressing the findings of our field research. We used
parametric statistical methods (Zar 1984) where samples were large (i.e., sample
sizes were twenty-five or more, or approached the absolute population) and data
appeared to be normally distributed when checked by eye against a normal dis-
tribution plot generated by SYSTAT 7.0 (SPSS Inc., Chicago, Illinois). When
we could not meet assumptions of normality or equal variances for parametric
tests, or if sample sizes were small, we used non-parametric tests (Daniel 1978).
Most statistical tests were performed using SYSTAT 7.0 or SAS 6.11 (SAS Insti-
tute Inc., Cary, North Carolina). Tests for differences between means or distri-
butions are typically 2-tailed unless specified. Experimental errors were con-
trolled at the 0.10 level of significance because we usually had small sample or
population sizes, which are characteristic of large-carnivore studies. The higher
alpha level reduces the probability of a Type II error (i.e., failing to reject a null
hypothesis that is false) and increases the probability of detecting a change. We
note statistical tests in the text by bracketed numbers (e.g., [1]) and report the
corresponding results in a special section at the end of each chapter. We typically
report mean statistics with ± 1 standard deviation (SD) to express variability of
the measurement. Sample size (n) is reported for readers wishing to calculate the
standard error. In addition, we provide a summary at the end of chapters in Parts
2–5.
P a rt I I
Puma Life History

Strategies and
Population Dynamics
Chapter 5
A Puma Population in the Desert
Reliable information on puma life-history traits and population dynamics has

been described for pumas in more-northern temperate and contiguous habitats
in North America (Hornocker 1970, Seidensticker et al. 1973, Lindzey et al.
1989, Lindzey et al. 1994, Ross and Jalkotzy 1992). But before our study, very
little was known of puma populations in the Southwest deserts of North Amer-
ica. Naturally, much of our information on puma population dynamics in the
desert is descriptive. In addition, we contrast puma population dynamics and
life-history traits in a Treatment Area, where we experimentally reduced the
number of pumas, with those on a Reference Area. The basin and range phys-
iography of much of the Southwest naturally fragments puma habitat and may
contribute to dynamics that are different than those observed in more contigu-
ous puma habitats. We discuss similarities and differences of pumas on our
desert study area with pumas in other populations in North American habitats,
and we try to explain them in the context of evolutionary ecology.
Research Hypotheses and Predictions

Meager quantitative information on life-history traits and population dynamics
of pumas and other solitary-living large felids, such as tigers, leopards, and
jaguars, constrained our hypotheses to a few published postulates and to natu-
ral phenomena that emerged during our study. We address these specific
research hypotheses in the appropriate sections in this chapter.
1. Puma mothers tend to have more litters during the period of maximum
63
64 PART II. PUMA LIFE HISTORY STRATEGIES AND POPULATION DYNAMICS
food abundance. A birth peak should coincide with the fawning period for
mule deer, the primary prey of pumas.
2. Pumas kill other pumas to benefit individual survival and reproductive suc-
cess. Thus, male pumas, who do not take part in raising young, should kill
other pumas for food, territory, and mates. But puma mothers, who solely
raise young and cannot afford debilitating injury from aggressive encoun-
ters, should avoid killing other pumas.
3. Intraspecies strife is density dependent. Frequency of intraspecies killing of
independent pumas should increase with density.
We examined three hypotheses (hypotheses 4–6 below) on the evolution
of the timing of progeny independence from mothers:
4. Timing of independence reduces competition for the mother’s time and
energy budgets required for mating, pregnancy, and rearing more progeny.
Thus, cubs should become independent when they are physically able to
survive on their own and about the time that their mother is ready to breed
again (i.e., when she comes into estrus).
5. Timing of independence reduces physical harm to cubs from aggressive
males seeking to mate with the cub’s mother. Cubs should become inde-
pendent before their mother comes into estrus and attracts dangerous
males.
6. Timing of independence reduces inbreeding between male offspring and
their mothers. Male progeny should reach puberty about the time they
became independent from their mothers.
7. Dispersal of puma progeny is independent of adult density (Seidensticker et
al. 1973). Thus, all progeny should disperse from their natal areas.
In addition, we investigated puma dispersal patterns so that we could test

three hypotheses pertaining to how dispersal affects fitness, discussed in the
Chapter 13 section “Why Do Pumas Disperse?”
Terms for Pumas

Terms we used for pumas in our study population depended upon their sexual
maturity, maternal dependence, site attachments, and place of origin. Adults
were pumas that bred or reached the average age of known breeders in the pop-
ulation. Subadults were independent of their mothers’ care but were apparently
not yet capable of successful breeding. Cubs were offspring that were dependent
on mothers for food and protection. Residents showed site attachment (i.e., con-
tinuous use of an area over time). The area where a puma restricted the major-
ity of its movements was considered its home range. Cubs that became subadults
and left the boundaries of their natal home range were considered dispersers.
CHAPTER 5. A PUMA POPULATION IN THE DESERT 65
Cubs that established home ranges post-independence that overlapped more

than 5 percent of their natal home ranges (based on the 90 percent Minimum
Convex Polygon) were considered philopatric (see Chapter 13). Immigrants were
pumas that dispersed from some other locality in New Mexico and moved onto
the San Andres Mountains; they subsequently established residency. Recruits
were either progeny born in the San Andres Mountains puma population or
immigrants that entered the resident adult portion of the population. An emi -
grant was an individual that dispersed completely outside of the San Andres
Mountains study area.
Counting Pumas
Through our intensive field efforts, we strove to make accurate estimates of the
puma populations in both the Treatment Area and the Reference Area for each
year. In essence, we tried to census the population. This enabled us to quantify
puma population dynamics and vital rates and to relate puma population char-
acteristics to traits of puma social organization and deer and sheep population
characteristics. But quantifying puma numbers is a formidable task. Their low
densities distributed over complex habitats requires extensive time in the field to
quantify numbers by capturing and marking individuals. Furthermore, puma
births, deaths, emigration, and immigration can occur year round. Conse-
quently, it was impossible to meet most of the assumptions for capture-mark-
recapture methods to estimate abundance.
Consider the simplest method of all, the Peterson Method, which assumes
(1) the population is closed so that the population number is constant during a
sampling period, (2) all animals have the same chance of getting caught in the
first sample, (3) marking individuals does not affect their probability of recap-
ture, (4) animals do not lose marks between the two sampling periods, and (5)
all marks are reported upon discovery in the second sample (Krebs 1999). First,
we could not trap for pumas in the Treatment and Reference Areas in a short-
enough sampling period to exclude population change; thus, we would violate
assumptions 1 and 2. Moreover, differences in survival, behavior, and area of use
of territorial males and maternal and non-maternal females would likely affect
probability of capture, again violating assumption 2. Second, we learned that
pumas became trap-shy after their first capture (Logan et al. 1999), thus violat-
ing assumption 3. We could only meet assumptions 4 and 5. For many of these
same reasons, we also could not meet assumptions for somewhat more complex
techniques for open populations (see Krebs 1999:49). Thus, we had to rely upon
more direct methods of quantifying the population and for gauging the accu-
racy of our estimates.
We developed a puma population chart that tracked the lives of all individ-
uals observed. We examined data on captured pumas, radiotelemetry, puma

tracks, and visual observations of untagged pumas. Initially, all adult and
subadult pumas captured for the first time were entered on the chart beginning
with their date of capture. Cubs were entered on the chart at their estimated
birth date. We extended the lifelines on each individual to their known or esti-
mated dates of death, emigration from the population, or disappearance. We
also refined the lifelines on adult pumas on the chart as we learned more about
when they might have entered the adult population.
We estimated absolute densities of adult pumas (i.e., breeders) based on
these assumptions and empirical ontogenetic data that we accumulated during
the research. First, we assumed that all independent pumas captured for the first
time and less than thirty-six months old had just entered the population. How-
ever, pumas captured for the first time that were thirty-six months or older were
assumed to have entered the population as adults at an earlier date. Thus, we
adjusted the estimates of the adult puma population to include those individu-
als that we had probably missed during previous years. We defined entry dates
for the latter group by (1) ages of known-age and approximately known-age
males and known-age females when they reached adulthood, and (2) the mean
ages of independent male and female pumas less than thirty-six months old cap-
tured for the first time and considered to be new recruits to the population. Four
female pumas that were not captured were charted into the population because
we were able to monitor their movements through a combination of tracks (all
four females), associations with radio-collared offspring (two females), associa-
tions with radio-collared males (one female), and visual observations of a female
and her cubs (one female). Two other adult female pumas, twenty-four and
thirty months old, were never radio-collared because they were dead when we
observed them. It is possible that the older female was marked, but because she
was severely decomposed, we did not find a tattoo. A woodrat, fox, or raven
could have easily hauled away her eartag. We charted these two females into the
population at their estimated date of death.
For female pumas thirty-six months or older, we used twenty-one months of
age to define when they entered adulthood in the population. Our criteria were
based on (1) the mean age of 21.4 ± 3.1 months for seven known-age females
when they first associated with adult males (see “Puberty and First Litters” in
Chapter 7), and (2) the mean age of 21.4 ± 2.6 months for eighteen approxi-
mately known-age female immigrants that were less than thirty-six months old
when first captured.
For male pumas thirty-six months or older, we used twenty-four months of
age to define when they entered adulthood in the population. Our criteria were
based on (1) the mean age of 24.3 ± 2.5 months for one known-age and four
CHAPTER 5. A PUMA POPULATION IN THE DESERT 67
approximately known-age male pumas when they first associated with adult
females, and (2) the mean age of 23.9 ± 6.3 months for seven known-age males
when they arrived in the area where they established their first territory.
In addition, we developed a percent accuracy (PA) from the data on the chart
to estimate the accuracy of our annual (January–December) counts of adult
pumas in the population. Our equation was: PA = [1 – BLyear i / (BLyear i + Cyear
i)] x 100 where BLyear i = the number of adult pumas backlogged into year i, and
C year i = the number of adult pumas actually counted (i.e., radio-collared or
observed) during year i. The proportion BLyear i / (BLyear i + Cyear i) x 100 esti-
mated the percentage of adult pumas we missed in annual counts. The PA
helped us to gauge how well we were meeting our operational objective of
counting all of the adult pumas on the study area.
The number of adult pumas thirty-six months or older when first captured,
and which we backlogged into the population to probable entry dates (i.e., dates
of sexual maturity), ranged from zero to three per year in the Treatment Area
and from zero to eight per year in the Reference Area (Table 5-1). Understand-
ably, the highest numbers of backlogged adults came during the first full years
of the study (1986, 1987) when we were developing our capture and search
techniques on the study area. By January 1988 we were thoroughly canvassing
the Treatment Area, but that stage did not come for the Reference Area until the
latter half of 1988. During the span 1988–1994, the numbers of backlogged
Table 5-1. Percent accuracy (PA) for the estimated number of adult pumas per
year, 1986–1994, in the Treatment Area (TA) and Reference Area (RA), San
Andres Mountains, New Mexico.
Area 1986 1987 1988 1989 1990 1991 1992 1993 1994
TA M:Fa 0:3 1:2 0:1 0:0 0:0 0:0 0:1 0:1 0:0
PAb 70 77 93 100 100 100 93 94 100
RA M:F 5:3 2:2 0:0 1:2 1:2 2:2 1:1 1:1 0:0
PA 33 66 100 87 88 86 93 94 100
aThe number of males (M) and females (F) signifies the number of adults in each sex that we appar-
ently missed in our capture efforts each year. We assumed that all pumas that were thirty-six months
or older when first captured had entered the adult population at an earlier date. Those pumas were
backlogged into the population chart to the month that females were twenty-one months old and
males were twenty-four months old (see text for explanation and empirical criteria).
bPercent accuracy = 1 – number of pumas backlogged / number of adult pumas counted plus adult
pumas backlogged per year ¥ 100. PA estimates the minimum proportion of the adult population
counted each year.
adults dwindled to 0–1 in the Treatment Area and 0–4 in the Reference Area.
We missed four adults in 1991, probably because we spent all of our efforts
removing pumas from the Treatment Area until June 1991. The average annual
PAs during the 1988–1994 span for the Treatment and Reference Areas were 97
percent and 93 percent, respectively. The population estimates that we report for
the Treatment Area begin in 1988, but our estimates for the Reference Area
begin in 1989. All our population tables and figures will show the adjusted esti-
mates for the adult puma population for each area. Consequently, our annual
estimates of adult pumas probably approached 100 percent of the adult pumas
actually present.
By charting the population, we were able to make three puma density esti-
mates: (1) We estimated adult and total puma density in January each year
(1988–1995 in Treatment Area, 1989–1995 in Reference Area); this was our
point estimate that coincided with New Mexico’s puma hunting season (Decem-
ber–March). (2) We estimated adult puma density for each annual period (Jan-
uary–December, 1988–1994 in Treatment Area, 1989–1994 in Reference Area);
this represented the number of pumas present throughout an entire year. (3) We
estimated annual adult puma density during August–July from 1987 to 1994 in
the Treatment Area and from 1988 to 1994 in the Reference Area to coincide
with the biological years for pumas and mule deer, which we began in August to
coincide with birth pulses for the two species.
Radio-collared pumas that lived along the boundary of the Treatment and
Reference Areas or that moved out of the San Andres Mountains at the north
and south ends of the study area were included in the Treatment Area or Refer-
ence Area based on the proportion of their aerial locations recorded in each area
during each twelve-month period. For example, a puma located fifteen times in
the Treatment Area and twenty-eight times in the Reference Area during one
year would be assigned values of 0.39 and 0.61 for each area, respectively. Cubs
were included in areas in identical proportions to their mothers. For the annual
adult density estimates, we also determined the proportion of time per year that
pumas were present on the areas. For example, a puma that apportioned its
annual activities by 75 percent in the Treatment Area and 25 percent in the
Organ Mountains (south of the study area) but was alive for 60 percent of the
year was assigned a value of 0.45 in the Treatment Area for that year. To get the
total annual adult density, we summed the proportions for all the adults in the
respective areas. We quantified these densities for each area per year and for each
area per year per 100 km2. All these methods of quantifying pumas were essen-
tial to describing puma population dynamics and how puma numbers related to
puma behavior and social organization and to prey population dynamics.
Chapter 6
Puma Population Structure
Sex Structure
The most distinguishing attribute of an individual within a population is its sex.
As we will explain later, a puma’s sex defines its behavior and how it interacts
with other pumas in the population. Examining the sex structure of the popu-
lation from birth to older life-stages helped us to understand consequences of
other population characteristics, including reproduction, survival, population
growth, and social organization.
Cubs
The sex ratio of puma offspring at birth represents the secondary sex ratio (pri-
mary sex ratio is at zygote formation, tertiary sex ratio is at puberty). Overall,
male and female cubs were born into the population in equal proportions, a
characteristic found in most sexually reproducing vertebrates (Ricklefs 1990).
We observed 210 cubs in seventy-six litters. Three other litters with about ten
cubs died shortly after birth, before we were able to examine the cubs. We han-
dled 148 nursing cubs in fifty different litters in which we examined all the cubs
when they were nine to forty-nine days old (x– = 31.6 ± 8.6); there were seventy-
five males and seventy-three females, the equivalent of a hypothetical 1:1 pri-
mary sex ratio. In another fifteen weaned litters with cubs 52 to 427 days old (x–
= 198.9 ± 121.6) there were fourteen males and twenty females, but the
male:female ratio did not differ from 1:1 [1]. In weaned litters, we suspected
69
that some postnatal mortality had occurred. Theoretically, equal sex ratios of off-
spring suggests there is a tendency for equal parental investment in each sex
(Trivers 1985).
The sex ratio of puma offspring on the San Andres Mountains was similar to
the sex ratio of 1.1 males to 1.0 females that Lopez-Gonzalez (1999) reported
for twenty puma litters in southern Idaho. It was also similar to more limited
information on cubs from other captive and wild populations. Robinette et al.
(1961) reported twelve male and twelve female cubs born at the National Zoo-
logical Park. Likewise, in the central Idaho wilderness, Seidensticker et al.
(1973) found twelve male and twelve female cubs that were various ages when
they were first observed. Information from other wild populations were also of
cubs representing a wide range of ages where postnatal mortality may have
affected observed sex ratios. Even though there were more males than females in
two studies (Ashman et al. 1983, Spreadbury 1989) and fewer males than
females in two other studies (Logan et al. 1986, Ross and Jalkotzy 1992), none
of the ratios was significantly different from 1:1.
A closer look at our data indicated sex ratio was female biased in first litters,
but more males were born in subsequent litters. In fifteen first litters of known-
age (n = 8) and approximately known-age (n = 7) mothers in which we sexed all
the cubs, there were seventeen males and twenty-eight females. In contrast, there
were forty-five males and thirty-five females in twenty-seven subsequent litters
produced by nineteen mothers. In the latter group, we did not have a complete
record of the reproductive history of the females; some of them might have pro-
duced litters prior to our research. Therefore, we discounted each female’s first
observed litter, in case it was actually her first, and used only her subsequent lit-
ters; these probably represented second to fourth litters. All cubs in both groups
were nurslings ranging in age from eighteen to forty-nine days old. Sex ratios in
first litters differed from sex ratios in subsequent litters [2]. We looked even
more carefully at the sample of mothers for which we had complete reproduc-
tive histories to compare sex ratios of their first and subsequent litters. In eight
first litters, there were eight male and nineteen female cubs; but in twelve sub-
sequent litters (i.e., second to fourth litters) there were twenty male and eight-
een female cubs. Not only was the sex ratio in the first litters different than that
in the subsequent litters [3], but also it was different from 1:1 [4]. The number
of cubs in first litters (x– = 3.4 ± 0.7) and subsequent litters (x– = 3.2 ± 0.6) were
similar.
The secondary sex ratio may be influenced by the physical state of female
pumas on the San Andres Mountains. We examined masses of nine females (for
which we had data) when they were eighteen to twenty-four months old (x– =
20.8 ± 2.2), near the approximate age of puberty (see Chapter 7, “Puberty and
CHAPTER 6. PUMA POPULATION STRUCTURE 71
First Litters”) and when they were twenty-eight to sixty months old (x– = 42.2 ±
10.7), and found that they gained mass (mean difference = 3.0 ± 2.5 kg) [5].
This suggests that puma mothers were still growing when they produced their
first litters. Hence, there may be a physiological trade-off involving allocation of
energy and nutrients for continued somatic growth of the mother, and investing
in less-costly female offspring while optimizing litter size (Sibly and Calow
1986, Stearns 1992). Not only are first-time mothers probably growing, but also
they likely are inexperienced in raising offspring, and relative to older estab-
lished females are not as familiar with resources (i.e., food, nurseries) in their
home ranges that are vital to the provisioning and safety of young. Without
these physiological and behavioral constraints, full-grown, older mothers can
invest more in male offspring. Hence, over their reproductive life span, mothers
may invest approximately equally in the sexes.
In fact, male puma cubs are generally larger than female cubs. When we
examined masses (kg) of 108 nursling cubs (eighteen to forty-nine days old) in
thirty-five mixed-sex litters and made eighty paired comparisons of masses
between male and female siblings within each litter, we found that males out-
weighed sibling females in forty-nine comparisons (61 percent); male and
female sibling masses were equivalent in twelve comparisons (15 percent); and
males weighed less than female siblings in nineteen comparisons (24 percent).
Masses of male and female cubs within litters diverged in favor of males at about
FIGURE 6-1. Masses of puma cubs, San Andres Mountains, New Mexico, 1986–1994.
twenty days of age (Fig. 6-1). Other puma biologists have also quantified that
male cubs grow larger than female cubs throughout the period of parental
investment (Robinette et al. 1961, Maehr and Moore 1992). Hence, male cubs
should be more costly for mothers to produce and rear successfully than females.
The actual proximate mechanism at work to adjust sex ratio in mammals is
not known (Clutton-Brock and Iason 1986, Hiraiwa-Hasegawa 1993, Krackow
1995, James 1996). But two physiological mechanisms have been suggested that
may work throughout pregnancy and even before zygote formation to skew sex
ratios (Krackow 1995). First, sex-specific embryonic mortality may occur, thus
favoring the survival of one sex at birth. This has been linked to steroid hormone
levels during pregnancy in laboratory rodents. We would expect that if this
mechanism were operating in pumas, then litter sizes for mothers producing
their first litters would be smaller than for older mothers. But we found that lit-
ter sizes for first litters and subsequent litters were about the same.
The second mechanism involves the relative time of insemination within the
estrus cycle. This has affected the sex ratio at birth in a variety of mammals (e.g.,
rabbits, white-tailed deer, humans), and no evidence of fetal loss has been found
to account for sex ratio variation. Levels of maternal gonadotropin and steroid
hormones at the time of conception and attendant uterine activity may mediate
sex-specific sperm selection (Clutton-Brock and Iason 1986, Krackow 1995).
Maternal gonadotropin includes two types of hormones, follicle stimulating
hormone (FSH) and luteinizing hormone (LH), which are secreted by the pitu-
itary gland located at the floor of the brain and act on the ovary. FSH induces
follicular development and secretion of estrogen. LH induces ovulation and
stimulates formation of the corpus luteum in the ovary, which manufactures
estrogen and progesterone. Steroid hormones include progesterone, estrogen,
and testosterone. James (1996, 2000) reviewed data on mammals (including
humans) showing that a high proportion of female offspring is associated with
high maternal levels of gonadotropin and progesterone at the time of concep-
tion. Based on our observations on relative litter sizes, lengths of estrus periods,
lengths of associations between breeding pairs, and the numerous copulations
exhibited by breeding puma pairs (Eaton 1976), the second mechanism seems
more plausible for pumas.
Our observations of differential sex ratios in first litters and subsequent lit-
ters might be an intra-population phenomenon. Future intensive puma research
that quantifies sex ratios of offspring in females with known histories in other
populations could test this hypothesis. Certainly, as other researchers ponder
explanations for adjustment in sex ratios, other hypotheses may emerge (see
Trivers and Willard 1973, Clutton-Brock and Iason 1986, Kruuk et al. 1999,
Nunn and Pereira 2000).
Subadults
Subadults comprised a smaller proportion of the total population probably
because of mortality and emigration, which also made it more difficult to quan-
tify this population segment. Consequently, we quantified subadult sex ratios
from the total number of subadults observed during January population esti-
mates from 1988 to 1995 in the Treatment Area and from 1989 to 1995 in the
Reference Area. Subadult sex ratios for the Treatment Area (n = 5–6 males,
10–11 females) and the Reference Area (n = 8 males, 10 females) were
1:1.17–1:2.2 and 1:1.2, respectively. None of the ratios was significantly differ-
ent from 1:1 [6], probably because of small sample sizes. But like those of the
weaned cubs, the sex ratio of subadults tended to have fewer males than females.
In this age class the divergence was probably linked to greater mortality and emi-
gration rates among males (see Chapters 8 and 9).
Very rarely has published literature on pumas clearly quantified the sexes of
independent subadults in a manner that allows comparisons with the pumas on
the San Andres Mountains, again reflecting the difficulty in quantifying these
data. A notable exception was Ross and Jalkotzy’s (1992) study of an increasing
and moderately hunted puma population in Alberta, Canada, where they
observed nine male and nineteen female subadults. Their findings generally
agree with our observations in the Treatment Area.
Adults
Adult males were outnumbered by adult females as well (Table 6-1). The male-
to-female ratio for adults observed each year (1988–1994) averaged 1:1.4 for the
Treatment Area, which was very near the average of 1:1.6 for the Reference Area
(1989–1994). Sex ratios of adults were not significantly different from a hypo-
thetical 1:1 ratio in all annual comparisons within the Treatment Area and the
Reference Area [7]. Adult sex ratios computed from January population esti-
mates from 1989 to 1995 produced almost identical results (Table 6-2). Mean
sex ratios for the Treatment and Reference Areas were 1:1.5 and 1:1.4, respec-
tively. Again, none of the annual comparisons of adult sex ratios in either area
were significantly different from 1:1 [8]. The sex ratio of the population of adult
pumas that we observed each year was representative of the sex ratio in the entire
population because we were observing 86 percent or more of the adult popula-
tion in each area each year. Before and after the removal of adult pumas from
the Treatment Area, adult females were favored numerically, regardless of
whether the sex ratio was quantified annually or each January. In addition, by
the time the Treatment Area recovered from the experimental removal thirty-one
months later, the exact adult sex ratio was restored (see Chapter 10). This sug-
gested that adult pumas that were removed were replaced in kind.
Table 6-1. Sex ratios of adult pumas in the Treatment Area and Refer-
ence Area each year, 1988–1994, San Andres Mountains, New Mexico.
Year Treatment Area a Reference Area a
Observed M:F b Operational M:F c Observed M:F b Operational M:F c
1988 1:1.4 1:0.9 1:1 1:0.9
1989 1:1.3 1:0.9 1:1.6 1:0.9
1990 1:1.4 1:1.3 1:1.7 1:1.1
1991 1:1.4 1:1.1 1:1.6 1:1.2
1992 1:1.8 1:1.2 1:1.6 1:1.2
1993 1:1.4 1:0.9 1:1.5 1:1
1994 1:1.1 1:0.8 1:1.3 1:0.7
aIndividuals that overlapped the Treatment Area-Reference Area boundary were counted in
both areas.
bSex ratios were determined for the number in each sex counted during an entire 12-month
period.
cOperational sex ratio represents the number of females available for breeding per male in a
12-month-period.
Ob s e rved adult puma sex ratio in our study population, as well as others
in North America, indicate a pattern of fewer males than females (Se i d e n-
sticker et al. 1973, Shaw 1977, Logan et al. 1986, Hopkins 1989, Maehr et
al. 1989, Sp re a d b u ry et al. 1996, Ross and Jalkotzy 1992, Beier and Barrett
1993, Lindzey et al. 1994). There are at least three biological reasons for the
pattern in adult pumas: (1) Males and females are born into the puma popu-
lation in equal pro p o rtions on average. But as subadults, male pumas have
higher mortality rates than females, hence, fewer males are available as poten-
tial re c ruits (see Chapter 8). (2) Females are recruited into the adult popula-
tion at higher rates than are males (see Chapter 10). (3) Pumas have a polyg-
ynous and promiscuous mating system where the dominant males breed with
s e veral females (see Chapter 7). Males engage in intense, agonistic competi-
tion for access to mates and territory, and males attempt to cover large areas
of terrain to pursue mates, both of which affect the spatial distribution of
males. This tends to space adult males out over larger areas in relation to
females (see Chapter 12).
Mate competition may be enhanced by the operational sex ratio—the ratio
of adult males to adult females available for breeding each year (Tables 6-1, 6-
2). We excluded females unavailable for breeding because they were rearing cubs
that were born in the previous year, unless those cubs reached independence or
Table 6-2. Sex ratios of adult pumas in the Treatment Area and Refer-
ence Area each January, 1989–1995, San Andres Mountains, New Mex-
ico.
Treatment Areaa Reference Areaa
Year Observed M:F b Operational M:Fc Observed M:F b Operational M:Fc
1989 1:1 1:0.7 1:0.9 1:0.1
1990 1:2 1:1.2 1:1.6 1:0.9
1991 1:1.4 1:0.3 1:1.6 1:0.6
1992 1:1.3 1:0.7 1:1.7 1:1.1
1993 1:1.8 1:0.6 1:1.4 1:0.6
1994 1:1.4 1:0.9 1:1.4 1:0.7
1995 1:1.3 1:0.4 1:1.4 1:1.1
aIndividuals that overlapped the Treatment Area–Reference Area boundary were counted in
both areas.
bSex ratios were determined for the number of adult pumas in each sex counted in January
each year.
cOperational sex ratio represents the number of females available for breeding per male in
January each year.
died and the female rebred and bore cubs in the tally year. Average operational
sex ratios in the Treatment Area and Reference Area were identical, 1:1. In gen-
eral there were as many years when there were slightly more adult males avail-
able per breeding female as there were years when there were slightly more
breeding females available per male. In reality, at any one point in time, the
operational sex ratio is even higher because the proportion of pregnant or cub-
rearing females is higher. For example, in January of each year, the average oper-
ational sex ratio in the Treatment and Reference Areas was identical, 1:0.7, and
highly variable (Table 6-2). Normally, there were more adult males available
than there were potential female mates. With these ratios, intense male-to-male
competition for mates should be expected in the polygynous, promiscuous mat-
ing system of pumas (see Chapters 7 and 8).
Age Structure
Another important distinguishing attribute of an individual is its age. Generally
speaking, the age of an individual puma influences its survival, reproductive
capacity, and social status. Thus, the age structure of the puma population influ-
ences the capacity for population growth.
FIGURE 6-2. Age structure of the puma population in the Treatment Area and Reference
Area each January, 1989–1995, San Andres Mountains, New Mexico.
The Population
We quantified the proportions of adults, subadults, and cubs in the Treatment
Area and Reference Area for each January from 1989 to 1995 (Fig. 6-2). The
greatest proportion of the puma population was composed of adults, followed
by cubs, and then subadults. In the Treatment Area (Table 6-3), adult males and
females comprised a mean proportion of 0.23 ± 0.07 and 0.33 ± 0.08 of the
population, respectively. Total adults (males plus females) comprised a mean
proportion of 0.56 ± 0.13, subadults a mean of 0.10 ± 0.09, and cubs a mean
of 0.34 ± 0.10 of the population. After discounting the first January population
estimate following the experimental removal of pumas in the Treatment Area
(1992), the mean proportions of the three major age classes in the population
were as follows: adult males = 0.25 ± 0.06, adult females = 0.36 ± 0.05, total
adults = 0.61 ± 0.08, subadults = 0.07 ± 0.07, and cubs = 0.32 ± 0.10. In the
Reference Area (Table 6-4), adult males and females comprised a mean propor-
tion of 0.26 ± 0.03 and 0.35 ± 0.07 of the population, respectively. On average,
adults together comprised 0.61 ± 0.09, subadults 0.06 ± 0.02, and cubs 0.33 ±
0.08 of the population. Based on the three broad age classes, the average annual
age structure in the Treatment Area and Reference Area were similar.
Table 6-3. Proportion of pumas in three broad age classes in the Treat-
ment Area population each January, 1989–1995, San Andres Moun-
tains, New Mexico.
Adults
Year Males Females Total Subadults Cubsa
1989 .32 .32 .64 .18 .18
1990 .20 .41 .61 .12 .27
1991 .25 .36 .61 .04 .36
1992 .14 .18 .32 .23 .45
1993 .17 .31 .48 .10 .41
1994 .30 .42 .72 0 .28
1995 .25 .32 .57 0 .43
Mean ± SD .23 ± .07 .33 ± .08 .56 ± .13 .10 ± .09 .34 ± .10
aProportion of cubs was calculated using the mean of the range of the estimated number of
cubs present.
Table 6-4. Proportion of pumas in three broad age classes in the Refer-
ence Area population each January, 1989–1995, San Andres Moun-
tains, New Mexico.
Adults
Year Males Females Total Subadults Cubsa
1989 .27 .24 .51 .07 .42
1990 .24 .38 .62 .06 .32
1991 .25 .34 .59 .03 .38
1992 .23 .38 .61 .05 .33
1993 .22 .30 .52 .10 .38
1994 .26 .35 .61 .09 .30
1995 .32 .46 .78 .05 .16
Mean ± SD .26 ± .03 .35 ± .07 .61 ± .09 .06 ± .02 .33 ± .08
aProportion of cubs was calculated using the mean of the range of the estimated number of
cubs present.
Unfortunately, almost all other literature that reports puma age structure
does not use similar age classes to ours; hence, comparisons with our study pop-
ulation were impractical. Only the study of a moderately hunted, increasing
Alberta puma population yielded similar data (Ross and Jalkotzy 1992). During
five years, the population age structure averaged about 48 percent adults, 19 per-
cent subadults, and 34 percent cubs.
Adult Pumas
We estimated the ages (in months) of adult pumas in the Treatment Area and
Reference Area in January each year for the seven-year period 1989–1995 (Table
6-5). In the Treatment Area, adult males were generally older than adult females
in the first five years but not in the last two years. The distributions of the ages
of males and females were identical in all years except 1990 [9]. In the Reference
Table 6-5. Mean ages (in months) of adult pumas in the Treatment
Area and Reference Area each January, 1989–1995, San Andres Moun-
tains, New Mexico.
Males Females All adults
–
Year n x ± SD n x– ± SD x– ± SD
Treatment Area
1989 7 76.3 ± 31.0 7 50.4 ± 22.3 63.4 ± 30.0
1990 5 83.0 ± 19.6a 10 52.2 ± 24.4a 62.5 ± 27.1b
1991 7 76.4 ± 33.7 10 58.9 ± 29.5 66.1 ± 32.5
1992 3 97.7 ± 40.1 4 80.5 ± 23.1 87.9 ± 32.7b
1993 5 78.0 ± 49.7 8 60.1 ± 36.3 67.0 ± 42.9
1994 7 55.3 ± 36.4 9 66.9 ± 37.3 61.8 ± 37.4
1995 7 46.7 ± 16.4 8 72.5 ± 34.0 60.5 ± 30.1
Reference Area
1989 8 50.9 ± 24.2 7 50.1 ± 19.7 50.5 ± 22.2
1990 8 56.6 ± 26.1 13 43.9 ± 24.1 48.8 ± 25.6b
1991 9 64.3 ± 27.5 12 53.6 ± 24.8 58.2 ± 26.5
1992 9 64.9 ± 32.2 15 47.1 ± 21.5 53.8 ± 27.4b
1993 11 68.2 ± 34.5 15 47.5 ± 18.1 56.3 ± 28.2
1994 11 62.8 ± 30.0 15 52.4 ± 20.5 56.8 ± 25.5
1995 12 64.8 ± 35.2 14 65.9 ± 20.5 65.3 ± 28.3
aDistribution of ages of males and females in the Treatment Area differed in 1990
(P = 0.05).
bDistribution of ages of all adults in the Treatment Area and Reference Area differed in
1990 and 1992 (P < 0.10).

Area, the distributions of the male and female ages were identical in all years
[10]. Still, ages of adults in the Reference Area had a similar profile to the Treat-
ment Area, with males being generally older than females in the first six years
but not in the last year (Table 6-5). This tendency for adult males to be older
than adult females reflects greater survival rates for males (see Chapter 8).
Ages of adult pumas (males and females combined) in the Treatment Area
and the Re f e rence Area we re identically distributed in all years except 1990 and
1992 [11] (Table 6-5). Still there was a trend in which adults in the Treatment
Area we re generally older than adults in the Re f e rence Area. This characteristic
was coincident with a lower density of adult pumas in the Re f e rence Are a ,
which we believe was a result of greater population reduction in that portion of
the San Andres Mountains during the puma control actions that preceded our
study. We would expect older-age adults where off-take was less seve re because
animals had a greater probability of living longer. Hence, it may be character-
istic of populations that are heavily exploited to be composed of relatively
young pumas as young-aged recruits composed of progeny and immigrants
replace killed adults.
We quantified the proportion of adult male and female pumas in three broad
age classes (Fig. 6-3). Young adults were pumas twenty-one to thirty-six months
old that had recently been recruited into the breeding population. Prime adults
were pumas thirty-seven to ninety-six months old that were well established in
the population and did most of the breeding. Old adults were pumas 97 to 156
months old that were rapidly declining in the population chiefly because of
deaths.
Pumas lived to old ages on the San Andres Mountains mainly because they
were protected from human exploitation during our study. Moreover, adult
males generally lived longer lives than adult females, reflecting their higher nat-
ural survival rates (see Chapter 8). The oldest male and female pumas we found
were estimated at 152 months and 146 months, respectively. The extreme
longevity for wild pumas is probably represented by two Florida panthers. A
male was about 180 months old when he was killed by another male panther.
And a female was about 158 months old when she was killed by a car (Maehr
1997a). Other old-age pumas were in a wild population in Alberta; there, a 132-
month-old male died while trying to kill a bighorn sheep, and a 108-month-old
female died while she was trying to bring down an elk (Ross et al. 1995). In a
puma population in California that had been protected from hunting for twelve
years, a 101-month-old male was killed by another puma and a 128-month-old
female died of undetermined causes (Hopkins 1989). In contrast, pumas sub-
jected to moderate to heavy hunting pressure in Wyoming very rarely reached
eighty-four months of age (Logan et al. 1986). Extreme ages for captive pumas
range from about 120 to 216 months (Anderson 1983).
FPO @ 75%
FIGURE 6-3. Adult puma age structure in the Treatment Area (TA) and Reference Area (RA)
in three broad age classes, San Andres Mountains, New Mexico.
LP > > KX
1. Through capture-mark-recapture and radiotelemetry techniques, we accu-
rately censused the puma population annually, accounting for 93–100 per-
cent of adults in the Treatment Area during 1988–1994 and 86–100 per-
cent of adults in the Reference Area during 1989–1994. By adding in the
individual adult pumas we apparently missed, we accounted for about 100
percent of the population each year. Adult pumas were animals that bred or
males that were twenty-four months or older and females that were twenty-
one months or older. Subadults were independent but did not breed. Cubs
were dependent on their mothers.
2. The secondary sex ratio averaged 1:1 for nursling cubs. But it apparently
shifted from favoring females in a mother’s first litter to favoring males in
her subsequent litters. We speculated that young first-time mothers may
produce more of the smaller female cubs and still have optimal litter sizes
as a physiological trade-off with somatic growth.
3. Sex ratios for subadults and adults favored females, owing to differential
survival rates in subadults and the influence of the social and mating behav-
ior in adults.
4. The operational sex ratio of adults was 1:1 and probably heightened male-
male competition for mates.
5. On average, the puma population on the San Andres Mountains comprised
61 percent adults, 6 percent subadults, and 33 percent cubs. The composi-
tion was similar in the Treatment and Reference Areas. The age (in months)
distribution of adult pumas was similar in the Treatment and Reference
Areas. But in general, adults in the Treatment Area were somewhat older
than adults in the Reference Area, probably because of past differential
exploitation pressure. Adults lived relatively long lives in this protected pop-
ulation. Generally, males lived longer than females. The oldest pumas we
found were a 152-month-old male and a 146-month-old female.
LN N5LN5)L
1. Male:female sex ratio for fourteen male and twenty female cubs 52 to 427
days old did not differ from 1:1. Chi-square test, c2 = 1.06, 1 d.f., P > 0.10.
2. Sex ratios in first litters (n = 15) and subsequent litters (n = 27) from nine-
teen mothers were different. Chi-square test, c2 = 4.675, 1 d.f., P = 0.03.
3. Sex ratios in first litters (n = 8) and subsequent litters (n = 12) of the same
females were different. Chi-square test, c2 = 5.901, 1 d.f., P = 0.02.
4. Sex ratios in first litters (n = 8) were different from 1:1. Chi-square test, c2
= 4.482, 1 d.f., P = 0.04.
5. Masses of nine females were different when they were eighteen to twenty-
four months old (x– = 31.6 ± 2.6, median = 31.3) compared to when they
were twenty-eight to sixty months old (x– = 34.7 ± 2.3, median = 35.4). Sign
test, P = 0.07.
6. Sex ratios of subadult pumas in the Treatment Area and Reference Area
were not different from 1:1. Treatment Area: Chi-square test, c2 = 1.0 or
2.25, 1 d.f., P > 0.10. Reference Area: Chi-square test, c2 = 0.22, 1 d.f., P
> 0.10.
7. Sex ratios of adult pumas observed per year in the Treatment Area and Ref-
erence Area did not differ from 1:1. Chi-square test, c2 < 2.71, 1 d.f., P >
0.10, all tests.
8. Adult sex ratios for January population estimates 1989–1995 for the Treat-
ment Area and Reference Area did not differ from 1:1. Chi-square test, c2
< 2.71, 1 d.f., P > 0.10 all tests.
9. Ages (months) of adult male (n = 3–7) and adult female (n = 4–10) pumas
in the Treatment Area, 1989–1995. Distributions of ages were identical for
all years (P > 0.10) except 1990 (Mann-Whitney U test, U = 9.0, P = 0.05).
10. Ages of adult male (n = 8–12) and adult female (n = 7–15) pumas in the
Reference Area, 1989–1995. Distribution of ages was identical for all years
(Mann-Whitney U test, P > 0.10).
11. Ages of adult male and female pumas combined in the Treatment Area (n =
7–17) and Reference Area (n = 15–26) were similar in all years (P > 0.10),
except 1990 (Mann-Whitney U test, U = 209.5, P = 0.095) and 1992
(Mann-Whitney U test, U = 131.0, P = 0.026).
Chapter 7
Reproduction
Natality
We documented the birth of 220 cubs from seventy-nine litters produced by
t h i rty-nine female pumas. Of those cubs, we captured and marked 174 (79
p e rcent). Of the forty-six cubs that we re not marked, we knew of ten cubs
from three litters that died shortly after birth, three others disappeared fro m
families before we could re c a p t u re them, and nine other cubs we re dependent
on their mothers at the close of our study. Hence, we did not have the oppor-
tunity to closely examine these cubs. Five nonviable cubs in two litters we re
born to the same female: one litter was apparently stillborn and the other was
a b o rted.
By examining cubs at various known ages during their development, we
learned that they suckled until they were six to eight weeks of age. During the
nursing stage, cubs were altricial, and mothers kept them at a single nursery or
moved them short distances, usually less than 200 m, to alternate nurseries.
Moreover, mothers usually tended to their cubs on a daily basis. It was during
this period of intensive maternal care that we estimated litter sizes at birth. Once
cubs were weaned, they were carnivorous and precocious. They moved to kills
their mother made and in the process were subjected to greater perils of their
environment. Mortality was expected to increase and thus reduce observed lit-
ter sizes.
Theoretically, in long-lived itero p a rous animals (i.e., animals that pro d u c e
more than one litter in their lifetime), natural selection favors individuals
83
that produce optimal litter sizes that maximize offspring survival and future
b reeding success (Lack 1947, Boyce and Perrins 1987, Freeman and He r ro n
1998). In fifty-three nursling litters in which we counted all the cubs nine to
f o rty-nine days postpartum (x– = 32.3 ± 8.9); there was an average of 3.0 ±
0.7 cubs. The distribution of litter sizes for the fifty-three litters was com-
posed of litters with two cubs (n = 13), three cubs (n = 26), and four cubs (n
= 14). Twenty-six of the fifty-three litters we re observed at birth nurseries
(i.e., where the cubs we re born); the other twe n t y - s e ven litters we re observe d
at secondary nurseries (i.e., nurseries used after mothers moved cubs away
f rom birth nurseries). For twenty-one other litters that we observed with
weaned cubs that we re 52–427 days old (x– = 175.7 ± 112.0), litter sizes ave r-
aged 2.2 ± 0.8 cubs. The smaller litter sizes probably reflects mortality that
o c c u r red after we a n i n g .
We estimated an overall natality rate of 0.17 by dividing the mean litter
s i ze of the fifty-three nursling litters (i.e., 3.0) by the birth interval that we
observed for sixteen intervals where at least one cub in the first litter sur-
v i ved to independence or to twe l ve months of age (i.e., 17.4 months; see
“ Mating, Gestation, and Birth In t e rvals”). Natality rate in the Tre a t m e n t
A rea was 0.16 in the pre - t reatment years (1988–1990) compared to 0.18 in
the post-treatment years (1992–1994). This was due to a slightly smaller
Slide @318%
PHOTO 16. Puma F147’s three cubs at five weeks of age.

CHAPTER 7. REPRODUCTION 85
Slide @318%
PHOTO 17. Puma F54’s birth nursery, where she had four cubs.
average litter size (x– = 2.7 ± 0.5, n = 6) in pre - t reatment years than in post-
t reatment years (x– = 3.2 ± 0.6, n = 11) [1]. This difference may have been
an artifact of the re l a t i vely small sample sizes of nursling litters in which we
observed all the cubs.
In addition, we quantified fecundity rates for the Treatment and Reference
Areas for biological years in the deer population increase phase (1989–1990 to
1991–1992) and decline phase (1992–1993 to 1993–1994) (see Chapter 17).
We defined fecundity rate as the average number of live cubs produced per adult
female monitored in each area during each biological year (August–July) (see
Caughley 1978:72). The average fecundity rate in the Treatment Area during
the deer increase phase was 1.6 ± 0.7; it was 2.0 in both years of the deer decline
phase. In the Reference Area, the average fecundity rate was 1.4 ± 0.2 in the deer
increase phase, and 1.3 ± 0.5 in the decline phase [2]. These statistics showed no
pattern suggesting that puma reproductive output was affected by the declining
prey base. However, the lowest fecundity rate of 0.9 occurred in the Reference
Area in the last year of the deer decline phase and may be a harbinger of what
could be expected during a severe and prolonged deer decline or population low.
Cub production apparently was not sensitive to the decline in the deer popula-
tion during our study. We speculated that the prey base had not yet reached a
threshold low enough to trigger nutritional deficiencies in reproducing females
and their offspring by late 1994. Hence, a time delay should be expected before
puma reproduction declines in response to reduced food resources. Decline in
productivity could be manifested in smaller litter sizes, longer birth intervals,
increased age at first reproduction, increased mortality, or a combination of
these properties.
In comparison with the reproductive output of other puma populations
studied in North America, mean postpartum litter size on the San Andres
Mountains was among the largest. The only one slightly larger was a mean size
of 3.1 ± 1.2 found by Spreadbury et al. (1996) in southeastern British Colum-
bia where they examined seven litters that were two to ten weeks old. Florida
panthers have produced the smallest litters, averaging 2.3 cubs in four litters that
were less than fourteen days old (Maehr and Caddick 1995). Most other stud-
ies quantified litter sizes when cubs were older, after greater postpartum mortal-
ity had probably occurred, as was likely the case with the older weaned litters
that we observed. In Idaho, where cubs were first observed at various ages, there
was an average of 2.6 cubs per litter (Hornocker 1970). Average litter size of 2.2
cubs in Alberta was estimated from track sets when cubs were less than four
months old (Ross and Jalkotzy 1992). In Wyoming, litters less than six months
old had, on average, 2.9 cubs and litters older than six months had, on average,
2.3 cubs (Logan et al. 1986). In Utah, where 65 percent of the cubs were older
than three months when examined, there were 2.4 cubs per litter (Lindzey et al.
1994). Again, Florida produced the lowest average litter size of 1.9 cubs when a
total of twenty-five Florida panther litters of various ages was considered (Maehr
and Caddick 1995).
Natality rates are not normally reported in technical puma literature because
data on mean litter size for nurslings and birth intervals are difficult to gather
for wild pumas. We could compare natality rates in the San Andres Mountains
puma population only after calculating rates from four other populations that
provided data on both statistics, mean litter size and birth interval. A population
in southeastern British Columbia (Spreadbury et al. 1996) had a natality rate of
0.17, identical to some of our rates. But natality rates for pumas in southwest-
ern Alberta (Ross and Jalkotzy 1992), south-central Utah (Lindzey et al. 1994),
and southern Florida (Maehr et al. 1991, Maehr and Caddick 1995) were lower,
at 0.11, 0.10, and 0.10, respectively [3]. Because average litter sizes for Alberta
and Utah used weaned litters that may have experienced greater mortality, we
believe the computed natality rates for those areas are actually biased downward.
However, the estimate for the endangered Florida panther used an average litter
size based on young nurslings and may indicate a relatively low natality rate.
Scant information on reproduction in other wild large solitary felids precludes
comparisons with our puma data, except for the tiger. We calculated a natality
rate of 0.14 for tigers by using the mean litter size of 3.0 and minimum mean
birth interval of 21.6 months that Smith and McDougal (1991) found in
Nepal’s Royal Chitwan National Park.
Puma hunters, predator control agents, and wildlife managers often believe
that first-time puma mothers normally produce smaller litters than experienced
mothers. We were interested in the validity of this notion because of its impli-
cations on natality rate in suppressed or rebuilding puma populations where
young-aged mothers may comprise a substantial proportion of breeding females.
Our data contradicted this widely held belief. We examined all of the nursling
cubs (nine to forty-six days old, x– = 30.7 ± 9.9) of eight known-age mothers and
eight approximately known-age mothers that produced cubs for the first time in
their lives. Mean litter size was 3.1 ± 0.7 cubs per litter, which was almost the
same as the mean of 3.0 ± 0.7 nursling cubs per litter (eighteen to forty-nine
days old, x– = 32.9 ± 9.0) in twenty-eight litters of nineteen females that pro-
duced their second to fourth litters. When we compared litter sizes for eight
matrilineal mothers for which we had complete reproductive histories, all of
which were progeny recruits that were philopatric (n = 6) or had home ranges
adjacent to their natal areas (n = 2), we found the mean size of their first litters
(3.4 ± 0.74) was about equal to the mean size of their twelve subsequent litters
(3.2 ± 0.58). Our observations indicate that productivity of first-time mothers
is about the same as experienced mothers. However, it is possible that older,
more-experienced mothers have greater success at rearing their young to inde-
pendence.
The eight matrilineal mothers tended to have larger first litters than six
known immigrant mothers (x– = 2.8 ± 0.75), although the difference was not sig-
nificant [4]. We would expect matrilineal mothers to have greater knowledge of
local resources (i.e., food, nurseries) and perhaps even greater social dominance
compared to immigrant females, which might result in a reproductive advan-
tage. If true, then philopatric females would be expected to have greater fitness,
particularly if their cubs also exhibited greater survival, which they apparently
do (see Chapter 8).
We also wanted to see if litter size changed with the age of the mother. Five
mothers that were 92–112 months old had six litters with an average of 2.8 ±
0.98 nursling cubs, which tended to be lower than the mean size of first litters
of the eight matrilineal mothers (above); however, the difference was not signif-
icant [5]. Apparently, litter size was not substantially affected by the biological
process of aging relative to the age distribution we observed in the San Andres
Mountains puma population.
Timing of Births
We determined the month of birth for seventy-eight puma litters on the San
Andres Mountains. Litters were born in every month of the year except Febru-
ary (Fig. 7-1). But there was a birth pulse during July, August, and September
when ten, eleven, and eleven litters (41 percent in aggregate) were produced,
respectively. The occurrence of puma litters in those three months was greater
than expected with a uniform distribution [6]. The timing coincided with the
birth period for mule deer fawns and the annual monsoon season, which occurs
during mid-July through September. During this time, food and cover is most
available for lactating does and their fawns, and, hence, probably enhances their
survival (see Sinclair et al. 2000). Similarly, food for puma mothers is the most
available and vulnerable at that time.
On the San Andres Mountains, mule deer comprised 91 percent of prey ani-
mals that we found were killed by pumas, and 92 percent of the relative biomass
of prey items consumed by pumas. Furthermore, fawns comprised more than
one-quarter of the puma-killed deer that we examined (see Chapter 17), and
they may be selected by puma mothers (Pierce et al. 2000a). During the fawn-
ing season, mule deer are more abundant than at any other time of the year, and
small fawns are the most vulnerable component of the population to predators.
The postparturition period is critical for puma mothers. Lactation is the most
energy-demanding part of the reproductive cycle, because mothers must main-
FPO @ 80%
FIGURE 7-1. Distribution of births of seventy-eight puma litters by month, 1986–1994, San
Andres Mountains, New Mexico.
tain themselves as well as produce energy-rich milk for their offspring (Bronson
1989). Mothers must also restrict their movements around nurseries while they
nurse and protect young. In addition, especially after the nursing period, the
energetic demands of the family unit increase dramatically, primarily due to the
rapidly growing cubs (Ackerman 1982), and must be satisfied by consuming
prey. Hence, puma births timed for periods when food is predictably abundant
and vulnerable would be an adaptive strategy. Mothers that produce litters at
those times probably incur survival advantages and relatively high net reproduc-
tive success; hence, a trait for giving birth when food is most abundant would
be expected to be favored by natural selection.
We examined variation in timing of birth for individual mothers by inspect-
ing the birth dates of twelve female progeny recruited as breeding adults on the
San Andres Mountains (i.e., F1 generation females) relative to when they gave
birth to nineteen litters themselves (i.e., the F2 generation). Ten (53 percent) of
the litters were born during the July–September birth pulse. Of the F1 genera-
tion mothers, six were born in the birth pulse themselves and produced six
birth-pulse litters. But four of those mothers also produced five litters outside of
the pulse. Of the six remaining mothers, three were born in March, two in May,
and one in December. Four of the females produced four birth-pulse litters, and
three of the females produced four litters outside the pulse (one female produce
one litter in and one litter outside the pulse). We also examined tendencies in
birth timing for twelve other mothers that had immigrated onto the San Andres
Mountains. Those females produced twenty-eight litters. Seven of the mothers
produced thirteen (46 percent) litters in the July–September birth pulse. Six
females were parents (i.e., P1 generation) to the twelve F1 mothers above;
together they produced nine birth-pulse litters. Yet, seven of the mothers that
had birth-pulse litters also produced thirteen litters outside of the pulse. Two
other mothers had litters in October and December.
If giving birth to cubs when food is abundant and vulnerable is so advanta-
geous, why don’t all females have cubs during that time? An answer may par-
tially lie with the fates of cubs and the fact that pumas are polyestrous, meaning
that they cycle into reproductive receptivity continually throughout the year
(Bonney et al. 1981). Some females that produce cubs lose the entire litter due
to disease, infanticide, or predation. Those females can recycle into breeding
condition within a few weeks or months (see “Mating, Gestation, and Birth
Intervals”), and thus they produce cubs during another part of the year. For five
mothers that lost their entire litters, then rebred successfully, we found that their
doomed litters were born in March, April, August, October, and November.
They had their subsequent successful litters in May (n = 4) and July. In addition,
females may reach reproductive maturity at different times of the year. Specific
mechanisms affecting this in pumas are unknown, but we presume they are sim-
ilar to those affecting other mammals. Fertility is influenced by phenotype and
environment. For example, genes influence somatic development and onset of
fertility; nutritional stress during adolescence could delay puberty; and repro-
ductive condition may be primed by pheromonal cues in the environment and
social situations could stimulate or suppress it (Bronson 1989).
A simple hypothesis may explain why there is an observed pulse in puma
births. Pumas breed and give birth to litters year-round, but litters born during
the period of food abundance have the greatest probability of survival. Lactation
poses the greatest energetic challenge to mothers (Bronson 1989). Besides pro-
ducing energy-rich milk, mothers must also sustain themselves and protect
young from the ambient climate and predators. After weaning, cubs eat meat
and grow rapidly; energetic demands (i.e., kcal / day) of the puma family unit
increase dramatically to as much as six times that of a nonproducing female,
depending upon the number and size of the cubs during adolescence (Ackerman
1982). Hence, litters born in periods of food abundance stand the greatest
chance of surviving beyond the first two to four critical weeks that it takes field
biologists to detect them. Comparatively, cubs born during other, less-favorable
times of the year would be expected to have lower survival. In this scenario, no
heritable trait for timing of birth is required. Considering the dispersal patterns
of pumas (see Chapter 9), it is more adaptive for them to be polyestrous year-
round instead of locking into a single birth season that has evolved from local
environmental conditions. Pumas with the former trait are more fit because they
can more successfully exploit changing environmental conditions. This has
allowed the puma to evolve into a highly adaptable and colonizing species.
The pattern of puma births on the San Andres Mountains conformed closely
with the pattern observed in Ne vada (Ashman et al. 1983), but there the birt h
peak occurred during June and July. In Wyoming, where 70 percent of puma cubs
o b s e rved we re born from August to November, Logan (1983) suggested that
mothers and weaned cubs took advantage of high concentrations of mule deer and
elk on winter ranges during November to March. Similarly, of observed puma
births, 78 percent occurred from June to November in British Columbia (Spre a d-
b u ry 1989), 77 percent occurred from August to October in Utah (Lindzey et al.
1994), and 60 percent occurred during June to August in Alberta (Ross and
Jalkotzy 1992). Anderson (1983) compiled data on fifty-three wild litters from
fourteen sources and showed that 66 percent of births we re observed during June,
July, September, October, and November. Florida panther repro d u c t i vecharacter-
istics support our hypothesis that observed birth pulses coincide with periods of
food abundance. Most panther litters we re recorded between March and July. This
overlapped the Fe b ru a ry to April white-tailed deer fawning season, which boosted

the deer population to its highest annual levels (Maehr 1997b).
Mating, Gestation, and Birth Intervals

By intensively monitoring radio-collared adult pumas, we were able to docu-
ment the mating of thirty-six pairs (Appendix 2). Two pairings resulted in non-
viable young from the same female; one litter was stillborn after eighty-five to
ninety days of gestation, and one litter was aborted after fifty-six days of gesta-
tion. Once we located a pair of pumas together, we homed on their locations
and found evidence of a potential breeding association. Evidence included tracks
of the pair traveling together, kills that they shared, vocalizations, and visual
observations of the pumas. Although we did not observe pumas actually copu-
lating, the ultimate proof of a breeding association was the subsequent birth of
cubs. We estimated the length of gestation by counting the number of days from
our first observation of the pair together to the estimated birth date of the cubs.
The mean gestation period that we calculated for the thirty-one pairings was
91.5 ± 4.0 days (range = 83–103). Considering the July–September birth pulse
that we observed on the San Andres Mountains, the corresponding breeding
period occurred during April–June. Quantitative information for length of
puma gestation in the published literature mostly pertains to pumas in captiv-
ity. The mean gestation period that we observed was practically identical to the
mean of 91.9 days that Anderson (1983) calculated from forty-two captive lit-
ters from eleven sources. Radio-tracking enabled Beier and Barrett (1993) to
estimate a mean gestation period of ninety-three days for only four wild litters
in Southern California.
Estrus, the period of the reproductive cycle when ovulation occurs and the
female puma is at maximum receptivity to the sexual overtures of males, ranges
from three to twelve days for captive pumas (see sources in Anderson 1983). For
pumas on the San Andres Mountains, we estimated estrus lasted one to ten days
based on our radiotelemetry data on consort pairs. Successfully breeding pairs
were together one to four days. We documented another pair in an apparent
breeding association for six days, but it did not result in viable cubs. Four
females associated with two different males during the same estrus period, indi-
cating that female pumas can be promiscuous, a behavior that was typically
exhibited by the male pumas. One of the females associated with one male for
one day, with a different male two days later, then with the first male again seven
days after that; this suggested a receptive period of ten days. A consort pair in a
wild puma population in central Idaho traveled together for up to sixteen days
(Seidensticker et al. 1973).
During mating, puma pairs copulate ve ry fre q u e n t l y. Captive pumas have

been observed to copulate as much as fifty to seventy times per day during
e s t rus (Eaton 1976). Female pumas are apparently induced ovulators (Bonney
et al. 1981), and thus, high copulation frequency may stimulate peak repro-
ductive condition. Experiments with captive pumas by Me h rer (1975) is sup-
p o rt i ve of this idea; the number of copulations contributed to ovulation. Eaton
(1976) speculated that the high copulation rates evolved as a measure of male
vigor and a means by which females can maximize repro d u c t i ve success. We
believe that besides the ability to repeatedly copulate, vigor of adult male
pumas is also rigorously tested during defense of territory or estrus females (i.e.,
access to mates) against other male interlopers (see Chapter 8). Thus, when an
e s t rus female monopolizes the sexual favors, and hence the sperm, of a success-
ful territorial male via an extended estrus period with frequent copulations, she
is in essence mating with the most fit male and acquiring “good genes” for her
offspring. Furt h e r m o re, these mating habits may enhance recognition betwe e n
the pair and awareness of paternity by the male, and may reduce aggression by
the male directed tow a rd the female or her cubs in subsequent encounters (see
Chapter 8).
The estrous cycle (i.e., reproductive cycle) in wild pumas is practically
impossible to observe directly. But because we could estimate the frequency of
estrus by using radiotelemetry, we estimated the length of the estrous cycle by
noting the frequency of apparent estrus periods. We did this by marking the
beginning of one estrus period to the beginning of the next as indicated by
twelve apparent breeding associations between radio-collared adult males and
five different females. They showed a mean estrous cycle length of 24.5 ± 6.8
days, with a range of fourteen to thirty-five days. Our estimated estrous cycle
length was similar to a captive female that exhibited nine estrus periods that
averaged 22.8 days apart (Rabb 1959). But estrous cycle lengths can be highly
variable as was exhibited by two other captive females that averaged 59.5 days
(range = 17–114) between a total of thirteen estrus periods (Mehrer 1975).
We also quantified the length of puma birth intervals (i.e., time span
between litter births). Birth intervals where at least one cub in the first litter sur-
vived to independence (n = 15) or to twelve months of age (n = 1) averaged 17.4
± 2.5 months (range = 12.6–22.1). This time span represented the amount of
time it took a mother to give birth, rear her cubs to independence, and then to
rebreed and have another litter of cubs. For fifteen other intervals in which we
did not know the fates of the first litter with certainty, intervals averaged 16.9 ±
3.4 months (range = 12.2–22.5). The close similarity in birth intervals between
these two groups suggests that in most cases mothers in the latter group raised
one or more cubs to independence. Because of inadequate samples, we did not
compare birth interval lengths in the mule deer increase phase (n = 13, range =
12.6–22.1 months) and the decline phase (n = 3, range = 16.8–21.6 months).
Five females that we monitored closely lost their first litters entirely, but they
successfully bred again an average of 100.4 ± 116.9 days later (range = 23–307).
The shortest interval of twenty-three days was for a female whose first cubs were
stillborn. Two of these females lost their entire litters to cannibalism by territo-
rial males; they rebred 74 and 307 days later (x– = 190.5 ± 16.5). The other three
females that lost their litters to other causes rebred twenty-three to fifty-four
days later (x– = 40.3 ± 15.8).
Birth interval length is another statistic on puma reproductive biology that
is difficult to document for wild pumas. But there is some limited information
from disparate parts of the puma’s range. Twelve female pumas in Nevada exhib-
ited a mean birth interval identical to what we found (i.e., 17.4 months), with
a range of 11.5–24.0 months (Ashman et al. 1983). Generally, birth intervals on
the San Andres Mountains were somewhat shorter than in other areas. In Utah,
Lindzey et al. (1994) reported a mean birth interval of 24.3 months for seven
intervals where one or more cubs survived twelve months or longer. In Alberta,
Ross and Jalkotzy (1992) calculated a mean birth interval of 19.7 months for
twelve intervals. However, they cautioned that in three cases (twenty-five,
twenty-five, and thirty-two months) interceding litters may have been born and
lost because of low monitoring frequency. If so, then actual birth intervals would
be expected to be somewhat shorter. Closely monitored endangered Florida pan-
thers exhibited a mean of 23.3 months in five intervals (Maehr et al. 1991).
Puberty and First Litters

We used the timing of first associations between male and female pumas with
known ages or approximately known ages to determine ages of puberty. We
define puberty as the beginning of the adult stage in life when pumas are phys-
iologically and socially ready to breed. Only females that eventually produced
cubs were included in our analysis. Seven females with known birth dates asso-
ciated with adult males for the first time when they were an average of 21.4 ±
3.1 months old (range = 18–27). Of those females, two produced litters that
resulted from those associations; they were nineteen and twenty months old.
Three other females had four subsequent associations with adult males that aver-
aged 22.8 ± 8.7 days (range = 13–34) after the first association, which suggested
the females were in estrus. In support of this, two of these associations resulted
in litters. Another six females with approximately known ages apparently asso-
ciated with adult males for the first time when they averaged 22.2 ± 7.6 months
old (range = 15–36). Three of these associations resulted in litters; the females
were about fifteen, seventeen, and twenty-five months old. Another female was
impregnated when she associated with the same male twenty-six days later, thus
suggesting that her associations were related to estrus periods. Based on these
data, we generalized that female pumas on the San Andres Mountains reached
sexual maturity at about twenty-one months old.
To determine the ages when female pumas first successfully produced cubs,
we analyzed data on twelve known-age females. These females conceived appar-
ently for the first time at an average age of 26.1 ± 6.0 months (range = 19–37).
They bore their first live litters at a mean age of 29.1 ± 6.0 months (range =
22–40 months). Another eight females of approximately known ages conceived
successfully apparently for the first time at an average age of 26.0 ± 9.4 months
(range = 15–42). They produced their first live litters at an average age of 29.0
± 9.4 months (range = 18–45). The youngest female to bear cubs was approxi-
mately eighteen months old. When we captured her for the first time at about
nineteen months of age, she had already given birth to three cubs, which were
five weeks old. Her tawny pelage showed obvious light brown dapples on her
forelegs, shoulders, hips, and thighs. This meant that she was about fifteen
months old when she conceived.
Philopatric females did not exhibit reproductive suppression as a result of
residing within the home range of their mothers and territory of their fathers.
Three philopatric females associated for the first time with adult males when
they averaged 19.7 ± 1.5 months old (range = 18–21). Five philopatric females
conceived for the first time when they averaged 26.8 ± 7.0 months old (range =
20–37) and produced their first litters when they averaged 29.8 ± 7.0 months
old (range = 23–40). Two of these females actually bred with their fathers, one
of them twice, for a total of ten cubs (five males, five females). Cubs from one
of the litters survived to nine months old before we lost track of them. Cubs
from the other two litters survived to the end of the study, when they were eight
and ten months old. Similarly, six approximately known-age immigrant females
associated for the first time with adult males when they averaged 22.2 ± 7.6
months old (range = 15–36). Three of those associations resulted in cubs; the
females were about fifteen, seventeen, and twenty-five months old. These six
females conceived for the first time at the average age of 26.0 ± 11.1 months
(range = 15–42) and produced their first litters when they averaged 29.0 ± 11.1
months old (range = 18–45). Hence, it appears that adult females, whether they
were recruited into the population as philopatric progeny or as immigrants,
become reproductively active at about the same age.
Information on age of first breeding from other parts of the puma’s range is
scanty owing to the extreme difficulty in obtaining this information from obser-
vations of wild pumas. The youngest known-age, reproducing female puma that
has been reported in the literature was a Florida panther; she conceived at six-
teen months of age. Two other known-age Florida panthers conceived for the
first time at eighteen and twenty-one months of age (Maehr 1997b). Ross and
Jalkotzy (1992) had six known-age females in Alberta that had their first litters
at an average age of 30.0 months, similar to our findings. In Utah, Lindzey et
al. (1994) had six known-age females that first gave birth at an average age of
twenty-six months.
Because most of our radio-collared male progeny emigrated from the study
area if they survived to independence and thus went on to occupy ranges where
other radio-collared pumas were absent, it was practically impossible for us to
determine when they became sexually mature. So, we had to rely mostly on
information from approximately known-age males that immigrated onto the
San Andres Mountains. One known-age male and five approximately known-
age males averaged 24.3 ± 2.5 months old (range = 21–27) when they associated
with adult radio-collared females for the first time. Males were slightly older
than females when they first associated with the opposite sex [7]. At least two of
those associations resulted in live litters ninety-one and ninety-two days later.
These males were twenty-four and twenty-seven months old. We generalized
that male pumas on the San Andres Mountains reached sexual maturity at about
twenty-four months old.
We suspect that timing of puberty in male and female pumas is probably
influenced by physiological, social, and resource factors that relate to reproduc-
tive success. In our study area, where the puma population was rebuilding from
low densities imposed by population control, reduction of puma density left
ample space and resources for female recruits to establish home ranges where
prey resources could support them during growth, maintenance, pregnancy, lac-
tation, and cub rearing. As density increased, competition between adult female
pumas for food likely also increased. Thus, their individual reproductive success
would be enhanced if they began socializing with potential mates and breeding
as soon as they were physiologically capable (Bronson 1989). This was charac-
teristic of the San Andres Mountains puma population during a time of mule
deer (i.e., their principal prey) increase and decline (see Chapter 17). However,
a reasonable hypothesis is that after a lag period following prey collapse, new
female recruits may delay sexually maturity to older ages.
In contrast, adult male pumas competed directly with each other for mates
(see Chapter 8). Reproductively successful males probably benefited from larger
body size and superior strength, attributes that would enable them to achieve
dominance over other males through fighting for access to mates and territory.
Hence, it would be advantageous for males to become sexually mature when
they had the physical ability to compete. In many other polygynous species, in
which males compete directly with one another for access to mates, male matu-
ration is delayed (i.e., relative to females) until they have attained greater mass
(Stearns 1992). Our data on measured male puma masses supported this
hypothesis. Independent males twelve to twenty months old had lower masses
than males twenty-one to twenty-seven months old that were in the age bracket
when males first associated with adult females [8]. In the first group, five males
were captured in areas where they eventually established a territory, but four dis-
persed and three others were killed by territorial males. Males in the second
group (n = 11) had masses only marginally lower than territorial males twenty-
eight to ninety-six months old (n = 16) [9]. Males in the second group demon-
strated their competitive abilities; all of them were captured in the areas where
they established territories.
Parental Investment
Expenditure of energy and time by a parent in the care of its offspring in a way
that increases fitness is called parental investment (Freeman and Herron 1998).
In pumas, the end of the mating interlude is also the end of the sire’s direct
parental investment in his offspring. In essence, the male has contributed only
so much sperm in multiple copulations over a span of a few days. He takes no
part in rearing the cubs. Instead, the male explores his territory seeking other
mating opportunities. Yet, territorial males may contribute to survival of their
offspring indirectly by dissuading activity of other males that could kill cubs (see
Chapter 8). In contrast, the female’s direct investment in offspring is extremely
high. Her reproductive success is dependent on her physical ability to produce
mature eggs for fertilization, sustain pregnancy, and feed and protect young after
birth (Bronson 1989). In time alone, she will invest about three months in ges-
tation and eleven to sixteen months in rearing the cubs (see Chapter 9). A great
deal of energy is expended in nursing, grooming, hunting and killing prey,
teaching cubs how to hunt, and protecting them from weather, predators, and
other hazards in life. Greater parental investment by female pumas relative to
males is exhibited by over 90 percent of mammal species (Woodroffe and Vin-
cent 1994). It is vital to the reproductive success of individuals in each sex and
to the evolution of sex differences (Trivers 1985), which we expand upon in the
next section.
Reproductive Success
Darwinian fitness of individuals is directly exhibited by their ability to survive,
breed, and successfully raise offspring so that they may breed and reproduce
themselves; therefore, it is principal to the concept of evolutionary biology

(Grafen 1988, Freeman and Herron 1998). More fit individuals leave propor-
tionately more of their genes in descendants than do other individuals in the
population. Genotypes of the more fit express phenotypes that are better
adapted to the rigors of the environment. A measure of fitness for an individual
is lifetime reproductive success, which is ultimately measured by the number of
offspring produced in a lifetime that survive to breed (Thomas and Coulson
1988). Obtaining information necessary to quantify lifetime reproductive suc-
cess for wild pumas would require monitoring the reproductive output for indi-
viduals throughout their lives as well as the number of their offspring that reach
maturity. Such feats are practically impossible to attain because of the difficulty
and costs of tracking adult pumas that can live over ten years and progeny until
they reach puberty. The latter task is particularly problematic because a large
proportion of puma progeny disperse long distances from natal areas. The best
we could do in our study was to estimate short-term reproductive success for
individual pumas by quantifying the number of litters and cubs they produced
during the cross-section of their lives that we studied them. In addition, we
looked at traits that might have affected their reproductive success. These traits
shed light on our interpretation of the evolution of puma social organization.
Quantifying short-term reproductive success in adult females was more reli-
able than in males. Females produced litters of altricial cubs that we were able
to count and tag if they survived the first two to four weeks after birth. In con-
trast, we estimated reproductive success in males based on their breeding associ-
ations with females, which we determined from radiotelemetry. Because our
opportunities to assess reproductive success in males was dependent on our
radiotelemetry monitoring schedule, we no doubt missed some breeding associ-
ations that were less than one week in duration; hence, reproductive success as
we quantified it for males was a relative index. In other words, we assumed that
the males we observed to be the most engaged in breeding associations (con-
firmed by cub births) were the most reproductively successful males in the pop-
ulation. The most powerful method for identifying puma fathers is paternity
exclusion, a technique that uses genetic material from tissue samples (e.g.,
blood, muscle, skin, hair) obtained from animals in the population. With the
knowledge of genotypes of mothers, their offspring, and potential fathers, the
observed inheritance of alleles by offspring (one from each parent) at highly
polymorphic loci is the metric by which fathers are included or excluded as
potential mates (Murphy et al. 1998a). Murphy’s research on puma reproduc-
tive success in Yellowstone National Park during 1988–1995 employed both
paternity exclusion and radiotelemetry methods and showed that the two meth-
ods yielded the same estimates of paternity. Consequently, based on his findings
and the gestation periods resulting from our observations of consorting pairs, we
were confident in our paternity assignments based on radiotelemetry.
Females
Most adult female pumas on the San Andres Mountains were raising cubs each
year during 1988–1994. On the Treatment Area, an average of 52 ± 18 percent
(range = 29–75) of adult females gave birth each year. Counting mothers with
dependent cubs born in previous years, an average of 76 ± 21 percent (range =
44–100 percent) of adult females were raising cubs each year. Percentages were
similar on the Reference Area. On average, 48 ± 18 percent (range = 21–73) of
adult females gave birth to cubs each year, and an average of 74 ± 13 percent
(range = 60–100 percent) were rearing cubs. In other parts of North America,
large fractions of adult female pumas were rearing cubs each year. In the Big
Horn Mountains of Wyoming, 55 percent and 86 percent of adult females
raised cubs during two consecutive winters (Logan 1983). In southwestern
Alberta, during each January 1981–1989, 42 percent of adult females were
accompanied by cubs less than six months old (Ross and Jalkotzy 1992). And in
south-central Utah, cubs were born to about half of the resident female popula-
tion each year (Hemker 1982).
Nonreproducing adult female pumas comprised 26 percent (n = 14) of the
fifty-three adult females we monitored. Of those, seven died, three were
removed from the Treatment Area (Ruth et al. 1998), and four were being mon-
itored at the end of our study. The seven that died averaged 28 ± 4.4 months old
(range = 21–34); we monitored them continuously via radiotelemetry for two to
ten months. The three adult females we translocated averaged 29.7 ± 8.6
months old (range = 22–39). Radiotelemetry on two females was continuous for
eight and twenty-two months. The third female was translocated upon her first
capture as a new immigrant to our study area when she was twenty-two months
old. Clearly, all these females were just reaching reproductive ages (see “Puberty
and First Litters”) and might have reproduced had they survived or not been
translocated from the study area. In fact, one of the translocated females (F31)
produced a litter of five cubs in her new home range in northern New Mexico.
The four females that we were still monitoring at the end of our study could also
have produced cubs after our departure. They averaged 35 ± 8.2 months old
(range = 26–42); we monitored them continuously for three to seventeen
months. It is also possible that some of these females produced viable cubs that
did not live long enough (i.e., two to four weeks) for us to detect them. Hence,
except due to death, we did not document any adult female pumas that clearly
demonstrated apparent physiological barriers to reproduction in the expanding
puma population on the San Andres Mountains.
We quantified short-term re p ro d u c t i ve success in the thirty-nine puma

mothers we studied (74 percent of the total females). Mothers bore one to
f i ve litters and a total of one to fifteen cubs each. Ten mothers (26 perc e n t )
had three or more litters during the study and produced 110 of the 220 cubs
(50 percent) we documented and seventeen of twenty-four (71 percent) of
the tagged progeny that we re re c ruited in the adult portion of the puma pop-
ulation on the San Andres Mountains. Productivity of three females was
reduced because we translocated one and two died as a result of capture -
related injuries. The translocated female had produced three litters (eight
cubs, two recruits) on the San Andres Mountains. Of the two females with
reproduction abbreviated by us, one produced two litters (seven cubs, one
recruit), and the other produced one litter (three cubs, ze ro re c ruits). Si x t y
p e rcent (n = 46) of the litters were produced by prime age females (range =
37–94 months), 33 percent (n = 25) were produced by young females (range
= 18–36 months), and 7 percent (n = 5) were produced by old females (range
= 97–127 mo.)[10]. We did not know the ages of two females that pro d u c e d
t h ree litters.
Mothers that produced the most litters, and hence had the greatest repro-
ductive success, were those that lived the longest lives. We found a highly sig-
nificant relationship between the number of litters produced per adult female
and the number of months that we monitored females by radiotelemetry.
Female longevity explained about 79 percent of the variation in the number of
litters produced (Fig. 7-2) [11]. The average number of litters produced per
month by the twenty-eight adult females that we monitored for twelve or more
months (range = 17–82 months) was 0.05 ± 0.02. Four mothers that each pro-
duced five litters were 92–112 months old (i.e., 7.7–9.3 years) at the birth of
their last litters. On average, those mothers produced 2.2 ± 0.3 cubs per year
(range = 1.8–2.5) during their reproductive life span that we monitored by
radiotelemetry (i.e., from twenty-nine months old to their death or end of the
study). The oldest reproducing female was about 127 months old at the birth of
her last litter.
Nine (23 percent) mothers produced seventy-nine cubs in twenty-seven lit-
ters, out of which twelve first filial generation (F1) females produced sixty-one
cubs in twenty litters (i.e., second filial generation, F2). Consequently, the nine
P1 females were related to at least 64 percent (i.e., 140 of 220) of the progeny
that we documented on the San Andres Mountains. All but two of the mothers
were matrilineal. The two included a P1 mother and her daughter, who dis-
persed 49 km south of her natal area before establishing a home range where she
bred in the San Andres Mountains.
Nineteen (49 percent) mothers that we monitored by radiotelemetry formed
FIGURE 7-2. Scatter plot of the number of litters produced by adult female and male pumas
monitored for at least twelve months by radiotelemetry, San Andres Mountains, New
Mexico.
eight matrilines composed of mothers and their female progeny that established
home ranges which overlapped or abutted their mother’s home ranges (see
Chapter 13). Those females produced more litters (n = 43) and cubs (n = 127)
than twenty other females that were not matrilineal or for which we did not
know genetic relationships (ns = 36 litters, 93 cubs). Although there was no dif-
ference in the proportion of litters produced by the two groups, the number of
cubs produced by matrilineal mothers relative to non-matrilineal mothers plus
mothers of unknown status was greater than expected [12]. In this instance, our
estimates of matrilineal mothers and their reproductive output are conservative
because of the high probability that some females we caught early in the study
were mothers or siblings of some of their female neighbors (i.e., matrilineal).
More importantly, when we quantified survival rates of cubs to age of inde-
pendence, we found that cubs of matrilineal mothers had a higher survival rate,
0.67, than cubs of non-matrilineal mothers, 0.56 (see Chapter 8). Hence, matri-
lineal mothers would be expected to have greater reproductive success than non-
matrilineal mothers. Furthermore, eight matrilineal P1 mothers, one philopatric
F1 mother, and one adjacent F1 mother produced twenty-one of twenty-four (88
percent) of the tagged progeny that were recruited into the San Andres Moun-
tains puma population. Three P1 mothers and one other philopatric F1 mother
also produced four of thirteen (31 percent) of the known tagged progeny that
emigrated to other puma subpopulations in southern New Mexico. Clearly, like
Murphy et al. (1998a) discovered in pumas in the Northern Yellowstone Ecosys-
tem, reproductive success in female pumas was highly variable. Yet, according to
our findings, reproductive success favored matrilineal mothers.
Female philopatry has been documented in puma populations in Colorado
(Anderson et al. 1992), Alberta (Ross and Jalkotzy 1992), Utah (Laing and
Lindzey 1993), and Idaho (Lopez-Gonzalez 1999). The Utah biologists, who
also studied a protected puma population, recognized that philopatry resulted in
“clusters of related females with overlapping ranges” (Laing and Lindzey
1993:1057). While studying how deceased residents were replaced, they found
that seven out of ten deceased females were replaced by at least one of their inde-
pendent daughters or a daughter of a neighboring resident female. However,
they did not report the reproductive histories of those females. We extended this
concept by recognizing that philopatric females and adjacent females were also
closely related, and called the group a matriline. Furthermore, we realized that
matrilineal mothers have the greatest reproductive success and are probably
favored by natural selection. Those females probably have higher survival rates
than dispersers (see Chapter 8), and upon reaching the physiological ability to
reproduce, they raise cubs among familiar surroundings where they have already
learned the locations of vital resources, such as food and nurseries essential for
their cubs’ survival and physical development. A large proportion of their female
progeny may then be recruited into the local population, while other surviving
progeny, particularly males, disperse and may become recruited into other dis-
tant puma subpopulations or colonize available habitat (see Chapter 13). In
addition, matrilineal females are socially familiar with the other adult pumas
with whom they share the landscape; this may enhance the survival of their off-
spring by reducing infanticide (see Chapter 8). Female pumas do not aggres-
sively exclude other pumas from their home ranges (see Chapter 12). Such
behavior would lower their fitness by increasing risks of mortality or debilitat-
ing injuries. Hence, direct and indirect fitness (i.e., inclusive fitness) should be
enhanced for females that share resources with closely related females (see
Armitage 1986).
We hypothesize that matrilines are naturally characteristic of nonexploited
puma populations. We would expect philopatry and matrilines to have similar
fitness benefits in tigers, African leopards, snow leopards, and jaguars, but extant
information linking these traits to reproductive success in those large, solitary-
living felids is sparse (Smith 1993, Bailey 1993). If philopatry in other large felid
lineages is the result of homology (i.e., similarity between species that results
from inheritance of traits from a common ancestor), we propose that philopa-
try and resulting matrilines could have provided a basis for the evolution of the
pride in African lions (see Chapter 15).
Matrilines that we documented had overlapping generations and longitudi-
nal structure. The longevity of puma matrilines is unknown, but theoretically
they could span several generations in unexploited puma populations. In six
matrilines, parental-generation (P1) mothers raised cubs at the same time that
they were neighbors with as many as three philopatric daughters and one adja-
cent daughter (i.e., home ranges abutted), which were born in as many as three
different litters (see Plate 5). P1 mothers and one daughter raised cubs concur-
rently on ten occasions. Three P1 mothers raised cubs concurrently with one
daughter on two occasions each. Two P1 mothers raised cubs concurrently with
two of their daughters; two daughters were from the same litter and two were
from different litters. The same two sisters from different litters raised cubs con-
currently twice. In another instance, siblings from the same litter raised cubs
concurrently. We documented two other matrilines in which daughters lived
within their deceased mothers’ home ranges. Those daughters extended mater-
nal lines by reproducing; two philopatric daughters resulted from one of the lit-
ters. Here again, the presence of older, closely related puma mothers did not
trigger reproductive suppression in philopatric or neighboring female progeny
and siblings. If anything, reproductive success may have been enhanced.
Because these closely related groups of females also breed with territorial
males overlapping their home ranges, presence of long-tenured territorial males
may facilitate incest between fathers and daughters (see “Puberty and First Lit-
ters”). Two of the three father-daughter matings we recorded involved M88 and
F107, both of which occurred in the Treatment Area after we experimentally
removed 53 percent and 58 percent of adult and independent pumas (i.e., adults
plus subadults) (see Chapter 10). Yet, it is possible that we partially facilitated
inbreeding in that area by removing potential male competitors and mates of
M88. In an unmanipulated puma population, we would expect intermittent
turnover in territorial males and occasional breeding success of neighboring ter-
ritorial males to increase variance in an individual male’s breeding success. In

addition, most replacement males would likely immigrate from far away, even
from other subpopulations, (see Chapter 9) and consequently promote out-
breeding within local matrilines.
Phenotypic traits, including a kink at the end of the tail and a distinctive
whorl of hair (i.e., cowlick) on the back, are thought to derive from inbreeding
in Florida panthers (Barone et al. 1994, Maehr and Caddick 1995, Maehr
1997b). Panthers also exhibit low fertility and heart defects thought to result
from inbreeding depression (O’Brien et al. 1990, Barone et al. 1994, Roelke et
al. 1993b, Hedrick 1995). On the San Andres Mountains, we found only three
pumas with kinked tails (i.e., bends involving the last few vertebrae). Two were
adult males, M14 and M153. M14 lived in the Reference Area until he was
killed by M19. M153, who was born in the Reference Area, dispersed to the
Treatment Area, where he established a territory. Female F96 was born on the
Treatment Area. She was killed by adult male M88 when she was a twelve-
month-old subadult. One female cub, F63, apparently was born without a tail.
She lived in the Reference Area, but died in adolescence. None of the live pumas
we examined had a cowlick on the back.
Non-matrilineal mothers and mothers with unknown relatedness to neigh-
bors produced a relatively small fraction (22 percent) of the tagged progeny
recruits on the San Andres Mountains. However, they produced nine of thirteen
(69 percent) of the tagged emigrants we documented. Again, the proportion of
emigrants attributed to this female group is probably biased upward because of
the likelihood that some of the mothers were actually matrilineal but we did not
know it. If this is the case, then non-matrilineal and matrilineal mothers may
produce about equal proportions of emigrant offspring.
Males
Reproductive success in male pumas depends on their ability to maintain phys-
ical condition, repel competitors, acquire territory, and mate successfully with
estrus females. Such feats are not easy; males compete vigorously for access to
mates. Sometimes the competition is lethal. In contrast, reproduction in adult
females carries less variability of success. Estrus females merely have to advertise
their favorable breeding condition and males appear to provide the requisite
service. After fertilization, females can devote their energy beyond maintenance
to parental investment.
Of thirty-four adult males we monitored during the study, fifteen (44 per-
cent) were documented in breeding associations with females that resulted in
viable litters. Our ability to detect pairing between radio-collared adult males
and females was influenced by the proportion of radio-collared adults in the
population and our monitoring frequency, which was mostly dependent upon
our weekly flight schedule. Hence, we treated our quantification of breeding
pairs as an index to male reproductive success. All successfully breeding males
were monitored as adults via radiotelemetry for six to eighty-seven months (x– =
48.5 ± 25.4). Only three of the males were monitored for less than twenty-four
months (i.e., six, twenty-two, and twenty-three months). The fifteen males sired
ninety-four live cubs in thirty-one litters, comprising 43 percent of all the cubs
and 39 percent of all the litters we documented. In all cases, males bred with
females within their territories. Territories of these males overlapped one to thir-
teen adult female pumas per year (x– = 5.0 ± 2.6, n = 27 puma-years). Six adult
males (18 percent of adult males, 38 percent of documented breeders) had nine
F1 offspring (eight females, one male) that produced forty-five F2 progeny in fif-
teen litters. Two adult males mated with their philopatric daughters on three
occasions (i.e., three of thirty-six, 8 percent of consort pairs), resulting in ten
cubs. The six P1 males were related to at least 45 percent (100 of 220) of the
progeny we documented; they sired at least 54 percent (thirteen of twenty-four)
of the recruits comprising tagged progeny on the San Andres Mountains, and
they sired three of thirteen (23 percent) of the tagged emigrants we documented.
Reproductive success of adult males was linked to their longevity, just as we
found with females, albeit the relationship was weaker. Male longevity explained
only 45 percent of the variation in the number of litters they sired (Fig. 7-2)
[13]. Variation in male reproductive success is also influenced by the number of
females within a male’s territory (Murphy et al. 1998a) and his ability to com-
pete for estrus females. The average number of litters sired per month by nine-
teen adult males that were each monitored for twelve or more months was 0.03
± 0.03 (range = 0–0.11). The high variation in the number of litters sired per
month by individual males and the relatively small proportion of reproductive
success explained by longevity was consistent with the idea that male reproduc-
tive success should be more variable for males than for females in a species with
a polygynous mating system, which is exhibited by pumas (Seidensticker et al.
1973, see below), where male competition for mate access is intense (Clutton-
Brock et al. 1988, Le Boeuf and Reiter 1988, Packer et al. 1988; see Chapters 6
and 8). More importantly, this pattern was congruous with puma reproductive
success determined by genetic markers for the puma population in the North-
ern Yellowstone Ecosystem (Murphy et al. 1998a).
On four occasions (11 percent of consort pairs), more than one male associ-
ated with an estrus female (see Appendix 2). Male M36 associated with estrus
females F41 and F68 two to five days after other territorial males (M29 and
M38) had consorted with the females. Two other sires, M88 and M5, also asso-
ciated with estrus females after another territorial male did on one occasion each
(M3 and M38, respectively). In each of the four cases when estrus females were
courted by two territorial males during the same estrus period, the males had
overlapping territories. Female promiscuity and sexually induced ovulation cre-
ates the possibility that some puma litters may have more than one sire. So far,
this has not been unequivocally tested in wild pumas because only one study has
adequately estimated paternity using genetic markers. In his study of pumas in
the Northern Yellowstone Ecosystem, Murphy et al. (1998a:95) observed that
“radio-collared males singly accompanied females and only one father of each
litter was identified using genetic methods.” Multiple paternity of offspring in
the same litter has been found for other mammals, including California ground
squirrels (Boellstorff et al. 1994), black bears (Schenk and Kovacs 1995), and
even humans (Smith 1984).
Although male pumas were polygynous and both sexes exhibited promiscu-
ity, we documented some mate fidelity. Fidelity seemed to be partially depend-
ent upon the position of the female in male territories and male longevity. It
may have also been partially influenced by the degree of aggressiveness or dom-
inance of a territorial male over other males. Two of female F21’s five litters were
sired by male M3, and her last one was sired by M88. Two of F37’s three litters
were sired by M7. F47 produced five litters: three were sired by M88, one by
M3, and her last by M193. M88 also consorted with F47 during the same estrus
that she first mated with M3. But M3 had died of old age four months before
M193 sired F47’s last litter. Two of F107’s three litters were sired by her father,
M88. Other females had different mates. Two of F15’s five litters were sired by
M14 and M19; M19 killed M14 and usurped his territory, which contained
F15. Two of F45’s five litters were sired by M52 and M151. Two of F41’s three
litters were sired by M38 and M46. F149’s two litters were sired by M5 and M3.
F27 had two nonviable litters, one sired by M20 and the other by M53. In all
cases where female infidelity was demonstrated with contemporary territorial
males, the male territories overlapped the female home ranges. Pumas in the
Northern Yellowstone Ecosystem exhibited strong mate fidelity (Murphy et al.
1998a).
In the protected puma population on the San Andres Mountains, prime
and old-age territorial male pumas we re engaged in 90 percent of the know n
breeding associations with females that resulted in viable litters. Prime-age
males thirt y - s e ven to eighty-eight months old we re in 71 percent (n = 22) of
associations. Old males 103–142 months old we re in 19 percent (n = 6) of
breeding associations. But young males twenty-two to thirty-five months old
we re in only 10 percent (n = 3) of them [14]. In heavily exploited puma pop-
ulations, we would expect the age distribution of breeding males to shift tow a rd
younger ages.
Male M88 sired the most litters (n = 6) even though we disrupted his terri-
torial tenure for 166 days by translocating him from the Treatment Area to
northern New Mexico (Ruth et al. 1998). He sired his first litter at twenty-two
months of age before we intervened. Thus, he might have sired even more lit-
ters as a young adult if we had not translocated him. After finding his way back
to his original territory, an excursion that covered 494 airline km, he sired the
other five litters when he was forty-five to sixty-seven months old. M88 was still
the dominant male in his area through the end of the study and could have pro-
duced more litters thereafter. Male M3 was the next-most productive male with
four litters that he sired when he was a prime to old male (i.e., 61–142 months).
He died of old age in his territory only twenty-four days after he sired his last
litter. Three other adult males (M7, M38, M46) sired three litters each. M7 and
M46 sired all their litters when they were prime age (M7 when he was fifty to
seventy-seven months old and M46 when he was thirty-five to eighty-six
months old). M38 sired his litters when he was prime to old age (45–103
months). Of these three males, M7 was translocated; thus, we eliminated his
further reproduction on our study area. M38 and M46 were still territorial at
the end of our study and could have produced more litters. Two males (M88 and
M5) also associated with estrus females after they had consorted with other
males (see Appendix 2). The males were about 36 and 135 months old, respec-
tively. During our study, seven male pumas with the greatest reproductive suc-
cess (i.e., had sired two or more litters) were those that maintained territorial
longevity for an average of 88.6 ± 30.5 months (i.e., 7.4 years; range = 41–128
months). And three of those were still vigorous territorial males at the end of the
study. Six males that sired one or more viable litters and died of natural causes
(i.e., three were killed by other territorial males, two died of complications of
aging, and one died of disease) during our research had maintained territorial
tenure for an average of 71.7 ± 43.4 months (i.e., six years; range = 21–128
months). In contrast, male pumas in moderately hunted populations may rarely
achieve long territorial longevity. For example, in a population in north-central
Wyoming, the oldest males were about seventy-two months old (Logan et al.
1986). Assuming that they acquired their territories at the age of twenty-four
months, their longevity was about forty-eight months.
The critical nature of longevity in a breeding range was perhaps best demon-
strated by M88 and M49, both territorial males that we translocated 490 and
465 airline km, respectively, to northern New Mexico (Ruth et al. 1998). M88
returned to his original territory 166 days later and resumed breeding. M49
returned to his original territory 469 days later, killed a mule deer, then associ-
ated with his previous territorial neighbor, M3. We believe this encounter may
have contributed to M49’s death about twelve days later. Both M88 and M49
had endured months of uncertainty and travail as they navigated their way back
to the areas where they had previously established social dominance and where
they were familiar with vital resources. Two other adult males that we translo-
cated also homed in the direction of the San Andres Mountains, but they died
in between (Ruth et al. 1998). These adult male pumas exhibited strong fidelity
to established territories, perhaps indicating that investing time and energy to
establish dominance in an area increases fitness, both in opportunities to breed
and in survival of offspring. Apparently it takes death or expulsion by a more
dominant male to move an adult male from his established territory (see Chap-
ter 12). It is likely that environmental conditions, such as severe restrictions in
food or mates or increased access to mates, may also prompt an adult male to
shift to a different area to improve his fitness, but empirical evidence for this is
so far lacking.
We did not document any breeding behavior involving males without terri-
tories. And in central Idaho, all breeding was done by resident adult males (Sei-
densticker et al. 1973). Similarly, all radio-collared sires in the Northern Yel-
lowstone Ecosystem were residents; however, biologists there suggested that
some untagged sires identified through genetic paternity analysis may have been
“roaming nonresidents that sired litters during temporary visits to the study
area” (Murphy et al. 1998a:90). If non-territorial male pumas (e.g., dispersers)
are involved in breeding, we expect the frequency to be relatively rare in pro-
tected populations such as ours. Conversely, we would expect such incidences to
increase in puma populations subjected to chronic severe exploitation where
male longevity and territorial structure is disrupted by humans killing male
pumas.
We observed three other adult males in breeding associations with females,
but the pairings did not result in viable litters. As previously mentioned, male
M36 paired with estrus females twice after the females had already consorted
with other males. Hence, we did not consider him to be the sire of those litters,
although he could have been. M36 was about forty-six and ninety-four months
old during those associations. Two other males, M20 and M53, consorted with
F27 at the north end of our study area and sired nonviable litters. M20 sired
F27’s first litter that was stillborn, and M53 sired the litter that F27 aborted just
prior to her death (see Appendix 2). Both males were in prime ages (seventy-four
and forty months old, respectively) when they sired the litters. M20 was moni-
tored for forty-nine months; he spent about 90 percent of his time on the
Oscura Mountains to the north of our study area. M53 was monitored as an
adult for sixteen months; he spent about 66 percent of his time also on the
Oscura Mountains. Consequently, both males could have sired other litters in
the Oscura Mountains, where we did not radio-collar pumas.
Sixteen other adult male pumas we re not documented as sires of viable lit-
ters on the San Andres Mountains. Most of the lack of documentation for
re p ro d u c t i ve success for these males was due to re l a t i vely short monitoring
periods and territorial tenure on our study area. In addition, our radioteleme-
try monitoring schedule apparently missed paternal connections for some lit-
ters. Nevertheless, we documented seven of these males in potential breeding
associations that did not result in viable litters. The number of times those indi-
vidual males associated with adult females ranged from one to six (x– = 2.1 ±
1.9, median = 1). They associated with one to three different females during
their tenure. The oldest male for which we did not document any associations
with females on our study area was M1. He was about 145 months old when
he died; but we monitored him for only forty-nine months. M1 spent about 40
p e rcent of his time on the Organ Mountains to the south of our study are a ,
where other pumas we re not monitored by radiotelemetry. M1 could have sired
litters there. M1’s time on our study area was reduced as a direct result of new
immigrant male M22, who usurped the part of M1’s territory there. M22 sired
one litter during his twenty-two-month tenure before he was killed by neigh-
boring territorial male M3.
The remaining fifteen males were monitored as adults for durations ranging
from one to fifty-one months (x– = 12.9 ± 12.0), considerably shorter than males
that were documented sires [15]. The seven males that we documented in poten-
tial breeding associations with females were monitored for six to fifty-one
months (x– = 17.7 ± 15.6). We inferred that all of these males had lower repro-
ductive success than documented sires because of their relatively short tenure.
Four of these males (M18, M23, M49, M114) were removed from the Treat-
ment Area; hence, we directly affected their reproductive success. Two other
males died. M73 was killed by another territorial male when he was 45.5
months old, and M192 died from capture-related injuries when he was about
thirty-eight months old. Of these fifteen males, six were young (twenty-four to
thirty-six months), six were young to prime (twenty-four to fifty-five months),
and three were prime (thirty-seven to ninety-six months) ages during the time
we monitored them. This relatively young age distribution tended to reflect low
observed reproductive success. But, nine of the males were still present on the
study area at the close of our research; thus they may have gone on to sire litters.
Only male M73 seemed physiologically incapable of reproducing. When we
initially marked him as a twenty-five-day-old cub we sexed him as a female
based on external genitalia. But when we recaptured him as a 27.5-month-old
adult, he was obviously a male. M73’s testicles had not descended into his scro-
tum; he was a bilateral cryptorchid. His body conformation reflected a lack of
testosterone in his blood stream; he was more feminine, like a large sleek adult
female than a robust male. And he probably lacked the aggressivity of normal
adult male pumas (see Bronson 1989). M73 weighed only 45.4 kg, at least 10
kg lighter than male pumas normally weigh at that age (see Appendix 1). More-
over, during the two years we monitored him as an adult, his home range did
not overlap any adult females. He was not competing with other territorial
males for access to estrus females. M73’s case was the only time we observed
cryptorchidism in adult male pumas on the San Andres Mountains (one of 33).
This 3.0 percent frequency of cryptorchidism in our population was similar to
the 3.9 percent that Barone et al. (1994) found in fifty-one male pumas sam-
pled from Texas, Colorado, Latin America, and captive stock of uncertain
genetic origin. However, those animals were unilateral cryptorchids (one testicle
not descended). Comparatively, endangered Florida panthers appear to have an
extraordinarily high frequency of unilateral cryptorchids, 43.8 percent, which
may be the result of inbreeding depression (Barone et al. 1994).
We inferred that reproductive success and hence fitness was greatest for ter-
ritorial males with the greatest longevity, a conclusion also reached for male
pumas in the Northern Yellowstone Ecosystem (Murphy et al. 1998a), African
lions (Packer et al. 1988), and tigers in Nepal (Smith and McDougal 1991). It
seems logical that traits that allow males to successfully compete for mates also
help them to live longer lives, particularly if their greatest cause of death is
intraspecies strife (see Chapter 8). We expect the largest, strongest, and most
aggressive males to be the fittest to compete with other males and thus maintain
long-standing dominance. This confers greater breeding opportunities with
adult females residing within their territories (see Weckerly 1998). Male com-
petition for mates is intense because low population density of adult females and
their long birth intervals actually make estrus females rare. Even though female
pumas are distributed at low densities in topographically and vegetatively com-
plex habitats, individual estrus females can be defended sequentially during the
puma’s year-round breeding season. This male-to-male competition for mates,
which results in variable reproductive success and traits like sexual dimorphism,
is known as sexual selection (Trivers 1985). Hence, sexual selection for traits,
such as greater mass, strength, aggressivity, and other related traits that enhance
survival and reproductive success, should improve the quality of genes passed by
males to their offspring (Trivers 1985).
Large body mass probably has been a strongly selected trait in male pumas
because of the advantages it awards for overpowering competitors during terri-
torial or mating challenges. In North America, adult male pumas, on average,
outweigh females by 40 percent (see Chapter 2). Intense competition between
males for mates has also been cited as a strong selective force for sexual dimor-
phism in other large mammals with polygynous mating systems, including large
primates (Mitani et al. 1996), red deer (Clutton-Brock et al. 1988), and ele-
phant seals (Le Boeuf and Reiter 1988, Hailey et al. 1994, Modig 1996). But if
bigness is so advantageous to male pumas, why aren’t they normally two or three
times larger than females? We suspect a trade-off favoring an optimal body size
is involved. Extremely large males would require more energy for growth and
maintenance. Consequently they would need to hunt and kill more frequently,
which would diminish the time and energy they could allot to searching for and
defending mates, engaging in copulation marathons to impregnate estrus
females, and defending territory. Hence, extremely large size would lose out to
optimally scaled males. Female pumas, on the other hand, have evolved an
energy-efficient size that allows them greater parental investment to raise more
offspring successfully. Part of that efficiency is the ability to kill large prey to feed
rapidly growing offspring. Next, we examine how rigors of puma life affect
pumas’ abilities to survive and breed.
LP > > KX
1. Nursling litter size averaged about three cubs. Weaned litter size averaged
about two cubs. Average litter size of nurslings was among the highest
reported for pumas. The natality rate (i.e., average litter size / birth interval)
averaged 0.17 and was also among the highest for puma. Fecundity rate
(i.e., average number of live cubs / adult female / year) in the Treatment
Area and the Reference area during the deer increase phase was 1.6 and 1.4,
respectively; during the deer decline phase, it was 2.0 and 1.3, respectively.
Fecundity was not obviously sensitive to the declining prey base. Sizes of
mothers’ first litters and subsequent litters did not differ, nor did litter sizes
of first-time mothers and old mothers.
2. Puma births occurred year-round but primarily during July–September,
coinciding with the mule deer fawning season. We hypothesize that cubs
born during a period of food abundance have the greatest chance of sur-
vival.
3. The puma gestation period averaged about ninety-two days. Birth dates
minus gestation periods put the corresponding breeding period during
April–June. We estimated estrus periods lasted one to ten days. Females
exhibited promiscuity. We estimated an average estrous cycle length of
about twenty-four days. Birth intervals on the San Andres Mountains aver-
aged 17.4 months, among the shortest reported for pumas. Mothers that
lost all their cubs successfully bred again an average of about one hundred
days later.
4. Puberty was reached at about twenty-one months in female pumas and
about twenty-four months in males. Females conceived successfully for the

first time at an average age of about twenty-six months and produced their
first litters at the average age of twenty-nine months.
5. Philopatric females did not exhibit reproductive suppression. In the grow-
ing San Andres Mountains puma population, philopatric female progeny
and female immigrants became reproductively active at about the same age.
6. Other than the actual breeding activity, puma sires have no other direct
parental investment. In contrast, mothers expend fourteen to nineteen
months in time and tremendous energy feeding and caring for the cubs.
7. The number of adult females giving birth to cubs each year in the Treat-
ment Area averaged 52 percent; an average of 76 percent were raising cubs
each year. In the Reference Area, these amounts averaged 48 percent and 74
percent, respectively.
8. Reproductive success was variable for females but greatest for mothers that
lived the longest lives. Mothers and recruited philopatric or neighboring
female progeny normally formed matrilines. Females in matrilines had the
highest reproductive success; hence, philopatry should be adaptive. Repro-
ductive success for males was even more variable, and again it was greatest
for fathers that lived the longest lives. Males with the greatest reproductive
success maintained territorial longevity that averaged over seven years. Vari-
ation in male reproductive success was also affected by the number of adult
females in his territory and his ability to compete with other males for estrus
females. Sires bred with females within their territories, and mates occa-
sionally exhibited fidelity. Adult males competed directly for territory and
mates. We propose that in pumas, sexual selection resulted in sexual dimor-
phism and male aggressiveness.
LN N5LN5)L
1. Test for difference in litter size in the Treatment Area, pre- and post-treat-
ment. Two-sample t-test, t = –1.7635, 15 d.f., P = 0.10.
2. Puma annual fecundity rates in mule deer population increase phase
(1989–1990 to 1991–1992) and decline phase (1992–1993 to 1993–1994).
Treatment Area: increase phase = 1.4, 1.0, 2.3, respectively; decline phase =
2.0, 2.0, respectively. Re f e rence Area: increase phase = 1.6, 1.2, 1.4, respec-
tively; decline phase = 1.6, 0.9, respectively.
3. Natality rates (mean litter size / mean birth interval in months): British Colum-
bia = 3.1 / 18.3; Utah = 2.4 / 24.3; Alberta = 2.2 / 19.7; Florida = 2.3 / 23.3.
4. Test for difference in sizes of first litters in eight matrilineal mothers and six
immigrant mothers. Two-sample t-test, t = 1.341, 12 d.f., P = 0.21.
5. Test for difference in sizes of litters of six old mothers and first litters of
eight matrilineal mothers. Two-sample t-test, t = 1.177, 12 d.f., P = 0.26.
6. Test for the occurrence of puma litters during July–September relative to
what is expected in those three months (19.5) and October–June (58.5).
Chi-square test, c2 = 10.68, 1 d.f., P = 0.0012.
7. Test for differences in ages of seven known-age female pumas and one
known-age and five approximately known-age male pumas at first associa-
tion with the opposite sex. Mann-Whitney U test, U = 8.5, P = 0.17.
8. Test for differences in mass (kg) in twelve independent male pumas eleven
to twenty months old (x– = 15.3 ± 3.2 months) that averaged 39.5 ± 8.4 kg,
90 percent CI = 35.1–43.8, median = 39.9, and eleven adult male pumas
twenty-one to twenty-seven months old (x– = 24.1 ± 2.0 months) that aver-
aged 53.1 ± 4.3 kg, 90 percent CI = 50.8–55.5, median = 52.2. Mann-
Whitney U test, U = 124, P = 0.0003.
9. Test for difference in mass (kg) in eleven adult male pumas twenty-one to
twenty-seven months old (see [8] above) and sixteen adult male pumas
twenty-eight to ninety-six months old (x– = 44.7 ± 17.8 months) that aver-
aged 56.4 ± 5.4 kg, 90 percent CI = 54.1–58.8, median = 56.4. Mann-
Whitney U test, U = 53.5, P = 0.09.
10. Number, percentage, and mean ages of mothers in young (eighteen to
thirty-six months), prime (thirty-seven to ninety-six months), and old
(97–156 months) age classes. Young: twenty-five, 33 percent, 26.9 ± 5.2;
prime: forty-six, 60 percent, 60.6 ± 17.4; old: five, 7 percent, 109.2 ± 11.5.
11. Linear regression of the number of litters produced per adult female puma
(monitored by radiotelemetry for twelve months or more each, n = 28) on
the number of months that each female was monitored (as an index to
longevity): Number of litters = –0.3733 + 0.0651 (number of months mon-
itored), P < 0.0001, r2 = 0.79.
12. Test for differences in number of litters and cubs produced by matrilineal
mothers and non-matrilineal mothers with unknown status mothers. Num-
ber of litters produced by matrilineal mothers versus non-matrilineal moth-
ers and mothers of unknown status was not different than an expected even
distribution. Chi-square test, c2 = 1.268, 1 d.f., P > 0.10. Number of cubs
produced by matrilineal mothers was greater than expected in an even dis-
tribution: c2 = 7.278, 1 d.f., P = 0.007.
13. Linear regression of the number of litters sired per adult male puma (mon-
itored by radiotelemetry for twelve months or more each, n = 19) on the
number of months that each male was monitored (as an index to longevity):
Number of litters sired = –0.4111 + 0.0427 (number of months moni-
tored), P = 0.002, r2 = 0.45.
14. Number, percentage, range, and mean ± SD ages of territorial male pumas
involved in thirty-one successful matings with female pumas: Young
(twenty-four to thirty-six months) – n = 3, 10 percent, range = 22–35
months, x– = 28.0 ± 6.6; Prime (thirty-seven to ninety-six months) – n = 22,
71 percent, range = 37–88 months, x– = 60.7 ± 14.5; Old (97–156 months)
– n = 6, 19 percent, range = 103–142 months, x– = 118 ± 17.3.
15. Test for difference in monitoring duration in fifteen sires and fifteen non-
sires. Mann-Whitney U test, U = 208.5, P < 0.0001.
Chapter 8
Mortality and Survival
Human-Caused Mortality
As we discussed previously, some puma deaths were caused by our own research
activities (see Chapter 4). Four adult pumas, three females (F27, F145, F87) and
one male (M192), died due to injuries they sustained while captured in our
foothold snares (F27 on 9 April 1990, F145 on 5 February 1992, F87 on 15
February 1993, M192 on 14 February 1994). The three females were successful
mothers ranging in age from sixty to sixty-nine months; they represented 5 per-
cent of all the adult females we documented on the San Andres Mountains dur-
ing our study. When F87 died, she was raising about four newborns that we
never got to see. At the time of F145’s death, she was raising three fourteen-
month-old cubs; at least two (both females) survived to adulthood. The adult
male represented 3 percent of all adult males we documented; he was about
thirty-eight months old. Nine cubs also died as a result of our research efforts,
representing 4 percent of all cubs we observed on the San Andres Mountains.
They ranged in age from newborn to six months. One was a male, two were
females; we did not handle the remaining six to determine their sex.
Even presumably inviolate wild lands like the San Andres Mountains are not
totally immune from poachers. They had a good excuse to be nearby; about 93
km2 of our study area on the northwest edge lay outside of White Sands Missile
Range boundaries and was subjected to legal sport-hunting and furbearer trap-
ping. Outfitters that came to hunt that tiny strip of land, even knowing that all
tagged pumas were legally off-limits, hoped to catch a rare untagged puma and
115
undoubtedly knew their dogs could end up within our study area where all
pumas were protected. Even if they did catch a tagged cat, at worst, they would
experience an exhilarating hunt. An unmarked female, about twenty-four
months old (we inspected the skull and skin) was killed by a puma hunter on
10 March 1988. We assumed this female had recently immigrated onto the
study area, perhaps coming from mountains to the west. Even though the out-
fitter took us to the very tree where the puma was shot, on Whites Sands Mis-
sile Range, neither he nor the hunter was prosecuted by authorities. Twice after
that, as we were locating radio-collared pumas from an airplane, we circled this
outfitter’s hunting party at puma locations. We figured his dogs had the cats
bayed in trees. In the second instance, he was clearly on White Sands Missile
Range. We even ground-tracked the hunting party and caught one of the dogs,
complete with a tag identifying the outfitter. According to the outfitter, the con-
tinual presence of our research team in the mountains finally discouraged him
and other puma hunters from hunting there for the duration of our study.
Before the crash in fur prices in the early 1990s, a few trappers worked the
northwest foothills of our study area. A trapper, no doubt trying for coyotes and
bobcats, caught radio-collared female F9 when she was thirty-one months old.
F9 escaped the steel foothold trap, but not before it ripped off two toes from her
left forefoot, exposing the remaining phalanges. She died, apparently from sep-
ticemia, on 14 January 1989. We knew F9 since she was a nursling. As an adult,
she had consorted with at least one male, but she did not produce cubs before
her death.
We also knew radio-collared female F181 since she was a nursling. On 12
November 1994 a mule-deer hunter snapped her spine with a .25 caliber bullet.
We found her prostrate body on the ground where she expired, clawing at the
grass. She was twenty-six months old and had not yet bred. During necropsy, we
found she had been living quite well with an abdominal hernia that we had ini-
tially diagnosed when we recaptured her at age 12.5 months. Apparently, this
condition was congenital because her sister (F183) also had an abdominal her-
nia, but their mother F68 did not. There was copious fat on her kidneys, peri-
cardium, mesenteries, and subcutaneous areas, signifying that she was well nour-
ished. These three females killed by humans represented another 5 percent of
the adult females we documented on our study area.
All but one of these human-caused deaths, that of a cub, occurred in the Ref-
erence Area. Deaths of the adult pumas occurred sporadically. One death
occurred in each of the years 1988, 1989, 1990, 1992, and 1993, and two adults
died in 1994. In Chapter 10, we address the potential effects that those losses
had on the dynamics of the puma population in the Reference Area.
CHAPTER 8. MORTALITY AND SURVIVAL 117
Natural Mortality
Natural mortality factors were independent of direct human influences. And
because we studied a protected puma population, most documented puma
deaths were from natural causes. By examining and quantifying natural puma
deaths, we were able to better understand how natural selection forces impacted
the puma population and individual reproductive success.
Cubs
We estimated cub mortality rates by quantifying changes in litter sizes over time.
We accomplished this by sneaking up on families with radio-tagged members to
count cubs and by finding track sets of radio-tagged families and noting attri-
tion in the number of cub tracks. Radio-tagged cubs provided us with direct
mortality data, but we relied on intensive ground tracking and luck to help us
find non-collared cubs that died. Most of the latter simply vanished from fam-
ily groups, with no clue of their demise. In a few instances, entire litters of suck-
ling cubs vanished before we even had a chance to examine them. We estimated
mortality and survival rates using data only from cubs that we initially recorded
as nurslings and that were born before January 1994; this way, the cubs could
have reached the extreme age of independence (i.e., sixteen months) before the
end of our study in March 1995.
Cub surv i val rate in the Treatment Area was slightly lower than in the Refer-
ence Area. Out of sixty-nine cubs born on the Treatment Area, about twenty-eight
died, twenty-seven from natural causes and one from research-related activities.
This produced a surv i val rate of 0.59 (i.e., mortality rate = 0.41). In the Re f e rence
Area, about thirty-four out of ninety-three cubs died (thirty from natural causes,
four from research-related causes), producing a surv i val rate of 0.63. When we
considered only natural deaths, surv i val rates increased slightly to 0.60 in the
Treatment Area and to 0.66 in the Reference Area. By combining cubs (n = 157)
subjected to natural death (n = 57) in the Treatment Area and Re f e rence Area, the
survival rate was 0.64. From this subset of cubs, we also estimated surv i val rates of
ninety-seven cubs we knew we re born to matrilineal mothers and fifty-seven cubs
born to mothers that we were fairly certain were not matrilineal mothers. Cubs of
matrilineal mothers had a higher rate of surv i val to independence, 0.67, than cubs
of non-matrilineal mothers, 0.56. Although the probability of surv i val of cubs in
each maternal category would be expected to be equal at P = 0.20 [1], we believe
a tendency for higher surv i val rates of cubs born to matrilineal mothers is biolog-
ically vital. Matrilineal mothers would tend to have greater fitness, and philopatry
as a trait, at least in females, would be strongly favored.
We further estimated cub survival rates in the Treatment and Reference Areas
during the mule deer increase (1989–1990 to 1991–1992) and decline
(1992–1993 to 1993–1994) phases while considering only cubs starting as
nurslings and subjected to natural causes of death. In the Treatment Area, cub
survival rate during the deer increase phase was 0.64 (i.e., twenty-three of thirty-
six), but it declined to 0.57 (i.e., thirteen of twenty-three) during the deer
decline phase. The Reference Area exhibited an upward change from 0.66 (i.e.,
thirty-one of forty-seven) in the deer increase phase to 0.72 (i.e., twenty-one of
twenty-nine) in the deer decline phase. Overall, cub survival did not appear to
be sensitive to the deer population dynamics during our research. We believe it
may take a sustained low deer population to trigger a steep decline in cub sur-
vival on the San Andres Mountains. These observations further support the
expectation for a time delay before the puma population declines in response to
a reduced prey base.
We combined cubs from the Treatment Area and Reference Area to develop
a survival curve from birth to thirteen months, approximately the age at which
cubs became independent of their mothers (see Chapter 9). Most of the mor-
tality (65 percent) occurred when cubs were less than or equal to three months
old (Fig. 8-1). This age span includes the time that cubs are suckling and con-
fined to nurseries, usually from six to eight weeks old, and the first month that
mothers are moving their cubs away from nurseries to prey kills that they made
to feed the now carnivorous cubs. Cubs are highly vulnerable during both of
these periods. Nurseries are the focal point of activity of mothers as they sortie
away to hunt and return, usually on a daily basis, to nurse and care for the cubs.
This high-frequency activity can work to the disadvantage of cubs since other
predators, including male pumas, detect visual, auditory, and olfactory cues that
can lead them to nurseries. Unless the small, helpless cubs can hide out of reach
of teeth and claws, they can easily become prey. As cubs leave the familiar sur-
roundings of the nursery to make their way to their mother’s kills, they still are
highly vulnerable to predation because of their small size and inexperience; and
they are vulnerable to accidents that may befall them while traversing rugged
terrain. Furthermore, their relatively immature immune systems make them vul-
nerable to diseases (Saunders 1970, Coe Clough and Roth 1998). Sex-related
survival rates (calculated by excluding cubs of unknown sex that were born and
disappeared) generated similar survival curves for males and females. But female
cubs seemed to be more vulnerable at older ages and had only a slightly lower
survival rate than males (Fig. 8-1).
Few other data sets on puma cub survival rates were available from other
parts of North America for us to compare. In an unhunted puma population
with a very low density of 0.5 pumas per 100 km2 in south-central Utah, sur-
FIGURE 8-1. Puma cub survival rates from birth to thirteen months, San Andres Mountains,
New Mexico.
vival of eighteen cubs monitored from contact (i.e., two to ten months old) to
dispersal had a survival rate of 0.67 (Hemker et al. 1986). Survival rates from
birth to independence were probably lower than this because survival of cubs
less than two months old was not estimated. In a human-dominated landscape
in Southern California that effectively isolated and fragmented puma habitat, an
annual survival rate for nineteen cubs was 0.52. But biologists in that study
pointed out that they normally counted cubs “several weeks” after birth of lit-
ters and that other cubs probably died; hence, they suggested that the actual sur-
vival rate was probably 0.45–0.52 (Beier and Barrett 1993). Puma cubs in nat-
urally fragmented habitat in southern Idaho had similarly low survival rates
(Lopez-Gonzalez 1999). There, survival for about sixty cubs studied from one
month old to dispersal was 0.42. The survival rate for males was higher, 0.58,
than for females, 0.45. In Alberta, forty-two cubs monitored from first contact
to independence had an extremely high survival rate of 0.98. But the biologists
cautioned that actual mortality levels were likely underestimated because some
cubs were not detected until they were at least 0.6 year old, and more postna-
tal mortality probably occurred at younger ages (Ross and Jalkotzy 1992). Sur-
vival rates in Florida panther cubs are likewise tenuous but are more serious for
this endangered population (see Maehr 1996: Appendix C). The differential
between mean litter sizes for four litters less than fourteen days old and twenty-
one litters four to twelve months old suggested a first-year survival rate of 0.84
(Maehr and Caddick 1995). However, this method underestimates mortality
because it excludes entire litters that might have been lost. Another estimate of
0.87 was based on thirteen survivors out of fifteen radio-collared panther cubs
“older than 4 months but less than 12 months” (Maehr and Caddick
1995:1295). Again, this underestimates cub mortality because probable losses of
cubs less than four months old were excluded. If Florida panther cubs had the
same distribution of deaths from birth to independence as we observed (Fig. 8-
1), then it is possible that only 26 percent of the total mortality occurred in pan-
ther cubs five to twelve months old. If these proportions are reasonable, we
could expect as much as 74 percent of the mortality to occur in cubs up to four
months old. Thus, about 60 percent of panther cubs may have actually survived
to independence.
Su rv i val curves from our study and Lopez-Gonzalez (1999) suggest a pos-
sible pattern of higher surv i val rates for male cubs than female cubs. If this
characteristic is real, higher surv i val in males may be due to greater maternal
i n vestment in male offspring. As mentioned previously, male cubs are gener-
ally larger than female cubs; furt h e r m o re, surv i val in the sexes begins to
d i verge in weaned cubs four or more months old in southern Idaho (Lopez-
Gonzalez 1999) and eight or more months old in our study. Larger male cubs
may dominate choice feeding sites at nipples and carcasses, avoid predation
better, and more efficiently traverse rugged terrain, all of which could give
them surv i val advantages over female siblings during the period of parental
investment.
Although thirty-five cubs disappeared from unknown reasons on our study
area, we were able to assign cause of death to twenty-seven other cubs (Fig.
8-2). Infanticide and cannibalism was committed only by male pumas; they
killed twelve cubs in six litters (44 percent). In all cases, plundering males appar-
ently were not sires of the cubs. We know that in one case, the sire was another
male. Ages of puma-killed cubs ranged from 1.1 to 5.0 months old (x– = 2.1 ±
1.1). Eleven of the cubs were eaten by male pumas, including two entire litters
with three and four cubs that were sixty-six and thirty-four days old, respec-
tively. In the former case, M88 consumed the nurslings of his daughter F107,
who was about 1.8 km away from the cubs at the time. In the latter case, M52
ate the nurslings of F45 while she was 1.3 km away feeding on a mule deer kill.
A 3.5-month-old cub was not eaten; she died as a result of canines penetrating
the braincase. In an instance where two five-month-old siblings were killed, the
mother (F61) was also killed by the male puma, probably as she futilely tried to
defend her cubs. Starvation apparently claimed ten cubs (37 percent). Starvation
of five cubs in two litters that were about one week and five months old, respec-
FPO @ 80%
FIGURE 8-2. Causes of death in puma cubs, San Andres Mountains, New Mexico. The
numbers represent the number of cubs that died in each category of mortality (death).
tively, occurred because their mothers (F15 and F86) were killed by male pumas.
Three 36-day-old nurslings starved eight days before their mother (F4) died of
an undiagnosed disease. Two five-month-old cubs probably starved not long
after their mother (F195) died from injuries sustained while trying to kill a deer.
We never found evidence of the cubs again. Disease that caused diarrhea and
dehydration killed three cubs in one litter when they were around fifty-seven
days old (11 percent). A fall from a cliff killed one 59-day-old cub (4 percent),
and coyotes killed one six-month-old cub (4 percent).
Two female cubs that were orphaned at ages 7.5 and 9.8 months survived.
The younger puma (F68) and her male sibling (M67) scavenged their mother’s
carcass after she died from undetermined causes. This provided them nourish-
ment that probably improved the cubs’ chances of survival. If M67 survived, he
apparently emigrated from our study area; we never found evidence of him
again. But F68 became a matrilineal mother by establishing her adult home
range on her natal area. Later, two of her own female offspring (F181 and F183)
exhibited philopatry after independence. The older orphaned female (F10) used
an area adjacent to her mother’s (F2) home range, and mostly in the Organ
Mountains south of our study area. But she was killed by a male puma (M22)
when she was eighteen months old, the same male that apparently killed her
mother.
Slide @318%
PHOTO 18. Orphaned puma F10 at ten months of age and nine days after her mother, F2,
died.
Subadults
We estimated survival rates for twenty-five radio-collared subadult pumas that
we monitored after independence from mothers and until they became adults or
died. Their ages ranged from eight to twenty-seven months. Nine males were
monitored for 1,797 days and sixteen females were monitored for 2,590 days.
Subadults males had a finite survival rate of 0.56, which was significantly lower
than the female finite survival rate of 0.88 [2]. Four males and two females died
when they were 14.3–18.0 months old (x– = 15.7 ± 1.5). All of them were killed
by male pumas. They all died as a result of bites that penetrated the brain case,
the frontal region of the skull, or the cervical vertebrae. Only one individual was
fed upon slightly, suggesting to us that most of these cats probably were regarded
as competitors.
Two male subadults were killed on their natal areas in the Treatment Area,
both by the same male. Male M64 was independent at twelve months old.
When he was 14.5 months old, he encountered a newly arrived immigrant,
twenty-four-month-old M88, at the cache of a yearling mule deer buck that
M64 had killed four days before. M88 was also the puma that apparently killed
M64’s mother (F60) sixteen days prior, also in an incident apparently involving
direct competition for food (see “Adults”). When we captured M88 three days
after M64’s demise, he still had wounds from battles with mother and son. His
head and neck were swollen from claw punctures and scratches, and he had both
fresh and mending claw scratches on his face and head. There were also claw
scratches on his left foreleg, shoulder, and hip. M64 still had hair of his foe stuck
to his foreclaws, but he had taken mortal punishment. Canine punctures to the
braincase were fatal. He also had massive bites to his forearms, triceps, shoul-
ders, and lumbar region. A canine punctured the right ileum. Claws had pene-
trated skin on his chest. M64 had been making a living well on his own. He had
moderate amounts of subcutaneous fat along the midline of his abdominal and
inguinal regions; there were dense fat deposits on his mesenteries, kidneys, and
pericardium. But he was clearly subordinate to the older and heavier M88 (56.7
kg vs. 41.7 kg).
Twenty months later, M88 again fought over another mule deer kill. This
time he encountered his own 14.3-month-old son M108, who had been inde-
pendent for only 1.7 months. Either M108 was investigating M88’s kill, con-
sidering the potential for a free meal, and triggered M88’s aggression, or M88
sensed M108’s kill and proceeded to usurp it by killing its inexperienced
defender. Canine punctures to the braincase were fatal. M88 did not get away
wholly unscathed; tufts of puma hair were scattered about, and on the battle-
ground we found M88’s eartag, which had been ripped from his pinna. In this
case too, M108 was no match for his experienced, larger father (63 kg vs. 39 kg).
In the encounters with M64 and M108, M88 was certainly directly com-
peting for food. In addition, since both subadults were killed in the core of his
territory, M88 may have also been exerting his dominance over these males in
the area where he acquired both food and mates. M88 had all the tactical advan-
tages to win these contests. He was older and more experienced, and he far out-
weighed the subadults 41 to 63 percent [3].
The other two males were killed after they dispersed from their natal areas.
M140 was 15.6 months old when his braincase was crushed and his spinal cord
severed between the atlas and axis by another male puma. He had been inde-
pendent for only nineteen days and in the interim had dispersed 42.2 km north
of his natal area, going from the Treatment Area to the Reference Area. His
death, too, may have been associated with food; M140’s stomach was full with
remains of a newborn calf oryx. We suspected, but could not prove, that M140
was probably killed by new immigrant male M192, who was about twenty-seven
months old at the time M140 died. Prior to his demise, M140 was healthy and
robust, weighing 50.8 kg. He had copious subcutaneous fat along the midline
of his thorax and abdomen all the way to the inguinal region, on his shoulders,
along the midline of his back, on his flanks, and along the entire length of his
tail. There were extensive fat deposits on his mesenteries and kidneys.
Subadult M122 was independent at 14.5 months old and almost immedi-
ately emigrated from the San Andres Mountains. During the next eighty-two
days he dispersed 134 km northeast of his natal area by traversing the compar-
atively desolate Tularosa Basin and onto El Capitan Mountain. When he was
seventeen months old, he encountered a male puma on this new terrain. In an
ensuing fight, M122 received canine punctures that penetrated the frontal area
of the skull, damaged the right orbit, and broke the right zygomatic arch.
Although we found tracks of the dominant male puma, we could not find any
evidence of a prey kill in the vicinity of the death site, and M122’s stomach was
empty. This direct competition may not have been over food but instead may
have been about dominance in that area. As a long-distance disperser, M122 was
apparently finding enough food; he weighed 49.9 kg at death and had subcuta-
neous fat along his back and abdomen.
As mentioned previously, subadult female F10 was orphaned at 9.8 months
of age, when her mother was killed by a male puma (probably new immigrant
M22). F10 was killed by M22 at the age of eighteen months as a result of a bite
that crushed the frontal region of her skull. She died 16.3 km southwest of her
natal area at the southern end of the Treatment Area. The male did not consume
F10 but bit chunks of skin from her back and left shoulder. F10’s stomach was
empty, and there was no evidence of a prey kill in the vicinity. However, F10 was
apparently at the onset of puberty. Her uterus was engorged with blood vessels,
suggesting that she was in proestrus or estrus. Vocalizations by F10 during either
period could have summoned the attention of M22 (see Mehrer 1975). Multi-
ple bites to the back of F10’s neck causing subcutaneous hemorrhaging sug-
gested that M22 attempted to mount her. Males of the larger species of the Fel-
idae usually use the neck bite as a vestigial behavior at the climax of copulation.
Canines are clearly lethal weapons, and presumably the delayed neck bite
evolved to minimize the danger of the mating act accidentally turning into an
attack. Selection would have favored males who did not harm their mates. In
pumas, the neck bite is variable (Ewer 1973) and may not be used at all during
mating (Mehrer 1975). We wondered if M22 may have used the neck bite far
too aggressively, triggering a defensive counterassault by the inexperienced F10,
which escalated to M22’s fatal attack.
The other subadult female, F96, was independent at about twelve months of
age. Her activities centered in her natal area in the Treatment Area until she met
her demise, at the age of 14.5 months, in the jaws of the notorious M88. This
was the third puma he had killed since his arrival on the San Andres Mountains
ten months previously. As in the prior two instances, the pumas directly com-
peted for a mule deer kill. Judging from F96’s stomach full of deer tissues, she
had fed on the deer at least once. M88 had crushed F96’s braincase and clawed
her back, stripping skin down the right side from the shoulders to the hips. He
consumed a small amount of muscle tissue along the spine. In both instances
that subadult females were killed by territorial males, the battles were probably
no contest. The males were 68–117 percent greater in mass and probably easily
overwhelmed the females [4].
Male M198 was born in the Treatment Area and dispersed north into the
Reference Area. His decomposed body lay in the bottom of a canyon 84 km
north of his natal area. He may have been dispersing or decided to try to estab-
lish a territory in the area when he died at about twenty-four months of age.
Damage to the left orbit of his skull indicated an injury that could have occurred
in a fight with another puma or by accident. Even his eye may have been avulsed
or destroyed. If the injury itself was not fatal, then infection may have killed
him.
An untagged twenty-month-old male puma had just immigrated to the
south end of the Treatment Area, perhaps from the Organ Mountains, when he
encountered territorial male M1. We followed tracks on the ground with both
pumas running or trotting where M1 chased the young male out of a rugged
canyon, and about 2 km across grass-covered flats dotted with bushy mesquites,
prickly pear, and ephedra. Finally, the subadult futilely sought to escape his
nemesis by climbing an electrical utility pole. It was either that or make a stand
on the desert floor; no boulder piles or crevasses were nearby to offer safety.
There was only one set of claw marks ascending the utility pole. Upon topping
the pole, the cat’s front feet and face completed the circuit, and he was electro-
cuted. The body fell to the ground, apparently at the feet of M1, who walked
around it, then traveled back to the canyon where the episode began. M1, about
115 months old, was certainly the most experienced and the most physically
dominant, weighing at least 27 percent more than the young male [5]. The
young male’s stomach was full of deer tissues, so it is possible that this aggressive
encounter was the result of direct competition for food.
We also found decomposed remains of two female pumas that died inde-
pendently of each other in the Reference Area when they were about twelve and
nine months old. Causes of death were unknown, but their skeletons did not
indicate trauma. No ear tags were found at the sites, but if present, they could
have been carried away by scavengers such as pack rats, foxes, or ravens. We had
noticed the disappearance of ear tags from carcasses of other marked pumas after
such scavenger activity.
Pumas endure a harsh life from birth to adulthood. As cubs, they must sur-
vive disease and accidents, and must avoid predation, including being eaten by
their own kind. It appears that cubs born to matrilineal mothers may have a
greater chance of survival than cubs born to mothers that are not members of a
matriline. If male cubs have a slightly better chance of survival than female sib-
lings, the scale changes during the stage of independence. As subadults, pumas
must be skilled enough to kill prey to sustain their rapidly growing bodies, and
if they disperse, must tolerate long-distance moves, sometimes across inhos-
pitable habitat, before they establish a new home. Most of all, they must avoid
encountering territorial male pumas, clearly their greatest threat as cubs and
subadults in populations unexploited by humans. This source of death provides
strong selective pressure for female pumas who avoid males outside of mating,
and for offspring to disperse from natal areas where a territorial male (usually
their father) resides. But dispersal has other costs, particularly for subadult
males, because practically all of them disperse long distances from natal areas
(see Chapter 9) and explore terrain with unpredictable resources and risks,
including other tenant male pumas determined to exert their dominance. We
suspect that survival rates of female subadults would have been lower than we
observed if females had the same propensity for long-distance dispersal as males.
In our rebuilding study population, only one of the sixteen radio-collared female
subadults dispersed clear of her natal area. All the other females either were
philopatric (n = 11) or dispersed only a short distance and established their
home ranges adjacent to their natal areas (n = 4). Considering that female dis-
persal may be partially density dependent (see Chapters 10 and 13), greater
subadult female mortality may occur at high densities. Male and female
subadult surv i val rates may be more equivalent in those conditions. If
philopatric females tend to have higher survival rates than dispersers, and cubs
of matrilineal mothers are favored, we would expect philopatry to be the favored
female strategy.
Quantitative data on survival rates of independent subadults in other puma
populations is practically nonexistent, primarily because most pumas monitored
in this age class dispersed from study areas and biologists lost contact with them.
Some information comes from Colorado’s Uncompahgre Plateau where Ander-
son et al. (1992) estimated an annual survival rate of 0.64 for pumas twelve to
twenty-four months old. He used data from twenty male and twenty-two female
pumas and combined the sexes to estimate the rate using the product limit (i.e.,
Kaplan-Meier) procedure (Pollock et al. 1989). Although this rate falls between
our estimated rates for subadult males and females from the San Andres Moun-
tains, a direct comparison is tenuous because the Colorado study did not indi-
cate the social status of the monitored pumas. Younger-aged pumas could have
been dependent cubs or independent subadults, whereas older-aged pumas may
have been dispersers or resident adults.
Life for a puma born on the San Andres Mountains was very risky, even
though the population was protected from human exploitation. The chance of
a male surviving to the age of adulthood was about 36 percent (i.e., 100 x 0.64
x 0.56) and that of a female with philopatric tendencies was about 56 percent
(100 x 0.64 x 0.88). If a female emigrated from the San Andres Mountains, we
would expect her chance of surviving to adulthood to be about the same as for
a male, 36 percent. These different survival rates partially explain the tendency
for fewer adult males than adult females in puma populations. Lower survival
rates for males further contribute to greater variability in reproductive success in
comparison to females, because a greater proportion of males do not even sur-
vive to have a chance to breed. As adults, life was still harsh, but the scales of
survival tipped again in favor of males.
Adults
Our information of survival and mortality in adults came from thirty-four male
and fifty-one female radio-collared pumas. They ranged in age from 18 to 152
months old. We used Micromort, a survival analysis software package (Heisey
and Fuller 1985a,b). This software takes input data on the number of days that
radio-collared pumas were monitored in an interval (in this case a year starting
with 1 January), the number of pumas that died of specific causes in the inter-
val, and then computes the survival rates and agent-specific mortality rates for
each interval. By using this technique, we were able to compare rates between
male and female adult pumas each year during the eight-year span from 1987 to
1994 (Table 8-1).
We considered the possibility that survival rates for adult pumas may differ
in the Treatment Area and the Reference Area, perhaps related to the population
flux caused by our experimental removal of pumas. But annual survival rates of
adult males living in the Treatment Area and Reference Area differed in only one
of the eight years (1989) [6]. Similarly, for adult females in the Treatment Area
and Reference Area, annual survival rates differed in two of the eight years
(1991, 1993) [7]. Because of the overriding tendency for comparable survival
rates of adults in the Treatment Area and Reference Area, we combined adult
pumas in each sex from both areas to examine survival rates for adults on the
San Andres Mountains as a whole. Consequently, we were able to analyze sur-
vival and agent-specific mortality rates for adults during 1987–1994 when we
monitored nine to twenty radio-collared males and seven to twenty-four radio-
collared females each year (Table 8-1). Annual survival rates for males and
females differed during the first four years (1987–1990), but not during the last
four years (1991–1994) [8] and in general favored males (five of the eight years).
Annual survival rates for males averaged 0.91 ± 0.08, and for females averaged
0.82 ± 0.14. These data (Table 8-1) did not suggest a declining trend in adult
survival rate during the mule deer decline that began in the 1992–1993 biolog-
Table 8-1. Annual survival rates for adult pumas, 1987–1994, San
Andres Mountains, New Mexico.a
Year Males Females
Puma-days Rate 95% C.L. Puma-days Rate 95% C.L.
1987b 2685 0.87 0.67–1.0 1689 0.52 0.25–1.0
1988b 3675 1.0 1.0–1.0 2735 0.76 0.53–1.0
1989b 4305 0.84 0.67–1.0 4672 1.0 1.0–1.0
1990b 4381 1.0 1.0–1.0 6179 0.79 0.63–1.0
1991 3340 0.90 0.72–1.0 5296 0.87 0.72–1.0
1992 4184 1.0 1.0–1.0 6353 0.94 0.84–1.0
1993 4656 0.79 0.61–1.0 6057 0.84 0.68–1.0
1994 4909 0.86 0.70–1.0 5199 0.87 0.72–1.0
aSurvival rates and 95 percent C.L. (Confidence Limits) were computed using Micromort
software (Heisey and Fuller 1985a,b).

bSurvival rates of males and females were significantly different (all P 0.06).
ical year, particularly in females, who affect population growth the most (see
Chapter 10).
Survival rates for adult pumas in other parts of their range are limited. Biol-
ogists studying a low-density puma population in southern Utah estimated sur-
vival rates for three to eight resident adult radio-collared pumas per year during
1980–1986 (Lindzey et al. 1988). Like us, they used Micromort software. They
defined “resident adults” as pumas “ 16 months of age that demonstrated con-
tinuous and predictable use of an area for 6 months” (Lindzey et al. 1988:665).
Annual survival rates for females averaged 0.71 and ranged from 0.45 to 1.0.
Because only one male could be monitored in some years, the biologists pooled
males and females to estimate adult annual survival rates with a mean of 0.72
and a range of 0.52–1.0. For a puma population in fragmented habitat in South-
ern California, biologists used the product-limit procedure (Pollock et al. 1989)
for twenty adult pumas (Beier and Barrett 1993). They pooled males and
females because of small numbers and estimated an adult annual survival rate of
0.75. In Colorado, biologists used that same method for twenty males and
twenty-two females, and estimated annual survival rates of 0.69, 0.92, and 0.80
for pumas in age classes twenty-four to thirty-six months, thirty-six to forty-
eight months, and forty-eight to sixty months, respectively (Anderson et al.
1992). In a puma population in southeastern Arizona subjected to extremely
heavy predator control and sport-hunting, biologists used Micromort software to
estimate survival rates for a small sample of four males and two to five females
each year over a three-year period. Annual survival rates ranged from 0.12 to 1.0
for males and from 0 to 0.81 for females. The biologists pointed out that female
survival rates were higher than males except during one year when only two
females were sampled. The higher male mortality rate was associated with a
greater number of males killed for depredation control (i.e., sixteen males versus
six females) (Cunningham et al. 1995).
Using Micromort software, we computed agent-specific mortality rates for
radio-collared adult pumas during the eight-year span 1987–1994. Results
showed that intraspecies strife (i.e., pumas killing other pumas) was the chief
agent of death for adult male and female pumas (mortality rates were 0.39 and
0.39, respectively). Accidents, chiefly during attempts to kill prey but including
one snakebite incident, was the next most important agent of death for females
(0.20). But males did not die from similar accidents. Disease was a greater agent
of death in females (0.16) than males (0.08). As would be expected, males were
more likely to die of old-age-related physiological problems (0.08) than females
(0.01). Although we could categorize all male deaths, we could not determine
agents of death for two females (0.04).
During our entire study from 1985 to 1995, we examined the remains of
thirty adult pumas, including eleven males and nineteen females, to try to deter-
mine causes of death (Table 8-2). On average, males died at older ages (90.0 ±
38.7 months) than did females (53.3 ± 32.6 months)[9]. Females that actually
produced litters during their lives (n = 13) died at the average age of 64.8 ± 33.7
months. Intraspecies strife was the greatest cause of mortality, comprising 46
percent of male deaths and 53 percent of female deaths. Ages of adults killed by
other pumas were highly variable (Table 8-2). Males killed fourteen of the
pumas; the sex of a puma that killed one adult female was unknown.
Through radio-tracking, then retracing pumas’ movements from tracks, as
well as through examining wounds on dead pumas and their perpetrators we
caught soon after battles, we were able to assess circumstances in which pumas
kill other pumas. Adult females were killed while defending their cubs, while
competing directly for food, and as prey themselves. Two females died appar-
ently while trying to defend their cubs. F2 (eighty-four months old) was prob-
ably killed by M22 (twenty-two months old), a new immigrant male to the
Treatment Area. F2 may have been protecting her three ten-month-old cubs at
the time of her encounter. M22 inflicted fatal bites that crushed F2’s braincase
and broke cervical vertebrae. At least two of the cubs survived the attack; female
F10 established a home range adjacent to her natal area (and later was killed by
M22), but the fates of her two brothers were unknown. F61 (twenty-five
months old) was probably trying to defend her two five-month-old cubs when
Table 8-2. Number, percentage, and ages of adult pumas that died of natural
causes on the San Andres Mountains, New Mexico, 1985–1995.
Males Females
Cause of death No. % Age (mo.) x– ± SD No. % Age (mo.) x– ± SD
Strife 5 46 80.2 ± 41.5 10 53 50.5 ± 33.6
Disease 3 27 74.7 ± 9.3 2 11 78.0 ± 15.6
Accidents 0 0 3 16 45.0 ± 22.5
Old age 2 18 147.5 ± 6.4 1 5 122.0
Unknown 1 9 81.0 2 11 30.0 ± 5.7
Snakebite 0 0 1 5 34.0
Total deaths 11 19
she was killed, probably by a new immigrant male puma (possibly M29, twenty-
eight months old) in the Reference Area. Canine punctures to her braincase
were fatal. The male had killed and almost entirely consumed at least one of the
cubs. The male did not feed on F61. Multiple scats on scrapes made by the male
puma in the vicinity instead suggested he probably ate both cubs.
Two other females were killed while competing directly with males for food.
F60 (forty-three months old) and a new immigrant male to the Treatment Area,
M88 (twenty months old), fought fiercely, probably over a mule deer killed by
F60. Massive multiple bites and claw punctures to F60’s legs and back eventu-
ally killed her. F15 (105 months old) was killed by a male puma in the Refer-
ence Area where she had killed a buck mule deer. At the time of her death, F15
had three suckling cubs; the nursery was only about 0.6 km away. F15 died from
canine punctures to the braincase. After her death, the male puma consumed the
remains of the deer. We suspect the male was M29 (fifty months old) who had
inflicted mortal wounds to M19 only three months prior in an area where their
territories overlapped.
Four adult females, and possibly a fifth, were clearly victims of puma preda-
tion. F40 (twenty-five months old) was a new immigrant to the Reference Area
when she was killed and eaten by resident male M19 (fifty-four months old).
M19 killed F40 by crushing her braincase, then proceeded to consume about 90
percent of her mass, leaving only her head, distal parts of the legs and tail, the
stomach, and part of the intestines. M19 covered F40’s remains with grass twice
between feedings, just as pumas normally do when feeding on ungulate prey.
F86 (thirty-nine months old) was killed and eaten by a male puma in the Ref-
erence Area. Apparently the male encountered F86 and her two five-month-old
Slide @318%
PHOTO 19. Puma F15’s skull, showing fatal bite wounds to the braincase and old damage to
the zygomatic arch (arrow).
cubs at their cached mule deer fawn. Tracks in the area indicated that the cubs
survived the attack, probably because F86 defended them. She succumbed to
canine punctures to the braincase. The male consumed about 90 percent of her
mass, leaving only the head, feet, tail, and a segment of the intestines. This male
too had covered F86’s remains with grass between feedings. We did not expect
the cubs to survive for long; inexperienced, and weighing only 11–14 kg each,
they probably would not be able to provision themselves with food sufficiently.
After a while, evidence of them disappeared. In another act of predation, F57
(104 months old) was killed and eaten by a male puma in the Reference Area.
We suspected M46 (60 months old), who was in the process of extending his
territory there. About 95 percent of F57’s mass was consumed; all that remained
were her feet, a strip of skin, a 2-m length of intestine, and bone fragments.
F185 (twenty-nine months old) was killed and eaten by resident male M29
(sixty-six months old). Canine punctures to the braincase were fatal. About 95
percent of F185’s mass was consumed. Only the anterior portion of the skull,
the feet, tail, stomach, pieces of skin, and tufts of hair remained. Again, the male
had covered the puma’s remains between meals. We noted that F185 was in
M29’s territory for the first time when she was killed. Another female may have
been the intended victim of predation in the Treatment Area. New immigrant
F39 (twenty-two months old) was apparently driven off a 6-m-high limestone
cliff to her death by advances of M22 (now thirty-five months old). We found
tracks of both cats on top of the cliff, but we could not fathom why F39 jumped
or fell. She died of massive head trauma. Her small undeveloped uterus sug-
gested she was not in proestrus or estrus, and competition for prey did not
appear to be involved.
In most instances where adult female pumas were killed by adult male
pumas, the relationships between the individuals were new. In particular, the
pumas did not have any previous breeding associations with each other. Both of
the females (F61, F86) that died defending their cubs were apparently killed by
immigrant males that eventually established territories there. Similarly, both
females (F60, F15) that died in food competition fought males with whom they
were unfamiliar. In five of the instances where females were preyed upon or pos-
sibly pursued as prey, the females were either new immigrants to areas occupied
by territorial males (F39, F40, F86, F185), or a territorial male was expanding
his territory into the home range of the female (F57). These females had little
chance of surviving violent attacks of males that were 60–150 percent greater in
mass [10]. Not surprisingly, we never found an adult male killed by a female
puma. These incidents stress the jeopardy of adult females in puma society, the
strong selective pressure for females that actively avoid male pumas, and that
local social transitions from immigration or territorial shifts can also have deadly
consequences.
Male pumas were killed by other males in disputes over territory or breeding
females. M22 (forty-five months old) was killed by M3 (eighty-one months old)
in the Treatment Area. At the time, M3 had been consorting with the apparently
estrus adult female F47 for four days in an area where the territories of both
males overlapped. M22 died of severe trauma caused by massive bite wounds
that penetrated sacral vertebrae and the right ilium, broke the radius of the left
foreleg and the fibula of the right rear leg, and punctured and tore muscles and
ligaments in those areas. In this fight, there was no injury to M22’s skull. Ear-
lier in life, he may have learned the hard way the virtue of guarding his head.
After cleaning his skull, we found old mended wounds, some with hair imbed-
ded, including two punctures to the right frontal, one puncture at the junction
of the left frontal, nasal, and maxilla, and one puncture on the posterior edge of
the left frontal—all wounds that were probably caused by canines of another
puma. Quite likely, when M22 immigrated onto the San Andres Mountains, he
vigorously defended his intentions to establish his territory there. Perhaps he
fought M1, the old male that he apparently displaced from the area, or his
neighbor M3 (see Chapter 12).
After M22’s demise, M1 started using that portion of his former territory
Slide @318%
PHOTO 20. Puma M22’s skull, showing mended canine puncture wounds.
again. But this time he localized his movements to a relatively tiny area of about
20 km2, which also overlapped M3’s territory. His small range probably was due
to a combination of his advanced age and an apparent infection of coccidioid-
mycosis that left numerous granulomatous lesions on his limb bones (discovered
after he died) and may have infected other vital organs (Adaska 1999). When he
was about 144 months old, M1 fought M3 (eighty-eight months old) and suf-
fered puncture wounds to the frontal region of the skull that also caused dam-
age to inner surfaces of both orbits. He died a few days after the battle, recum-
bent in the shade of juniper trees beside a mountain spring. He apparently
succumbed from infection that caused necrosis of the damaged skull bones.
M14 (seventy-one months old) was killed by M19 (forty months old) in the
Reference Area. The males fought in an overlap zone of their territories. M14
died at the edge of the churned-up battle ground from fatal bite wounds that
punctured the frontal of his skull and caused severe trauma and bleeding in both
forelegs. M19 did not get away unscathed; he received multiple claw scratches
to his face, head, shoulders, and thorax, and he had canine punctures to his left
foreleg above the elbow. But he lived to usurp all of M14’s former territory.
M19 (ninety-six months old) was killed by M29 (forty-eight months old),
apparently over breeding female F15 (see Chapter 14). He probably died from
infection that attacked his body via canine punctures to both frontals and the
left maxilla and nasal. We found his body just beyond the north edge of his ter-
ritory, but not far from where he had been spending time in an overlap zone
with M29’s territory (see Fig. 14-3). M29 also suffered, but much less severely,
with multiple claw scratches on his face and forelegs and canine punctures to
both elbows.
M73 never was to leave a legacy. He was an unusually small, feminine-look-
ing adult male, traits that were probably affected by his bilateral cryptorchidism.
When he was forty-five months old, he was killed by another male puma at the
southern edge of his home range. There were a few suspects, M52 (seventy-eight
months old), M161 (sixty months old), or new immigrant M219 (thirty
months old), whose territories M73’s overlapped. M73 died at the edge of the
battle ground from canine punctures to the braincase. Other bites penetrated
the frontals, avulsed the left eye, and pierced the right zygomatic arch.
In four of these five cases of male-to-male contests, victorious pumas were
10–30 percent greater in mass than the losers [11], supporting the hypothesis
that larger males have greater fitness. Only M1 (64.4 kg) was larger than M3
(60.3 kg), but he was disadvantaged by old age and a fungal disease that proba-
bly affected his physical prowess. Two of the losing males, M1 and M19, were
past their prime. Of the three in their prime, one (M73) was physically out-
matched and incapable of reproducing.
As puma deaths from intraspecies strife began to accumulate in our study, we
hypothesized that their frequency would be related to puma population density.
That is, as population density increased, then direct competition among adult
and subadult pumas also would increase as they vied for essential resources such
as food, mates, and territory. We predicted that evidence for this hypothesis
would be a strong monotonic association between the annual frequency that
independent pumas (i.e., adults plus subadults) were killed by other pumas and
year-round adult puma density. In the Treatment Area (January–December,
1988–1994) the frequency of deaths was associated with adult male density but
not with total or female adult density [12]. In the Re f e rence Are a
(January–December, 1989–1994), there was no association between deaths and
any of the three adult density categories [13] (Fig. 8-3). We rejected the hypoth-
esis that the frequency of intraspecies strife is affected by adult puma density. In
addition, intraspecies strife occurred during years of mule deer population
increase and decline. We recorded seventeen adult and subadult puma deaths on
the San Andres Mountains from 1987 to January 1992 when the deer popula-
tion increased (see Chapter 17) and three such deaths during 1993–1994 when
the deer population declined. The actual frequency with which intraspecies
killing in pumas occurs is probably largely influenced by chance events such as
FIGURE 8-3. Relationship of adult puma density to frequency of independent puma deaths
from intraspecies strife in the Treatment Area (TA) and Reference Area (RA), San Andres
Mountains, New Mexico.
estrus periods of females, timing of immigration of male competitors, contests

between neighboring competing males, and opportunistic competition for prey
carcasses. All this suggests that intraspecies strife should be expected in any
puma population, regardless of population density and prey trends. And since
this pattern held for both our Treatment Area and our Reference Area, deaths
from intraspecies strife should be expected to occur in exploited and nonex-
ploited populations.
Pumas killing other pumas was clearly the dominant mortality factor in the
adult portion of the population, just as it was with the cubs and subadults.
Infanticide, cannibalism, and intraspecies strife in pumas is not unique to the
San Andres Mountains. In fact, these modes of death have been documented
across the puma’s North American range, including Florida (Maehr 1990a),
Texas (McBride 1976), Colorado (Anderson et al. 1992), Idaho (Hornocker
1970), Montana (Williams 1992), Alberta (Ross and Jalkotzy 1992), British
Columbia (Spreadbury et al. 1996), California (Beier and Barrett 1993), Ari-
zona (Cunningham et al. 1995), and Utah (Hemker 1982). Although the toll of
these deaths on puma populations has not been quantified in most cases,
intraspecies strife was identified as the single-greatest natural cause of death of
Florida panthers (Maehr 1997a) and of pumas in California (Beier and Barrett
1993) and Utah (Lindzey et al. 1988). Furthermore, fighting has occurred in
puma populations with a wide range of adult puma densities among them, such
as in Alberta (1.5–2.2 / 100 km2, Ross and Jalkotzy 1992), British Columbia
(0.93–1.1 / 100 km2, Spreadbury et al. 1996), Wyoming (1.4–1.5 / 100 km2,
Logan et al. 1986), and Utah (0.3–0.6 / 100 km2, Lindzey et al. 1988).
Disease was the second most important mortality factor in adult pumas on
our study area. The pathogen could not be determined in four of five deaths,
primarily because of tissue autolysis. Dry, hot desert conditions made it imper-
ative that tissues from dead animals be collected and preserved in less than
twenty-four hours to be useful in diagnosis. Septicemic plague (Yersinia pestis)
was diagnosed in F68 (sixty-seven months old). We found her only about twelve
hours after she died beside a spring-fed pool in which she was observed lying the
day before. The septicemic form is a direct result of the bacteria Y. pestis multi-
plying rapidly in the victim’s blood and quickly producing blood poisoning. An
animal with septicemic plague can pass from a healthy state to collapse and
death within a few hours (Rail 1985). Besides this occurrence of Y. pestis in a
puma on our study area, the disease was also diagnosed in two female pumas
found dead within 100 m of each other in the Sandia Mountains in central New
Mexico in 1994 (New Mexico Department of Agriculture Veterinary Diagnos-
tic Services, Albuquerque). It has also been diagnosed in pumas in California
(W. Boyce, DVM, and B. Chomel, DVM, University of California, Davis, per-
sonal communication). The disease is not endemic to the Western Hemisphere;
apparently it was introduced in about 1899 by Chinese immigrants traveling
from Hong Kong to San Francisco, California (Gregg 1985). By the middle
1930s, plague had spread from California eastward across the Rocky Mountains
and has been periodically active in fifteen of the seventeen contiguous Western
states (Thorne et al. 1982).
Effects of Y. pestis on puma populations is presently unknown, but there is a
real probability that significant morbidity and mortality may occur in localized
areas, especially in New Mexico, which has the highest incidence of the disease
in North America (Rail 1985). Y. pestis is especially pathogenic in felids (Thorne
et al. 1982). The disease may become increasingly infectious in pumas during
low populations of their principal ungulate prey, when smaller prey items, such
as rodents and rabbits, become more important in the diet. In these conditions,
pumas may be more susceptible to infection by ingesting plague-infected ani-
mals or from bites from disease-carrying fleas on those hosts (Rail 1985). Thus,
it may be significant that the two females from central New Mexico and F68
died in 1994 and 1995 at the height of a drought that triggered a mule deer
population crash on our study area and elsewhere in the state. Occurrence of the
disease in deer apparently is rare. Biologists handling plague-afflicted animals
should be aware that they can contract the disease through the bite of an
infected flea or through direct contact with an infected carcass (Thorne et al.
1982). We suspected, but could not confirm, that Y. pestis was also involved in
the death of M29 (sixty-seven months old). He too may have been trying to cool
a fever in a water trough. But we examined him six or seven days after death;
warm spring temperatures had already taken their toll on tissue autolysis. M29
died exactly two years earlier than F68 and 7 km away. His territory included
F68 and he even sired a litter with her. We wondered if that area of the San
Andres Mountains could have been a focus for the disease.
Although we got to F4 (eighty-nine months old) only twelve to twenty-four
hours after death, a veterinary diagnostic laboratory could not isolate a
pathogen. F4 had suffered severe wasting. She lost about 34 percent of her nor-
mal body mass. Her condition, rendering her mammary glands nonproductive,
no doubt caused her to abandon her three nursling cubs two weeks before she
died.
Killing large prey is a dangerous way for pumas to make a living. Three acci-
dental deaths to adult females occurred apparently during botched attempts to
bring down mule deer. All three appeared to be in excellent physical condition
prior to death. Two of these, F13 (thirty-one months old) and F128 (seventy-
one months old), suffered massive body blows that severely bruised the chest
and thorax and caused the lungs to bleed. F13 also had three broken ribs that
punctured the right lung. We suspect that these pumas may have been kicked,
slammed against rocks or trees, or pinned beneath falling prey. F195’s (thirty-
three months old) injury was extraordinary. Apparently, in a defensive counter-
attack, a mule-deer buck’s antler impaled F195’s left eye and penetrated the left
side of her braincase.
Sometimes pumas survive broken bones. In adults that died from a variety
of causes, we found two females (F128, F195) and three males (M1, M3, M53)
that each had one to four ribs that had been broken earlier in life but had since
mended. Male pumas on our study area rarely suffer fatal injuries of this type,
probably because their greater mass enable them to more effectively subdue deer
before defensive blows can be effectively landed. We would expect puma moth-
ers to be more susceptible to this type of mortality because they have to kill prey
more frequently to satisfy the energetic demands of the family (see Ackerman
1982). Inexperience in subadults probably also makes them vulnerable. Chances
of death for both male and female pumas probably increase where principal prey
are larger, such as elk.
Pumas fatally wounded in struggles with prey have also been documented
in other populations in Utah, Idaho, and Alberta (Gashwiler and Ro b i n e t t e
1957 and Lindzey et al. 1988, Hornocker 1970, Ross and Jalkotzy 1992,
respectively). In Alberta, three females and one male died in bungled pre y - c a p-
ture attempts. These deaths comprised 27 percent of the natural mortality. The
three females suffered injuries while killing a mule deer, an elk, and a bighorn
sheep. The male puma, while attacking a bighorn sheep, slipped over a cliff
with its prey.
The most unusual death we documented was of F141 (thirty-four months
old). She was apparently bitten by a rattlesnake on her left foreleg. Three species
of rattlesnakes, including the western diamondback, black-tailed, and rock rat-
tler, are fairly common on the San Andres Mountains. F141 was bitten in March
when rattlesnakes were emerging from winter dens.
Particularly in protected populations, a few pumas have the opportunity to
live out their entire biological life span until processes of aging bring them
down. Three pumas on our study area died when they were about 122–152
months old. We attributed their deaths to complications of old age, although we
could not isolate the exact cause. M3 killed his last mule deer when he was 143
months old. He had already eaten the deer’s liver and had started plucking hair
from another part of the carcass when he suddenly collapsed beside the deer. M5
was 152 months old when he reposed for the last time under the shade of a
desert willow tree. F21 died at 122 months of age, orphaning her three nine-
month-old cubs. But she may have raised them just long enough for some of
them to survive. One of the last signals we got from F21’s radio-collared cub
F194 was when she was dispersing eastward across the Tularosa Basin at the age
of eleven months.
Three adult pumas died of unknown causes. F30 was thirty-four months old
when she died in the back of a narrow cave. Due to abnormal Army restrictions
on the area at the time, we could not reach her for about three weeks after her
death. By that time, tissue autolysis was extreme. F66 was twenty-six months old
when she died. Although we got to her about three days after death, her two
eight-month-old cubs, M67 and F68, had almost completely scavenged her car-
cass, leaving only part of the skull, the feet with the stripped leg bones attached,
and the lumbar vertebrae and pelvis. M49 died at the age of eighty-one months
and shortly after he had returned from a 469-day absence from the San Andres
Mountains resulting from translocation to northern New Mexico (Ruth et al.

1998). About nine days after his return, M49 killed a mule deer in his original
territory. There, he associated with M3, a territorial male who was M49’s neigh-
bor before translocation and who took over the open area left by M49’s depar-
ture. M49 immediately vacated the area of the encounter and traveled north
about 45 km, where he died about twelve days later. Unfortunately, we found
him about five days after death, much too long, for the intensely hot midsum-
mer sun had mummified his carcass into skin and bones. We could only specu-
late that he may have been fatally wounded in the encounter with M3.
Why Do Pumas Kill Other Pumas?

Because intraspecies killing was the single most important cause of death in all
classes of pumas, we asked if such a behavioral trait could be adaptive. What
benefits are there to males that kill and eat cubs or females? What is the advan-
tage to males that engage in mortal combat? Are such aggressive interactions
merely pathological? Not surprisingly, there are fitness benefits to individual
male pumas. But in some instances, such as with M22 and F10, the killings may
be the result of a fatal mixture of social miscues and puma weaponry.
There is no mystery as to why pumas might fight over a deer carcass; the
winner acquires substantial nourishment. But competition for food does not
seem to be an overriding reason for combat; otherwise, we would have expected
greater frequency of killing during the deer decline phase. Because the loser may
be killed or fatally injured in fights, natural selection should favor pumas that
actively avoid other pumas and cover their prey to avoid detection, traits that are
common in pumas. Another downside is that the aggressor puma might also be
injured, perhaps even debilitated to the extent that he cannot effectively provi-
sion himself. Hence, when pumas engage in direct competition for food, the
cost of fighting should be less than the alternative, which is to ignore the food
of another puma, and instead hunt and kill another prey animal. In our study,
we learned that whenever male pumas competed with other pumas for prey, the
other puma was always considerably smaller, and likely easily overwhelmed. We
would expect incidences where adult male pumas of similar mass compete
directly for a carcass to be quite rare.
Certainly, pumas that prey on other pumas are also acquiring nourishment,
particularly if they devour entire carcasses, such as those we saw. In every case,
large males consumed smaller females or cubs. Still, on the surface, it does not
make sense that an adult male should kill and eat a female puma with which it
might sire offspring. His fitness would seem to be reduced. But if the male is
hungry and the interloping female is not a recognized breeder or recognized kin
(i.e., as in a matriline), if she competes with him for food, and if she competes
for food with mothers that are raising his offspring or his daughters in a local
matriline, then this type of cannibalism could enhance the male’s fitness. The
male that eats cubs is gaining nourishment, but more importantly he may also
increase his reproductive success if the cubs are not his offspring. By killing
unrelated cubs, the male can stimulate the female to cycle into breeding condi-
tion earlier than if the female raised her young to the normal age of independ-
ence (i.e., about thirteen months). For example, F45 and F107 lost their entire
litters to cannibalism when cubs were thirty-four and sixty-six days old, respec-
tively. These mothers bred again successfully 307 and 74 days later. On average,
the loss of their litters accelerated the time for the females to breed again by
about five months. If the average of 100 days to rebreed is representative (for lit-
ters lost to all causes, see Chapter 7), then the loss of litters with newborns to
two-month-old cubs may accelerate breeding access to females by eight to ten
months. By providing a male an earlier opportunity to sire a litter, this strategy
should enhance the male’s fitness. In fact, both males that killed and ate entire
litters in our study subsequently sired litters with the mothers of the ill-fated
cubs. Similar observations have been made for the leopard (Bailey 1993). Con-
vincing evidence for this hypothesis in felids comes from African lions where
coalitions of males take over prides and kill unrelated cubs less than six months
old and generally evict older cubs. As a result of this genocide, pride females
return to sexual receptivity, on average, eight months sooner, bear offspring of
the new males, and thus enhance the males’ fitness (Packer and Pusey 1983,
Packer et al. 1988).
On the other hand, infanticide should be expected to lower the female’s fit-
ness by reducing the number of offspring she raises to independence during her
lifetime, especially if she is killed in an attempt to defend her cubs. In f a n t i c i d e
by new immigrant males may even trigger a period of infertility until the
female becomes familiar with the new male, similar to African lion females
when their prides are taken over by new male coalitions (Packer and Pusey
1983). Our data suggest this may occur in puma, though our samples are lim-
ited (see Chapter 7). We did not observe any cases in which female pumas
killed cubs.
Puma mothers we studied exhibited behaviors that could increase their fit-
ness by reducing the threat of infanticide. As we quantify and discuss later in
Chapter 15, avoidance of other pumas by females seemed to be the most impor-
tant strategy. We will never know how many times mothers sensed other pumas
and simply trailed their families away, completely evading potentially dangerous
encounters. Also, promiscuity during estrus would be adaptive if it cultivated
amicable social relations between males and females, and if it confounded pater-
nity and thus made it less likely for males to kill cubs. Nonviolent associations
fostered through breeding courtships seemed vital to reproductive success and
stability in puma society.
Occasionally mothers opted to defend their cubs. Clearly, this choice was
extremely risky, sometimes causing the death of mothers and cubs alike. But
there were at least two instances when defense may have saved cubs and moth-
ers. F90 may have defended her three forty-five-day-old cubs, sired by M38,
from territorial male M46. One cub (F204) was killed, but the other two sur-
vived. Similarly, F147 may have defended her three eighty-day-old cubs from
territorial male M161. Again, one of the cubs (F207) was killed, but two sur-
vived. Aggressive defense, whether intended to protect cubs or food, can still
bear severe physical costs. We examined twenty-four adult female pumas (age
range = 22–122 months, x– = 52.2 ± 30.4) after death. This sample included two
that we translocated from the Treatment Area (Ruth et al. 1988). We discovered
two (8 percent) had suffered severe cranial injuries earlier in life. F15 and F37
were productive mothers; they bore five and three litters, respectively. Old skull
injuries, now healed, were consistent with massive bites inflicted by other
pumas, probably males. F15’s skull was actually laterally lopsided from trauma
to both frontals, the left maxilla, and both zygomatic arches. F37 had canine
punctures to both frontals. It was remarkable that they had survived such
injuries. In addition, we noted that 24 percent (eleven of forty-five) of adult
females we captured had scars, apparently from aggressive encounters with other
pumas. The relatively low frequency of such injuries in these samples suggested
to us that adult female pumas generally choose to avoid confrontation, a strat-
egy that would increase their fitness. When they do defend, females seldom sur-
vive such encounters. Natural selection is against those phenotypes. Adult male
pumas, however, are a different story.
Fitness of adult males is closely linked to their longevity; hence, mate and
territory defense should be strongly conserved strategies. Individual males that
successfully defend their right to breed with an estrus female can influence the
fitness of themselves and other male pumas directly, such as when M3 fought
M22. Indirectly, territorial males that successfully dissuade activity of other
males may also contribute to survival of their own offspring by reducing the risk
of infanticide. But there are also physical costs to the winners of battles. We
examined thirteen territorial males (age range = 28–152 months, x– = 88.6 ±
40.2) after their deaths. This sample included three males translocated from the
Treatment Area (Ruth et al. 1998) and one that, as a subadult, emigrated north-
eastward across the Tularosa Basin and established a territory on the Carrizo
Mountains. We found that six (46 percent) had survived cranial injuries that
were consistent with bites from other male pumas. These injuries all appeared to
be canine punctures that penetrated the anterior part of the skull involving
mostly the frontals, but also the nasals and maxillae. Moreover, of the adult
males we captured, 97 percent (twenty-eight of twenty-nine) had wounds
inflicted during fights with other pumas. Again, we submit that a trait such as
large mass, which would confer an advantage to a male in combat, should be
strongly selected. No one has been able to document behavioral interactions
such as posturing in unfettered pumas challenging each other for food, mates,
or territory. But we suspect that large mass conveys advantages to males before
a single blow is struck. It is likely that in some cases prospective challengers
retreat from perilous encounters because they perceive their nemesis to be too
big and powerful. In those instances, victors win without a fight and potentially
debilitating injuries.
Extended longevity of territorial males should benefit male and female repro-
ductive success by encouraging recognition among resident adults and by dis-
couraging activity of new males. But, we would expect population perturba-
tions, such as sport-hunting, that disrupt social relationships by reducing tenure
length of territorial males to increase chances that females will encounter unfa-
miliar males that threaten their fitness. Biologists studying pumas in Alberta
came to a similar conclusion after they observed that stability (i.e., long tenure)
of adult males was apparently associated with decreased infanticide (Ross and
Jalkotzy 1992). In addition, observations made in British Columbia support
this; non-sires were involved in two separate infanticide incidents (Spreadbury
et al. 1996). Survival of tiger cubs has also been linked to adult male stability.
Smith and McDougal (1991) estimated 90 percent of cubs survived to dispersal
and no infanticide occurred when resident males were stable. But only 33 per-
cent of cubs survived and infanticide was rampant when new males were taking
over territories of former residents.
LP > > KX
1. Although we tried to minimize human impacts to our study population, 5
percent of all the pumas we studied died as a result of poaching or research
accidents. Survival rates of cubs (including natural and human-caused
deaths) were 0.59 in the Treatment Area and 0.63 in the Reference Area.
Excluding human-caused deaths, survival rates for each area were 0.60 and
0.66, respectively. Cubs of matrilineal mothers had higher survival rates
(0.67) than cubs of non-matrilineal mothers (0.56). Cub survival rates were
not sensitive to the deer population decline. Cubs are most vulnerable to
mortality when they are less than or equal to three months old. Survival
rates of male cubs may be slightly better than female cubs. Infanticide and
cannibalism combined as the principal causes of cub death, followed in

importance by starvation, disease, accidents, and coyote predation.
2. Subadult females had a higher survival rate (0.88) than subadult males
(0.56). All the subadult pumas that died were killed by male pumas. We
propose that females survive better because of their tendency to be
philopatric or disperse short distances, in contrast to males that disperse
long distances.
3. Adult male pumas had a higher average annual survival rate (0.91) than did
adult females (0.82). Intraspecies strife was the greatest cause of mortality
for adult males and females. In order of importance, accidents, disease, and
aging also caused female deaths. For males, other causes of death included
disease and aging. The most unusual cause of death was snakebite.
4. Male pumas in particular kill other pumas and enhance reproductive suc-
cess because they acquire food, mates, and dominance over territory. Female
pumas avoid aggressive encounters with other pumas and enhance their
reproductive success by enhancing the survival of themselves and of their
cubs.
LN N5LN5)L
1. Test for difference in finite survival rates (i.e., number of pumas surviving /
number of pumas monitored) for ninety-seven cubs from matrilineal moth-
ers and fifty-seven cubs from non-matrilineal mothers. Chi-square test of
homogeneity (2 ¥ 2 contingency table): c2 = 1.82, 1 d.f., P = 0.20.
2. Test for difference in finite survival rates for nine radio-collared male and
sixteen radio-collared female independent subadult pumas. Chi-square test
of homogeneity (2 ¥ 2 contingency table): c2 = 3.22, 1 d.f., P = 0.08.
3. Puma masses: M64 at death = 41.7 kg, M88 three days after killing M64 =
59.0 kg, M108 at death = 39.0 kg, M88 nine months before killing M108
= 63.5 kg and two months after = 62.6 kg.
4. Puma masses: M22 = 52 kg, M88 = 63 kg, F10 = 31 kg, F96 = 29 kg.
5. Puma masses: M1 = 64.4 kg, subadult male M300 = 50.8 kg.
6. Survival rates for adult male pumas in the Treatment Area (n range = 5–9)
and Reference Area (n range = 5–14) were compared using Z-tests (Heisey
and Fuller 1985a). Rates differed only in 1989: Z = 1.688, P = 0.05. All
other years: Z range = 0.68–1.145, P range = 0.13–0.5.
7. Survival rates for adult female pumas in the Treatment Area (n range =
4–12) and Reference Area (n = 3–17) were compared using Z-tests. Rates
differed in 1991: Z = 1.573, P = 0.06, and in 1993: Z = 1.992, P = 0.02.
All other years: Z range = 0–1.038, P range = 0.15–0.5.
8. Survival rates for adult male and adult female pumas were compared using
Z-tests (refer to Table 8-1). Rates differed for 1987: Z = 1.535, P = 0.06;
1988: Z = 1.621, P = 0.05; 1989: Z = 1.560, P = 0.06; 1990: Z = 2.221, P
= 0.01. Rates for all other years: Z range = 0.057–1.018, P range =
0.15–0.48.
9. Test for differences in ages of adult male (n = 11) and adult female (n = 19)
pumas that died. Two-sample t-test: t = 2.853, 28 d.f., P = 0.008.
10. Masses (kg) of adult pumas involved in intraspecific killing. Females: n = 7,
x– = 31.8 ± 3.9, range = 27.2–38.6; males: n = 5, x– = 58.6 ± 7.0, range =
52.2–68.9.
11. Masses (kg) of adult male pumas involved in intraspecific killing. Winners:
n = 6, x– = 61.2 ± 4.9, range = 54.9–68.9; losers: n = 5, x– = 56.4 ± 8.9, range
= 43.5–64.4.
12. Test for association between annual frequency of intraspecies killing (adult
plus subadult pumas) and year-round (January–December) density of total
adults, adult males, and adult females in the Treatment Area (1988–1994,
n = 7). Spearman correlation coefficients (1-tailed test): total adult density
rs = 0.09, P > 0.25; male density rs = 0.62, P = 0.08; female density rs =
–0.09, P > 0.25.
13. Test for association between annual frequency of intraspecies killing (adult
plus subadult pumas) and year-round (January–December) density of total
adults, adult males, and adult females in the Reference Area (1989–1994, n
= 6). Spearman correlation coefficients (1-tailed test): total adult density rs
= 0.06, P > 0.25; male density rs = 0.29, P > 0.25; female density rs = 0.06,
P > 0.25.
Chapter 9
Independence of Puma
Progeny, and Philopatry,
Emigration, and Immigration
By tagging a large number of dependent progeny (i.e., offspring produced by

mothers on the San Andres Mountains) we were able to discover for the first
time how behaviors of philopatry, dispersal, emigration, and immigration
affected puma population dynamics. We published our findings earlier in Con -
servation Biology (Sweanor et al. 2000). Therefore, in this chapter we only reit-
erate the results that pertain to population dynamics and how these behaviors
affect fitness. We discuss the implications of these behaviors to puma conserva-
tion further in Chapter 20.
We defined philopatric pumas as progeny whose adult home ranges over-
lapped their natal area by 5 percent or more (based on the 90 percent minimum
convex polygon method of home range estimation; see Chapter 13). Dispersers
were progeny that established independent home ranges that overlapped less
than 5 percent of their natal area. Dispersal began when a subadult made its first
movement outside its natal home range and did not return. Dispersers that
departed from the puma population on the San Andres Mountains study area
altogether were emigrants. Immigrants were pumas that originated from popula-
tions outside of the San Andres Mountains before establishing residency on our
study area. Hence, recruits into our study population consisted of progeny born
on the San Andres Mountains and immigrants that entered the resident adult
portion of the population. Fundamentally then, philopatry, emigration, and
145
immigration were vital components of population growth. All of the informa-

tion that we gleaned on philopatry, dispersal, and emigration came from pumas
that we captured and tagged while they were still dependent on mothers. After
cubs achieved independence from mothers, we followed their movements by
radio-tracking them and by receiving news of their fates from hunters, predator
control agents, or automobile drivers who killed them, and we identified indi-
vidual pumas by their ear tags, tattoos, or radio collars. In addition, our inten-
sive and extensive capture efforts on the San Andres Mountains helped us to
detect the arrival of new immigrants that were recruited into the population.
Independence of Progeny
We studied fates of forty-three tagged progeny that reached independence. We
estimated age at independence for thirty of those. For six males and six females
that we radio-tracked before and after becoming independent of their mothers,
age at independence averaged 13.7 ± 1.6 months (range = 11.1–16.0). Our sam-
ple size was small, but males appeared to be dependent on mothers (x– = 13.8 ±
1.8) for about the same duration as females (x– = 13.5 ± 1.6). Siblings became
independent within zero to twenty-eight days of each other (x– = 10.0 ± 14.0, n
= 5 comparisons of seven siblings in three litters). For eighteen other progeny,
our data were less accurate because we could not radio-track the animals during
separation; either the cubs were not radio-collared or their mother’s radio collar
had malfunctioned at the time. Hence, we estimated a puma’s age at independ-
ence based on the age when we recaptured it as an independent subadult (n =
5), the age at which radio-collared siblings became independent (n = 3), the age
at which mothers appeared to no longer associate with cubs based on attrition
in tracks (n = 2), the age at which siblings separated from one another (n = 3),
and the timing that their mother gave birth to her next litter minus the gesta-
tion period (n = 5). For this last group, we calculated a range of ages based on
the range of days that offspring associated with mothers before and after esti-
mated conception dates. Ranges of average ages at independence were 13.7 ±
2.6–14.5 ± 2.4 months for seven males and 13.7 ± 3.0–14.8 ± 2.4 months for
eleven females. Because of the five offspring (three males, two females) that were
already independent when we radio-collared them, these ages are biased upward.
In general, we considered offspring to be independent of their mothers after
they were about thirteen months old.
Independence seemed to be initiated by the mother’s inattentiveness to her
young, either by abandoning them at the edge of natal range or by not return-
ing to them at a rendezvous site within the natal range (see Chapter 12). Simi-
lar behavior at separation has been reported for pumas in central Idaho (Sei-
CHAPTER 9. INDEPENDENCE OF PUMA PROGENY 147
densticker et al. 1973) and Southern California (Beier 1995). After independ-
ence, we observed siblings associating for up to 1.7 months. Puma littermates
have been known to associate for up to three months after independence in
Alberta (Ross and Jalkotzy 1992). By the age that puma cubs are becoming inde-
pendent, they are physically capable of killing deer-size prey; females are
approaching the size of adult females and males may exceed it (see Appendix 1).
In addition, their permanent teeth are in place and canines are almost fully
extended. Of seventeen offspring for which we determined the month of their
separation from mothers, eleven (five males, six females) became independent
during the July–August mule deer fawning season when their principal prey
were most abundant and vulnerable; three others were independent shortly
thereafter in October. This pattern would benefit survival of young, but it may
be an artifact of our small sample size. Information on the age at which puma
offspring separate from mothers in other populations is limited. Comparable
data come from a hunted puma population in southern Alberta where thirty-six
offspring became independent throughout the year at the average age of 15.2 ±
3.0 months (Ross and Jalkotzy 1992). Other studies did not differentiate ages at
independence and dispersal.
Timing of independence in nine cubs from six litters that we followed closely
with radiotelemetry was linked with resumption of breeding behavior in their
mothers. Offspring became independent within one to forty-four days of the
mother consorting with an adult male puma (x– = 19.1 ± 15.5). In four events
cubs were independent thirteen to twenty-one days (x– = 16.0 ± 3.6) before the
mother consorted with a male, and in five events cubs were independent one to
forty-four days ( x– = 21.6 ± 21.2) after the mother consorted. As a result of those
unions, three of the six mothers produced new litters one gestation period later.
Two other mothers bred with males eleven and forty-seven days after the first
association, then produced new litters one gestation period later. The sixth
mother consorted with adult males on two occasions twenty-five days apart, but
apparently was not impregnated. In the first intensive field study of pumas in
the early 1970s, biologists observed that “In the months before and just after the
young become independent, the female associates with (tolerates?) adult males
and even adult females more frequently and for longer periods than at any other
time.” They postulated that “this is related to hormonal changes associated with
the onset of estrus” (Seidensticker et al. 1973:40,41). Our data support their
conclusions.
Hypothetically, independence of offspring at this critical time may be adap-
tive for at least three reasons: (1) large cubs would not be direct competitors for
the mother’s time and energy required for mating, pregnancy, and rearing new
cubs; (2) offspring would not be close to their estrus mothers and potential tar-
gets for aggressive male mates; (3) male offspring would avoid inbreeding with
their mothers. Of these, the first hypothesis seems the most plausible; this tim-
ing of independence would contribute to the lifetime reproductive success of
puma mothers by ending maternal care for offspring that no longer require it
and enabling her to devote her energy to raising a new litter. In the second
hypothesis, we would expect a large majority of the cubs to become independ-
ent before the mother began to attract adult males that could easily kill the cubs.
But results from our small data set were equivocal. Less than half of the offspring
became independent before their mother resumed breeding activity and more
than half became independent afterward. The third hypothesis does not seem
likely because we found that young males generally reach puberty at about
twenty-four months of age, several months after becoming independent of
mothers. In males, inbreeding is avoided principally by dispersal (see Chapter
13). Moreover, this hypothesis does not explain timing of independence for
female cubs.
Philopatry and Dispersal

Following independence, progeny either remained within the natal area (i.e.,
they were philopatric) or they dispersed. Only a fraction of females dispersed;
the remainder were philopatric. Thirteen females exhibited philopatry and four
dispersing females established home ranges adjacent to their mothers. Together
these females formed eight matrilines (see Chapter 13, Plate 4). Three other
subadults did not disperse from their natal areas. As we discussed in Chapter 8,
two subadult males (M64, M108) and one subadult female (F96) were killed by
male pumas on natal areas about seventy-five, fifty-two, and eighty-seven days,
respectively, after independence. Based on the histories of tagged subadults, we
suspected that the males would have eventually dispersed if they had survived;
but the female may have been philopatric.
Dispersal from natal areas for eight known-age progeny occurred at the aver-
age age of 15.2 ± 1.6 months. Our small samples suggested that males tended to
disperse at older ages (n = 6, x– = 15.7 ± 1.4, range = 14.0–17.0) than females (n
= 2, x– = 13.6 ± 0.9, range = 13.0–14.3). Both sexes dispersed from natal areas
before reaching puberty (see Chapter 7). During the subadult life-stage, energy
is invested in growth, movement away from natal areas, and socializing directly
and indirectly with other pumas, not in reproduction. For pumas, lower fitness
would be expected if dispersal occurred during the reproductive stage of life
when energy and time budgets should principally be allotted to mating and
parental investment. Ages of pumas from other populations fit a pattern of dis-
persal prior to puberty. In southwestern Colorado, a slightly larger sample size
Slide @171%
PHOTO 21. Three eleven-month-old

siblings F107, M108, and F109 with
Ken Logan and Frank Smith (right).
M108 (held by Ken) was killed by his
father, M88, as a subadult. Both
daughters were philopatric, and F107
(held by Frank) bred with her father
twice.
indicated that eight males and four females dispersed at mean ages of 11.8 ± 1.0
months and 20.0 ± 8.9 months, respectively (Anderson et al. 1992:63). Twelve
females and five males dispersed from natal areas in an Alberta population at the
average age of 16.0 months (range = 10–22) (Ross and Jalkotzy 1992). In a Utah
population with a relatively low density of pumas (mode = 0.4 total pumas / 100
km2) twelve cubs dispersed when they were sixteen to nineteen months old
(Hemker et al. 1984). In a puma population in Southern California that was
practically surrounded by urbanization, six known-age males dispersed at a
mean age of 18.0 ± 2.8 months (Beier 1995). This was practically the same as
the mean of 17.9 ± 4.0 months for seven male Florida panthers in a population
constricted by their limited habitat (Maehr et al. 1991). In the Florida popula-
tion, dispersal for only one female was described, but it was peculiar for pumas
because she dispersed after reproducing. The panther raised her first litter within
her natal home range, then at about thirty-three months old she dispersed 16
km away (Maehr et al. 1989).
We traced straight-line distances for twenty-seven offspring that dispersed
from their natal home ranges (Table 9-1). Of those, six males and six females
moved minimum distances of 47.1 and 5.6 km, respectively, before establishing
adult home ranges within the San Andres Mountains study area. Another ten
males and three females dispersed so far that they emigrated from our study
population (Fig. 9-1). Two other males were dispersing within the San Andres
Mountains when a male puma killed one and the other shed his radio collar at
the southern border of our study area. We assumed the latter puma either died
or emigrated because we never recaptured him on the study area. On average,
male offspring dispersed 8.1 times the distance of females from their natal home
Table 9-1. Dispersal distances for independent pumas born on the San Andres
Mountains, New Mexico (n = 40), and for pumas with origins outside the San
Andres Mountains (n = 3), 1986–1994 (adapted from Sweanor et al. 2000).
Distance Male dispersal distance (km) Female dispersal distance (km)

estimatora n x– ± SD Range n x– ± SD Range
Progeny
NAC to IACb 8 101.3 ± 57.7 47.1–192.5 7 28.3 ± 26.1 5.6–78.5
NAC to IAC (P) 0 12 3.2 ± 2.6 0.7–9.9
NAC or CS to 5 139.8 ± 68.7 56.0–214.9 0
AMS
CS to IAC 0 1 36.6
CS to IAC (P) 0 1 2.2
NAC or CS to 5 67.2 ± 37.5 53.1–133.8 1 76.6
LLS or SMSc
TOTALd 13 116.1 ± 62.5 47.1–214.9 21 13.1 ± 19.5 0.7–78.5
Origin outside
study area
CS to LL or 1 175.7 2 96.3 ± 31.0 74.4–118.2
IAC
aOrigins and endpoints of distance estimators: NAC—natal home range arithmetic center; IAC—
independent home range arithmetic center; P—philopatric; CS—capture site; AMS—adult mortality
site; LLS—last location as a subadult; SMS—subadult mortality site; LL—last location, status
unknown. CS for progeny was within the natal home range.
bMales dispersed significantly farther than females (Mann-Whitney U test: U = 49.0, P = 0.02).
cNot included in analysis of sex-related dispersal distances.
dMales dispersed significantly farther than females (Mann-Whitney U test: Z = 4.56, P < 0.001).
Total includes only pumas that probably completed their natal dispersal moves and those that were
philopatric into adulthood. We assumed that pumas that had reached adulthood prior to death or loss
of radio contact had completed their dispersal moves, whereas subadults had not.
FPO @ 65%
FIGURE 9-1. Dispersal moves of pumas from the San Andres Mountains to other habitats in
New Mexico.
ranges to adult home ranges. Effective dispersal for thirteen male progeny aver-
aged 7.3 home range diameters, with extremes of 3.0–13.6 (based on home
range estimates in Chapter 12). None of these males settled adjacent to its
mother or siblings on the San Andres Mountains. In contrast, twenty-one
female progeny (including dispersing and philopatric females) effectively dis-
persed an average of 1.4 home range diameters, with extremes of 0.1–8.5. Dis-
persal directions of male progeny were bimodally distributed with a diameter
line oriented at 347 degrees, while directions for females were random (Sweanor
et al. 2000). In general, males and females tended to disperse along the north-
to-south axis of the San Andres Mountains, reflecting the pumas’ affinities to
favorable habitats during initial stages of dispersal and emigration. Some pumas
traversed broad sweeps of non-puma habitat extending 45–65 km across desert
basins to the east and west, making unidirectional moves to other large patches
of mountainous terrain in periods of fewer than seven days. Pumas have been
known to travel long distances in relatively short time spans. A male dispersing
in Southern California covered about 29 airline km in 48 hours (Beier 1995),
and an orphaned eight-month-old female translocated in southwestern Col-

orado covered 43 airline km in about 10 hours (Anderson et al. 1992).
In our situation it appeared that dispersing pumas we re simply moving to
other mountain ranges that they spied across the basins. They used smaller
patches of puma habitat to link their dispersal moves to more distant habi-
tats (see Chapter 13). Pumas dispersing from the San Andres Mo u n t a i n s
reached puma habitats throughout the southern half of New Mexico (Fig. 9-
1). In addition, we caught one male and two female subadult pumas that
we re apparently born elsew h e re and had immigrated into our study area tem-
porarily, then left it as they continued their dispersal moves. Their behavior
demonstrated how pumas from other patches sometimes use the San Andre s
Mountains as a “stepping stone” to other habitat patches as well as the tran-
sient nature of some individuals that really we re not members of the local
population. The longest dispersal movement yet re c o rded for pumas was
about 483 airline km for a male tagged as a cub in the Big Horn Mo u n t a i n s
in north-central Wyoming and killed at thirty months of age by a hunter in
Black Hawk Canyon, west of De n ve r, Colorado (Logan and Sweanor 2000).
Dispersal of male subadults from natal areas was obligatory even at low pop-
ulation densities. Ac c o rd i n g l y, male dispersal was density-independent. Bu t
because some female progeny dispersed and some did not (i.e., they we re
philopatric), we interpreted female dispersal to be partially density dependent.
In the rebuilding puma population on the San Andres Mountains, there appar-
ently was ample food and room for female progeny to establish their adult
home ranges overlapping or adjacent to their natal areas, as well as for immi-
grant female re c ruits. But because of the programmed termination of our study,
we we re unable to quantify how female dispersal or philopatry changed in re l a-
tion to puma population density and prey abundance. Additional long-term
re s e a rch is needed to understand these relationships. A reasonable hypothesis is
that rates of dispersal in female progeny increase as per capita food resources
decline. If dispersal of female progeny is affected in this way, then competition
b e t ween pumas for food may be a population-regulating mechanism. We
would expect a greater dispersal rate to slow the population growth rate (see
Chapter 10).
Hornocker (1970) first observed that puma progeny on a winter study area
dispersed from it. He reported dispersal distances for three tagged progeny that
were killed by hunters: a male moved 161 km and two females moved 64 and
113 km. While extending that study, Seidensticker et al. (1973) observed that
dispersal of subadult puma progeny was independent of adult density: no prog-
eny seemed to settle as adults within their study population. Immigrant pumas,
which they called “transients,” replaced adults that died in the study population.
Immigration was also noted as an important source of recruits in puma popula-
tions in Alberta (Ross and Jalkotzy 1992), Utah (Laing and Lindzey 1993), and
Wyoming (Logan et al. 1986). Numerous studies have now shown that disper-
sal in puma progeny is sex-biased; male dispersal is generally obligatory, and
males disperse consistently farther than females (see Anderson et al. 1992), char-
acteristics that are common in polygynous or promiscuous mammals (Dobson
1982 see Chepko-Sade and Halpin 1987, Stenseth and Lidicker 1992). In his
summary on dispersal patterns of sixty-five North American pumas, Anderson
et al. (1992:66) found that on average males dispersed 2.7 times (n = 33, x– =
85.0 km, range = 29–274 km) farther than females (n = 32, x– = 31.4 km, range
= 9–140 km). Although the importance of dispersal to the dynamics of local
puma populations was previously recognized by other puma biologists, its effects
on rates of emigration and recruitment were not quantified until our study. In
Chapter 13, we examine the adaptive significance of dispersal in pumas (see
“Why Do Pumas Disperse?”).
Emigration
As pumas dispersed beyond the San Andres Mountains, they emigrated from the
local population. We estimated total successful emigration spanning a 5.1-year
period from 1 February 1990 to 23 February 1995 by using our capture infor-
mation on tagged progeny that survived to adulthood and estimating cub and
subadult survival rates. Total successful emigration from our study area during
that time span was about 27.5–43.7 progeny, including about 19.1 males and
8.4–24.6 females (depending upon the survival schedule used for females, i.e.,
0.56 or 0.88). This would equate to 5.3–8.5 emigrants per year, about 3.7 males
and 1.6–4.8 females. Of the estimated total progeny that would have survived
to adult ages, we estimated that 83 percent of males and 33–59 percent of
females emigrated from the San Andres Mountains (Sweanor et al. 2000). These
were pumas that were raised by mothers on the San Andres Mountains, emi-
grated from the area, and survived long enough to be recruited into other puma
subpopulations in New Mexico. Based on the reproductive success of immigrant
pumas to the San Andres Mountains, we surmised that emigrants became pro-
ductive members of other subpopulations as well. While pumas were leaving the
San Andres Mountains and moving to other subpopulations in southern New
Mexico, pumas were emigrating from those subpopulations and making their
way to our study area.
Recruitment of Progeny and Immigrants

We estimated recruitment during 1 February 1990–23 February 1995 by count-
ing the tagged progeny and immigrants that reached adulthood within the San
Andres Mountains. Immigrants were identified as such, because they did not
have tags when we initially captured them. About twenty-one progeny, includ-
ing four males and seventeen females (1 male:4.2 females), were recruited, for
an average rate of 0.8 male and 3.3 females per year. During the study, twelve
females but no males (57 percent in aggregate) produced offspring themselves.
It is possible that three of the males sired litters without us detecting their con-
tribution. The fourth male was infertile (M73). For the same time period we
estimated that about twenty-two immigrating pumas, including fourteen males
and eight females, were recruited into the San Andres Mountains population, at
an average rate of 2.7 males and 1.6 females per year. Of these immigrant
recruits, at least five males and five females (45 percent in aggregate) produced
offspring. Again, it is likely that more males sired litters without our knowledge.
These statistics indicated that recruitment into the growing San Andres Moun-
tains puma population was dependent equally upon both progeny (4.1 / year)
and immigrants (4.3 / year). In addition, they showed recruitment of males and
females was affected by differential dispersal patterns in the sexes. We calculated
that 78 percent of male recruits were immigrants, whereas 68 percent of female
recruits were progeny. Together, progeny and immigrant recruits (8.4 / year) bal-
anced or exceeded the estimated number of progeny that successfully emigrated
(5.3–8.5 / year). The contribution of immigrants to population growth exem-
plifies the importance of considering puma distribution in the context of a
source-sink metapopulation structure. This subject, along with puma density
and population growth, will be discussed in our next chapter.
LP > > KX
1. Puma offspring became independent of their mothers at an average age of
about fourteen months. Mothers sometimes initiated independence by
abandoning the offspring. Independence was associated with the resump-
tion of breeding behavior in the mothers. Independence seemed to occur at
a time when cubs could be self-sufficient and the mother could devote time
and energy to more reproduction.
2. Some female progeny were philopatric, while others dispersed. Mothers,
philopatric daughters, and daughters that established adjacent home ranges
formed eight matrilines on the San Andres Mountains.
3. Puma offspring that dispersed left their natal areas at the average age of
about fifteen months. Both sexes dispersed before reaching puberty. On
average, males dispersed about 101 km, and females about 28 km, from
natal areas. Effective dispersal was about 7.3 home range diameters for
males and about 1.4 home range diameters for females. Male dispersal was
density independent. We hypothesize that female dispersal is partially den-

sity dependent.
4. Surviving puma progeny emigrated from the San Andres Mountains at the
rate of about 5.3–8.5 pumas per year, of which about 3.7 were males and
1.6–4.8 were females. Of the total surviving progeny, about 83 percent of
males and 33–59 percent of females emigrated and were potential recruits
to other puma populations.
5. Growth of the adult segment of the puma population was dependent upon
progeny and immigrant recruits. Progeny were recruited at the rate of about
4.1 per year and immigrants were recruited at the rate of about 4.3 per year.
The male:female ratio of progeny recruits was about 1:4, while for immi-
grant recruits it was roughly 2:1.
Chapter 10
Puma Population Density, Growth,

and Metapopulation Structure
Experimentally Removing Pumas

A powerful way of learning how quickly a puma population can grow, and about
the patterns of replacement of breeding adults, is through experimentally remov-
ing animals. We were already learning valuable things about those properties by
studying the dynamics of a rebuilding population. But by experimentally reduc-
ing a portion of the population in the 703-km2 Treatment Area, we could test if
the patterns we initially saw in the Treatment and Reference Areas were real and
also gain estimates of variation. Consequently, we removed thirteen pumas from
the Treatment Area in a 6.5-month period from 9 December 1990 to 22 June
1991. Five were adult males (mean age = 64.0 ± 32.8 months, range = 30–102),
six were adult females (mean age = 46.0 ± 28.9 months, range = 22–102), and
two were subadult females (mean age = 16.5 ± 0.7 months, range 16–17). We
translocated those pumas to northern New Mexico (Ruth et al. 1998). During
the same period, two 2- to 3-month-old siblings (M117 and F118) died of nat-
ural causes on the Treatment Area. We believe another female cub (F102), which
disappeared at about 10.5 months old, also died.
Of the pumas we translocated, two adult males returned to their original ter-
ritories on the Treatment Area. Puma M88 was back on 21 July 1991, 166 days
after removal. He remained the dominant male there through the end of the
study. M49 returned on 7 July 1992, 469 days after removal. But his stay on his
157
original territory was brief, only about nine days. Possibly driven from the area
by an encounter with M3, M49 subsequently traveled north into the Reference
Area and died. Consequently, M88 contributed to the post-treatment (i.e., after
removal) puma population on the Treatment Area, but M49 did not.
At the end of the experimental removal, the entire puma population in the
Treatment Area had been reduced by 47 percent. Adult pumas had been reduced
by 53 percent, and all independent pumas (adults plus subadults) had been
reduced by 58 percent (Table 10-1). By January 1994, thirty-one months post-
treatment, the adult segment of the puma population had practically recovered
to the 1991 pre-removal level, with a difference of –0.27 pumas (Table 10-2).
The actual rates of increase for adult pumas in the Treatment Area during pre-
and post-treatment periods and in the Reference Area will be discussed in detail
Table 10-1. Puma population, number of pumas removed, and post-removal

population in the Treatment Area, December 1990–July 1991, San Andres
Population Adults Subadults Cubs

description Male Female Male Female Male Female Unk. Sex Total
Pre-treatment
(Dec. 1990)
Pumas present 7 10 0 2 4 5 2 30
Treatment
(Dec. 1990–
June 1991)
Pumas removeda 5 6 0 2 1c 2c 0 16
% Removedd 71 60 100 25 40 53
Post-treatment
(July 1991)
Pumas present 3b 5 0 0 3 3 2 16
% Removed d 57 50 100 25 40 0 47
aIndependent pumas were removed alive and translocated to northern New Mexico from 9 December
1990 to 22 June 1991 (see Ruth et al. 1998).

bOne adult male (M88) that was removed returned to his original home range 166 days later (21 July
1991). He remained as a member of the Treatment Area population through the end of the study.
cDuring the removal period, two cubs (one male, one female) from the same litter died, and one
female cub from another litter disappeared.

dPercentage of pumas removed from each category. All adults were reduced by 53 percent. All inde-
pendent pumas (adults and subadults) were reduced by 58 percent. The total puma population was
reduced by 47 percent.
CHAPTER 10. PUMA POPULATION DENSITY, GROWTH, AND METAPOPULATION 159
later in this chapter. Sex ratios of the adult pumas in January 1991 and January
1994 were identical (i.e., 1 male:1.4 females), suggesting that adult pumas were
replaced numerically by same-sex recruits. Our findings further confirm a pat-
tern of replacement of adult pumas in-kind that has been observed by other
biologists studying puma populations in Idaho (Seidensticker et al. 1973) and
Utah (Laing and Lindzey 1993). This tendency points to the strong influence of
the mating system and social behavior in the structure of puma populations (see
Chapters 12 through 15).
Recruits into the Treatment Area after the removal originated from three
sources. Three females (F107, F109, and F149) were progeny born to Treatment
Area females (F47 and F21) that were not removed during the treatment. Of
those, two were philopatric siblings (F107 and F109), and one (F149) estab-
lished a home range adjacent to her natal area. Three pumas, including two
males (M124, M153) and one female (F103), were progeny of females (F65,
F28, and F87) in the Reference Area. Six pumas, including five males and one
female, immigrated to the Treatment Area from outside the San Andres Moun-
tains. The origin of one other recruited female was unknown. In the last year of
the study, we traced her movements by ground-tracking her and visually
observed her three large cubs. But we never got to see if she was tagged. In gen-
eral, proportions of female recruits from progeny born on the Treatment Area
and immigrants (either from the Reference Area or from elsewhere in New Mex-
ico) were about the same as we saw in the San Andres Mountains puma popu-
lation as a whole. However, all of the male recruits immigrated even though at
least eight tagged male offspring that were born in the Treatment Area reached
independence during the treatment and post-treatment period. One of those
(M108) was killed in his natal area by his father (M88). Two others died as they
dispersed north through the Reference Area. One of them (M140) was killed by
a male puma, and the other (M198) apparently died of a head injury. All the
rest, if they survived, apparently emigrated from the San Andres Mountains.
Biologists studying a puma population in south-central Utah were the only
others to experimentally reduce a population. Their objective was “to monitor
the response of a [puma] population with known characteristics to removal of
individuals, simulating a harvest” (Lindzey et al. 1992:224). Six pumas, includ-
ing two male and one female that were 1.5 years old, and one male and two
females that were 3.5 years old, were removed from a 1,900-km2 central core
area within a 4,500-km2 study area. In addition, two other adult pumas
(unspecified sex and age) died from natural causes. The combination of
removals and deaths comprised 36 percent of the “harvestable population,”
which the biologists defined as the number of pumas older than one year. With
the possible exception of one adult male, the adult segment of the population
recovered in nine months. The rapid recovery was probably facilitated by pumas
living in the large adjacent protected area (i.e., hunting was banned). By the sec-
ond year after the removal, the number of adult males had recovered fully, but
the number of adult females had declined by one as a result of three deaths that
were replaced by two recruits. In that manipulated population, recovery of adult
female pumas was from progeny and immigrants, and all of the males were
replaced by immigrants, just as we had observed for our study area.
Density
Pumas are large, warm-blooded vertebrates that are obligate carnivores. They
occupy the third trophic level, above producers and herbivores. Because energy
is inefficiently transferred between trophic levels (see Ricklefs 1990), pumas
must acquire sufficient energy for maintenance, behavioral activity, reproduc-
tion, and rearing of young by pursuing mobile vertebrate prey in large areas. A
consequence of these energetic constraints and movement patterns is that pumas
live at relatively low population densities.
The puma population on the San Andres Mountains increased during the
study. Our January population point estimates for the puma population in the
Treatment Area and Reference Area are given in Table 10-2. Density of adult
pumas in the Treatment Area increased from 1.16 to 2.10 per 100 km2 during
the pre-treatment and treatment years (1988–1991) and from 0.84 to 1.99 per
100 km2 in the post-treatment years (1992–1995; Table 10-3). By January
1994, thirty-one months after the removal of 53 percent of the adults, the Treat-
ment Area population had almost recovered to its pre-treatment density. How-
ever, by January 1995, adult puma density on the Treatment Area declined by 4
percent because of the death of one female. Density of adults post-treatment still
increased to the pre-removal level even though the deer population declined pre-
cipitously (see Chapter 17). Density of all pumas, including adults, subadults,
and cubs, in the Treatment Area ranged from 2.01 to 3.91 per 100 km2 in the
pre-treatment and treatment years and from 2.78 to 4.25 per 100 km2 in the
post-treatment years. January population estimates for the Reference Area indi-
cated that adult puma density increased from 0.94 to 2.01 per 100 km2 between
1989 and 1995 (Table 10-3). Density of all pumas ranged from 1.72 to 3.90 per
100 km2. For our entire San Andres Mountains study area during January each
year from 1989 to 1995, the number of adult pumas ranged from twenty-five
to forty-two, and the total number of pumas ranged from forty-two to eighty-
two (Table 10-4).
We examined the potential effects of unintended removals of pumas from the
Treatment and Re f e rence Areas on January point estimates of adult pumas. These
Table 10-2. Estimated puma population each January in the Treatment

Area (1988–1995) and Reference Area (1989–1995), San Andres
Mountains, New Mexico.a
Adults Subadults
Year Males Females Males Females Cubs Total
Treatment Area
1988 5.14 3.02 0 0 6 14.16
1989 5.94 6.04 1 2.59 2.89–4.67 18.46–20.24
1990 4.47 8.82 1 2 4.19–9.01 20.48–25.30
1991b 6.34 8.47b 0 1b 7–11.68 22.81–27.49
1992 2.67 3.22 0.68 4 10 20.57
1993 4.08 8.20 2–3c 0–1c 6.20–14.60 21.48–29.88
1994 5.32 9.20 0 0 5–8 19.52–22.52
1995 5.78 8.20 0 0 9–15 22.98–28.98
Reference Area
1989 6.81 5.96 1 0.41 9.11–12.33 23.29–26.51
1990 7.04 11.18 1 1 10–14 30.22–34.22
1991 7.84 10.53 0 1 9–17.32 28.37–36.69
1992 8.33 14.28 0.32 1 8.50–14.50 32.43–38.43
1993 9.66 14.80 2 3 13.80–23.40 43.26–52.86
1994 8.97 14.54 2 2 9–16.48 36.51–43.99
1995 11.16 16.10 1 1 6 35.26
aRadio-collared pumas that lived along either the Treatment Area–Reference Area boundary
or the study area boundary were included in the Treatment Area or Reference Area based on
the proportion of their aerial locations in the area during each year. Cubs were included in
areas in identical proportions as their mothers (see text for details).
bPumas were experimentally removed from the Treatment Area from 9 December 1990 to
22 June 1991. The January 1991 estimate reflects the absence of one adult female and one
subadult female that were removed in December 1990.
cSubadults present in January 1993 consisted of either three males, or two males and one
female.
removals included one adult male that was translocated from the Treatment Area
on 6 April 1989 because he killed three desert bighorn sheep, and seven human-
caused puma deaths (i.e., one male, six females) in the Re f e rence Area during
1988–1994 (see Chapter 8). We did this by assuming that each puma would have
survived re l a t i veto the annual surv i val probabilities (i.e., rates) that we calculated
(using Mi c ro m o rt software) for adult males and females in the respective area. In
Table 10-3. Estimated density of pumas each January in the

Treatment Area (1988–1995) and Reference Area (1989–1995),
San Andres Mountains, New Mexico.
Pumas / 100 km2
Year Adult Males Adult Females Total Adults Total Pumasa
Treatment Area
1988 0.73 0.43 1.16 2.01
1989 0.84 0.86 1.70 2.63–2.88
1990 0.64 1.25 1.89 2.91–3.60
1991 0.90 1.20b 2.10 3.24–3.91
1992c 0.38 0.46 0.84 2.93
1993 0.58 1.17 1.75 3.06–4.25
1994 0.76 1.31 2.07 2.78–3.20
1995 0.82 1.17 1.99 3.27–4.12
Reference Area
1989 0.50 0.44 0.94 1.72–1.96
1990 0.52 0.82 1.34 2.23–2.52
1991 0.58 0.78 1.36 2.09–2.71
1992 0.61 1.05 1.66 2.39–2.83
1993 0.71 1.09 1.80 3.19–3.90
1994 0.66 1.07 1.73 2.69–3.24
1995 0.82 1.19 2.01 2.60
aTotalpumas includes adults, subadults, and cubs.
bThe January 1991 estimate reflects the absence of one adult female that was
removed in December 1990.
cPumas were experimentally removed from the Treatment Area from 9 December
1990 to 22 June 1991.
other words, we estimated when these pumas would have been members of the
population if they were only subjected to natural causes of mort a l i t y. Each adult
puma was added to the January point estimate for each year if the product of its
survival rates was greater than 0.50 for that ye a r. The puma was considered dead
and expunged from the estimate when the product of its survival rates was less
than 0.50. We also assumed that the lone translocated male in the Treatment Area
would have been removed with the rest of the pumas after January 1991. Given
these assumptions, adult male density in the Treatment Area during 1988–1991
ranged from 0.73 to 1.04 males per 100 km2, while total adult density ranged
from 1.16 to 2.39 adults per 100 km2 ( c o m p a re with Table 10-3). In the Re f e r-
ence Area, four of the females would have died in the interval, while the male and
Table 10-4. Estimated puma population each January,

1989–1995, San Andres Mountains, New Mexico (2,059 km2).
Adults
Year Males Females Subadults Cubs Total
1989 13 12 5 12–17 42–47
1990 12 20 5 14–21 51–58
1991 15 19 2 16–29 52–65
1992 11 17 6 20–26 54–60
1993 14 23 7 20–38 64–82
1994 15 24 4 14–25 57–68
1995 17 25 2 15–21 59–65
three females would have surv i ved to January 1995. By adjusting adult puma
numbers in the Re f e rence Area, Ja n u a ry point estimates during 1989–1995 we re
0.50–0.90 per 100 km2 for males; 0.51–1.41 per 100 km2 for females; and
1.02–2.31 per 100 km2 for all adults. Hypothetically then, maximum adult male
densities in the Re f e rence Area (January 1995) and realized densities in the Tre a t-
ment Area (January 1991) would have been equivalent (0.90 males / 100 km2).
Howe ver, in 1995 the female density in the Re f e rence Area (1.41 females / 100
km2) would have exceeded the maximum female density realized in the Tre a t-
ment Area in 1994 (1.31 females / 100 km2), and the maximum total adult den-
sity in the Re f e rence Area (2.31 adults / 100 km2) would have exceeded the max-
imum adult density realized in the Treatment Area (2.10 adults / 100 km2). Still,
the maximum hypothetical adult puma densities in the Treatment and Reference
Areas were ve ry similar. We believe these hypothetical Re f e rence Area puma den-
sities are sensible given that the population there was not experimentally reduced,
as was the Treatment Area population, and thus had a longer uninterrupted
period to grow. These human-caused deaths we re also additive because they
apparently suppressed adult puma density.
Our estimates of adult puma density, based on the number of adult pumas
present year-round (January–December) in a 100-km2 area, increased over time.
The annual density on the Treatment Area increased from 1.36 to 2.01 adults
per year per 100 km2 during the pre-treatment years (1988–1990) and from
1.09 to 1.87 adults per year per 100 km2 during the treatment and post-treat-
ment years (1991–1994). In the Reference Area, density of adult pumas
increased from 1.13 to 1.79 during 1989–1994 (Table 10-5).
As would be expected, our adult puma density estimates for the biological
Table 10-5. Yearly (January–December) adult puma density

estimates for the Treatment Area (1988–1994) and Reference Area
(1989–1994), San Andres Mountains, New Mexico.
Pumas / yeara Pumas / year / 100 km2
Year Males Females Total Males Females Total
Treatment Area
1988 5.14 4.46 9.60 0.73 0.63 1.36
1989 4.91 7.48 12.39 0.70 1.06 1.76
1990 5.55 8.57b 14.12b 0.79 1.22 2.01
1991 2.67 4.96 7.63 0.38 0.71 1.09
1992 3.66 6.96 10.62 0.52 0.99 1.51
1993 4.45 8.62 13.07 0.63 1.23 1.86
1994 4.82 8.33 13.15 0.69 1.18 1.87
Reference Area
1989 7.14 8.10 15.24 0.53 0.60 1.13
1990 7.62 10.49 18.11 0.56 0.77 1.33
1991 8.41 11.25 19.66 0.62 0.83 1.45
1992 8.77 13.35 22.12 0.65 0.98 1.63
1993 9.28 14.67 23.95 0.68 1.08 1.76
1994 9.10 15.20 24.30 0.67 1.12 1.79
aPumas / year = sum of proportions of a given year that each adult puma was present
on a particular area (see Chapter 5 for details).
bAssuming that the one adult female we removed on 9 December 1990 would have
been present to 31 December, females / year would equal 8.63, and the total would
equal 14.18. We used these values in rates-of-increase calculations for 1990 to con-
trol for removing the puma.
year (August–July) also showed increasing trends. In the Treatment Area, bio-
logical year density increased from 1.24 to 1.89 adults per year per 100 km2 in
pre-treatment years (1987–1988 to 1989–1990), and from 1.12 to 1.83 adults
per year per 100 km2 in post-treatment years (1991–1992 to 1993–1994). Ref-
erence Area biological year density estimates increased from 0.94 to 1.73 adults
per year per 100 km2 during 1988–1989 to 1992–1993, and declined slightly
to 1.71 adults per 100 km2 in 1993–1994 (Table 10-6).
Clearly, puma deaths caused by predator control and sport-hunting during
1979–February 1985 severely suppressed the puma population on the San
Andres Mountains. Of the fifty pumas that were killed, about forty were adults.
In the Treatment Area, fourteen were killed for predator control and three by
Table 10-6. Biological year (August–July) adult puma density

estimates for the Treatment Area (1987–1988 to 1993–1994)
and Reference Area (1988–1989 to 1993–1994), San Andres
Pumas / yeara Pumas / year / 100 km2
Year Males Females Total Males Females Total
Treatment Area
1987–88 4.77 3.96 8.73 0.68 0.56 1.24
1988–89 5.44 6.84 12.28 0.77 0.97 1.74
1989–90 4.72 8.58 13.30 0.67 1.22 1.89
1990–91 4.20 7.12 11.32 0.60 1.01 1.61
1991–92 2.91 5.02 7.93 0.41 0.71 1.12
1992–93 4.34 8.20 12.54 0.62 1.17 1.79
1993–94 4.17 8.75 12.92 0.59 1.24 1.83
Reference Area
1988–89 6.07 6.63 12.70 0.45 0.49 0.94
1989–90 7.66 10.19 17.85 0.56 0.75 1.31
1990–91 8.23 10.48 18.71 0.61 0.77 1.38
1991–92 8.34 13.17 21.51 0.62 0.97 1.59
1992–93 9.40 14.10 23.50 0.69 1.04 1.73
1993–94 8.57 14.62 23.19 0.63 1.08 1.71
aPumas / year = sum of proportions of a given year that each adult puma was present
on a particular area (see Chapter 5 for details).
sport-hunters. And in the Reference Area, twenty fell to predator control and
three to sport-hunters. Overall, this kill rate of 11.2 adult pumas per year
exceeded the recruitment rate of 8.4 pumas per year that we observed during
February 1990–February 1995. The rate of removal by predator control alone,
9.5 adults per year, also exceeded the observed recruitment rate. Moreover, after
February 1985, when the puma population was protected for our research, the
puma population increased. It took from three years (in the Treatment Area) to
ten years (in the Reference Area) for the puma population to reach the relatively
high density of about two adults per 100 km2.
We believe the pattern of a lower puma density in the Reference Area rela-
tive to the Treatment Area was affected principally by three factors: intensive
predator control on the Oscura Mountains, habitat quality, and human-caused
mortality. During October 1980–October 1981, a total of thirteen pumas,
including eleven adults, were killed for predator-control reasons on the 673-km2
Oscura Mountains, which are just to the north of the Reference Area. The kill
rate of 1.6 adults per 100 km2 per year probably severely reduced the puma pop-
ulation on those mountains; consequently, dispersal of subadults from the
Oscura Mountains and the attendant immigration of pumas into the Reference
Area would have been reduced. The northern portion of the Reference Area,
from Silvertop Mountain north to Mockingbird Gap, consists of a fragmented
assemblage of mountains and low hills separated by desert flats and basins. In
contrast, the Treatment Area is relatively contiguous. In addition, the southern
portion of the Reference Area, from Hembrillo Canyon north to Cottonwood
Canyon, appeared to have a lower abundance of mule deer than we observed in
other parts of the San Andres Mountains. We believe this was influenced pri-
marily by the high distribution of tall and low mixed desert shrub and grass
communities in those canyons that offered relatively low-quality deer foods.
Both fragmented habitat and lower prey abundance would lower habitat quality
for pumas. As we explained above, the deaths of seven adult pumas at the hands
of humans during our research also probably lowered the adult puma density.
Trying to compare puma densities on the San Andres Mountains with den-
sities estimated in other puma populations is risky. Different studies have
applied different estimation methods and varying degrees of intensity in research
efforts, and therefore yield a varying degree of accuracy. For example, biologists
in Alberta used intensive capture-mark-recapture and radiotelemetry techniques
to quantify the puma population (Ross and Jalkotzy 1992), while biologists in
Southern California relied about equally on radiotelemetry and tracks presum-
ably of non-tagged pumas to estimate the total population (Neal et al. 1987). In
addition, puma study areas are usually contiguous with other puma habitat,
causing population density estimates to be affected by individuals having partial
spatial and temporal use of the study area (Neal et al. 1987). Studies that exerted
intensive capture-mark-recapture and radiotelemetry techniques on study areas
where puma movements were restricted by deep snow in winter, or where habi-
tat conditions caused natural constraints in the distribution of pumas (e.g., the
San Andres Mountains surrounded by desert basins), have provided the most
accurate estimates of population composition and density. Therefore, we limited
our comparisons to those.
Ranges of adult puma density estimates made in January each year (adults /
100 km2) on the Treatment Area and Reference Area were most similar to the
range of adult densities reported for a lightly hunted puma population in cen-
tral Idaho (Table 10-7). A moderately hunted population in north-central
Wyoming had densities at about the middle of our range. An Alberta popula-
tion with light to moderate hunting pressure with three interceding years of zero
off-take had densities in our upper range. Yet in southern British Columbia,
where pumas were not hunted but suffered significant accidental mortality from
automobiles, densities were in our low range. Only the unhunted population in
south-central Utah exhibited a density well below our range and that of all other
studies. Our total puma density ranges include the ranges for the Idaho and
British Columbia puma populations. The Alberta and Wyoming populations
reached slightly higher extremes. Again, the Utah population had a lower range
for total population density than all other studies.
Some similarities in puma densities were apparent even though the environ-
ments where studies were conducted were different. The puma population on
the San Andres Mountains was not influenced by severe winter weather that
caused seasonal population concentrations or migrations by pumas and their
prey. Thus, these factors did not trigger within-year fluctuations in abundance
of adult pumas. However, puma populations in Idaho, Wyoming, Alberta,
British Columbia, and Utah were censused in winter when pumas and their
ungulate prey were concentrated on low-elevation winter ranges because deep
snow constricted their movements at higher elevations. As snow receded in sum-
mer, pumas followed ungulate prey to higher elevations. As a result, puma home
ranges either enlarged or pumas occupied summer home ranges distinct from
winter home ranges (Seidensticker et al. 1973, Hemker 1982, Logan and Irwin
1985, Ross and Jalkotzy 1992, Pierce et al. 1999). Puma population densities
have not been estimated on summer ranges in those environments.
Identifying reasons why puma populations are similar or different in dis-
parate areas is difficult. Yet, such efforts are important to help us understand fac-
tors affecting density. In general, it is logical to consider that densities would be
affected by variations in prey biomass—essential energy required to fuel pumas.
For example, studies of wolves (Fuller 1989, Messier 1995) and leopards
(Stander et al. 1997) have shown that prey density (i.e., prey biomass) is a very
important predictor of densities of these carnivores. But because of complica-
tions in attaining estimates of puma prey biomass, comparative data do not
exist. Therefore, we could not make inferences about its effect on density across
the sample of study areas. Other limiting factors such as predator control and
sport-hunting that occurred around practically all of these study areas to various
degrees might have affected density of pumas thereon by influencing the num-
ber of dispersers available as potential recruits. In addition, pumas use habitat
features selectively to attain food and security for offspring (Logan and Irwin
1985, Belden et al. 1988, Laing and Lindzey 1991). Structure of puma habi-
tat—its continuity or fragmentation, its physiography and vegetation, and how
these characteristics influence prey distribution, abundance, and vulnerability to

predation—also affect density.
Rates of Population Increase

We were able to estimate realized rates of puma population growth because we
obtained annual censuses of a puma population that was protected after it was
severely reduced by predator control actions during the early 1980s. We also
experimentally removed 53 percent of the adult pumas and 58 percent of inde-
pendent pumas (i.e., adults plus independent subadults) from the Treatment
Area, and then monitored recovery. At the same time, we quantified rate of pop-
ulation increase for the protected Reference Area. The severe reduction in the
puma population before our research and our experimental removal in the Treat-
ment Area, coupled with protection of the puma population and its prey during
our research, should have given us a good indication of the maximum realized
rate of increase of the puma population in the prevailing environmental condi-
tions. We calculated observed exponential rates of increase for adult pumas in
the Treatment and Reference Areas separately using three population density
estimators: (1) January point estimate, (2) each annual period January–Decem-
ber, and (3) each biological year August–July (Table 10-8). Observed exponen-
tial rate of increase (r) was calculated for each estimator by linear regression of
the natural log of the estimated number of adult pumas each time period (i.e.,
the dependent variable) on the units of time numbered sequentially starting
with zero (i.e., the independent variable) (Caughley 1978, Van Ballenberghe
1983). The slope of the regression line is r; it represents the average rate at which
the puma population changed per year. The puma population grows if r is pos-
itive; it declines if r is negative.
Puma population growth rates are an index to how interactions of popula-
tion density and habitat quality affect reproduction, mortality, immigration,
and emigration. Several patterns in r emerged that we explain with biological
and environmental phenomena. We also ventured some predictions based upon
our knowledge of puma population dynamics and behavior:
1. The puma population grew relatively rapidly after it was severely reduced,
then protected, during a time when per capita food resources (i.e., princi-
pally deer) increased (refer to Table 10-8). Rates of increase for all adults
were highest during time spans beginning with low puma density, regard-
less of the area or estimator. This indicates that puma populations have a
high potential to increase when intraspecies and interspecies competition
for resources and other limiting factors (e.g., human-caused mortality) are
weak. In addition, given the high puma population growth rate in the Treat-
ment Area during the deer decline, the puma’s numerical response lagged
behind the numerical response of the prey (see Chapter 17).
2. Maximum observed rates of increase in the Treatment Area and the Refer-
ence Area were variable. January point estimates for the Treatment Area pre-
(r = 0.21) and post-treatment (r = 0.28) and the Reference Area during
1989–1992 (r = 0.17) averaged 0.22 ± 0.06 (Fig. 10-1).
3. The Reference Area growth rate during 1989–1995 (r = 0.11) was for the
longest time span for which any puma population growth has been quanti-
FIGURE 10-1. Observed exponential rates of increase for adult pumas in the Treatment Area
(TA) and Reference Area (RA) using January population estimates, San Andres Mountains,
New Mexico.
fied (Fig. 10-1). Moreover, it represents a period with varied ecological con-
ditions. Relatively high growing-season precipitation and an increasing
mule deer population characterized years 1985–1991, but growing-season
drought and a declining deer population characterized years 1992–1995
(see Chapter 17). Additionally, potential immigration of progeny recruits
was reduced due to the experimental removal of pumas from the Treatment
Area.
4. Re f e rence Area growth rate during 1992–1995 declined (r = 0.05). We
believe the reduced r was affected primarily by at least three factors: a decline
in food re s o u rces, reduced re c ruitment of immigrants from the Treatment
Area, and seven human-caused deaths. We would expect intraspecies com-
petition for food to be the highest during times of declining prey resources
and increasing puma density. The puma population continued to increase,
albeit at a slower rate, creating a time lag. Eventually, we would expect the
puma population to decline and be subjected to mechanisms resulting in
lower puma surv i val rates, lower reproductive rates, greater subadult female
dispersal, and possibly lower settlement rates of immigrants. Unfortunately,
our re s e a rch ended too soon for us to observe how the puma population
responded. No tagged progeny born in the Treatment Area were recruited
into the Re f e rence Area during this period, while three tagged progeny born
in the Re f e rence Area were re c ruited in the Treatment Area. This indicates
that severe reduction of pumas in one area can affect population growth in
an adjacent area. Without population reduction in the Treatment Area
(including six adult females), we would have expected more progeny to be
available as potential re c ruits into the Re f e rence Area.
5. To estimate the effect of seven adult puma deaths (one male, six females)
due to human causes on the Reference Area, we recalculated rates of
increase using hypothetical January point estimates assuming that the seven
pumas would have been subjected only to natural causes of death. With this
adjustment, the r for all adults during 1989–1995 was 0.11 ± 0.02 (SE ),
identical to the realized rate (compare with Table 10-8). In each four-year
segment, rates increased slightly. During 1989–1992, r for all adults was
0.19 ± 0.05, and during 1992–1995 it was 0.06 ± 0.04. For adult males
during 1989–1995, r was 0.09 ± 0.01. Male r during 1989–1992 remained
exactly the same because there was no change in density, but during
1992–1995, r i n c reased to 0.11 ± 0.05. For adult females during
1989–1995, r declined slightly to 0.13 ± 0.04. During 1989–1992, female
r increased slightly to 0.28 ± 0.09, and during 1992–1995 it remained the
same at 0.03 ± 0.04. Although the seven deaths reduced density, they appar-
ently occurred too infrequently to change the rate of population increase
substantially. Still it is possible that the deaths of the adult females in par-
ticular might have affected r more than we realized. Had they lived, some
of their progeny, especially females, might have been recruited into the Ref-
erence Area population. This could have boosted density and rates of
increase, unless density-dependent mechanisms increased in strength (e.g.,
greater female emigration).
6. Adult male puma rate of increase based on January point estimates for the
Treatment Area during 1988–1991 was affected by the removal of one adult
male in 1989. Assuming that he would have survived to January 1991,
adult male r would have increased substantially to 0.10 ± 0.05 (compare
with Table 10-8) and would be closer to the r for adult male pumas in the
Reference Area during 1989–1992. However, r for all adults in the Treat-
ment Area increased only slightly to 0.23 ± 0.04.
7. Adult female rates of increase were generally higher than for males. This
phenomenon is due to the tendency for some female subadults, which have
relatively high survival rates, to become philopatric adults. Inversely,
subadult males have lower survival rates and most emigrate. Adult male
recruitment into the local population relies primarily upon immigrants
coming from some other population.
8. Female rates of increase appear to be sensitive to puma density and trends
in deer numbers. Female rates were highest during the period of low puma
density and increasing deer numbers. However, female rates declined dur-
ing deer loss. Decline in the rate was particularly precipitous in the Refer-
ence Area. The greatest changes in puma population density appear to occur
in the adult female segment. We hypothesize that during times of food
abundance, philopatry in surviving female progeny is relatively high, and
dispersal is low. But when food is limiting, female dispersal rates increase.
Hence, competition for food may regulate dispersal in surviving female
progeny.
9. Male rates of increase in the Treatment Area did not parallel the declining
deer numbers, but in the Reference Area male rates of increase were about
equivalent in the January point estimate and declined in the other two esti-
mators. The tendency for male r in the Treatment Area population to be less
sensitive to changes in food abundance may be related to two factors. First,
the adult male takes no part in raising offspring; hence, his total energy
demand may be up to one-third that (estimated in kcal / day) of a female
raising cubs (Ackerman 1982). Secondly, males were establishing in a pop-
ulation with a drastically reduced puma density, and hence competitors,
which resulted from removal. In the Reference Area, we hypothesize that
the decline in male rates of increase for the two year-round estimators may
be the result of increased male-to-male competition for mates.
10. The January–December and biological year (i.e., August–July) estimates of
r, which represented the number of adult pumas present throughout each

twelve-month period, indicated declining rates of increase for all adult cat-
egories in the Reference Area during deer decline years. In the Treatment
Area, the rate of female population growth declined during the deer popu-
lation decline. This would be expected if puma density was ultimately lim-
ited by food.
11. For all adult categories, rates of population increase generated by the Janu-
ary point estimator were fairly consistent with rates from the biological year
estimator ending five months prior.
Other than our estimates, quantitative data on rates of increase of puma pop-
ulations are practically nonexistent in scientific literature. However, in examin-
ing population estimates from other intensive, long-term studies on puma pop-
ulation dynamics, we found that biologists in southern Alberta (Ross and
Jalkotzy 1992) and south-central Utah (Lindzey et al. 1994) presented data
showing increasing populations. We used their data to calculate observed expo-
nential rates of increase for adult pumas in each population to compare with our
findings. In Alberta, population estimates were made on 31 March each year;
biologists defined adults as pumas that were older than 2.5 years. During three
consecutive years (1984–1986) the number of adult pumas on the 780-km2
study area increased from twelve or thirteen to fourteen. The r ranged from 0.04
± 0.02 (SE ) to 0.08 ± 0.002. This was a hunted population where three pumas
were killed the first year, representing 16–20 percent of the estimated number
of adults and subadults (i.e., pumas 1.6–2.5 years old). But no pumas were
killed the following two years. Density of adult pumas during the three years
ranged from 1.5 to 1.8 adults per 100 km2, which was within the high range of
densities that we found on the San Andres Mountains (Table 10-7). In Utah,
population estimates were made during January–March each year. Biologist
there defined adults as pumas more than 1.5 years old that showed continuous
use of a predictable area for more than six months. In four consecutive years
(1984–1987) during which no adult pumas were removed, the number of adults
on the 1,900-km2 study area increased from six to twelve, and r was 0.24 ± 0.03
(SE ). Density of adult pumas during the four years ranged from 0.32 to 0.63
per 100 km2, well below the lowest densities on the San Andres Mountains.
Given the different criteria for defining an adult puma by the Alberta and
Utah researchers, direct comparisons of rates of increase with those we calculated
for the San Andres Mountains are tenuous. There may have been some
“subadults” in the Alberta population that were sexually mature and hence met
our definition of adult. On the other hand, some individuals in the Utah study
may have been subadults. Still the range of ages of adult pumas on the San
Andres Mountains certainly included the range of ages of pumas classed as

adults in both the Alberta and Utah studies. It is at least noteworthy that the
rates of increase were inversely related to puma density. In other words, the rel-
atively high-density Alberta puma population exhibited the lowest rate, while
the very-low-density Utah population had the highest rate. This was the basic
relationship that we found in the San Andres Mountains. As we mentioned pre-
viously, the lower rate of population increase associated with higher puma den-
sities in the Reference Area was partially affected by a declining mule deer pop-
ulation, low recruitment of pumas from the Treatment Area, and human-caused
deaths.
It is important to realize, however, that some puma populations apparently
do not grow during extended periods even though their prey base does. The
lightly hunted puma population studied over eight years in the central Idaho
wilderness declined slightly due to puma deaths and emigration and coincident
with increasing mule deer and elk populations, which constituted the puma’s
major prey (Hornocker 1970, Seidensticker et al. 1973). The biologists hypoth-
esized that the puma social organization, via a land tenure system, limited the
density of adult pumas (Seidensticker et al. 1973). The adult segment of the
puma population in Utah was relatively constant for the first seven years
(1979–1985), changing by ± 0–2 adults per year even though the deer popula-
tion increased throughout the study. But the magnitude of the deer population
increase was not ascertained directly. Biologists found only a weak relationship
between puma numbers and a fecal pellet group index to deer abundance. They
suggested this might support the hypothesis initially established by the Idaho
researchers, or it was possible that the deer population did not increase enough
to cause a numerical response in the puma population (Lindzey et al. 1994).
Metapopulation Dynamics
A fundamental concept we developed about the puma population on the San
Andres Mountains is that it is a subpopulation (i.e., a local population) in a con-
stellation of puma subpopulations in the Southwest (Sweanor et al. 2000). The
basin and range physiography of the landscape naturally fragments puma habi-
tat and the attendant puma population. Pumas live mostly in the mountains and
foothills where there are large prey, including desert mule deer, desert bighorn
sheep, javelina, white-tailed deer, elk, and a variety of smaller prey, such as wild
turkey and porcupine. But in the broad intervening areas of non-habitat, or
matrix, the desert is so harsh that prey occurs in extremely low densities or not
at all. Hence, puma longevity there is low, as is their ability to successfully raise
cubs. By studying the movements of dispersing pumas, we learned that emi-
grants were a critical link between puma subpopulations and they contributed
substantially to subpopulations. In the San Andres Mountains, about 78 percent
of male recruits and 32 percent of female recruits were immigrants that made
their way from other subpopulations. We documented that at least 45 percent
of immigrant recruits produced offspring, including males and females. Simi-
larly, emigrants from the San Andres Mountains were contributing recruits to
other puma subpopulations. Because of these dynamics, we inferred that the
puma population structure in southern New Mexico generally met the defini-
tion of a metapopulation—“a network of semi-isolated populations with some
level of regular or intermittent migration and gene flow among them, in which
individual populations may go extinct but can then be recolonized from other
populations” (Meffe and Carroll 1997:678).
Even though each subpopulation is dependent upon the immigration of
pumas for numeric and genetic augmentation, we expect that the dynamics of
each subpopulation is not necessarily correlated with other subpopulations.
Pumas on the San Andres Mountains were principally dependent upon mule
deer for food; consequently, a steep decline in deer abundance, triggered initially
by severe drought, would likely result in a decline in puma numbers, albeit after
a lag period (see Chapter 17). The same drought would not necessarily affect
puma numbers in the same way in the Sacramento Mountains across the
Tularosa Basin to the east, where the climate was somewhat wetter and the envi-
ronment supported both mule deer and elk as principal puma prey. The same
could be said for the Black Range and the wilderness mountains of the Gila
River drainage in southwestern New Mexico where mule deer also declined, but
elk and javelina numbers seemed stable or even increased. In those areas, pumas
could alter their hunting patterns to kill the more locally abundant prey. If
pumas switched to using elk and javelina more frequently than they did when
mule deer were more abundant, the puma population might not decline at all.
Pumas could also switch to killing domestic livestock, and thereby maintain
their numbers (Cunningham et al. 1995).
Furthermore, management practices impact puma population dynamics.
Pumas and large prey on the San Andres Mountains were mostly protected from
human off-take for the sake of our research, and therefore the population was
mostly affected by natural dynamics. But sport-hunting pressure in the Sacra-
mento Mountains and especially in the Black Range–Gila complex could have
been impacting puma population dynamics by altering mortality rates, although
the extent was unknown. In some regions, management objectives are to pur-
posely reduce puma density to minimize predation on domestic livestock, such
as in the Guadalupe Mountains in southeast New Mexico. On the other hand,
local puma population densities can be reduced inadvertently by chronic annual
heavy hunting. Yet other puma subpopulations may be stable or growing as a

result of relatively light hunting pressure.
Differential dynamics where some subpopulations are increasing while oth-
ers are declining create a source-sink metapopulation structure. Source subpop-
ulations are increasing or stable; recruitment via local reproduction and immi-
gration outpace mortality. Source populations produce emigrating progeny that
immigrate into other subpopulations, augmenting them numerically and genet-
ically, and can rescue declining populations or colonize empty habitat patches.
Stable populations, especially if they have high densities, can be sources of emi-
grants that can be recruited into other subpopulations. Source populations, par-
ticularly if they are large or persistent, stabilize the metapopulation. In contrast,
sink populations are those where mortality outpaces recruitment; thus, puma
numbers decline. Comparatively, they contribute few emigrating progeny as
potential recruits for other subpopulations. If not rescued, sink subpopulations
could go extinct. Large sinks could affect the dynamics of nearby subpopula-
tions by reducing potential recruits that might otherwise offset or exceed mor-
tality, and contribute to population decline over a larger region. Furthermore,
immigrants to sink populations have lower survival rates and reduced chances of
reproducing thereon or escaping to other more secure subpopulations. Sink sub-
populations could destabilize the metapopulation. Whether they do or not will
probably depend upon their size, number, and distribution.
As emigrants move from one subpopulation to another and breed, genetic
structure of pumas in the metapopulation is affected. High rates of gene flow
among populations are possible in carnivores that exhibit high rates of dispersal
and disperse long distances (Wayne and Koepfli 1996). Male pumas in particu-
lar, because of their tendency to disperse long distances and across matrix,
enhance nuclear gene flow between widely separated subpopulations, including
ones with vastly different habitats (e.g., from the Chihuahua Desert to conifer-
forested mountains and vice versa). Our radiotelemetry data on a sample of
study area progeny indicated that most (seventeen of twenty females, eight of
thirteen males) did not disperse more than five home-range diameters, the dis-
tance that may be necessary to disrupt genetic adaptations to regional environ-
mental conditions (Shields 1982). Yet, the few progeny that did disperse beyond
those distances could promote an outbreeding population structure (Templeton
1987). Outbreeding can introduce new alleles to a population, thereby increas-
ing genetic variation, with the prospects of enhancing fitness of individuals in
contemporary or future environments. Outbreeding may be a vital reason why
pumas have adapted so well to a broad range of habitats in the Western Hemi-
sphere. Puma immigration on the San Andres Mountains greatly exceeded the
one-to-five per generation (Sweanor et al. 2000:Table 2), which is suggested by
genetic theory to ensure effective transfer of genetic variation (Crow and Kimura
1970, Lacy 1987). With such high rates of gene flow, we expect little genetic
subdivision among puma subpopulations in southern New Mexico.
In a source-sink structure, dispersing pumas have critical consequences to
the persistence of subpopulations. They can reduce the size of each subpopula-
tion needed to maintain viability (Beier 1993). Furthermore, environmental
variation, such as weather patterns that trigger severe prey depletion, or localized
severe puma exploitation, is not as powerful an extinction force when popula-
tions are spread across space (Shaffer 1987, Noss et al. 1996). We estimated the
San Andres Mountains chain (about 3,000 km2 of puma habitat), which
includes the San Andres Mountains plus the Organ Mountains to the south and
the Oscura Mountains to the north, supported thirty-six to sixty adult pumas
during 1985–1995 (Sweanor et al. 2000) and a genetically effective population
size (Ne—based on the actual number of breeding individuals and the distribu-
tion of offspring among families, Meffe and Carroll 1997) as low as half the
adult density (Nunney and Elam 1994). This area easily exceeds the
1,000–2,000 km2 critical patch size needed for a 98 percent probability of per-
sisting for one hundred years (Beier 1996). Not surprisingly, we learned through
model simulations that the San Andres chain should be able to withstand the
threat of extinction for one hundred years as long as it supports the puma demo-
graphics that we documented (Sweanor et al. 2000). Yet, to achieve recruitment
rates we observed, the population relied upon both progeny born within the
subpopulation and immigrants from other subpopulations. Smaller subpopula-
tions inhabiting smaller habitat patches, such as the Caballo (about 200 km2)
and Fra Cristobal (about 100 km2) Mountains, would have much smaller Ne.
Without gene flow via immigrants, such populations could eventually lose
genetic variability by genetic drift or inbreeding and hence the ability to adapt
to altered environmental conditions (Chepko-Sade et al. 1987). Eventually, they
could go extinct (Saccheri et al. 1998), particularly populations that undergo a
rapid decline in size and that lack histories of severe fluctuations (Gilpin 1987,
Lande 1988). Even with gene flow, smaller habitat patches have high probabili-
ties of puma extinction. We found that twelve of one hundred simulated puma
populations went extinct during a 100-year period for patch areas the size of the
Organ Mountains (about 270 km2), even when using the productive population
parameters and high immigration rate for pumas on the San Andres Mountains
(Sweanor et al. 2000).
Still, this does not mean that large puma habitat patches, such as the Sacra-
mento Mountains (about 6,200 km2), are immune to extinction. Certainly, if
negative population growth rates persist in large patches and surrounding
patches to the extent that immigration does not offset mortality, then pumas in
large patches can disappear. This may have occurred in the Sacramento Moun-
tains in recent history. By the 1940s, state-sanctioned bounties and unregulated
killing had confined pumas in New Mexico mainly to natural refuges in the
remote mountains west of the Rio Grande (Young 1946) where hunters could
not kill out the last pumas as they were able to do with the Mexican wolf and
grizzly bear (Brown 1984, Brown 1985). However, regulations on puma killing
since 1972 probably allowed remaining pumas to increase in numbers and to
produce dispersers that recolonized or stabilized patches east of the Rio Grande.
On a much grander scale, we know that pumas went extinct from almost all of
their eastern North American range during the early 1900s. And even more
remarkable, if Melanie Culver (1999) is right, pumas in North America may
have gone extinct at the end of the Pleistocene; then immigrants coming from
South America recolonized the continent.
Puma population dynamics and life history strategies are inextricably linked.
Clearly they contribute to the puma’s high adaptability. As we have pointed out
in this chapter, they are influenced greatly by the environment and the way
pumas interact with one another. In Part 3 we discuss adaptations in puma
behavior and social organization.
LP > > KX
1. We experimentally removed pumas from the 703-km2 Treatment Area dur-
ing 9 December 1990–22 June 1991. The entire population was reduced by
47 percent. Adults were reduced by 53 percent, and independent pumas
(i.e., adults plus subadults) were reduced by 58 percent. Post-treatment, it
took thirty-one months for the adult segment of the population to recover
to its pre-removal density. Adult pumas were replaced by same sex recruits,
thus restoring the pre-removal sex ratio. Recruits came from within the
Treatment Area, the Reference Area, and from other puma populations.
2. We estimated puma density using three estimators: (1) a point estimate of
adult pumas and total pumas in January of each year, (2) an estimate of the
number of adult pumas present year-round from January to December, and
(3) the number of adult pumas present in each biological year from August
to July. January point estimates for the Treatment Area pre-treatment and
treatment years increased from 1.16 to 2.10 adults per 100 km2 and total
pumas ranged from 2.01 to 3.91 per 100 km2. In the post-treatment years
pumas increased from 0.84 to 1.99 adults per 100 km2 and total pumas
ranged from 2.78 to 4.25 per 100 km2. Reference Area density increased
during the seven-year period 1989–1995 from 0.94 to 2.01 adults per 100
km2 and total pumas ranged from 1.72 to 3.90 per 100 km2. Densities of
adult and total pumas were in the moderate to high range for pumas in
other western North America puma populations. The predator control and
sport-hunting that occurred during 1979–1985 prior to our research had
suppressed the puma population. Off-take exceeded the recruitment rate
that we observed.
3. Except for our experimental removal of pumas from the Treatment Area,
the puma population on the San Andres Mountains increased. Using Janu-
ary point estimates in the Treatment and Reference Areas, the average max-
imum observed exponential rates of increase (r) for adult pumas was 0.22 ±
0.06. For a seven-year span in the Reference Area that included the deer
increase and decline phases, r was 0.11. During the mule deer population
decline, r in the Reference Area was 0.05. Adult females had a tendency to
grow at a faster rate than adult males. Female recruitment was more sensi-
tive to the deer decline than male recruitment.
4. We inferred that pumas living in the basin and range configuration of habi-
tat in the Southwest form a source-sink metapopulation structure. Subpop-
ulations are linked to one another via dispersing pumas. Immigrants are
vital for the numeric and genetic augmentation of subpopulations. Large
source populations contribute to metapopulation persistence, while sink
populations contribute to instability. Actual dynamics of the metapopula-
tion will probably depend upon the size and distribution of source and sink
populations. Gene flow between subpopulations probably enhances the
puma’s ability to adapt to a wide range of environmental conditions and
should result in little inter-subpopulation genetic subdivision.
P a rt I I I
Puma Behavior and

Social Organization
Chapter 11
How Should Desert Pumas Behave?
Social organization is the manner in which individuals of the same species inter-
act and are arranged in space and time relative to one another. Although pumas,
and all other known species of cats except the African lion and cheetah (Schaller
1972, Caro 1994), live a solitary existence, this mode of life does not equate
with absence of social structure (see Leyhausen 1965). Solitary simply means
individuals do not cooperate with one another to rear young, forage, achieve
matings, or defend against predators (Sandell 1989).
Besides our descriptive study of puma social organization in the desert, we
examined two hypotheses that explain the function of the puma’s social system.
It is widely believed that the mating and spacing patterns we observe evolve from
the different behavioral strategies that males and females employ to maximize
individual reproductive success (Trivers 1972, Wrangham 1980, Sandell 1989,
Caro 1994). We call this the two-strategies hypothesis. It has also been postulated
that the social system functions to limit puma numbers within a given popula-
tion (Hornocker 1969, 1970, Seidensticker et al. 1973). We term this the self-
limiting hypothesis.
Two-Strategies Hypothesis
How should pumas behave to maximize individual reproductive success? In
pumas, as in other solitary carnivores, females typically provide greater parental
investment (i.e., pregnancy, lactation, uni-parental care) than males. Thus, a
female’s re p ro d u c t i ve success should principally be limited by access to
resources, her own reproductive condition and survival, and the survival of her
183
184 PART III. PUMA BEHAVIOR AND SOCIAL ORGANIZATION
cubs. Female pumas should compete more with one another than with males for
resources that are necessary to successfully rear offspring, while at the same time
avoiding potentially dangerous pumas. In contrast, unencumbered males are
free to seek breeding opportunities because female pumas can breed year-round.
But because receptive females are relatively rare at any point in time, a male
should compete with other males for mates. His reproductive success is limited
by his own reproductive condition and longevity, the number of females to
whom he has access, and by other males with whom he has to compete (Emlen
and Oring 1977; Wrangham 1980, Macdonald 1983, Erlinge and Sandell 1986,
Clutton-Brock 1989, and Chapters 7 and 8).
If male and female pumas on the San Andres Mountains were conforming
to these two strategies to maximize individual reproductive success, the follow-
ing predictions should hold:
1. Each adult male should attempt to breed with as many females as possible.
Males should maintain large home ranges to increase encounter rates with
females.
2. Males should treat females as a critical defendable resource because breed-
ing opportunities are rare (see Chapter 6). Hence, males should fight for
breeding opportunity.
3. Removal of male competitors should allow remaining males to expand their
home ranges to breed with more females. Removal of females should cause
males to expand or shift their home ranges to seek breeding opportunities.
4. Females should attempt to successfully raise as many offspring as possible.
To do this they must maximize their reproductive attempts and survival of
themselves and their young (see Chapter 7) by avoiding competitors and
evading predation.
5. Female home range fidelity should be high even when other adults are
experimentally removed. This way, females cultivate knowledge of vital
resources and minimize dangerous encounters with other predators and
competitors.
6. If resources allow, females should share portions of their range with female
offspring. This would increase inclusive fitness.
7. We expect reduced intersexual competition because critical resources are
different for the sexes. Therefore, adult pumas should exhibit more spatial
exclusion within than between sexes.
Self-Limiting Hypothesis
Hornocker’s and Seidensticker’s research in central Idaho laid the foundation for
what is understood about puma social organization, and it introduced the con-
CHAPTER 11. HOW SHOULD DESERT PUMAS BEHAVE? 185
cept of self-limitation for the species (Hornocker 1969, 1970; Seidensticker et

al. 1973). After five years of study, Hornocker described a relatively peaceful,
polygamous society, where adult pumas occupied territories (loosely defined by
Hornocker as an attraction to a particular locality) that were relatively exclusive
in males and overlapping in females. Mutual avoidance behavior, aided by visual
and olfactory cues, helped to distribute pumas in both space and time. This
peaceful mechanism for spacing individuals was adaptive, Hornocker argued,
because solitary predators cannot risk debilitating injuries that may result from
fighting. Hornocker (1970:37) concluded that “intraspecific relationships, man-
ifested through territoriality, acted to limit numbers of [pumas] and maintain
population stability.”
After three more years of research on the same puma population, during
which radiotelemetry techniques were employed, Seidensticker et al. (1973)
shied away from the term “territoriality,” instead using the less ambiguous, more
descriptive term “home area” to characterize the area over which resident pumas
roamed. Tenure of these home areas, they argued, was based on prior rights. A
puma could claim a home area only when it became vacant, and once it estab-
lished a home area, its simple occupancy would deter new individuals from set-
tling there. They surmised that the presence of adult resident males was the pri-
mary factor limiting adult male density, and the presence of resident, breeding
females limited the female breeding population. These conclusions were sup-
ported by the following observations: a resident male’s death resulted in the
establishment of a home range by a new immigrant male as well as a shuffling
of space by the remaining resident males; transient males rarely bred; no new
males moved in to claim ranges made vacant by the deaths of resident adult
females; and breeding in young adult females was apparently suppressed. Since
Seidensticker and colleagues did not observe a positive correlation between
puma density and prey numbers during their eight-year study (overall, the
puma’s major prey increased while the puma population declined slightly), they
also surmised that “the land tenure system maintains the density of breeding
adults below a level set by food supply in terms of absolute numbers of mule
deer and elk” (1973:59). After further examination of the social structure of
pumas, bobcats, and leopards, Hornocker and Bailey (1986:218) concluded that
the social “system’s primary function was population regulation through a spa-
tial distribution of individuals.” In other words, the social structure was adap-
tive because the predator did not harm itself by overpopulating and depleting its
food supply.
The idea that Hornocker (1969, 1970) and Seidensticker et al. (1973) devel-
oped has been the model for how many wildlife biologists and managers alike
view puma social organization and its influence on prey. Puma studies in
Alberta, British Columbia, and Utah (Ross and Jalkotzy 1992, Spreadbury et al.
1996, Lindzey et al. 1994, respectively) have described similar behavioral pat-
terns, although evidence of fighting (i.e., scarring in males and puma deaths)
indicated other puma societies were not as peaceful as the Idaho research sug-
gested. These studies did not refute Seidensticker et al.’s (1973) thesis that the
social system maintains the density of breeding adults below a level set by the
food supply. However, none could adequately test it. In Utah, Lindzey et al.
(1994) documented an increase in prey numbers while puma numbers remained
somewhat stable; unfortunately, they could not determine the magnitude of the
increase. In fact, they believed the increase in prey numbers may have been
insufficient to provide an adequate test of the self-limiting hypothesis.
Only one study, conducted in the Sierra Nevada of California on the distri-
butions of pumas and their major prey (Pierce et al. 2000b), has seriously chal-
lenged the self-limiting hypothesis. Pierce and colleagues surmised that for a
land tenure or territorial system to limit a puma population below the level set
by the prey, territorial individuals would have to sequester more prey than nec-
essary for reproduction, as well as limit the availability of prey to other pumas.
They tested this primarily by examining the distribution of pumas in relation to
the distribution of their major prey. They concluded that the puma’s land tenure
system did not limit the population of pumas in their study area via partition-
ing of prey; instead, the puma population was limited by food supply.
Because we were able to manipulate the puma population and we could
ascertain the trend in the deer population, we were able to develop some pre-
dictions for pumas on the San Andres Mountains if they conformed to the self-
limiting hypothesis.
1. The puma population exhibits stability when the prey base is increasing.
Stability is maintained through a peaceful, land-tenure mechanism. The
presence of resident adult males limits the density of resident males, and the
presence of resident breeding females limits the female breeding population.
Consequently, transient males should not successfully breed, and breeding
in young adult females should be suppressed. All progeny emigrate unless
space is made available within the population by the death of a same-sex
adult. There should be no increase in density, decrease in home range size,
or increase in spatial overlap to accommodate more adult pumas.
2. Given a stable or increasing prey base, new pumas will be able to claim
vacant ranges after experimental removal. New adults should reoccupy the
area in the same numbers, arrangement, and sex ratio as before the removal.
3. A substantial decline in the prey base should trigger a socially induced
adjustment to puma population density, such as emigration of resident
adults. Some vacated ranges (because of death or translocation) should not
CHAPTER 11. HOW SHOULD DESERT PUMAS BEHAVE? 187
be re-occupied by new pumas. Otherwise the puma population will hasten

the prey’s decline.
In the next four chapters, we examine the social organization of pumas in the
San Andres Mountains, and we test the ideas presented here. In the final chap-
ter of Part 3, we synthesize the information from studies across the puma’s range
to obtain a clearer understanding of the adaptive significance of the puma’s
social structure. We also examine the puma’s social structure in relation to other
large felids.
Chapter 12
Adult Home Range Characteristics
An animal’s home range is the area in which it chooses to live because it provides
vital services, including food, mates, and a secure place to rear young (Burt
1943). Home range implies predictable use of an area over time. However, home
range can vary seasonally as an animal moves between feeding or breeding areas;
it can also change as the animal matures. The size, constancy of use, and orien-
tation of the home range can give valuable insights into how pumas are influ-
enced by other pumas as well as their environment. In this chapter, we present
information on home range size and degree of home range fidelity in adult
pumas.
Delineating the Home Range

Although the definition of home range seems simple enough, delineating the
actual boundaries, and hence area, is difficult. We used two methods to estimate
home range: the minimum convex polygon (MCP; Hayne 1949) and the adap-
tive kernel (ADK; Worton 1989) methods in CALHOME software (Kie et al.
1994). We chose the MCP because of its graphic simplicity, wide historical use,
and ease of calculation (Jennrich and Turner 1969, White and Garrott 1990),
and because of its comparative value to the many other studies that have used
the MCP to describe puma home ranges. In the MCP method, the peripheral
locations of an animal are connected in such a way that the internal angles of
the polygon thus generated do not exceed 180 degrees. Because home-range size
calculated using the MCP is greatly influenced by outlier locations (Ackerman
et al. 1989, White and Garrott 1990), we calculated a 90 percent MCP (the
189
smallest area encompassed by 90 percent of the points) in addition to the 100

percent MCP. We also used the ADK, a non-parametric method for estimating
an animal’s utilization distribution (UD; Worton 1989, Kie et al. 1994), because
it was a flexible, probabilistic model that was considered to provide a more real-
istic representation of a home range (Worton 1989). The UD estimates where
an animal is during the times between locations as well as the amount of time it
spends in any particular place (Seaman and Powell 1996). Contours connecting
areas of equal densities of observations can describe any usage area of the home
range. For this study, we defined the home range and core area as the smallest
area containing 90 percent and 60 percent of the UD, respectively. Although
many studies report the 95 percent UD, we found that the distribution of puma
locations in the long, narrow San Andres Mountains caused the 95 percent con-
tour to balloon outside the study area boundary where pumas normally did not
range.
Of the 126 radio-collared pumas, we monitored eighty-six as adults on the
study area. On average, we obtained 137 locations (range = 1–569) over a mon-
itoring period of 28.8 months (range = 0–102) on each of these adults. Pumas
were known to be traveling during 5 percent of locations for which we could
identify activity; the rest were at day beds, nurseries, or prey caches. We gener-
Slide @318%
PHOTO 22. Aerial telemetry was critical to estimating puma home ranges and determining
puma interrelationships.
CHAPTER 12. ADULT HOME RANGE CHARACTERISTICS 191
ally located pumas that utilized areas surrounding one of our camps more fre-
quently than ones residing in other parts of the study area. Because home range
estimators are strongly affected by the number of locations used and time
between locations, we selectively chose location data to ensure successive loca-
tions were spatially independent and reliable comparisons between individuals
were made. Consequently, we calculated home ranges using locations from the
weekly flights (during which all radio-collared individuals were generally
located) augmented with ground locations, as long as all locations were more
than three days apart. This way we could bolster our aerial sample with ground
locations during the weeks we were unable to fly. Tests to determine independ-
ence of animal-movement data (Schoener 1981; Swihart and Slade 1985, 1986)
still indicated serial autocorrelations in some cases. We disregarded this for two
reasons: individual movements likely were dependent on past experience and
knowledge of resources in the home range, and pumas were capable of travers-
ing their home range in a single day. Home ranges can provide unbiased esti-
mates of space-use patterns when successive points are not independent statisti-
cally, as long as they are biologically independent (Lair 1987).
Seasonal and Annual Home Range

We did not calculate seasonal home ranges because our earlier analyses (Sweanor
1990) did not indicate puma home ranges were highly influenced by seasonal
changes. We found that an adult puma’s home range size did not change in any
predictable pattern based on the two seasons (spring-summer: April to Septem-
ber; fall-winter: October to March) that we delineated. Furthermore, pumas did
not exhibit a tendency to use seasonal home ranges. Instead, they typically used
the same area year-round. We expected these results, because the puma’s prey did
not migrate and winter movements were not restricted by deep snow.
Annual home ranges were calculated for each adult puma that was present
for at least ten months of any particular twelve-month period (1 January to 31
December). We used thirty-two to fifty-three (x– = 43.9 ± 3.9) locations to cal-
culate each annual home range. Although the ADK home range estimator
required a greater number of locations than did the MCP method, sample sizes
appeared adequate to depict annual home range sizes (Appendix 3, Fig. A3-1).
We obtained seventy-two annual estimates of home range size for twenty-
four adult male pumas and seventy-one annual estimates for thirty adult female
pumas (Appendix 3, Table A3-1). Annual home range size (90 percent ADK) for
males and females averaged 193.4 km2 (range = 59.3–639.6) and 69.9 km2
(range = 13.1–287.4), respectively (Fig. 12-1). Females used between 33 and 38
percent of the average area used by male pumas, depending on the home range
FIGURE 12-1. The means and standard deviations of annual, birth interval, and lifetime
home range sizes, based on the 90 percent adaptive kernel home range estimator, for adult
pumas on the San Andres Mountains, New Mexico, 1986–1994. Annual home range is
mean of means.
estimator used [1]. The larger home range size of the male was reflected in his
daily movements. Our earlier analyses (Sweanor 1990) found that males and
females traveled, on average, 4.1 km and 1.5 km (linear distance) between loca-
tions one day apart, respectively.
Birth-Interval Home Range

For females, we calculated home range over a biological time scale: an entire
birth interval (the birth date of one litter to the birth date of the next litter). The
birth interval (BI) was then split into three consecutive cub-rearing periods, each
six months long. During each period, the female was raising young cubs (six
months or younger), large cubs (seven to twelve months) or no cubs (i.e., cubs
were, or became, independent and the female had not yet produced another lit-
ter). Because fewer locations were obtained in these shorter time periods, we
only used the MCP to calculate cub-rearing home range size.
We calculated home ranges for nine females over eleven birth intervals where
females were radio-monitored without interruption and at least one cub from
the first litter survived to independence. The average BI was 17.6 ± 1.9 months
long and BI home range size averaged 64.9 ± 37.8 km2 (90 percent ADK; Table
12-1). The annual home range size for the same nine females averaged 58.9 ±
29.2 km2. Although the BI involved a longer time period, only one of the four
home range estimators (100 percent MCP) indicated BI home range size was
larger than annual home range size [2]. When we further examined female home
range size in relation to the presence and age of cubs, we found a female’s home
Table 12-1. Female puma home range size during a birth interval (birth of
one litter to the birth of subsequent litter, n = 11), San Andres Mountains, New
Mexico.
Home range estimator (km2)d
Minimum
Birth No. No.
Adaptive kernel convex polygon
Puma no. yeara monthsb locationsc 90% (–) 60% (–) 100% (+) 90% (–)
4 1986 16.9 66 30.8 11.0 50.8 27.5

15 1987 18.3 68 56.0 17.6 74.2 38.1
21 1991 19.0 75 115.6 45.4 116.1 88.0
37 1988 16.0 61 131.2 62.8 116.5 84.8
37 1989 18.3 69 105.0 33.2 105.0 33.2
41 1988 18.0 71 63.6 25.5 75.1 47.2
45 1991 17.8 66 26.0 6.7 35.6 20.0
47 1990 14.0 57 17.7 5.1 22.7 14.7
47 1993 16.8 69 40.4 13.8 47.2 31.7
65 1990 17.1 60 70.9 24.0 79.6 24.0
91 1992 21.6 81 57.0 24.4 66.2 41.1
Mean ± SD 17.6 ± 1.9 64.9 ± 37.8 24.5 ± 17.4 71.7 ± 31.5 40.9 ± 24.3
aBirth year = birth year of first litter.
bNumber of months from the birth of first litter to the birth of the next litter.
cAll aerial locations augmented with ground locations that were more than three days apart.
dADK = adaptive kernel; MCP = minimum convex polygon. A (+) indicated the cyclic home range
size was significantly larger than the annual home range size (P = 0.01) and a (–) indicated that there
was no significant difference in cyclic and annual home range sizes (P > 0.1).
FIGURE 12-2. Home range size (mean and standard deviation based on the minimum con-
vex polygon) for female pumas (n = 8) during three consecutive cub-rearing periods on the
San Andres Mountains, New Mexico, 1986–1994. During the no-cub period, progeny were
thirteen to eighteen months old; they typically became independent at the start of, or early
in, this period. Females did not give birth to new litters during this period.
FIGURE 12-3. The home range (90 percent minimum convex polygon) of puma F15 during
three consecutive cub-rearing periods, San Andres Mountains, New Me x i c o. F15 gave birth to
a litter of three cubs on 17 May 1987 (star), and she bore her next litter 18.3 months later.
F15’s cubs from her 1987 litter became independent at ten to fourteen months of age; conse-
quently, she was solitary for at least four of the six months that comprised the “no-cub”
period. Her home range sizes for the three consecutive periods were 17.5, 30.6, and 45.0 km2,
respectively.
range size generally increased as the age of her cubs increased and was largest
during the period when she regained solitary status (Fig. 12-2)[3]. Puma F15’s
behavior after the birth of her litter in May 1987 is a clear example (Fig. 12-3).
Lifetime Home Range

Cumulative or “lifetime” home ranges were obtained for pumas monitored for
continuous periods in excess of forty-eight months as resident adults. We chose
forty-eight months because it represented such a substantial portion of an adult
puma’s life. Based on the average age of death of thirty radio-collared adult
pumas (nineteen males, eleven females) on the study area (see Chapter 8), a
forty-eight-month period would typically comprise 73 percent or more of an
adult male’s life and 100 percent of an adult female’s life.
Lifetime home ranges were estimated for sixteen pumas (Table 12-2).
Although each puma’s lifetime home range was larger than the average area it
utilized over an annual period, the differences were less pronounced using the
ADK versus MCP home range estimator [4]. Based on the 90 percent ADK
method, ten males monitored for an average of 71.1 ± 18.2 months utilized
areas that were 70 percent ± 42 percent larger than the average areas they occu-
pied on an annual basis. The difference was not quite as large for females; the
six females followed for periods averaging 63.8 ± 12.7 months used areas about
45 percent ± 23 percent larger than their average annual home ranges. However,
some pumas used areas during individual annual periods that approached the
size of their lifetime home range (this was not observed using the MCP estima-
tor). For example, ten pumas used lifetime home ranges that were three to eight
percent larger (n = three males, three females), or 1 to 14 percent smaller (n =
two males, two females), than their largest annual home ranges (e.g., Plate 1).
The average lifetime home range size for females (n = 6) was 22 percent the size
of lifetime ranges used by males (n = 10).
What Factors Influence Adult Home Range Size?

In the San Andres Mountains, there was wide variation in the size of the area a
puma used from year to year (as indicated by the large standard deviations in
Appendix 3). Annual variation in a puma’s range size was probably influenced
by a number of factors, including (1) changes in energy demands, (2) changes
in food availability, and (3) responses to the presence or absence of other pumas.
The first factor pertains most to females. A positive relationship between the size
of a female’s range (or at least her mobility) and the age of her cubs has been
observed in other puma populations, notably Alberta (Ross and Jalkotzy 1992)
and Utah (Hemker et al. 1984). This behavior has been linked both to increas-
ing energy demands (Ackerman 1982) and to the improved traveling capabili-
Table 12-2. Cumulative home ranges for ten adult male and six adult
female pumas that were radio-monitored for forty-eight consecutive
months or longer on the San Andres Mountains, New Mexico.
Home range estimator (km2)
Adaptive kernel Minimum convex polygon
Puma Months No.
no. monitored locationsa 90% 60% 100% 90%
Males
1 48 169 279.7 99.4 364.3 220.2
3 101 400 317.5 122.0 459.2 300.4
5 86 304 227.3 60.9 341.4 192.7
7 57 215 353.2 129.7 452.8 228.8
18 50 181 677.4 192.7 772.5 487.7
19 55 190 274.8 106.3 382.2 205.4
36 86 286 559.2 185.8 1077.0 526.0
38 85 314 451.5 189.7 815.4 398.3
46 69 269 444.4 167.5 575.8 366.1
52 74 253 132.8 48.9 195.3 117.5
x– 71.1 258.1 371.8 130.3 543.6 346.0
SD 18.2 72.0 163.1 52.6 268.2 210.9
VS ANNUALb 1.7x 1.6x 2.5x 2.2x
Females
15 63 229 76.5 29.2 119.2 67.2
28 50 192 127.0 69.2 209.4 108.3
41 62 247 84.6 40.0 125.5 70.1
45 81 301 58.6 16.2 87.3 54.6
47 76 307 44.2 12.8 93.6 51.0
54 51 197 105.2 29.8 182.4 82.4
x– 63.8 245.5 82.7 32.9 136.2 72.3
SD 12.7 49.7 30.2 20.4 49.2 21.0
VS ANNUALb 1.5x 1.7x 2.4x 1.9x
aIncluded are locations from all flights as well as any ground locations that were obtained
more than three days from any other location.
bAverage lifetime home range size relative to mean annual home range size.
ties of older cubs. Even during the “no-cub” phase, the size of the female’s range
may be influenced by her offspring, since many cubs actually become inde-
pendent during the early part of this period. However, it is also possible that the
female is utilizing a larger area in an attempt to distance herself from newly inde-
pendent cubs and to locate prospective mates. The influences of the second and
third factors (food availability and other pumas) on an individual’s annual home
range size are examined later in this chapter.
In the course of an adult puma’s life, it may traverse a much larger area than
it will cover over any particular annual period. We believe this occurs for two
main reasons. First, home range size increases with an increasing number of
locations, even though we tried to minimize this effect on our study animals.
For example, F47’s cumulative home range size based on the 90 percent adap-
tive kernel estimator and 307 locations (about fifty-one locations per year) was
44.2 km2 (Plate 1). If we included all radio locations that were at least one day
apart (n = 462 locations or about 77 per year), F47’s cumulative home range was
12 percent larger, or 49.3 km2. However, an individual’s lifetime home range
size was nearly equivalent to, or smaller than, its largest annual home range size
for ten out of sixteen pumas. As we will show when we examine fidelity, some
of the increases in home range size probably resulted from a second cause: home
range expansions or shifts over time.
We also noted wide within-gender differences in annual home range size .
Although the majority of adult males (n = 14) used average annual home
ranges of 150–250 km2, the rest used larger (n = 4) or smaller (n = 6) ranges.
Si m i l a r l y, most females (n = 16) used average annual ranges of 50–100 km2,
but the rest used ranges that we re larger (n = 5) or smaller (n = 9). These dif-
f e rences in range size we re undoubtedly amplified by variations in habitat
quality throughout the mountain range. The southern third and north-central
p o rtions of the study area contained re l a t i vely contiguous segments of ru g g e d
terrain and abundant mountain shrub and piñon-juniper cover. As a conse-
quence, we believe it provided better stalking cover and supported more deer.
Conversely, the central portion of the range (Hembrillo and Su l p h u r
Canyons) appeared to be drier and contain more desert shrub cover (i.e.,
poorer-quality deer habitat). Our observations, though not quantitative, sup-
p o rted this notion; we saw fewer deer and less deer sign in these areas. The
n o rth end of the study area had mountainous patches fragmented by inter-
vening desert flats that also provided less cover and seemed to support fewe r
deer. Pumas use features of the habitat selectively (Logan and Irwin 1985,
Belden et al. 1988, Laing and Lindzey 1991), and specific habitat features
confer advantages to pumas in hunting prey (Logan 1983). Consequently,
larger home ranges probably are essential in areas with poor or fragmented
habitat patches. We documented our largest home ranges (M18, M20, M151,
M161, F147, F160; see Appendix 3) in the central (poor) and northern (frag-
mented) portions of the study area, as well as in the Malpais (a rugged lava
flow east of the study area with extremely poor deer habitat). Si m i l a r l y, in
female tigers (Smith et al. 1987) and leopards (Bailey 1993), large variation in
home range size is believed to be associated with habitat quality, part i c u l a r l y
as it is influenced by prey abundance.
Although habitat variability may partly explain variations in range size of
same-sex individuals, what accounts for the notable differences in home range
size between the two sexes? Studies across the West have consistently reported
larger home ranges for male pumas (Table 12-3). Within the same population,
male home ranges generally average 1.5 to 5 times the size of female ranges. The
same trend is found for other large, sexually dimorphic felids, including the
leopard (Bailey 1993; Stander et al. 1997), jaguar (Schaller and Crawshaw
1980), and tiger (Schaller 1972, Sunquist 1981). In carnivores, there is a strong
linear relationship between home range size and body mass, indicating individ-
uals use areas that satisfy their metabolic demands (Gittleman and Harvey 1982,
Lindstedt et al. 1986). Consequently, larger-bodied or pack-forming animals
should have greater metabolic needs, and should generally require larger areas,
than smaller-bodied or solitary animals. But in sexually dimorphic felids, larger
males do not necessarily have greater metabolic needs than smaller females (Ack-
erman et al. 1986, Sandell 1989). Ackerman (1982) found that puma mothers
with two or three yearling cubs needed to procure 12 kg of food per day, whereas
a solitary male needed only 4 kg. Considering energetics alone, females with off-
spring should have larger home ranges than males.
Why then are male ranges so much larger? Home range size differences are
probably related to disparate strategies to maximize reproductive success. Trivers
(1972) was the first to suggest that female behavior should be directed toward
raising young successfully, while male behavior should be directed toward mate
acquisition. Because only female pumas rear young, they must singly acquire
critical resources such as food, cover, and security for cubs. Unencumbered male
pumas are free to seek out breeding opportunities. In species that occur in low
densities, have no set breeding season, and are well dispersed over the landscape
(all of which apply to pumas), males must traverse large areas to increase their
chances of finding mates. Hence, the home range of a male in a polygynous
mating system should be larger than that required to meet energetic demands
(Sandell 1989, Lott 1991, Frank and Heske 1992, Minta 1993). For example,
male badgers increase home range size during the breeding season but reduce it
to meet energetic demands during the non-breeding season (Goodrich and
Buskirk 1998). Since pumas breed year-round, male puma home range size
should remain large.
Although many studies have used radiotelemetry to produce home range
estimates on pumas, differences in the methods used to analyze home range data
(Hopkins et al. 1986, Neal et al. 1987), as well as sample size and duration of
observation, make comparisons difficult. Puma home range sizes reported in the
literature vary widely. Some of the largest home ranges have been reported in
Utah; in this extremely low-density puma population, annual home range size
exceeded 800 km2 for one male and averaged over 600 km2 for four females
(Hemker et al. 1984). In contrast, and based on the same method of estimating
home range (100 percent MCP), some of the pumas on the San Andres Moun-
tains used some of the smallest documented home ranges. Three adult males and
two adult females in our puma population utilized areas under 80 km2 and 20
km2, respectively, during annual periods.
O verall, puma home ranges in the San Andres Mountains we re intermedi-
ate to small in size compared to those re c o rded elsewhere in North America
(Table 12-3) and we re most similar to those re p o rted for the Diablo Range in
California (Hopkins 1989) and the Guadalupe Mountains in New Mexico
(Smith et al. 1986). Two environmental factors probably had an influence on
the smaller range size. First, in all three areas, the major prey source did not
migrate. Annual home ranges that must encompass different summering and
wintering areas should typically be much larger. For example, annual home
range size for four females that followed migrating prey in Idaho averaged 268
km2 (100 percent MCP; Seidensticker et al. 1973), which was almost four
times the size of the annual female range size in the San Andres Mountains.
Second, the San Andres Mountains are a long, narrow band of puma habitat
(where stalking, nursery and resting cove r, and adequate numbers of a large
prey species are available) surrounded on two sides by wide, re l a t i vely inhos-
pitable desert basins. It is possible that the isolated nature of the mountain
range forced individuals to use smaller areas in an effort to both utilize the best
habitat and avoid one another. Only when habitat conditions became poor
(during the drought and subsequent deer decline in 1993 and 1994) did a few
pumas (generally males in the Re f e rence Area) extend their movements fart h e r
into the adjacent desert flats (see Fig. 4-1).
Home Range Size, Prey Abundance, and Puma Density

Assuming that male and female pumas employ different strategies to achieve
reproductive success, we should see different behavioral responses by each sex to
changes in prey abundance. Because a female puma’s reproductive success is
dependent on her ability to secure adequate food for her young, her home range
size is expected to be directly affected by the density, dispersion, richness, and
vulnerability of prey (Seidensticker et al. 1973, Macdonald 1983, Sandell 1989).
The relationship of male home range size to food abundance should not be as
clear-cut, because although a male must obtain adequate food resources for sur-
vival, his level of reproductive success is dependent on securing matings. Con-

sequently, we hypothesized that in the San Andres Mountains, female home
range size would be inversely related to prey abundance, whereas there would be
no clear relationship between male home range size and prey abundance.
Does Prey Abundance Affect Home Range Size?

Correlations between prey abundance (generally density or biomass measures)
and home range size have been found in some felids, including bobcats (Litvaitis
et al. 1986), Canadian lynx (Ward and Krebs 1985) and African lions (Van Ors-
dol et al. 1985). Unfortunately, the difficulty of obtaining detailed information
on both variables simultaneously has precluded such an analysis in pumas. How-
ever, a strong negative relationship between population density and female
home range size has been observed across species of solitary carnivores, indicat-
ing a relationship between both of these parameters and food abundance
(Sandell 1989). Since a few studies can provide reliable estimates of both puma
home range size and puma density, we tried a two-level approach to examining
puma home range size and prey abundance. First, within the San Andres Moun-
tains, we compared puma home range size to relative deer abundance based on
the mule deer population model we developed (see Chapter 17). The model
indicated relatively low deer abundance during 1987, 1993, and 1994, and high
deer abundance during 1990 and 1991; thus, we compared puma home range
sizes between these two sets of years. This approach had limitations because we
could only estimate the relative abundance of deer on the study area; we could
not determine deer densities within individual puma ranges. We also looked for
a relationship between puma home range size and puma density across popula-
tions in western North America.
There was no relationship between the home range size of pumas (male or
female) and relative deer abundance in the Treatment or Reference areas of our
study population (Table 12-4)[5]. Our home range estimates each year often
included different sets of individuals (i.e., some pumas died, new individuals
were caught, and radio collars on others failed), which may have contributed to
the lack of a relationship. Still, we obtained home range estimates on the same
six individuals during at least four of the five years, and they did not show any
consistent changes relative to deer abundance. For example, M38 used home
ranges of 199 and 296 km2 in 1990 and 1991 (years of high deer abundance),
and 168 and 264 km2 during 1993 and 1994 (years of low deer abundance),
Similarly, F47 used home ranges of 44.8 and 24.4 km2 in 1990 and 1991, and
16.8 and 42.8 km2 in 1993 and 1994. She was also raising cubs during each of
those years. Obviously, prey abundance did not explain the variation we
observed in home range size. It is likely other factors, especially changes in deer
Table 12-4. Annual home range sizea (mean ± SD) for adult pumas
during years of relatively low and high deer densities, San Andres
Low deer densityb High deer densityb
Sex n HR size (km2) Sex n HR size (km2)
Treatment Area
M 10 140.8 ± 51.3 M 5 136.4 ± 42.4c
F 10 56.0 ± 29.0 F 8 69.0 ± 29.0
Reference Area
M 15 236.4 ± 86.6 M 10 197.4 ± 87.6
F 13 52.6 ± 27.3 F 13 68.6 ± 36.1
aHome range (HR) size based on 90 percent adaptive kernel method.
bLow deer densities occurred during 1987, 1993, and 1994. High deer densities occurred
during 1990 and 1991. Mean home range sizes of pumas for each of these years are
reported in Table 12-5.
cWe only estimated home range size for one male (M3) in the Treatment Area in 1991.
Because his home range in 1991 was strongly affected by the removal of all other males, we
did not include it in the overall average.
distribution, negated any relationship that might occur between prey abundance
and puma range size. For example, if deer concentrate around critical water
sources during drought, a female puma’s home range size might shrink, even
though overall deer density has declined. It is possible this occurred with F21.
In 1993 (drought year) her home range size was 58 percent the size it was in
1991 (non-drought year). But during 1993, 63 percent of her locations occurred
within 1 km of active springs, whereas only 36 percent of her locations were that
close to water sources in 1991. An apparent lag in the pumas’ response to
changes in prey abundance also undermined the relationship. Puma densities in
the San Andres Mountains continued to increase during 1993 and 1994, even
as drought took hold and prey abundance declined (see Chapter 17). This is
contrary to the expectation of a positive correlation between these two variables
(Sandell 1989). Such a positive relationship has been demonstrated in the eco-
logically similar African leopard (Stander et al. 1997). We suspect it might take
a severe reduction in overall prey abundance to trigger a notable increase in the
average home range size of pumas.
When we plotted home range size on density for five North American puma
FIGURE 12-4. Adult puma home range size correlated with adult puma density in five study
areas (Alberta, British Columbia, Idaho, New Mexico, and Utah) in North America. Winter
home range sizes are used in Idaho and Utah because that was when the best density esti-
mates were obtained.
populations with the most reliable estimates of both variables—Alberta, British

Columbia, Idaho, New Mexico, and Utah (Ross and Jalkotzy 1992, Spreadbury
et al. 1996, Seidensticker et al. 1973, our study, Hemker et al. 1984, respec-
tively)—we found no relationship between puma home range size and puma
density (Fig. 12-4)[6]. Home range sizes were notably larger only at the lowest
reported densities (the Utah data point). In a similar analysis of leopard popu-
lations from ten different regions in sub-Saharan Africa, Stander et al. (1997)
found home ranges were notably larger only at very low leopard densities and
rainfall; there was no linear relationship between the parameters. Unfortunately
in our analysis on puma, the minimal numbers of data points and the tendency
to average density and home range estimates across years (during which time
prey abundance may fluctuate) may obscure any relationship.
Does Puma Density Affect Home Range Size?

Our analysis of puma densities and range sizes across populations suggests that
puma home ranges become notably larger only when puma densities (and thus
apparently prey densities) are very low. But what if we examine this relationship
within a single puma population? Although a negative relationship between
puma home range size and its population density may imply a negative rela-
tionship between range size and prey abundance, it may also connote a response
by one puma to the presence or absence of another. We examined this possibil-
ity in a two-step manner. First, we looked for correlations between mean home
Table 12-5. Mean annual home range size (90 percent adaptive kernel) and
density for adult male and female pumas on the San Andres Mountains, New
Mexico, 1988–1994.
Males Females
Home range Density Home range Density
Year n 2
(km )a (no./100 km2)b n 2
(km ) a (no./100 km2)b
Treatment Areac
1987 3 184.4 ± 59.8 2 81.4 ± 60.4
1988 3 189.4 ± 36.8 0.73 1 127.1 0.63
1989 4 150.1 ± 65.6 0.70 4 93.4 ± 64.0 1.06
1990 5 136.4 ± 42.4 0.79 5 70.4 ± 24.3 1.22
1991 1 295.8 0.38 3 66.5 ± 41.8 0.71
1992 4 170.1 ± 29.6 0.52 4 50.6 ± 23.2 0.99
1993 3 138.1 ± 9.9 0.63 4 46.4 ± 25.0 1.23
1994 4 110.1 ± 47.7 0.69 4 53.0 ± 11.0 1.18
Reference Areac
1988 2 312.2 ± 71.6 1 41.2
1989 7 164.2 ± 50.5 0.53 5 61.0 ± 11.3 0.60
1990 6 217.8 ± 87.6 0.56 7 51.8 ± 26.2 0.77
1991 4 167.0 ± 28.4 0.62 6 88.2 ± 38.1 0.83
1992 6 194.0 ± 91.7 0.65 11 57.8 ± 19.6 0.98
1993 7 229.6 ± 90.9 0.68 7 51.6 ± 30.6 1.08
1994 6 219.0 ± 85.5 0.67 5 56.3 ± 27.8 1.12
aMean ± SD.
bObtained from Table 10-5.
cExcludes any individual whose home range was not at least 60 percent within the designated area.
range size (male and female) and adult puma density (male, female, and total)
in the study area each year. Second, we plotted average puma range sizes, puma
densities, and deer population trends in the study area each year. We examined
the Treatment Area over a seven-year period (1988–1994) and the Reference
Area over six years (1989–1994). Each year one to five males (x– = 3.4 ± 1.3) and
one to five females (x– = 3.6 ± 1.3) provided the home range estimates used to
examine the relationship of home range size to density in the Treatment Area.
Another three to seven males (x– = 5.8 ± 1.5) and five to eleven females (x– = 7.0
± 2.1) provided the home range data for the Reference Area (Table 12-5). We
used puma densities reported in Table 10-5 and deer population trends from
Chapter 17 (Fig. 17-6, Table 17-7) in the comparisons. Based on population

modeling, the mule deer population was increasing during most of the study
but then declined sharply in 1993 and 1994, even as puma numbers remained
high.
FEMALE RESPONSE TO CHANGES IN PUMA DENSITY

We found negative relationships between female home range size and female
density in both the Treatment and the Reference Areas, albeit these relationships
were not significant [7]. A weak, negative correlation also occurred between
female home range size and male density in the Reference Area, but this char-
acteristic was reversed in the Treatment Area.
The consistent negative relationships between female range size and density
suggested, unlike our initial analysis, that overall prey abundance might have
had some effect on female range size. However, it could not tell us whether or
not intraspecific interactions could have had a compounding influence. For
example, if female pumas compete most with other females, it is plausible they
may try to avoid one another by reducing their respective range sizes. This way,
they will not compete for the same prey, spoil one another’s hunt, or subject
their cubs to undue risk. Accommodation of related females would also increase
inclusive fitness if food were abundant and risks of dispersal were high. Chap-
ters 9 and 13 describe at least eight matrilines that formed on the study area.
Similarly, reductions in female densities may result in increasing home range
sizes as the potential for encounters decreases. Consequently, we might conclude
that a negative relationship between female density and home range size is the
result of intrasexual interactions as well as changes in prey abundance. How can
we separate these two effects? If females try to accommodate and avoid one
another by reducing range size, we would expect female ranges to be smaller
when both prey densities and puma densities are high than when prey densities
are high and puma densities are low. By removing pumas during a period of high
prey abundance, we could look for this relationship. Additionally, if prey densi-
ties declined without a corresponding increase in female range size, we could
suspect that females were exhibiting a degree of avoidance.
After the removal of 71 percent of the adult males and 60 percent of the
adult females from the Treatment Area during December 1990 through June
1991, only three of the original females and one of the original males remained
exclusively in the Treatment Area. Home range sizes of the remaining females
actually declined slightly during the year of removal (Fig. 12-5). During that
same period, deer densities continued to increase. Hence, the removal of com-
petitors did not cause female home range expansions; in fact, the increasing deer
densities may have allowed females to use even smaller areas. It is likely that
appreciable increases in home range size did not occur at low female densities
because females already had access to adequate resources to raise their young.
Any increase in home range size would only increase the potential for unwanted
intraspecific encounters with unfamiliar females and males (prediction 5 of the
two-strategies hypothesis). However, once puma densities were high, a drop in
prey densities (beginning in 1992–1993) did not produce an immediate increase
in female range size. At this point, females may have been responding more to
the presence of other pumas (i.e., trying to avoid them) than to the declining
density of prey. Consequently, females may have been at risk of using too small
of an area to supply the minimum amount of food and cover necessary to suc-
cessfully raise young. An alternate hypothesis (as discussed earlier) is that deer
distributions became more clumped in vulnerable areas (e.g., at springs) as the
FIGURE 12-5. Changes in annual home range size (90 percent adaptive kernel) and density
of adult pumas in the Treatment Area, San Andres Mountains, New Mexico. Superimposed
is the mule deer population trend over biological years, based on model 2 in Chapter 17
(gray line with open boxes, scale not given). The decline in puma density in 1991 corre-
sponds to the removal of thirteen independent pumas.
drought took hold. Females could use similar-sized areas and still obtain ade-
quate food, at least to a point. Once overall prey densities declined far enough,
these clumps would not be sufficient. We may have been seeing this response in
1994; female puma densities were starting to taper off (Reference Area) or
decline (Treatment Area), and female home range sizes were starting to increase.
All the information suggests that female pumas were influenced more by prey
abundance and distribution than the numbers of other females. This supports
the two-strategies hypothesis. However, the relationship was not a direct one,
because the deer decline did not generate an immediate and opposite response
from the pumas. Additionally, avoidance of other females by reducing home
range size may not be feasible if prey resources are clumped. Instead, females
may use shared areas at different times. This will be addressed in Chapter 14.
MALE RESPONSE TO CHANGES IN PUMA DENSITY

Analyses of how male home range size related to puma density produced con-
flicting results. The only significant relationships were found in the Treatment
Area, where male home range size was inversely related to both female and total
density. A weak, negative relationship was also found between male home range
size and male density [8]. No strong correlations were found in the undisturbed
Reference Area, but the pattern of correlation was reversed from what we found
in the Treatment Area. Male home range size increased with increasing puma
density (male, female, and total) [8].
Although a male puma also requires food for survival, his reproductive suc-
cess is ultimately determined by the availability of estrus females and the num-
ber of other males with whom he competes. Consequently, we might expect
changes in male range size to better reflect responses to changes in numbers of
other pumas than prey abundance. If so, which affects males more, the number
of mates or the number of competitors? To try to determine this, we developed
three alternative hypotheses where male home range size is influenced most by
(1) deer density, (2) female mates, or (3) male competitors. If the first hypothe-
sis is correct, then male range size should decrease with increasing deer density,
even if concurrently the number of mates and competitors is decreasing. Simi-
larly, male range size should increase with declining deer density, even in the face
of a growing puma population (and the increasing number of mates and com-
petitors it implies). In hypotheses 2 and 3, deer densities should be secondary.
In hypothesis 2, a male’s range size should decrease with increasing densities of
females, more so than with increasing densities of males, whereas in hypothesis
3, his range size should decrease with increasing densities of males, more so than
with increasing densities of females.
In the Treatment Area, one male (M3) had a profound effect on the result-
ing negative correlations between home range size and density (Fig. 12-5). After
the experimental removal from December 1990 through June 1991, only M3
remained exclusively in the Treatment Area. When all male competitors as well
as most potential mates were removed, M3 greatly expanded his home range
(Fig. 12-6). In the three years prior to puma removal, M3’s annual home range
size contracted from 227 km2 to 141 km2, a reduction of about 22 percent per
year. During this same time interval, both puma and mule deer densities were
increasing, suggesting either could have influenced his home range size (Fig. 12-
5). Then in 1991, the year we removed pumas, M3’s annual home range size bal-
looned by 110 percent, to 296 km2. That same year the deer population con-
tinued its increase (contrary to hypothesis 1). M3 also mated successfully with
two of the remaining females (both familiar to him) and associated with two
other females with whom he had no prior contact. This was his best breeding
FPO @ 80%
FIGURE 12-6. Puma M3’s annual home range (90 percent adaptive kernel), 1988–1990 (a),
and 1991–1993 (b), in the Treatment Area, San Andres Mountains, New Mexico. M3’s
home range declined in size each year until 1991. We removed all of his male neighbors (n =
5, black dots) as well as eight independent females from the Treatment Area from De c e m b e r
1990 through June 1991. That year M3’s range expanded by 110 percent, and he courted
four females (white dots); two produced his cubs, and the other two we re translocated in
June, after they we re caught courting with him. M3’s home range declined in size and
shifted north during the following two years. One of his neighbors (M88) returned to his
former home range south of M3 in July 1991, possibly encouraging M3’s nort h w a rd shift.
success in the eight years for which we had annual home range data. As pumas
repopulated the Treatment Area in the following years, M3’s home range con-
tracted again. In 1993, when M3 was twelve years of age, his home range size
was almost identical to what it was the year prior to puma removal (146 km2).
That same year female density had rebounded to a level equivalent to that found
prior to puma removal (1.23 per 100 km2 in 1993, 1.22 in 1990), but deer den-
sity was declining (Fig. 12-5). If food supply was of primary importance to M3,
his range size should have decreased during times of plenty and increased as the
deer population entered its decline. Instead, M3 responded most to the presence
and absence of other pumas. Because we reduced the density of male and female
pumas at the same time, we could not determine which sex had the most influ-
ence on M3’s home range size. M3’s mating behavior and the strength of the
correlations [8] suggest that M3 was most affected by the density of females (i.e.,
potential mates); however, his best breeding success occurred when he had the
least male competition. These results support the two-strategies hypothesis.
We suspect a negative association was not found between male home range
size and density in the Reference Area because interactions were more complex
than in the Treatment Area. There were more male pumas in the Reference Area,
and each male had to respond to the behaviors of all of his neighbors. As we will
see when we examine fidelity, males may exhibit shifts in their home ranges in
response to other males. Such shifts may result in a larger annual home range
size because the annual home range includes areas used before and after the shift.
For example, average male home range size in the Reference Area (n = 6)
increased from 164 to 238 km2 between 1989 and 1990. During that same
period, male density also increased from 0.53 to 0.56 males per 100 km2. As we
will later illustrate, two of the males with the largest home ranges in 1990 (M38,
M46) exhibited home range shifts that year. Excluding these males, average male
home range size for 1990 is 159 km2, slightly lower than the mean home range
size for 1989. If males are shifting in response to increasing puma densities, then
a positive correlation between home range size and density may occur.
Does Home Range Size Reflect an Attempt at Population

Self-Limitation?
If pumas are self-limited, as Seidensticker et al. (1973) hypothesized, we would
expect both males and females within the puma population to stabilize at a den-
sity well below a limit set by the prey population. Under these circumstances, all
puma home ranges should become occupied and no longer constrict to accom-
modate more pumas. If prey densities subsequently decline, pumas should
respond in kind; otherwise, their high densities and attendant predation will
further push the prey population downward.
In the Treatment Area, the absence and subsequent reappearance of com-
petitors may have first freed M3 to claim vacant areas, then forced him to relin-
quish them. Consequently, adjacent males may have restricted M3’s movements,
which suggests that the presence of adult male pumas does influence adult male
density. However, M3 also appeared to expand and contract his sphere of influ-
ence in direct response to availability of potential mates. These combined behav-
iors suggest males were attempting to maximize breeding opportunities. Thus,
we concluded that males were simply attempting to maximize individual repro-
ductive success, not seeking to protect the food supply from being overutilized.
The absence of any clear response by females to the removal of other pumas, as
well as the tendency for pumas to accommodate more individuals even as the
prey density declined, also indicated that the pumas in our study population
were not limiting themselves below the level set by the prey as Seidensticker et
al. (1973) reported for pumas in Idaho.
It is important to emphasize that many of the relationships we found
between puma home range size and adult density were weak, probably because
of the dynamic nature of the society (e.g., each individual is responding to
arrivals, deaths, and breeding condition of its immediate neighbors). We clarify
some of these dynamics as we examine home range fidelity later in this chapter
and puma interactions in Chapter 14. Additionally, habitat quality, prey density,
and prey distribution among individual home ranges vary and were not meas-
ured in our study. These variables undoubtedly affect home range size. Prey dis-
tribution was found to be an important factor determining the spatial distribu-
tion of pumas in the Sierra Nevada in California (Pierce et al. 2000b).
Adult Home Range Fidelity

Fidelity is the degree to which an individual uses the same area from one time
interval to the next. It is an important way of gauging how adult pumas may be
affected by a changing environment, changing physiological needs (as with
females raising cubs), or other pumas. We hypothesized that female pumas
would exhibit strong home range fidelity because the main prey, mule deer, did
not migrate, and familiarity with an area should enhance cub survival. Fidelity
would remain high unless shifting resulted in increased inclusive fitness (i.e.,
more young are produced by that individual female or her close kin). Abandon-
ment should occur only under the most severe conditions. In contrast, we
hypothesized that male pumas would exhibit greater variability in fidelity
because the resource they seek (receptive females) is widely dispersed in space
and time. Home range shifts, or even roaming behavior, should result as a con-
sequence of poor mating probabilities within the home range or increased com-
petition for mates.
We assessed fidelity over annual and six-month intervals as well as during the
different phases of a female’s reproductive cycle. We did not examine fidelity on

a seasonal basis because the puma’s major prey was nonmigratory (Sweanor
1990) and pumas do not have a specific breeding season (see Chapter 7).
Fidelity was determined using two methods. In method 1, we calculated a
fidelity index (FI) for each adult puma during pairs of years (one, two, and three
years apart) where annual home range estimates (90 percent ADK and 90 per-
cent MCP) were obtained. We modified the simple ratio of Ginsberg and Young
(1992) for quantifying association patterns:
FIx = n1 + n2 / N1 + N2 ¥ 100
where x is the time span between pairs of years, n1 and n2 are the number of
locations recorded within the overlap zone of the two years in question, and N1
and N2 are the total number of locations recorded within each annual home
range (Fig. 12-7). For individual pumas with greater than one FI1, FI2, or FI3
(i.e., pumas monitored for more than two, three, or four consecutive years), we
treated each FI value as an independent measure of fidelity. The indices were
grouped based on sex and length of time between pairings. We assumed a puma
made a home range shift if the overlap zone for the pairing was less than 40 per-
cent. We plotted sex-specific frequency distributions of FIx and then tested for
FIGURE 12-7. A simplified example of a fidelity index (FI). The polygons and dots represent
a hypothetical puma’s home range and locations over consecutive annual periods. In this
example, FI = 13 / 21 x 100 = 62 percent.
normality in each of the distributions. We also tested for differences between the
distributions of FIx for male and female pumas.
In method 2, we used the location data to calculate the mean x and y UTM
coordinates for an individual during specific time intervals and then measured
the distances moved between successive mean locations. This method elimi-
nated the biases associated with home range estimators (White and Garrott
1990). First we examined changes in mean locations between consecutive years.
If, from one year to the next, an individual moved a distance greater than one
standard deviation above the mean distance moved by members of the same sex,
we calculated the distance moved between mean x and y coordinates during suc-
cessive six-month intervals. This helped to pinpoint the time of a suspected
home range shift. We also calculated distances between mean locations for the
phases of a female’s reproductive cycle. The cycle was divided into four consec-
utive periods when the female was: (1) raising cubs from birth to six months of
age; (2) raising cubs seven to thirteen months of age (which included cubs to the
average age of independence or to known independence; (3) without cubs; and
(4) raising her subsequent litter of young (birth to six months) cubs. These peri-
ods were not identical to ones used to calculate home range size because we did
not have the sample size constraint inherent to using home range estimators. For
example, a female that was without cubs (period 3) for less than six months
would have too few locations for adequate home range analysis but sufficient
locations to generate an arithmetic center of home range use. Once a shift was
detected, we looked for possible explanations for the shift.
Method 1—Fidelity Index

We monitored seventeen adult male and twenty-one adult female pumas from two
to seven years each and obtained eighty-seven measures of FI1, sixty-one measures
of FI2, and twenty-nine measures of FI3. Similar patterns of fidelity were found
using both the 90 percent ADK and the 90 percent MCP home range estimators,
except that the 90 percent MCP produced slightly lower estimates of fidelity. For
simplification, we re p o rt only results obtained using the 90 percent ADK. Distri-
butions of FI2 and F I3 did not differ from normal for either males or females, nor
did the distribution of FI1 for females [9]. Although the distribution of FI1 for
males differed from normal [9], a visual exam of the frequency distribution did not
indicate a bimodal distribution (Fig. 12-8). A bimodal distribution would suggest
two distinct behaviors: high fidelity, and abandonment or strong home range
shifting. Such dual behaviors have been documented in female martens (Phillips
et al. 1998). When distributions of male and female pumas we re compared, FI1
distributions we re almost identical (Fig. 12-8)[10]. Both sexes showed high
fidelity to their home ranges; F I1 for males (n = 48) and females (n = 39) averaged
FPO @ 80%
FIGURE 12-8. Frequency distribution of the fidelity index (FI1) for adult male and female
pumas, based on the 90 percent adaptive kernel home range estimator, San Andres Moun-
tains, New Mexico, 1986–1994. Each 10 percent increment in fidelity corresponds to a
range (e.g., 60 percent includes the range 60.0–69.9 percent).
75.1 ± 14.2 percent and 74.4 percent ± 16.6 percent, respectively (Fig. 12-9 and
Fig. 12-10). However, as the length of time between annual home range estimates
widened, fidelity declined (Fig. 12-9). For example, M36 slowly moved nort h w a rd
over time. Although his consecutive annual home ranges overlapped, within four
years he was utilizing an area completely outside his first estimated annual home
range (Fig. 12-11). Although differences in male and female distributions for FI2
and F I3 were marginally nonsignificant [10], the decline in fidelity was more dra-
matic with males. Be t ween F I1 and FI3, fidelity declined, on average, by 32.0 per-
cent in males and only 8.9 percent in females. There was also greater variability in
the fidelity of males (Fig. 12-9). Home range shifting occurred three times in F I1
(4.2 percent of male F I, 2.6 percent of female FI; Fig. 12-12), nine times in FI2
(25.8 percent of male F I, 5.0 percent of female F I), and twenty times in FI3 (42.1
percent of male F I, 10 percent of female FI). There were three cases (9.7 percent)
in FI2 and five cases (26.3 percent) in FI3 where males utilized areas that ove r-
lapped by less than 10 percent (Fig. 12-11).
Method 2—Distances between Mean Locations

A total of eighty-nine (forty-nine male, forty female) distances between consec-
utive mean annual locations were calculated for thirty-six pumas (sixteen males,
twenty females). Distances between mean annual locations averaged 5.3 ± 4.8
km for adult males and 2.6 ± 2.1 km for adult females. Males moved greater dis-
FIGURE 12-9. Fidelity index (mean and standard deviation) for adult pumas on the San
Andres Mountains, New Mexico. Fidelity was estimated between annual home ranges (90
percent adaptive kernel) that were one, two, and three years apart. Sample sizes are indi-
cated at the top of each line.
FIGURE 12-10. Examples of strong annual home range fidelity in adult pumas on the San
Andres Mountains, New Mexico. Between consecutive years shown above, pumas M19 (a)
and F45 (b) had fidelity indexes of 82 percent and 75 percent, respectively. Home ranges
are depicted using the 90 percent adaptive kernel estimator. For simplicity, only four out of
six of F45’s annual home ranges are shown. Grey dots indicate mean locations for each year.
FPO @ 70%
FIGURE 12-11. Puma M36’s annual home ranges (90 percent adaptive kernel) during six of
seven consecutive years, San Andres Mountains, New Mexico. M36’s fidelity indexes during
pairs of years one, two, and three years apart averaged 71 percent, 51 percent, and 47 per-
cent, respectively. However, his mean location each year (depicted sequentially) slowly
moved northward, and after five years (1988 to 1993), his annual home ranges no longer
shared common area.
tances than females [11]. Five males (10.2 percent) and two females (5.0 per-
cent) moved distances greater than one standard deviation above the mean dis-
tance moved for their respective sex; four pumas (three males, one female)
moved distances greater than two standard deviations above the mean. This
analysis identified annual home range shifts for two of the same individuals that
FI1 did (M18, F21; Fig. 12-12); however, it also indicated possible annual shifts
in five pumas that were not identified in FI1 (M3, M7, M38, M53, F28), and
it did not indicate shifting in one male that FI1 did (M46). This method also
did not identify pumas that may have taken longer periods of time (two to three
years) to make a home range shift.
Home Range Shifts in Pumas

Based on the two fidelity indices, eight individuals (eight males, two females)
made home range shifts within one annual period. For these individuals, we cal-
FIGURE 12-12. Examples of home range (90 percent adaptive kernel) shifts exhibited by
adult pumas over consecutive years, San Andres Mountains, New Mexico. Puma M18 (a)
shifted south between 1987 and 1988 as indicated by a fidelity index (FI1) of 37 percent
and a linear distance between home range centers (gray dots) of 23.3 km (greater than three
standard deviations above the mean distance moved by all males). M18’s FI between 1987
and 1990 (FI3) was 0 percent. Puma F21 (b) shifted north between 1991 and 1992, then
shifted back south between 1992 and 1993. Her FI1 for those years were 41.2 and 22.1
percent; corresponding distances between home range centers were 9.9 and 10.0 km
(greater than four standard deviations above the mean distance moved by all females).
culated distances moved between mean locations over consecutive six-month

intervals. Of the six males, four shifted one time and two shifted twice. One of
the two females shifted three times, while the other shifted only once.
MALE SHIFTING BEHAVIOR

The six males shifted during specific six-month periods when immigrant adult
males arrived within or adjacent to their home ranges (M7, M18, M38, M46,
M53), when the only resident female within the male’s territory died (M53)
and/or when neighboring adult males died or were experimentally removed
(M3, M46). Five males shifted away from new male arrivals, suggesting that new
individuals can usurp space from established residents. For example, M7’s home
range center from January to June 1990 was in San Andres Canyon. In October
1990, new immigrant M114 was captured about 1 km north of this location,
and he spent the next 2.5 months in close proximity to San Andres Canyon.
M7’s apparent response was to shift 13.3 km south during the period
July–December 1990. Similarly, M161 arrived within M53’s home range in
May 1990 and began establishing his own home range. During the following
six-month period (June to December 1990), M53 shifted 14.7 km north and
spent the majority of his time north of the study area boundary. M53’s shift may
also have been precipitated by the lack of females within his home range. F27,
the only known female to reside within the north end of the study area in 1990,
died in April of that year. Consequently, M53 may have been searching for other
potential mates. M18 showed a substantial southward shift (11.9 km) during
the six-month period (January to June 1988) after two new males (M36, M38)
arrived in the northern part of his home range. He made an even larger south-
ward shift during the next six-month interval (24.8 km), possibly in continued
response to those new males. Likewise, M192 arrived within the southern end
of M46’s home range in April 1993 and may have induced M46 to shift 14.6
km north during June–December 1993. M46 was able to shift into an area that
had become vacant after M29’s disease-related death in May 1993. M38 shifted
17.1 km north after the arrival of M173 in midsummer 1992. Four males
(M18, M36, M173, M192), which were captured or recaptured during or soon
after the times of documented shifts, bore recent scratches and bite wounds that
we believe were inflicted as males sorted out dominance relationships through
fighting.
As we indicated earlier in this chapter, M3 shifted in response to the
re m oval of neighboring residents in the Treatment Area (Fig 12-6). He shifted
11.7 km to the north during the six-month interval (June to December 1991)
after all of his male neighbors we re re m oved, and he began frequenting areas
formerly used by four of those neighbors. Initially, this may have been an
expansion of his home range rather than a true shift, because he continued to
visit the southern portions of his home range and to breed with the females
that resided there. When M88, M3’s southern neighbor, returned to his for-
mer home range at the end of July 1991 (166 days after being translocated to
n o rthern New Mexico), M3 may have continued to shift due to his presence.
The two males apparently fought in October 1991 at which time M3 bit into,
and destroyed, the radio transmitter on M88’s collar. We do not know what
injuries M3 sustained, but in a later recapture his head, neck, and forearms
we re grizzled with white flecking, an indication that teeth and claws had
found their mark. From January to June 1992, M3’s home range center shifted
another 8.5 km north. Although his new home range center was less than 10.1
km north of his prior center (less than 1 S D + mean distance moved), M3 was
only located two times within M88’s 1992 home range (90 percent ADK).
Two other home range shifts probably occurred, but we could not accurately
measure them. M1, who lived at the south end of the study area, apparently
made a shift after the arrival of new immigrant male M22 inside his home range
(Plate 2). Because M1 began using an area south of the study area boundary,
where air space and ground access were restricted by White Sands Missile Range,
we could not adequately monitor his movements. M22 was later killed by his
northern neighbor, M3, during a time when M3 and F47 were in a suspected
breeding association. M1 returned north to his former haunts within one month
of M22’s death. The other suspected shift was by M46; he apparently made a
southward shift after a fight with resident M19 along a shared home range
boundary in July 1991. Unfortunately, his collar was ripped off in the fight and
we could not obtain the necessary relocations over the subsequent six months to
measure the shift. However, during the year after his recapture in 1992, he used
an area 14.5 km to the south of the area he had used during the two years prior
to the fight (see Chapter 14).
We found when short-term home range shifts occurred, they were associated
with the arrival of new immigrant males, the removal of other resident males
(either through death or translocation), or the reduction in available mates
within a male’s home range. Male home range shifts after the death or removal
of an adjacent male resident have been reported in other puma studies (Seiden-
sticker et al. 1973, Hopkins 1989, Maehr et al. 1991), as well as in lynx
(Schmidt et al. 1997), bobcats (Anderson 1988) and leopards (Bailey 1993). In
Florida, a male panther’s northward shift may have been influenced by the pres-
ence of a new male within the southern end of his range as well as the presence
of two adult females within the northern half of his range (Maehr et al. 1991).
We suspect more gradual shifts have similar causes, except that possible disputes
between male neighbors may take longer to settle (also see Chapter 14). Multi-
ple responses to various stimuli (e.g., new immigrants, deaths of residents, and
estrus females) may complicate and slow the shifting process. It is important to
note that we were looking for significant shifts in home range use; such meas-
urements do not reflect subtle responses between individuals or the fact that
some individuals do not shift for the simple reason that they are killed by other
males. For example, M3 did not shift to accommodate less-tenured resident
M22 along his southern border; instead, M3 killed M22. New immigrants M19
(see Sweanor 1990) and M219 both usurped area from resident males they
killed. Almost all adult males exhibited scars from fighting at one time or
another during the course of the study, evidence that they occasionally tested
one another and directly competed for space and mates.
FEMALE SHIFTING BEHAVIOR

Only two females showed strong shifting tendencies between consecutive annual
periods. F28’s home range center shifted 7.3 km north between the first and sec-
ond six-month periods of 1991, coinciding with the independence of her cubs.
She successfully bred in January 1992 at a location 8.2 km north of what was
her home range center during the first half of 1991. We did not document the
arrival of any new females to the south of F28 during the period she shifted.
However, we do know her mother, F6, whose range overlapped the south end of
F28’s, produced a new litter of cubs in November 1991. The other female, F21,
made shifts over three consecutive six-month intervals. In 1992, as her cubs aged
from two to fourteen months, F21 made two northward shifts in her home
range center totaling 12.1 km. These shifts coincided with the arrival of an unre-
lated female (F103, daughter of F87), who, in September 1992, produced her
first litter within the boundaries of F21’s 1991 home range. In January 1993,
F21 left her cubs at the northern extreme of her home range, where her daugh-
ter from a previous litter was just beginning to take up residency. Afterward, F21
shifted 11.8 km back south, where she shared the western flank of her range
with F103 and successfully bred with the resident male.
This prompts the question, do females usually shift their home range as their
reproductive status changes? We radio-tracked ten females over fourteen cycles
as they raised young cubs (period 1) and older cubs (period 2), regained solitary
status (period 3), and then produced new litters (period 4). We found that, over-
all, females did not show any greater shifting tendencies in their mean locations
based on their reproductive status [12]. Distances between mean locations aver-
aged 2.6, 2.8, and 1.9 km between periods 1 and 2, periods 2 and 3, and peri-
ods 3 and 4, respectively. We documented five apparent shifts (i.e., distance
moved was greater than the mean + 1 SD for the period) out of forty-two dis-
tance measures (11.9 percent). Two shifts were by F21, discussed above. F37
also shifted twice. First she shifted south at the time her cubs became inde-
pendent, similar to the behaviors of F21 and F28. As she raised older cubs from
her next litter, she shifted back north. This shift coincided with the time that her
daughter, F44, established an independent home range and produced her first
litter within the southern end of F37’s home range. The only other shift was by
F107. She shifted 8.7 km north between the time M88 killed and ate her two-
month-old cubs in December 1993 and the time she produced her next litter in
May 1994, which M88 sired (Plate 3).
An adult female moved, on average, 3.7 ± 2.8 km between mean locations
from the time she was raising one young litter of cubs (period 1) to the time she
was raising her consecutive young litter (period 4) (Table 12-6). This distance
was greater than the distances moved over shorter intervals (i.e., periods 1–2,
Table 12-6. Distance between a female puma’s mean locations (n = 10 females

over 14 birth intervals) during consecutive six-month periods as she raised
young cubs (period 1), older cubs (period 2), regained solitary status (period 3),
and raised her subsequent litter of young cubs (period 4), San Andres Moun-
tains, New Mexico.
Distance between mean locations (km)
Female Period 1 to Period 2 Period 2 to Period 3 Period 3 to Period 4 Period 1 to Period 4
15 0.4 2.3 1.6 3.5

21 10.2 10.1 3.8 4.8
37 1.5 7.5 0.5 8.8
37 6.0 1.4 1.9 2.8
41 2.3 2.2 1.8 2.9
41 1.3 2.1 0.8 1.7
47 0.8 0.1 1.3 2.0
47 3.2 3.0 1.7 2.0
65 1.9 5.3 1.6 3.1
91 2.7 0.5 0.8 2.9
54 1.5 2.0 0.4 3.7
54 0.6 1.1 0.2 0.6
103 1.9 1.5 1.7 1.8
107 1.4 0.4 8.7 10.4
Mean ± SD 2.6 ± 2.6 2.8 ± 2.9 1.9 ± 2.2 3.7 ± 2.8
2–3, and 3–4) [13] and indicated some females may not raise small cubs from
consecutive litters in the same localities. However, we documented two cases in
which females gave birth to consecutive litters at the same nursery. Additionally,
the few females we could follow continuously over long periods generally did
not shift in one direction over time but instead returned to places they had pre-
viously occupied (see Fig. 12-12).
On the San Andres Mountains, where prey did not migrate, female home
range shifts appeared to benefit fitness. Reproductive success may be enhanced
through avoidance of other females; for example, a shift may result when a
female with vulnerable young accommodates an unrelated female (e.g., F21’s
shifting behavior after the arrival of F103). Shifts can also increase inclusive fit-
ness when resources are abundant, dispersal risks are relatively high, and the
individuals that are accommodated are close relatives (e.g., F37 accommodating
daughter F44, the matrilineal structure described in Chapter 7). Similarly, Hop-
kins (1989) speculated that maternal partitioning may have caused a shift in one
of his resident female pumas in a coastal range of California. Such mother-

daughter behavior also has been documented in tigers (Sunquist 1981, Smith et
al. 1987). A female puma may also shift away from conditions that produced
poor reproductive success. We suspect F107 shifted because her young cubs
(father unknown) were cannibalized by resident male M88 three days after he
was in a fight with a relatively new immigrant, M193. The heightened state of
aggression probably exhibited by the two males during this period, and more
importantly the loss of her litter, probably prompted F107 to move (Plate 3).
She successfully produced her subsequent litter (fathered by M88) within an
area located deeper within M88’s home range. Since she did not return to her
former home range after her cubs were born, this could be considered home
range abandonment. Although other studies have documented shifts of adjacent
resident females into vacated areas (Maehr et al. 1991, Laing and Lindzey 1993,
Spreadbury et al. 1996, Maehr 1997b), complete abandonment of an estab-
lished home range is apparently quite rare. We could only find two probable
cases in the literature: an adult female in British Columbia abandoned her home
range and moved about 15 km south to an area within prime elk winter range
that was vacated by another female (Spreadbury 1989), and an adult female in
Florida moved to an adjacent but vacated home range created by the removal of
a nonreproductive female (Maehr 1997b). A female may also shift away from
maturing offspring, possibly to force independence (e.g., F21, F28, F37 in our
study; Beier 1995).
SHIFTS IN RELATION TO A CHANGING FOOD BASE

Although a declining food resource may eventually stimulate home range shifts
or possibly abandonment as pumas seek more productive habitats, we did not
observe any significant shifts that could be attributed to changes in food abun-
dance. Only two home range shifts (M46, F21) occurred during the years of
dramatic deer declines, and both could be attributed to intraspecific interac-
tions. Even during the last full year of the study, both male and female pumas
showed strong fidelity. For females, the deer decline apparently was not severe
enough or did not occur over a long-enough period of time to force abandon-
ment. Males with access to mates apparently had no reason to abandon an estab-
lished territory.
Homing by Translocated Pumas

Perhaps the most dramatic exhibitions of home range fidelity were displayed by
pumas we translocated from the Treatment Area to northern New Mexico (Ruth
et al. 1998). Eight of fourteen pumas (57 percent) exhibited homing behavior
after translocation, and two (M49, M88) actually returned to their original
home ranges. Of the translocated pumas, the ones that exhibited the greatest
movements and homing tendencies after release had been prime, reproducing
residents on the San Andres Mountains. The considerable value of the home
range is indicated by the attempts of these residents to return to it. Familiarity
with resources (including mates), dominance status over neighboring pumas,
and the ability to protect dependent progeny from those neighbors may all be
factors that drive a puma to return home. The two returning males had been
adult residents on the study area for one to three years prior to being translo-
cated 465–490 km to northern New Mexico. M88 and M49 took 166 and 469
days, respectively, to make the return trip to their former ranges. Although M88
maintained his home range through the end of the study and successfully bred
at least six times with three different females (once before translocation and five
times after returning), M49 was not as fortunate. Within two weeks of his
return, M49 made a 45-km northward movement outside his former home
range and died. M49’s home range abandonment and death coincide with an
association he had with M3, who had claimed most of M49’s home range after
his long absence. It is possible M3 inflicted mortal injuries, but, because M49’s
carcass was badly decomposed, we could not determine cause of death. M88
may have been more successful at reclaiming his former range because he was
not absent for as long a period as M49. Consequently, he was probably not as
physically stressed, and it is possible his male neighbor still recognized his dom-
inance.
Benefits of Fidelity and the Two-Strategies Hypothesis

Prior to the New Mexico research, only limited analyses of home range fidelity
in pumas had been conducted. Seidensticker et al. (1973:17) stated that it was
“clear that resident adults reused definite areas, although there were seasonal pat-
terns, changes over time, and individual variations.” Strong fidelity by adults
would be expected under the land tenure system described for central Idaho
pumas. Other biologists who examined fidelity did so to determine whether
pumas shift seasonally in response to migrations of prey. Seidensticker et al.
(1973) found that pumas left winter home ranges to follow prey to summer
ranges and then returned to the same winter home ranges the following year. In
a more-detailed analysis of pumas in southwest Colorado, Anderson et al.
(1992) found that directional shifts in a puma’s center of activity from one sea-
son to the next mimicked the directional movements of migrating mule deer
and elk. Pierce et al. (1999) found two patterns for pumas and prey that
migrated in California’s Sierra Nevada. Some pumas slowly moved with prey but
did not utilize distinct winter and summer home ranges; others showed distinct
seasonal shifts. Two females that made seasonal shifts exhibited strong fidelity:
over a combined total of twelve seasons, summer and winter home range over-
laps averaged 63.4 percent and 86.4 percent, respectively. Directional home
range shifts of African leopards have also been attributed to seasonal (wet and
dry) density, distribution, and behavior of impala (Bailey 1993).
Puma studies in areas where ungulate prey were generally nonmigratory
(Hopkins 1989, Ross and Jalkotzy 1992, Maehr 1997b, our research), found no
such seasonal shifts. Resident adult Florida panthers appeared to exhibit home
range stability, with the exception of shifts owing to panther deaths; however,
fidelity was not quantified (Maehr et al. 1989, Maehr 1997b). Pumas in Cali-
fornia (Hopkins 1989) maintained relatively stable home ranges; out of thirteen
distance measurements between consecutive annual harmonic mean centers
(three distances for two males, ten distances for five females), there were only
two apparent shifts (33 percent of male, 10 percent of female measurements).
The amount of annual home range shifting in the San Andres Mountains (with
a larger sample size of eighty-nine distance measurements between mean annual
locations) was even lower (10.2 percent of male, 5.0 percent of female measure-
ments).
Most adult female pumas show strong site fidelity over long periods. The
a rea used by a resident adult female was apt not to change much, even after
three years. Female tigers also exhibited strong site fidelity (Sunquist 1981,
Smith et al. 1987). The long-term stability of female home ranges is adaptive ,
since familiarity with the home range is advantageous in the exploitation of
food re s o u rces, especially for females raising cubs. Females also become more
knowledgeable about the best nursery sites and possibly escape routes, which
can be important when facing aggre s s i ve male pumas and other predators. In
addition, fidelity results in increased recognition between the female and local
resident males. These males are probably less apt to harm females or their cubs
than new immigrants (see Chapter 8), and they may deter potentially danger-
ous males from settling in the area. All these reasons should enhance fitness,
and they explain why the translocated female pumas showed such strong hom-
ing tendencies.
Most adult male pumas also showed strong home range fidelity, at least
between consecutive years. Fidelity must provide advantages, considering the
arduous journeys undertaken by M88 and M49 to return to their former
home ranges after translocation. Long-term fidelity should enhance the male’s
recognition of potential mates and paternity. It also provides opportunities
for resident males to become familiar with one another and thus establish
clear dominant-subordinate rolls. This may reduce the number of potentially
debilitating fights between neighbors as well as enable the resident to quickly
identify and challenge new immigrants (who could potentially harm the re s i-
dent male’s current mates or offspring). Conve r s e l y, roaming behavior subjects

males to more potential fights with unknown males, does not ensure paternity,
and puts both females and progeny at greater risk. Although there was a
g reater decline in male fidelity over time (in our sample, 42 percent of males
and 10 percent of females made home range shifts between annual periods
that we re three years apart), such slow shifts would allow most females time to
raise a given male’s litter before his area of influence shifted completely out-
side her home range.
Fidelity in Desert Pumas and the Self-Limiting Hypothesis

Some of our observations on puma home range fidelity and shifting behavior do
not support the self-limiting hypothesis. First, a new immigrant male sometimes
contested an established male’s prior rights to an area, forcing the resident to
yield by shifting his home range. Second, resident females sometimes shifted to
accommodate other female pumas, including daughters and new immigrants.
These shifts facilitated an overall increase in the puma population over time,
even as the deer population declined precipitously. Consequently, the behaviors
we observed did not ensure that the puma population would keep its numbers
below the levels set by the prey.
LP > > KX
1. We radio-monitored eighty-six adult pumas on the San Andres Mountains
for an average of 28.8 months (range = 0–102) each, and obtained an aver-
age of 137 locations (range = 1–569) on each individual.
2. Adult pumas did not exhibit seasonal movements, probably because their
prey did not migrate and winter movements were not restricted by deep
snows.
3. The annual home ranges of adult male pumas ranged from 59.3 to 639.6
km2. Annual home range size for twenty-four males averaged 193.4 km2, or
about 2.8 times the size of the average adult female’s home range. This sup-
ports prediction 1 of the two-strategies hypothesis. Within their ranges,
males moved about 4.1 km (linear distance) each day.
4. Adult female pumas used annual home ranges of 13.1–287.4 km2. Annual
home range size for thirty females averaged 69.9 km2. Females tended to
move shorter distances (1.4 km) each day than males.
5. Females used home ranges over annual periods similar in size to those used
during entire birth intervals. However, females’ home ranges enlarged with
the increasing age and mobility of their cubs and were largest during the
periods when they were without cubs.
6. An adult male’s lifetime home range was about 70 percent larger than his
average annual home range. Females also used larger areas over their life-
times than those they used over annual periods, but the increase was not as
large (45 percent) as for males. The larger lifetime ranges were attributed,
in part, to individuals making home range shifts.
7. Habitat quality, sex, reproductive status, and responses to other pumas
influenced home range sizes of adult pumas.
8. Although there was no direct relationship between puma (male or female)
home range size and deer density, a weak negative relationship between
female density and female home range size suggested that females used
smaller areas when prey was more abundant. The relationship was probably
clouded by uneven distributions in the deer population, especially as the
drought took hold and deer began to concentrate around limited water
sources.
9. Home ranges of remaining adult female pumas did not expand in response
to the removal of six female competitors and five potential mates from the
Treatment Area. Expansions were apparently unnecessary because the
females were obtaining all of the necessary resources within their home
ranges prior to competitor removal. They also did not have to search for
mates because the remaining adult male located and mated with all estrus
females.
10. In the Treatment Area, adult male home range size was strongly and
inversely affected by puma density. Behavior of the one remaining male
before and after the removal of six potential mates and all male competitors
indicated that he responded to the presence and absence of both male and
female pumas but not to changing deer densities. Although the correlations
suggest that the male was most affected by the density of his potential
mates, his best breeding success occurred when he had the least male com-
petition. This supports prediction 3 of the two-strategies hypothesis.
11. Adult female pumas exhibited strong home range fidelity over long periods.
Between consecutive years each female used ranges that overlapped by an
average of 74.4 percent. Even after three years, the proportion of a female’s
initial range that was still utilized averaged 65.5 percent. The distance
between the arithmetic centers of a female’s consecutive annual home
ranges averaged 2.6 km. This supports prediction 5 of the two-strategies
hypothesis.
12. Adult male pumas also exhibited strong home range fidelity between con-
secutive years ( x– = 75.1 percent); however, male home range fidelity
declined more rapidly over time than did female fidelity. In the fourth year,
males were still using, on average, only 43.1 percent of the same area uti-
lized in the first year. The average distance between arithmetic centers of a
male’s consecutive annual ranges was also greater than that found for
females (5.3 km).
13. Females were less prone to make home range shifts than males. Identified
causes of shifts included avoidance of neighboring females raising new lit-
ters, abandonment of cubs that had reached independence, accommodation
of a philopatric daughter, and abandonment of an area where dependent
progeny were cannibalized.
14. Males were more likely to exhibit home range shifts than females we re. Causes
of shifts that occurred over short intervals (six months) included avoidance or
re t reat from new immigrant males, a low density of available mates within the
present home range, and the death or removal of a neighboring resident male.
A few males also exhibited directional shifts over much longer periods. We
suspect these shifts may have resulted from a combination of causes, includ-
ing neighborly disputes that simply took longer to settle.
15. Fidelity was displayed by translocated pumas. Eight of fourteen exhibited
homing behavior, and two adult males returned to their original home
ranges. One of these males maintained this same home range for the next
three years (through the end of the study).
LN N5LN5)L
1. Annual home range sizes (90 percent and 60 percent ADK, 100 percent and
90 percent MCP) of adult males (n = 71 estimates for twenty-four males)
and adult females (n = 72 estimates for thirty females) were different.
Mann-Whitney U tests, Z 9.36, P < 0.001, all tests.
2. Annual and birth interval home range sizes (90 percent and 60 percent
ADK, 100 percent and 90 percent MCP) for the same females (n = 11 esti-
mates for nine females) were only different based on the 100 percent MCP.
Wilcoxon signed-rank tests, Z = 0.889, P = 0.374 for the 90 percent ADK;
Z = –0.889, P = 0.374 for the 60 percent ADK; Z = –2.223, P = 0.026 for
the 100 percent MCP; Z = –0.356, P = 0.722 for the 90 percent MCP.
3. A female’s home range sizes (100 percent and 90 percent MCP) during each
of three consecutive six-month periods within a birth interval (n = seven
females during eight birth intervals) were different. Kruskal-Wallis test, H
= 13.934, 2 d.f., P < 0.001 for the 100 percent MCP; H = 13.139, d.f. = 2,
P = 0.001 for the 90 percent MCP. For six other females during seven peri-
ods when they were raising small cubs and seven consecutive periods when
they were raising large cubs, home range sizes were larger when they were
raising large cubs. Wilcoxon signed-rank tests, Z = 3.30, P < 0.001 for both
the 100 percent and the 90 percent MCP.
4. Mean annual and cumulative home range sizes for ten adult males moni-
tored from 48 to 101 months ( x– = 71.1 ± 18.1 months) each, were differ-
ent. Wilcoxon signed-rank test, Z = 3.92, P < 0.001, based on all home
range estimators (90 percent and 60 percent ADK, 100 percent and 90 per-
cent MCP). Mean annual and cumulative home range sizes for six adult
females monitored from fifty to eighty-one months ( x– = 63.8 ± 12.7
months) each, were different. Wilcoxon signed-rank tests, Z = 3.06, P =
0.002, for all four home range estimators.
5. Test for difference in puma home range size (90 percent ADK) during years
of low deer abundance (n1) and years of high deer abundance (n2), using
Mann-Whitney U tests. For Reference Area males, n1 = 15, n2 = 10, U =
96.0, P = 0.24; for Treatment Area males, n1 = 10, n2 = 4, U = 22.5, P =
0.77; for males in the Reference and Treatment Areas combined, n1 = 25,
n2 = 14, U = 197.5, P = 0.43. For Reference Area females, n1 = 13, n2 = 13,
U = 65.0, P = 0.32; for Treatment Area females, n1 = 10, n2 = 8, U = 29.0,
P = 0.33; for Reference and Treatment Area females combined, n1 = 23, n2
= 21, U = 186.0, P = 0.19.
6. Test for linear relationship between puma home range size (100 percent
MCP) and puma density across populations of pumas. Linear regression of
female home range size on female density, n = 5, female home range =
156.0612 – 38.7590 (female density), r2 = 0.09, P = 0.62. Linear regression
of male home range size on male density, for n = 4, male home range =
493.2500 – 158.2692 (male density), r2 = 0.31, P = 0.45.
7. Test for associations between adult female annual home range size (90 per-
cent ADK) and year-round density of adult males, adult females, and total
adults in the Reference Area (RA, 1989–1994) and Treatment Area (TA,
1988–1994). Spearman correlation coefficient (2-tailed test) for the RA:
female range size and male density, rs = –0.543, P > 0.20; female range size
and female density, rs = –0.429, P > 0.20; female range size and total den-
sity, rs = –0.429, P > 0.20. For the TA: female range size and male density,
rs = 0.643, P = 0.125; female range size and female density, rs = –0.536, P
= 0.2; female home range size and total density, rs = –0.214, P > 0.2. Com-
parisons using the 90 percent MCP resulted in slightly weaker but similar
correlations in both the RA and the TA; they are not reported here.
8. Test for associations between adult male annual home range size (90 percent
ADK) and year-round density of adult males, adult females, and total adults
in the Re f e rence Area (RA, 1989–1994) and Treatment Area (TA,
1988–1994). Spearman correlation coefficient (2-tailed test) for the RA:
male range size and male density, rs = 0.429, P > 0.20; male range size and
female density, rs = 0.371, P > 0.20; male range size and total density, rs =
0.371, P > 0.20. For the TA: male range size and male density, rs = –0.429,
P > 0.20; male range size and female density, rs = –0.821, P = 0.03; male
range size and total density, rs = –0.964, P = 0.002. Comparisons using the
90 percent MCP resulted in slightly weaker but similar correlations in both
the RA and the TA; they are not reported here.
9. Lilliefors test (Lilliefors 1967) to determine if distributions of FI differed
from normal. For males: FI1, D = 0.14, n = 48, P = 0.07; FI2, D = 0.12, n
= 31, P = 0.27; FI3, D = 0.15, n = 19, P = 0.34. For females: FI1, D = 0.12,
n = 39, P = 0.15; FI2, D = 0.17, n = 20, P = 0.15; FI3, D = 0.17, n = 10, P
= 0.61.
10. Test for differences between the distributions of FI for male and female
pumas. Two-sample Kolmogorov-Smirnov test: for FI1, nmale = 48, nfemale =
39, D = 0.09, P = 0.98; for FI2, nmale = 31, nfemale = 20, D = 0.32, P = 0.11;
for FI3, nmale = 19, nfemale = 10, D = 0.45, P = 0.13.
11. Test for differences in distances moved between mean annual locations of
males (n = 49 distance measures for 16 males, x– = 5.3 ± 4.8 km, range =
0.2–23.3 km) and females (n = 40 distance measures for 20 females, x– = 2.6
± 2.1 km, range = 0.3–10.0 km). Mann-Whitney U test, U = 1435,
P < 0.001.
12. Test for differences in distances moved between mean locations by females
during consecutive periods (1, 2, 3, and 4; see text) of their reproductive
cycle. Distances between mean locations: period 1–2 = 2.56 ± 2.60 km;
period 2–3 = 2.83 ± 2.90 km; period 3–4 = 1.90 ± 2.15 km, n = 14.
Kruskal-Wallis test, H = 1.41, 2 d.f., P = 0.49.
13. Test for differences in distances moved by females between mean locations
over consecutive periods of their reproductive cycle (i.e., periods 1–2, 2–3,
and 3–4) and distances moved between mean locations while raising young
cubs of two consecutive litters (i.e., periods 1–4), n = 14. Mann-Whitney
U test: for periods 1–2 and 1–4, U = 55.0, P = 0.048; for periods 2–3 and
1–4, U = 67.0, P = 0.15; for periods 3–4 and 1–4, U = 36.5, P = 0.005.
Chapter 13
Subadult Ranging Behavior
Subadults are important to the establishment, maintenance, and growth of

puma populations as well as to the spread of genetic material between popula-
tions. Consequently, it is important to understand how they behave. In our
analysis, we used the following definitions. Natal home range was the area occu-
pied by a cub’s mother for the twelve-month period after the cub’s birth. Inde -
pendent home range was the area occupied by an independent progeny during the
twelve-month period after site attachment. Site attachment was exhibited when
a puma stopped making one-way directional movements and used a predictable
area into adulthood. Subadult home range only included the area used by a puma
after it showed site attachment and before it was classified as an adult; therefore,
it was typically of shorter duration than the independent home range. A
philopatric puma used an adult home range that overlapped its natal range by 5
percent or more, based on the 90 percent minimum convex polygon. A transient
home range was an area that an animal used after leaving its natal range but later
abandoned (Beier 1995). Duration of transient behavior was the time between
independence and subsequent site attachment with no further dispersal.
Subadult and transient home ranges generally spanned short periods and were
represented by minimal numbers of radio locations; consequently, we only
depicted them using the MCP home range estimator. To avoid confusion, all
home range sizes in this chapter are based on the 90 percent MCP.
We documented the fates of forty-three progeny after independence (twenty
males, twenty-three females). Of these, thirteen remained philopatric into adult-
hood, twenty-seven dispersed, and three died within their natal home ranges
shortly after independence. Obtaining detailed information on ranging behav-
231
ior during the entire period a puma was a subadult was difficult because only
fourteen pumas wore radio collars from the time they became independent until
the time they either died as subadults (n = 5) or reached adulthood (n = 9). We
learned about the fates of tagged (but not radio-collared) cubs either by recap-
turing them on the study area post-independence (n = 23) or from tag returns
(n = 6). Tags were returned to us by the New Mexico Department of Game and
Fish (from pumas killed by hunters), by depredation control officers, and by
drivers involved in automobile-puma collisions.
Philopatry in Females
All thirteen philopatric pumas were females, and ten of those were sister pairs.
We radio-tracked three philopatric females from independence to adulthood:
F183 was born on the Reference Area and became independent during the deer
decline; siblings F107 and F109 became independent about three months after
we stopped removing pumas from the Treatment Area and at the apex of the
deer population. These three females became independent between 11.1 and
15.1 months of age and were classified as adults when eighteen to twenty-eight
months old. Although they were philopatric, at least one of these females (F109)
made a dispersal attempt. F109 dispersed 66 km north in a twenty-one-day
period immediately after independence. However, she then spent the next
twenty-eight days traveling back to her natal home range, where she remained
into adulthood. F183 made one foray 12 km south of the southern border of
her natal range when she was twenty-three months old, but she returned within
seven days. It is possible both F109 and F183 were assessing the prospects of dis-
persal and philopatry, which suggests the behaviors are somewhat flexible and
not completely innate. F107 did not show any dispersal behavior (see Plate 3).
These three females occupied subadult home ranges for an average of 8.0 ± 4.0
months that encompassed areas averaging 35.6 ± 16.0 km2 (90 percent MCP).
Their subadult home range sizes were similar to the average annual home range
for adult females (44.6 ± 18.5 km2, see Appendix 3). F107 and F109 subse-
quently produced cubs in areas encompassed by their subadult home ranges.
Although we do not know whether other philopatric females attempted
subadult dispersal moves before they were recaptured and radio-collared as
subadults or young adults, they subsequently exhibited strong natal range
fidelity. The average distance between the arithmetic centers of a philopatric
female’s natal range and independent home range was 3.2 km (see Table 9-1),
similar to annual distances moved by resident adult females (see Chapter 12).
Home range sizes of newly established adult females were also consistent with
those of older adult females. Seven philopatric females radio-tracked during the
CHAPTER 13. SUBADULT RANGING BEHAVIOR 233
twelve-month period immediately after adult onset had a mean home range size
of 45.6 ± 15.4 km2, whereas the annual home range size of different females that
were at least three years old (n = 55 annual measurements from twenty-seven
females) averaged 42.9 ± 28.7 km2 [1].
Philopatry resulted in the formation of eight matrilines (see Chapter 7 and
Plate 4). We documented as many as five related females (i.e., mothers, daugh-
ters, sisters) that inhabited abutting or overlapping home ranges in a given area
of the mountain range at the same time (Plate 5).
Female Dispersal
Some female progeny dispersed. In fact, of the females that survived to adult
ages, we estimated 33 to 59 percent actually emigrated from the San Andres
Mountains (see Chapter 9). Nine dispersing females traveled an average of 34.6
km from their natal ranges to their last locations or independent home ranges
(see Table 9-1). Of those, three (F9, F10, F54) were radio-monitored during the
entire period between independence and adult onset. F9 and F10 were born in
the Treatment Area in 1986; F54 was born in the Reference Area in 1988. These
females became independent between 9.8 and 14.3 months of age; the youngest,
F10, was orphaned when her mother was killed by an immigrant male. F10 and
F54 dispersed an average of 11 km within seven days of independence, then dis-
played site attachment to areas adjacent to their natal home ranges. F9 spent
about twelve days in the vicinity of her natal range post-independence, then dis-
persed 81 km north in under forty-one days and began to show site attachment
near the western boundary of the Reference Area. The distance between the cen-
ters of her natal and independent home ranges (78.5 km) was almost identical
to the distance she traveled during her dispersal move, suggesting that she set-
tled quickly. For the 6.5 ± 1.2 months that these three females showed site
attachment as subadults, they used home ranges averaging 66.6 ± 43.1 km2
(range = 24.5–110.6 km), all within the limits of home range sizes found for
adult females. All three females showed subsequent breeding activity (i.e., asso-
ciations with resident males) within the areas encompassed by their subadult
home ranges. Although F9 and F10 died before producing cubs, F54 bore three
litters and was still alive at the end of the study.
Four other natal dispersers provided information to suggest female dispersal
was often of brief duration and ceased prior to adulthood. F160 was captured
and radio-collared within her natal home range at about eighteen months of age,
four months after her mother (F145) died. Genetic testing supported our con-
tention that she was indeed F145’s daughter. Within five days of capture, F160
emigrated from the study area. Less than forty-six days later, she showed site
Slide @171%
PHOTO 23. Brian Spreadbury with
four-month-old puma F54. As an
adult, F54 established a home range
adjacent to her mother and bore three
litters.
attachment to an area 37 km to the east of her capture site. She became preg-
nant at the age of twenty-five months, 182 days after showing site attachment.
Another female, F150, whom we suspect was also orphaned (her mother disap-
peared), began dispersing at nine months of age. She moved 27 km to the west
edge of the San Andres Mountains within ten days, and then spent about twelve
days at this western border before trekking 51 km across the Jornada del Muerto
to the Caballo Mountains. Until she shed her radio collar 119 days later, F150
used a 57-km2 area (n = 13 locations) in the Caballo Mountains. Because of the
short monitoring period, we cannot be sure whether this was a transient home
range or she was showing site attachment. Two other tagged females (F28, F149)
had completed dispersal moves of 17–20 km before they were recaptured and
radio collared at 17.7 to 18.5 months of age. Both displayed strong site fidelity
post-capture and produced their first litters when twenty-three to thirty-six
months old.
One female displayed dispersal behavior over a long period (Plate 6). F148
was captured and radio-collared near the study area’s southern border at eight-
een months of age. We suspect she was born in the adjacent Organ Mountains.
F148 used two overlapping transient home ranges during a 185-day period after
capture. Five other adult female home ranges overlapped F148’s transient
ranges; two gave birth to cubs during F148’s short tenure, and the litters of two
others had recently become independent. F148 then dispersed 35 km north over
ten days (and through the home ranges of two other adult females). In the fol-
lowing seven days she abandoned the San Andres Mountains and traveled 56 km
southwest, across the desert flats to the Red House Mountains. After having
exhibited transient behavior for 202 days, F148 finally showed site attachment.
She established an independent home range of 108.6 km2 over the next year that
included naturally fragmented puma habitat in the Red House and Caballo
Mountains. Ten months after showing site attachment, and at the age of thirty
months, F148 localized in a 1- to 2-km2 area for forty-two days. Her behavior
suggested she had given birth to cubs.
Duration of transient behavior in subadult females (n = 9: F9, F10, F54,
F107, F109, F148, F150, F160, F183) was typically short, averaging 49 ± 79
days (range = 0–202). However, the average independent home range size of six
progeny that remained within the study area (F9, F10, F54, F107, F109, F183)
was much smaller (x– = 48.1 ± 31.2 km, range = 19.8–101.1) than the inde-
pendent home range size of two females that emigrated (F148, F160; x– = 161.6
± 75.0, range = 108.6–214.7). F160 established a home range in the confines of
the Malpais, an expanse of sharp igneous rock and deep crevices formed by an
ancient lava flow between the Oscura and Sacramento Mountains. F148’s home
range encompassed mountainous habitat patches fragmented by desert basins.
We had explored both areas and found them to be relatively prey-poor as com-
pared to the San Andres Mountains.
The rapidity with which most females established independent and adult
home ranges on the study area, as well as the high degree of philopatry, sug-
gested that females were quickly finding sufficient resources to survive and pro-
create and also that competition from other pumas was generally insufficient to
deter settling. During the early stages of the study, when puma density was low
and deer numbers were increasing, it was easy to understand how females could
settle quickly. The behavior of F148 also suggests that it was becoming more dif-
ficult to settle as the area filled up with females and their young. However, we
documented philopatry as late as August 1994, when two siblings (F181, F183)
in the Reference Area were just reaching adulthood. F181 was illegally killed
within her natal area in December 1994, but her sister remained philopatric
through the end of the study. During this time, female puma density in the Ref-
erence Area was at its highest level and the deer population was well into its
decline.
So, why did F183 stay and F148 leave? Both dispersal and philopatry are
adaptive strategies for female pumas. Philopatry may be a better strategy under
some circumstances because it has the potential of increasing individual as well
as inclusive fitness (see Chapter 7). F183 may have put herself at greater risk by
dispersing through unfamiliar and potentially hostile country instead of staying
in a food-poor environment. A dead subadult leaves no progeny. Her mother
should also be willing to share resources with her daughter if her inclusive fit-
ness increases. A similar benefit would not accrue to a female that shared mini-
mal resources with an unrelated female such as F148 (who was not born on the
study area). Although we did not document fighting between females, and we
found no greater level of associations between independent females and their
mothers as between unrelated females, some of the largest spatial overlaps of
annual home ranges were between related females (see Chapter 14). Two sisters
(F107, F109) remained philopatric and produced litters within one month of
each other at nurseries that were less than 600 m apart. This suggests related
females may be more tolerant of one another. The behavior of three pumas (F10,
F150, F160) that dispersed even though their natal ranges were vacated by the
deaths of their mothers indicates that philopatric and dispersal behaviors also
have innate components. A more thorough discussion of why pumas disperse is
presented later in this chapter.
Male Dispersal
No males that survived to the average dispersal age of about fifteen months
exhibited philopatry. Male progeny dispersed an average of 102.6 km (n = 18;
see Table 9-1). We radio-tracked three progeny (M26, M82, M92) during the
entire period from independence to adulthood. Both dispersal onset and dis-
persal duration were quite variable in these three males. M26 dispersed at four-
teen months, immediately after independence. In contrast, M82 and M92, who
became independent at twelve to sixteen months of age, remained within their
natal areas 52 to 238 days before successfully dispersing. M26, M82, and M92
showed site attachment 145, 259, and 43 days, respectively, after initiating suc-
cessful dispersal moves and emigrating from the study area. M26 and M92 were
younger (18.8 and 19.2 months old, respectively) than M82 (28.7 months)
when they began establishing independent home ranges. Distances between
natal and independent home range centers averaged 167.2 ± 29.9 km (range =
134.2–192.5) for the three males.
The use of transient home ranges also varied between males. M92 made one
swift dispersal movement to the area where he established an independent home
range northeast of the study area. In contrast, M26 and M82 each used one to
three transient ranges prior to showing site attachment. At 15.6 months of age,
M82 made a circuitous dispersal attempt south of his natal area that spanned
twenty-eight days and covered 114 km. He started a second, successful disper-
sal move at 20.2 months of age. After dispersing 90 km to the south end of the
study area over a thirty-eight-day period, M82 spent ninety-seven days in a 46-
km2 transient home range in the Organ Mountains. Further dispersal was
impeded to the south by minimal mountainous terrain and the town of El Paso,
whereas dispersal to the east and west entailed crossing wide desert basins. We
were unable to radio-locate M82 for eighteen days; then, by expanding our
flight search eastward, we found him in the Sacramento Mountains. He had dis-
persed 74 km across the stark expanse of the Tularosa Basin. During the subse-
quent 106 days, M82 traveled 73 km east-southeast through broken mountain-
ous terrain, where he initiated site attachment in the Guadalupe Mountains (see Plate 7).
M26 used three transient home ranges prior to showing site attachment. He
spent forty-nine days in the Organ Mountains (immediately adjacent to his
natal area), fourteen days in the Doña Ana Mountains, and twenty-seven days
in the Caballo Mountains during his 145-day dispersal move to the Black
Range. All three transient ranges were in mountainous patches surrounded by
flat desert basins. M26 may have abandoned his transient range in the Doña
Ana Mountains because of its size (about 25 km2), which was considerably
smaller than the home ranges of resident males on the San Andres Mountains.
Our cursory survey also indicated a sparse prey base, an absence of resident
pumas, and considerable recreational use by people from Las Cruces. Although
the Caballo Mountains (his third transient range), encompassed about 200 km2
of mountainous habitat, M26 was only located there three times as he moved
north, then west to the Black Range (Sweanor et al. 2000, Fig. 3). M82 and
M26 both used transient home ranges when they reached mountainous patches
surrounded by flat desert basins, suggesting that either the males examined the
sites, found them unsuitable, and continued their dispersal moves, or the desert
basins imposed temporary dispersal barriers.
Evidence also suggests resident males may encourage continued dispersal.
We suspect M26 was persuaded to leave his first transient range in the Organ
Mountains by M1, a resident that used an area overlapping M26’s natal range.
For three days the two males confronted each other in a jumble of boulders on
Antelope Hill, where the mountains terminate and the Tularosa Basin begins.
On the fourth day, M26 evaded M1 and dispersed west to the Doña Ana Moun-
tains. The seriousness of the encounter can be conveyed by recounting the fate
of a less-fortunate subadult six months previously. M1 pursued subadult M300
for about 2 km along the southern flank of Black Mountain. In an attempt to
escape, M300 bounded up a power pole and was promptly electrocuted.
Another subadult, M221, who was born outside the study area, used a small
transient range of about 10 km2 for two months after his capture. Within three
days of being located near resident M124 and a fresh deer kill, he made a 14 km
northward movement. Over the next seven days, M221 traveled another 40 km
further north.
Detailed information from M92, as well as less-complete data from two
other male progeny (M23, M122), indicated that subadult males could disperse
long distances and begin showing site attachment over a short time period. M23
became independent and dispersed between the ages of 10.8 and 14.4 months.
After his recapture at 14.4 months and a dispersal move of 73 km, he showed
site attachment. He utilized a 115-km2 subadult home range until we classified
him as an adult at twenty-two months of age. Less than a month later, we
translocated him. A second male, M122, dispersed at 14.5 months, seven days
after independence. In twenty-six days, M122 dispersed 118 km northeast
across the Tularosa Basin and onto the Capitan Mountains. Over the next fifty-
six days and until he was killed by another male puma, he used a 22-km2 area
(n = 6 locations), indicating he was either using a transient home range or show-
ing site attachment. The duration of transient behavior in four males (M23,
M26, M82, M92) averaged 211 ± 192 days (range = 95–497). Males took
longer than females to settle because they typically dispersed longer distances
and probably had to locate an area where they were competitive with other
males for breeding females. The competition in particular would require that
subadult males grow in mass and experience (see Chapter 8), which would also
take time.
The independent home range size of three males (M26, M82, M92) that
emigrated from the study area averaged 490.7 ± 217.5 km2 (range
356.5–741.7). This was much larger than the 115-km2 area used by M23, a
non-emigrating male, over the 8.5 months he was monitored after site attach-
ment. The behavior of two other male progeny (M73, M124) also suggested
that dispersers remaining within the study area had smaller home ranges than
newly established emigrants. M73 and M124 were recaptured and radio-collared on
the study area at an average age of 28.2 ± 1.1 months, and they had already com-
pleted dispersal moves of 47.0–69.9 km. During the year following recapture,
they used home ranges of 61.3 and 129.5 km2, which were within the limits
found for adult males on the study area (see Appendix 3). We suspect the larger
areas used by emigrant males were most likely caused by the presence of fewer
competitors or mates, or fights that resulted in home range shifts. The behavior
we observed in resident M3 after our experimental removal in the Treatment
Area indicates that a reduced density of pumas (males and/or females) can cause
the expansion of a male’s home range. In Utah, the extremely large home range
size of the male puma, and the corresponding density that was one of the low-
est reported for pumas (Hemker et al. 1984), also suggest low densities may
cause (or allow) males to use large areas. We also documented several cases in
which fights resulted in large home range shifts. The behavior of M82, the only
emigrant we followed for two full years after site attachment, suggests that his
large home range may have been the result of a fight and subsequent shift into
better habitat (i.e., more mates). M82 showed site attachment for about four
months; then, over the next seven months, his home range center (mean x and
y coordinates) shifted 13.3 km to the east. During the year after that, his home
range center shifted only 3.9 km (less than the average distance between con-
secutive annual home range centers of resident males on our study area; see
“Adult Home Range Fidelity” in Chapter 12) and his range size shrank from
356.5 to 96.6 km2 (Plate 7).
Frustrated Dispersal
Male philopatry appears to be exceptional in pumas. It has only been docu-
mented in Florida, where extensive agricultural and urban development isolated
the panther population. Philopatry was essentially forced because panthers
could no longer freely disperse north out of southern Florida. All male subadult
panthers attempted to disperse, but most (53 percent) were killed by other
males. Although some of the subadult males temporarily resided in poor-qual-
ity habitat apparently devoid of adult females, survivors eventually returned to
the vicinity of their natal areas (Maehr 1997a).
Similarly, dispersal of male pumas born in the Santa Ana Mountains of
Southern California has been frustrated by a burgeoning human population
along the periphery of those mountains (Beier 1995). All eight male progeny
studied there attempted dispersal. Male dispersers moved rapidly between one
and four small, elongate, transient home ranges along the wildland–urban inter-
face. One male returned to establish a transient range abutting his natal range,
and two others returned to their natal ranges for seven and ten days, then
departed without returning. Although Beier suggested that these behaviors were
probably due to attempts by dispersers to avoid conflict with territorial adult
males, we believe that they may also have been influenced by urban barriers to
dispersal. We also documented the use of transient home ranges, but they always
occurred along the mountain–desert basin interface and were used prior to dis-
persal movements to mountainous patches on the other side of the basin. If the
Santa Ana Mountains were not surrounded by urbanization, we think the males
would have dispersed farther than they did, and probably most of them would
have emigrated from the entire mountain range toward more distant puma habi-
tats before attempting to establish home ranges. Only two of the males (25 per-
cent) survived to adulthood and established home ranges 56 to 75 km from their

natal areas. One was along the eastern edge of the Santa Ana Mountains study
area; the other was mostly on the northern end of these mountains but included
the small Chino Hills to the north (Beier and Barrett 1993). Both the San
Andres (2,059 km2) and Santa Ana Mountains (2,070 km2) study areas were of
similar size; correspondingly, the six males that remained within the San Andres
Mountains into adulthood dispersed distances similar (x– = 66 ± 15 km) to those
of the two California males. However, seven other male progeny from the San
Andres Mountains dispersed an average of 159.5 ± 53.5 km to areas outside the
San Andres Mountains. In contrast to Southern California, human development
surrounding our study area posed few artificial barriers to dispersal (Sweanor et
al. 2000). It took the two surviving California males an average of 434 days to
establish stable home ranges, whereas the average transient phase for four male
progeny born on the San Andres Mountains was 211 days. This suggests males
from the San Andres Mountains could cover more distance and consequently
find suitable habitat more quickly.
In Beier’s study, dispersal of the only radio-collared female probably was also
thwarted by the extensive human development. She covered 342 km or more
and changed directions at least five times upon encountering urban areas. Four
months after initiating dispersal, she returned to her natal area. She bedded near
her mother, then died near that location five days later from undetermined
causes (Beier 1995).
Why Do Pumas Disperse?

Theoretically, the ultimate adaptive significance of dispersal in pumas is to
increase individual reproductive success (Morris 1982). There are three leading
hypotheses on proximate causes of dispersal in large mammals: (1) competition
for mates, (2) competition for resources, and (3) avoidance of inbreeding (Sin-
clair 1992). We address each of these in turn as they pertain to pumas.
If competition for mates influences dispersal, then it is theorized that subor-
dinate individuals of the sex in which competition is most intense should dis-
perse (Dobson 1982). In naturally functioning puma populations, adult males
vigorously compete for a limited number of estrus females, whereas there are
ample mates available for females (see Chapter 7). Subadult males also tend to
disperse greater distances than females, and only females commonly exhibit
philopatry; consequently, mate competition appears to be a probable explana-
tion for male, but not female, dispersal. It seems beneficial for male progeny to
disperse away from areas that are dominated by their fathers. The cost of stay-
ing in the natal area can be extreme, since sons, who may be indistinguishable
from other male competitors, can be killed by fathers. We documented just such
a fight between M88 and his subadult son M108. Although dispersing subadults
risk being killed by unrelated males encountered during dispersal moves (e.g.,
the deaths of subadult males M140 and M122), it is probably most adaptive for
them to continue dispersing until they are physiologically competitive (i.e.,
larger, more experienced, and sexually mature). The possibility of altercations
with larger, dominant adult males could be strong motivations to keep moving.
Settlement might be triggered by finding suitable mates in an area with relatively
little competition from other males, and the onset of puberty. This all supports
the competition for mates hypothesis; however, other findings do not. Males
should disperse only as far as necessary to avoid competition and find suitable
mates. Since new males were continually coming into our population and estab-
lishing residency, there were apparently adequate resources and tolerable com-
petition. Consequently, most of the male recruits in our growing population
should have been progeny born on the San Andres Mountains. They were not.
Also, once males established their adult home ranges, direct competition did not
trigger the kind of long-distance dispersal exhibited by subadult males. Instead,
the loser of a conflict, if he survived, tended to shift his home range just far
enough to avoid the victor (see Chapters 12 and 14).
Competition for food does not adequately explain dispersal in males either.
Most male progeny were not just dispersing beyond their natal areas to the near-
est vacant territory but were emigrating entirely from the growing puma popu-
lation on the San Andres Mountains. We know there was adequate space and
food for male progeny, particularly when the deer population was growing,
because male immigrants were recruited into the population. If dispersing males
were striving to reduce competition for food with their own mother and
philopatric sisters, which would enhance inclusive fitness, we would expect
males to exhibit a pattern of dispersing just beyond the ranges of related females.
Hence, males would reside adjacent to close relatives. But so far, there is little or
no evidence for this in puma populations unbound by human development.
Even so, males would have to compete with other nonrelated pumas. There
must be some other explanation compelling them to engage in long-distance
moves with the relatively high risk of mortality. Unless the fitness of dispersing
males is benefited directly by some other means, we would expect the dispersal
trait to be selected against.
On the other hand, competition for resources may adequately explain dis-
persal in some female pro g e n y, particularly if other females are philopatric
and cause a rapid increase in the local density of adult females (see Chapter
10). Re p ro d u c t i ve success of some females could be enhanced if they left
their natal areas and dispersed just far enough to find suitable habitat with
reduced competition. This may explain the shorter dispersal distances for
females re l a t i ve to males, and the tendency for some females to disperse just
b e yond the natal area. Re s o u rce competition may also explain why some
females disperse long distances (i.e., for females). They may have to keep
m oving until they find an area with adequate habitat and where competition
is tolerable.
Clearly, dispersal behavior in female pumas is more flexible than that exhib-
ited by males. Dual behaviors (i.e., philopatry and dispersal) have been docu-
mented in female puma siblings in Alberta (Ross and Jalkotzy 1992). Although
we could only assume that sisters of some philopatric females dispersed (because
we never recaptured them as adults), we observed where female cubs from dif-
ferent litters but the same mother (n = 2) showed alternate behaviors. The tac-
tic adopted is probably influenced by local environmental conditions. Female
dispersal may be due to resource competition with a partial density-dependent
effect. Considering that dispersing females probably have lower survival rates
(see Chapter 8) and lower reproductive success (see Chapter 7), we would expect
females to become philopatric or settle adjacent to their natal area if there was
an adequate food supply. But as competition for food intensifies, and especially
when food is limited, dispersal costs may be less than the costs of philopatry.
Hence, dispersal could become a more successful tactic. Some threshold of com-
petition may have to be perceived by individual female progeny to trigger dis-
persal. The payoff could come later if the female were able to settle in a more
favorable environment for rearing cubs. Competition with other carnivores may
similarly affect dispersal in subadult female pumas. A puma’s reproductive suc-
cess may be higher in areas with reduced interspecific competition. This rela-
tionship may be manifested on ranges where pumas and wolves are sympatric;
wolves have been known to usurp puma prey kills and to aggressively pursue and
even kill pumas (White and Boyd 1989, Boyd and Neale 1992, T. K. Ruth, per-
sonal communication). It is possible that female dispersal is socially mediated in
some way. For example, dominant female progeny may tend to be philopatric
while subordinate female siblings tend to disperse. Because of the near-impossi-
bility of observing behavior among family members, there are no empirical data
to support this notion.
Whatever the cause, female dispersal would have the added benefit of
expanding puma populations, particularly with obligatory male dispersal. Puma
natural history has provided ample opportunities for such expansions (see Chap-
ter 2). Climatic changes during interglacial periods certainly created more habi-
tat in North America. Moreover, drying of the isthmus linking North and South
America opened an entire continent to puma colonization. If Culver’s (1999)
scenario about a late Pleistocene extinction of puma in North America is cor-
rect, then population expansion into the continent may have occurred more
than once. Much more recently, beginning about the mid-1960s, pumas appar-
ently recolonized historical habitats in western North America following pro-
tective legislation that ended the era of indiscriminate killing that extirpated
local puma populations.
Avoidance of inbreeding seems to be the best explanation for dispersal in
male pumas. If female matrilines are naturally characteristic of puma popula-
tions as we believe they are, then opportunities for inbreeding would abound for
philopatric males. They could breed with their mother, sisters, daughters, and
granddaughters. Breeding with close relatives can reduce reproduction and sur-
vival of offspring in two ways: (1) by increasing homozygosity in individuals,
which can result in the expression of deleterious, recessive alleles, and (2) by has-
tening the loss of heterosis—the superiority of a heterozygote over either
homozygote for a specific trait (Pusey and Wolf 1996). Ultimately, fitness of ani-
mals is reduced. This condition, called inbreeding depression, has repeatedly
been shown to be a cost of inbreeding in animals and can increase extinction risk
(Jiménez et al. 1994, Saccheri et al. 1998, DeRose and Roff 1999). The Florida
panther—with its reduced genetic variation, its males having an extremely high
rate of cryptorchidism and poor seminal characteristics, and high rate of heart
defects—may be the living expression of inbreeding depression in pumas
(O’Brien et al. 1990, Barone et al. 1994, Roelke et al. 1993b, Hedrick 1995).
Culver et al. (2000) considered the Florida panther to be a “highly inbred”
species because it had eight fixed microsatellite loci out of ten used to study
genetic variation in the puma across its entire Western Hemisphere range. Sim-
ilarly, two other small, isolated, inbred populations of big cats—African lions in
Tanzania’s Ngorongoro Crater (about 75 to 125 animals) and Asiatic lions in
India’s Gir Forest (about 250 animals)—have reduced genetic variation, high
rates of sperm abnormalities, and reduced circulating testosterone, all of which
are traits that could reduce fitness (Packer et al. 1991, O’Brien et al. 1987, Wildt
et al. 1987). We believe the long-distance dispersal of male pumas is an adaptive
strategy to avoid inbreeding because the male moves away from his mother and
philopatric sisters, and usually well beyond his dispersing sisters. Reciprocal
effects are just as vital; this dispersal pattern promotes outbreeding, particularly
when immigrants from other subpopulations and environments are recruited
into local breeding populations. This avoids inbreeding depression and can
enhance viability of offspring by increasing their genetic variability (Shields
1987). Fitness benefits from inbreeding avoidance and outbreeding may have
been so powerful as to fix dispersal as a trait in male pumas. Of course, disper-
sal in males, just as in females, would have the added benefit of population
expansion.
Transient Behavior in Pumas

Transient is synonymous with wanderer, nomad, floater, and roamer. It implies
that the individual travels widely and has no stable home. The term fits subadult
pumas that are dispersing from natal ranges in search of a permanent place to
live. Transient behavior has been documented in some adult felids, including
cheetahs and African lions. Male cheetahs that traveled widely, rarely marked in
conspicuous places, and only spent short periods in any given area before mov-
ing on were considered nonterritorial or floaters, by Caro (1994). Floating
appeared to be an alternative behavioral tactic (to territorialism) to obtain breed-
ing success. In lions, individuals that wandered widely, often following the
movements of migratory prey, were considered nomadic. Adult males could shift
from nomadic to resident behavior and vice versa during their lifetimes, depend-
ing on whether they could exert dominance over other pride males or whether
they were ejected themselves (Schaller 1972).
Four conditions might result in transient behavior in adult pumas: (1) pop-
ulation isolation; (2) an extremely low, patchy, or migratory food base; (3) an
extremely diffuse puma population; and (4) inability to compete. If a popula-
tion experiences the first condition, natal dispersers are unable to emigrate. In
isolated puma populations in California and Florida, subadults typically
s h owed transient behavior over longer periods but either established residency
or died prior to adulthood (Beier 1995, Maehr 1997a). Some surviving yo u n g
adult males in the Florida panther population apparently showed site attach-
ment to areas that we re within poor puma habitat and along the periphery of
resident male ranges. Maehr (1997a) described them as transients, nonre s i-
dents, or subordinates because they apparently did not successfully breed, and
adjacent resident males did not shift home ranges in response to their deaths.
Howe ve r, since these males showed site attachment, it may be less equivocal to
avoid the transient label. The second and third conditions may often be inter-
related. Long prey migrations could trigger seasonal transiency, where a s
extended prey population lows could cause adults to abandon home ranges in
s e a rch of more pro d u c t i ve habitats. An extremely diffuse puma population
resulting from either poor habitat or ove rkill could cause adults to wander
widely in search of mates (Sandell 1989). Fo u rth, males may be unable to com-
pete due to their small mass, inexperience, or infirmity caused by illness, injury,
or old age. Hence, they may move until a time when they can compete or find
a more secure habitat. In the process they could avoid challenges by resident
males and possibly encounter a few estrus females. Such a strategy has been
o b s e rved in male lions and cheetahs that failed to form (or lost members of )
coalitions, and in male cheetahs that we re in poorer physical condition
(Schaller 1972, Caro 1994). Although this behavior may occur in old, possibly
infirm pumas, we did not observe it in the San Andres Mountains. Instead, old
males lived out their last days within their home ranges. For example, M1, an
old male who apparently suffered from coccidiodmycosis, used a small area
(i.e., about 20 km2) within his home range for a six-month period prior to
being killed by neighboring male M3. Likewise, in the six months prior to his
old-age-related death, M5 used an area that was within, but only 60 percent the
s i ze of, the area he had used in the year before that.
We believe that transient behavior in adult pumas is rare. Although transient
adults have been reported in other populations, this may often be the result of
differences in the term’s definition. Transient has sometimes been used to
describe any puma of unknown origin or fate; consequently, an adult “transient”
may in fact be (1) a resident from outside the study area boundaries that either
made a foray or shifted his home range into the study area, (2) a resident that
was not observed during subsequent field seasons, not because it left the study
area but because it died, or (3) an incorrectly aged subadult. In the latter case,
such an error is most likely if the animal is independent but remains in its natal
area for a long period before dispersal (similar to M82 in our study population).
Although transient adults have been reported in Idaho (Hornocker 1970, Sei-
densticker et al. 1973), Wyoming (Logan et al. 1986), and British Columbia
(Spreadbury 1989), we suspect most fit one of the above categories and were not
true nomads.
LP > > KX
1. Female progeny often remained philopatric into adulthood. Although
philopatric females sometimes made brief exploratory dispersal moves after
independence, their subsequent subadult home ranges were similar in size
to the annual home ranges of adult females. The distance between the cen-
ters of a philopatric female’s natal and independent home ranges averaged
3.2 ± 2.6 km and was similar to the annual distances moved by resident
adult females (2.6 ± 2.1 km). Philopatry resulted in the formation of eight
matrilines.
2. Females that dispersed traveled, on average, 34.6 km from natal ranges to
their independent home ranges or last locations. Dispersal duration in
females was typically brief and ceased prior to adulthood. Only one female
displayed dispersal behavior over a long period and used short-term tran-
sient home ranges. Females that dispersed but remained within the San
Andres Mountains used independent home ranges that were similar in size
(x– = 48.1 km2) to those of resident adult females but much smaller than
those of females that emigrated (x– = 161.2 km2). Subadult females (both
dispersing and philopatric) exhibited transient behavior over shorter periods

(x– = 49 days) than subadult males.
3. All surviving male progeny dispersed, and they dispersed much longer dis-
tances (x– = 102.6 km) than subadult females. Dispersal onset and duration
were highly variable, occurring from 0 to 238 days after independence, then
lasting from 43 to 259 days. Subadult male transient behavior lasted an
average of 211 days. Males appeared more likely to use small transient
ranges during dispersal moves than females. Transient ranges were typically
used when the male encountered the mountain–desert basin interface.
Independent home ranges of males that emigrated from the San Andres
Mountains were larger (range = 356.5–741.7 km2) than the home ranges of
subadult or young adult males that remained within the study area (range
= 61.3–115.0 km2).
4. We deduced that male progeny disperse from natal areas principally to
avoid inbreeding and that female progeny disperse to avoid excessive com-
petition for resources. Dispersal in both sexes hastens colonization of avail-
able habitats.
5. No adult pumas exhibited transient behavior.
LN N5LN5)
1. Mean home range size for seven philopatric females during the twelve
months immediately after reaching adulthood (minimum age 18.5–21
months; maximum age 30.5–33 months) was 66.4 ± 20.1 km2, 27.0 ± 13.9
km2, 72.0 ± 19.8 km2, and 45.6 ± 15.4 km2 for the 90 percent ADK, 60
percent ADK, 100 percent MCP, and 90 percent MCP home range esti-
mators, respectively. Comparable home range sizes for twenty-seven mature
females (thirty-six months or older at start of annual period) over fifty-five
annual periods based on the same home range estimators were 67.4 ± 43.4
km2; 21.9 ± 13.0 km2; 68.9 ± 41.2 km2; and 42.9 ± 28.7 km2.
Chapter 14
Interactions between Pumas
An understanding of puma social organization and its function requires that we

examine how pumas use space in relation to one another. We can visualize this
social arrangement by determining the degree of spatial overlap in puma home
ranges, whether or not they use shared areas at the same time, how frequently
pumas associate and why, and how pumas communicate their presence and
intentions to one another.
Spatial Relationships
Pumas appear to show some flexibility in their spatial arrangement. Although
puma studies generally report extensive overlap between females and little or no
overlap between males (Table 14-1), there is variation. In British Columbia and
the Diablo Range of California, female pumas exhibited minimal spatial overlap
(Spreadbury et al. 1996, Hopkins et al. 1986), whereas male pumas in Monterey
County and the Sierra Nevada of California, as well as in Colorado, have dis-
played extensive spatial overlap (Sitton and Wallen 1976, Neal et al. 1987, and
Anderson et al. 1992, respectively). Unfortunately, a variety of problems plagued
most of these studies, including small sample sizes, the use of different home
range estimators, high human-caused mortality, and the inability to capture all
adult individuals within the study population. These problems most likely result
in reports of exclusivity when perhaps some overlap is the norm. For example,
although relatively exclusive ranges are reported for a sample of four male pumas
in the Guadalupe Mountains along the Texas–New Mexico border, population
247
Table 14-1. Spatial relationships between adult same-sex pumas in other North
American study areas.
Male Female
Location Overlapa nb Overlap nb Methodc Reference
Alberta <O 4–5 O 8–12 MCP Ross and Jalkotzy 1992
British Columbia E 1–2 E 4 MMA Spreadbury et al. 1996
California
Monterey Co. O 4 O 2 MMA Sitton and Wallen 1976
Diablo Range O/E 2 O/O 4 MCP/MMA Hopkins 1989
Sierra Nevada O 5 O 9 CSL Neal et al. 1987
Colorado O 5 O 9 .95 HM Anderson et al. 1992
Florida <O 4 O 9 MCP Maehr 1997a
Idaho E 2 O 4 MCP Seidensticker et al. 1973
New Mexico O 10–12 O 11–18 .90 ADK This study
Utah U 1 O 4 MCP Hemker et al. 1984
Wyoming E 2 O 4 MCP Logan et al. 1986
aOverlap between pairs of pumas: O = overlap; E = exclusive; <O = less overlap than for pairs of the
opposite sex; U = unable to determine. When percentages are available, exclusive indicates pair overlaps
that were less than 10 percent.
bNumber of adults radio-monitored during the same annual or seasonal period.
cMCP = minimum convex polygon; MMA = modified minimum area polygon; CSL = connected
sequential locations with resulting outer lines depicting the home range; ADK = adaptive kernel; HM =
harmonic mean.
turnover was high and only two males were monitored concurrently (Smith et
al. 1986). When pumas were first studied in the Diablo Range of California, the
two collared females had exclusive ranges (Hopkins et al. 1986). However, sub-
sequent research on a larger female sample (n = 7) indicated as much as 50 per-
cent overlap (Hopkins 1989). Beier and Barrett (1993) noted that studies with
sample sizes of fewer than four males generally documented no overlap in male
ranges whereas studies with larger numbers of males did document overlap.
Even in the Idaho research, where relatively exclusive male ranges were first
reported, only one to three adult males were radio-tracked simultaneously (Sei-
densticker et al. 1973).
We examined spatial distribution during four years (1989, 1990, 1992, 1993)
CHAPTER 14. INTERACTIONS BETWEEN PUMAS 249
when an average of 95 percent of the adults were tagged and the majority of
adults also had functional radio collars. Puma densities varied over these years,
not only because of puma removal in the Treatment Area from December 1990
through June 1991, but also because the entire San Andres puma population was
recovering from exploitation prior to our re s e a rch. Consequently, we hoped to
determine the effect of puma removal and puma density on spatial overlap. We
analyzed spatial distribution of adult pumas using two methods: home range
overlap indices and nearest-neighbor distances between arithmetic centers.
Home Range Overlap Indices

We depicted 123 annual home ranges for four years (1989, 1990, 1992, 1993)
by using the 90 percent MCP and 90 percent and 60 percent ADK home range
estimators. Of these, ninety-five annual estimates were based on adult individu-
als radio-tracked for at least ten months of a given twelve-month period (the
method used to determine home range sizes in Chapter 12). However, we also
depicted home ranges for pumas present on the study area as adults for as few
as eight months of a given year. This allowed us to include pumas that arrived
on the study area as adults or that had attained adult status by 1 May of a given
year (n = 11 annual home range estimates). Additionally, if an adult puma’s radio
collar became nonfunctional during the year but sign and subsequent recapture
indicated the animal was present the entire year, we estimated a “pseudo-annual”
home range using the locations from a twelve-month period closest to the time
period being analyzed (n = 17 annual home range estimates). For example, if a
puma’s radio collar became nonfunctional 1 November 1992, then we recap-
tured and reinstrumented the puma in the same area 1 May 1993 and subse-
quently tracked it through the end of the study, we could estimate its 1993
home range using locations from 1 May 1993 through 30 April 1994. “Radio
blackouts” were most common in females (n = 13 of 17 home range estimates).
Because females generally exhibited strong home range fidelity, we believed that
the use of these pseudo-annual home ranges in our overlap measurements would
not heavily bias our results. Similar to our measurements of fidelity, we used a
simple ratio to quantify association patterns:
n1 + n2 / N1 + N2 ¥ 100
where n1 and n2 are the number of locations recorded within the overlap zone
for a pair of pumas in a given year, and N1 and N2 refer to the total number of
locations within each puma’s home range in that same year. Neighboring pumas
with nonoverlapping home ranges were included when home range boundaries
were within 0.5 km of each other. Puma pairs, or dyads, were classified into
three groups: male-male (M-M), female-female (F-F), and male-female (M-F).
We also calculated the percentage of a puma’s home range that was over-
lapped by all other pumas of the same sex (total overlap):
n1 / N1 ¥ 100
w h e re n1 is the number of an individual puma’s locations re c o rded within all
overlap zones, and N1 refers to the total number of locations within that indi-
vidual’s home range in that same ye a r. For example, M46 shared parts of his
home range with three other males in 1989. M-M dyad overlap between M46
and the three males was 1.2 percent, 5.8 percent, and 40.5 percent. Howe ver,
total overlap (M46-M total) equaled the number of M46’s locations within
the area overlapped by all three males (n1 = 26) divided by the total number
of M46’s locations within his 1989 annual home range (N1 = 45), or 57.8
p e rcent. We calculated total annual overlap for a puma only if we could ade-
quately re p resent the home ranges for all of that individual’s neighbors.
When we depicted the home ranges of male and female pumas on an annual
basis, we found that the San Andres Mountains was well covered with a series of
overlapping polygons (Plates 8 and 9). Each year each adult puma shared part
of its annual home range with, on average, 1.8 to 3.8 same-sex adults, and each
adult male shared parts of his home range with an average of 3.3 to 4.9 females
(Table 14-2).
Because overlap indices obtained using the 90 percent MCP always fell
between the indices obtained using the 90 percent and 60 percent ADK home
range estimators, and the trends we re identical, we only report results using
the ADK estimators. Home range overlap occurred within the M-M and F-F
dyad classes in all years. Average annual overlap based on the 90 percent ADK
home range estimator ranged from 18 to 28 percent for M-M dyads and fro m
17 to 26 percent for F-F dyads, although there was high variability in the
amount of overlap between individual pairs (Table 14-3). Average annual
overlap dropped considerably between core use areas (60 percent ADK), rang-
ing from 3 to 13 percent for M-M dyads and from 4 to 9 percent for F-F
dyads. Overlap between M-M dyads was greater than for F-F dyads during
one of the four years [1]. Although we looked for changes in the amount of
overlap from year to year (which might occur because of changing puma den-
sities), we found no differences in the amount of overlap each year for M-M
or F-F dyads in the Treatment Area, in the Re f e rence Area, or in the Sa n
A n d res Mountains as a whole [2].
However, when we examined the percent overlap between specific pairs over
consecutive years (1989–1990 and 1992–1993), we found fourteen of twenty-
four male pairs and eighteen of twenty-four female pairs showed declines in the
Table 14-2. Number of adult pumas of the same and opposite sex with which
each puma shared parts of its home range, San Andres Mountains, New Mex-
ico. Overlap was determined based on the 90 percent adaptive kernel home
range estimator.
Male Female Male Female
with males with females with females with males
Year n No. shared n No. shared n No. shared n No. shared
Treatment Area
1989 4 2.5 ± 1.3 9 3.1 ± 1.5 4 4.25 ± 0.5 9 2.7 ± 0.9
1990 6 3.0 ± 1.2 8 2.8 ± 1.6 6 5.0 ± 1.2 8 3.1 ± 0.6
1992 4 2.8 ± 1.3 3a 2.0 ± 0 4 3.8 ± 1.0 6a 1.7 ± 0.8
1993 3 1.3 ± 1.5 – 3 4.0 ± 1.7 4 1.5 ± 0.6
Reference Area
1989 7 2.7 ± 1.1 5 1.6 ± 1.5 7 2.7 ± 1.0 5 2.8 ± 1.3
1990 5 2.8 ± 0.4 10 1.7 ± 0.7 5 4.4 ± 1.5 10 3.0 ± 0.9
1992 8 2.9 ± 1.0 13 4.2 ± 2.0 8 5.5 ± 3.9 13 3.3 ± 0.8
1993 9 3.4 ± 0.7 11a 3.4 ± 0.9 9 4.0 ± 3.0 12a 2.8 ± 0.9
Total for both areas
1989 11 2.8 ± 1.0 14 1.8 ± 1.0 11 3.3 ± 1.1 14 2.7 ± 1.0
1990 11 2.9 ± 0.9 18 2.2 ± 1.2 11 4.7 ± 1.3 18 3.1 ± 0.8
1992 12 2.8 ± 1.0 16a 3.8 ± 2.0 12 4.9 ± 3.3 19a 2.8 ± 1.1
1993 12 2.9 ± 1.3 11a 3.4 ± 0.9 12 4.0 ± 2.7 16a 2.5 ± 1.0
x– of x– 4 2.8 ± 0.1 4 2.7 ± 0.9 4 4.2 ± 0.7 4 2.8 ± 0.2
aSample sizes were greater for the female-with-males category than the female-with-females category in
some cases, because we could not always depict the home ranges of all of the females that a female’s
range overlapped but we could depict the home ranges of all the males that her range overlapped.
amount of home range overlap. For example, spatial overlap between F6 and
F28 declined from 28 percent to 15 percent during consecutive years 1989 and
1990. The only comparison that did not produce a clear declining trend in over-
lap was between pairs of males for 1992–1993 [3]. Four of eight male pairings
showed declines in overlap and four showed increases. It is possible that the
small sample size as well as great changes in the male component of the popula-
tion (three males died and three new males established residency during 1993)
had a strong influence on the amount of overlap between neighboring males.
Because each puma typically shared parts of its home range with more
Table 14-3. Percent home range overlap (mean ± SD) between pairs of adult
pumas in the San Andres Mountains, New Mexico. The overlap zone was iden-
tified using the 90 percent and 60 percent utilization distribution of the adap-
tive kernel (ADK) home range estimator.
% overlap, Male–Male dyad % overlap, Female–Female dyad
Year No. pairs 90% ADK 60% ADK No. pairs 90% ADK 60% ADK
Treatment Area
1989 6 29.7 ± 19.4 5.3 ± 7.1 10 25.6 ± 19.9 4.8 ± 7.3
1990 6 37.8 ± 13.8 16.1 ± 11.9 12 23.2 ± 31.1 11.4 ± 26.1
1992 4 17.8 ± 12.8 2.2 ± 4.4 6 44.0 ± 33.6 16.6 ± 21.9
1993 5 14.1 ± 29.6 9.7 ± 21.6 7 32.2 ± 30.5 20.6 ± 20.0
Reference Area
1989 8 23.3 ± 20.4 5.3 ± 7.9 5 10.3 ± 16.9 1.5 ± 3.3
1990 8 20.9 ± 12.3 9.5 ± 10.0 12 11.2 ± 14.2 1.6 ± 3.2
1992 14 17.7 ± 13.5 3.8 ± 7.3 30 21.9 ± 22.1 7.3 ± 12.4
1993 14 28.7 ± 21.5 13.7 ± 20.9 21 19.1 ± 14.2 4.9 ± 7.0
1989 14 26.1 ± 19.5 5.3 ± 7.3 16a 21.0 ± 19.2 4.3 ± 6.6
1990 14 28.2 ± 15.2 12.4 ± 10.9 25a 17.0 ± 23.9 6.3 ± 18.5
1992 18 17.7 ± 13.0 3.4 ± 6.7 36 25.6 ± 25.2 8.9 ± 14.4
1993 19 24.8 ± 24.0 12.7 ± 20.6 28 22.4 ± 19.7 8.0 ± 13.4
‡
Ç of Ç‡ 4 24.2 ± 4.6 8.4 ± 4.8 4 21.5 ± 3.6 6.9 ± 2.1
aThe total includes a female pair along the boundary of the Treatment and Reference Areas that was
not included in the respective areas.
than one other same-sex individual each year, the total amount of area it
s h a red was much greater than that found for individual M-M or F-F dyads.
On average, 50 to 68 percent of a male’s annual home range (90 perc e n t
ADK) was overlapped by other male pumas, and 43 to 64 percent of a
female’s home range (90 percent ADK) was overlapped by other females
(Table 14-4). The percentage of a puma’s core area (60 percent ADK) share d
with all other same-sex pumas was much smaller, averaging 14 to 38 perc e n t
for males and 10 to 32 percent for females. In one year (1992) females share d
significantly more of their home ranges with other females than males share d
with other males. No significant differences could be found for the other
t h ree years [4].
Table 14-4. Percentage of a puma’s home range (mean ± SD) shared by all
other adult pumas of the same sex, San Andres Mountains, New Mexico. Over-
lap was identified using the adaptive kernel (ADK) home range estimator.
Total % overlap, Male by Males Total % overlap, Female by Females
Year n 90% ADK 60% ADK n 90% ADK 60% ADK
Treatment Area
1989 4 54.7 ± 29.3 16.8 ± 13.4 9 60.3 ± 28.0 13.5 ± 5.2
1990 6 65.0 ± 15.1 38.1 ± 20.2 8 63.0 ± 37.4 34.5 ± 35.9
1992 4 34.2 ± 20.5 7.6 ± 11.4 3 48.0 ± 37.7 26.8 ± 43.2
1993 3 47.1 ± 41.2 32.2 ± 28.0 – – –
Reference Area
1989 7 61.8 ± 18.2 21.0 ± 13.5 5 16.8 ± 17.8 3.0 ± 4.1
1990 5 62.5 ± 17.4 34.7 ± 32.1 10 27.6 ± 23.7 3.9 ± 5.4
1992 8 57.6 ± 19.2 17.5 ± 14.0 13 68.3 ± 23.1 33.2 ± 23.4
1993 9 75.4 ± 13.7 39.9 ± 24.4 11 59.4 ± 22.7 15.3 ± 10.8
1989 11 59.2 ± 21.7 19.4 ± 12.9 14 44.7 ± 32.3 9.8 ± 13.2
1990 11 63.7 ± 15.4 35.8 ± 24.2 18 43.4 ± 34.6 17.5 ± 28.1
1992 12 49.8 ± 22.0 14.2 ± 13.5 16 64.5 ± 26.1 32.0 ± 26.3
1993 12 68.3 ± 24.7 38.0 ± 24.2 11 59.4 ± 22.7 15.3 ± 10.8
‡
Ç of Ç‡ 4 60.2 ± 7.9 26.8 ± 11.8 4 53.0 ± 10.6 18.7 ± 9.5
We examined percent spatial overlap between adult males and females only
during 1992, the year we had the most measurable M-F pairings (Table 14-5).
Generally, overlap between M-F pairs was higher than for same-sex pairs [5].
Additionally, the percentage of a puma’s home range that was shared by all
neighboring same-sex individuals was typically much less than the amount
shared with members of the opposite sex [6]. These relationships suggested that
competition within the sexes was greater than between the sexes and supports
the two-strategies hypothesis we presented in Chapter 11. Thirteen out of nine-
teen females’ home ranges (90 percent ADK) were completely engulfed by sur-
rounding resident males. The other six females shared an average of 91 percent
of their ranges with males. In contrast, a generous portion of a male’s range
(x– = 36.6 percent) was not shared with females. Although a male’s range could
entirely surround the range of a female (this occurred in five of the nineteen
females), other males often shared portions of that same female’s range.
Table 14-5. Percent home range overlap (mean ± SD) between adult
pumas on the San Andres Mountains, New Mexico, during 1992.
Home range estimatorb
Category of overlap na 90% ADK 60% ADK
Dyad pair
M-M 18 17.7 ± 13.0 3.4 ± 6.7b
F-F 36 25.6 ± 25.2 8.9 ± 14.4
M-F 56 32.1 ± 25.7 17.2 ± 22.4b
Total overlap
Male by males 12 49.8 ± 22.0 14.2 ± 13.5c,d
Female by females 16 64.5 ± 26.1e 32.0 ± 26.3e
Male by females 10 63.4 ± 28.8 43.2 ± 23.4c
Female by males 19 97.4 ± 5.5d,e 68.5 ± 26.6d,e
aSample size for dyad pairs is number of pairs; for total overlap, it is number of individuals.
bADK = adaptive kernel.
cDistributions of M-F and M-M dyad overlap were significantly different, P = 0.026.
dA larger portion of a male’s core area was overlapped by females than by other males, P =
0.003.
eLarger portions of a female’s home range and core area were overlapped by males than by
other females, P < 0.001.
A visual exam of puma home ranges in the Treatment Area in 1990 (just
prior to puma removal), and again in 1993 (after the population had almost
recovered) also indicated that pumas on the San Andres Mountains did not
reoccupy home ranges in the same spatial arrangement (Plate 10). This is con-
trary to one of the predictions of the self-limiting hypothesis (see Chapter 11).
Range boundaries were apparently not well defined, a conclusion supported by
our examination of fidelity in Chapter 12.
Nearest-Neighbor Analysis
We calculated the arithmetic center of each individual’s locations for six-
month subsets (January–June and July–December) of the same four annual
periods (1989, 1990, 1992, 1993). Because some pumas were prone to shift
home range use over time, we believed the shorter time periods would better
reflect the relationships between neighboring pumas. We used the Clark and
Evans (1954) test based on nearest-neighbor distances to determine the spa-
tial distribution of arithmetic centers. Clark - Evans ratios (R) of 0, 1, and
2.15 re p resent maximum aggregation, random distribution, and perfect uni-
formity, re s p e c t i ve l y. This means a ratio approaching 2.15 would indicate

s t rong avoidance between pumas, whereas a ratio near ze ro would indicate
s t rong attraction. To determine M-M distributions, we started with an effec-
t i ve area of 2,059 km 2, which defined the area incorporated by 99 percent of
the radio locations of all adult pumas (n = 8,490) during the course of the
study (see Fig. 4-1). Ad d i t i o n a l l y, a boundary length of 274.4 km (120 km
long by 17.2 km wide) was incorporated into the calculations to account for
edge effects (Sinclair 1985). But because the northern end of the study are a
was composed of more fragmented habitat and was void of a resident female
for almost two of the four years, we excluded this area from the analysis of F-
F and M-F distances. The resulting area was 1,859 km2 with a boundary
length of 228.4 km.
Clark-Evans ratios of arithmetic centers indicated pumas generally occurred
in random distributions (Table 14-6). However, female R values were typically
less than one, showing a tendency toward aggregation, whereas male R values
were typically greater than one, indicating a tendency toward avoidance. The
distance between centers of closest pairs averaged (mean of means) 8.9 km for
males, 5.1 km for females, and 4.0 km for pumas of the opposite sex. The only
time males showed a tendency toward aggregation was from January to June
1990; this was caused by the arrival of three new males. During this period, the
arithmetic centers of two of these males (M29, M161) were 0.9 to 1.7 km from
the arithmetic centers of long-established residents (M46, M53). The third new
male (M114) apparently encouraged resident M7 to shift to within 1.5 km of
resident M49. In the subsequent six-month period, only M29 and M46
remained in close proximity (centers 1.7 km apart). The arithmetic centers for
the other two pairs of males were 14.7 and 16.2 km apart. Although we could
find no relationship between male distribution and density, female distributions
became more aggregated with increasing density, and male-female distributions
became less aggregated with increasing density [7]. Increased aggregation
between females makes sense given the matrilineal structure we observed (see
Plate 4). Less aggregation between males and females with increased density also
suggests that males are spacing themselves to maintain breeding opportunities
with a large number of females, and females may be doing their best to avoid
males, who may threaten them or their offspring.
Spatiotemporal Relationships
Our examination of spatial relationships is only the first step in the analysis of
interrelationships, because it does not address the temporal order of locations
(e.g., two pumas may share the same space but use it at different times). To
address the possibility that pumas of the same sex may temporally segregate their
shared use of space to relieve competition, we determined the proportion of

time that pairs of pumas used the same area at the same time. We did not exam-
ine M-F dyads here because the majority of a female’s core area was typically
shared with neighboring males, and we could get a clearer understanding of
male-female relationships by examining direct interactions (see the next section,
“Direct Interactions”). We defined simultaneous locations as those obtained
within one hour of each other from fixed-wing aircraft. A pair of pumas was
considered to be using the same area at the same time if simultaneous locations
occurred within the overlap zone for that pair. Overlap zones were determined
for annual home ranges (90 percent ADK) and annual core use areas (60 per-
cent ADK). We calculated spatiotemporal overlap indices, or the proportion of
simultaneous locations within the overlap zone, for M-M and F-F dyads for the
same four years spatial overlap was calculated.
We evaluated spatiotemporal relationships between overlapping and adja-
cent pairs of male (n = 16–19 pairs each year) and female (n = 8–32 pairs each
year) pumas. Although there was considerable variation among dyads, pumas
with overlapping home ranges generally used those overlap zones at different
times. Each year, spatial overlap for both M-M and F-F dyads ranged betwe e n
17 and 28 percent, whereas mean spatiotemporal overlap averaged only 3 to 10
p e rcent (Table 14-7; Fig. 14-1). Of the thirteen F-F dyads that exhibited more
than 15 percent spatiotemporal ove r l a p, 69 percent we re closely related. Sp a-
tiotemporal overlap between all mother-daughter and sister-sister pairs (n = 18)
averaged 20.1 percent, whereas spatiotemporal overlap for unrelated pairs (n =
60) averaged 4.5 percent. T h e re was even less simultaneous use of ove r l a p p i n g
core areas, with indices averaging 2 percent or less each year for both M-M and
F-F dyads. Only three male pairs exhibited 3 percent or more spatiotemporal
overlap of core areas. Fights occurred between two of these pairs (M1-M3 and
M19-M46), resulting in one death (M1) and one home range shift (M46). The
other pair of males (M3, M88) was courting F47 during a single estrus cyc l e .
Of the eight female pairs that exhibited some degree of spatiotemporal ove r l a p
of core areas, all but one we re closely related. Patterns of spatiotemporal ove r-
lap indices we re similar between M-M and F-F dyads during all years exc e p t
1990, when males showed greater simultaneous use of overlap zones based on
the 90 percent ADK [8].
Direct Interactions
The most direct measure of sociality we quantified were the frequency and pur-
pose of interactions between pumas. Interactions were primarily determined
from simultaneous radio locations. Because each location had a measure of error
FIGURE 14-1. Mean and standard deviation of the annual spatial overlap (solid circles) and
spatiotemporal overlap (open circles) between pairs of same-sex adult pumas on the San
Andres Mountains, New Mexico. Home ranges were delineated using the adaptive kernel
(ADK) estimator.
associated with it, we assumed pumas that we located within overlapping error
polygons were in association. When possible, we attempted to confirm the asso-
ciation, either by obtaining subsequent visuals or by finding sign of the individ-
uals at the site. We also attempted to determine the association’s purpose. Breed-
ing activity was confirmed by the birth of cubs approximately ninety-two days
later. We classified an association as probable breeding activity given the follow-
ing conditions: (1) a male was documented in association with an adult female
that was not known to be pregnant and was either without cubs or was caring
for cubs that were nine months or older; and (2) we obtained visual observations
of the pair, heard vocalizations that suggested the female was in estrus, located
the pair together for consecutive days with no resulting female mortality, found
tracks of the pair traveling together, and/or documented estrus-cycling behavior
in the female. Locating a male and female pair together, as in condition 1 above,
without corroborating evidence from condition 2 was considered as possible
breeding activity. We confirmed aggressive encounters by finding mortalities or
observing fresh wounds on captured individuals. We also obtained an index of
association frequency between independent pumas during each of four years
(1989, 1990, 1992, 1993) by calculating the percentage of simultaneous loca-
tions when dyad pairs with overlapping home ranges were in association. Some
individuals were only in the study area as adults for a few weeks or months of a
given year, yet they sometimes associated with other adult pumas. In order to
include these animals in our index, we delineated their 90 percent MCP home
ranges based on the locations we did have for that particular year, then paired
them with pumas whose home ranges touched or overlapped theirs. We then
compared the association frequency among adult F-F, M-M, and M-F dyads
during each year. Behavior between females and their cubs was documented by
visually observing the family groups, finding their tracks, and radio-tracking
thirty-eight radio-collared cubs.
Associations between Independent Pumas

Independent pumas rarely associated with other pumas. Of the 12,490 locations
we obtained on independent pumas, only 5.1 percent (n = 643) involved asso-
ciating pairs. The rest were of solitary individuals or mothers with dependent
cubs. We observed 269 separate associations between independent pumas, the
majority of which (78.4 percent) were between males and females (Table 14-8,
Fig. 14-2). M-M and F-F associations occurred at similarly low frequencies
(10.4 percent and 11.1 percent of all associations, respectively). When we exam-
ined associations between adult dyad pairs, we found low frequencies of associ-
ations between M-M and F-F dyads for all four years (average of 0–1.4 percent
of simultaneous locations each year, Table 14-9). There were a greater number
of associations between F-F dyads than M-M dyads in only one of the four years
(1992) [9]. Although still rare, associations between adult M-F dyads averaged
2.6 to 6.2 percent of simultaneous locations each year and were much greater
than those of same sex dyads [9]. We describe specific associations in the next
sections.
MALE-MALE
Males (M-M pair or M-M-F group) associated on thirty-five occasions; ten
resulted in deaths. Associations may not have been quite as rare as our data sug-
gested, because we generally located individuals after they had bedded down for
the day, and all but one of the adult males we captured bore scars from fighting.
Of the associations, seven were between two adult males and an estrus female
(two resulted in male deaths), two were between adult males apparently search-
ing for a female known to be in estrus, six were between adult males involved in
suspected territorial disputes (three males died), four were between an adult and
a subadult at a food cache (three subadults were killed), two were possible terri-
torial disputes between an adult resident and an intruding subadult, though
food may also have been involved (one subadult died), and thirteen were for
Table 14-8. Associations (n = 269) between independent pumas on the

Association typea No. associationsb No. mortalities
AM-AFO 154 (192) 5 (Females 39, 40, 57, 60, 185)
AM-AFC 40 (46) 4 (Females 2, 15, 61, 86) and cubs
AM-AM 19 (20) 3 (Males 1, 14, 73)
AFO-AFO 12 (12)
AFO-AFC 9 (10)
AM-SM 9 (11) 5 (Males 64, 108, 140, 198, 300)
AM-AM-AFO 7 (7) 2 (Males 19, 22)
AM-SF 4 (6) 2 (Females 10, 96)
AFC-AFC 4 (4)
AFO-SM 3 (7)
AFC-SM 3 (3)
SF-SF 3 (3)
AFO-SF 2 (2)
AFC-SF 0 (0)
Total 269 (323) 21 independent pumas
aA = adult, S = subadult, M = male, F = female, O = female without cubs or cubs nine
months or older, C = female with cubs younger than nine months or known to be preg-
nant.
bAssociations are listed in descending order of frequency. They include the number of sepa-
rate associations, with number of location days in parenthesis. Two associations resulting in
mortalities are not included because we could not categorize the association: thirty-month-
old puma F304 was killed by an unknown puma, and a litter of four cubs was killed by res-
ident M52 when the radio-collared mother was not with them. We included six associa-
tions, where eleven cubs from six litters were killed by males, in the AM-AFC type,
although we were certain the mother was present in only one of the associations (she was
killed along with the cub).
unknown reasons (one subadult was killed). Adult mortalities (M1, M14, M19,
M22, M73) always occurred within the victim’s core home range (60 percent
ADK) (Fig. 14-3). In two cases, the victor was a new immigrant who usurped
most of the victim’s home range after his death. In at least three of five subadult
deaths (M64, M108, M300), the fight occurred within the resident adult’s core
area.
We do not know if all associations resulted in fights, but we suspect that
fights that did not result in the death of a combatant helped to clarify an indi-
vidual’s dominance ranking and to consequently explain some of the subsequent
shifts we observed in individual ranging patterns. For example, new immigrant
FPO @ 90%
FIGURE 14-2. Number of associations (n = 269) between independent pumas (M = male; F

= female) that were related to breeding, non-breeding, or unknown activities on the San
Andres Mountains, New Mexico, 1985–1995. Twenty-one associations resulted in the death
of an independent puma.
M193 left the San Andres Mountains immediately after a fight with resident
M88 in M88’s core area (Plate 11). He spent eight weeks in the Doña Ana
Mountains, a small, isolated mountain range about 18 km west of the study
area, before returning to M88’s home range. Within three weeks of his return,
M193 and M88 apparently fought again. Puma M193 spent the next twenty-
one days within 1 km of the site where the association took place, which sug-
gested he was recovering from injuries. His tenacity apparently paid off. Within
six months of this second association, M88’s center of activity shifted 4.3 km to
the north. Although this was not a statistically significant shift (based on our cri-
teria in Chapter 12), it was biologically significant; during this period, M88’s
southernmost movements were north of any movements made by M193.
FEMALE-FEMALE
Females associated on thirty separate occasions, none of which resulted in death
or known injuries. Nine of the associations were between siblings or between
mothers and daughters. Considering the large number of matrilines we docu-
mented in our population (see Plate 4), related females did not appear to asso-
ciate more frequently than unrelated females. Three associations may have been
FIGURE 14-3. Home ranges (90 percent and 60 percent adaptive kernel) for adult pumas
M19 and M29 in 1991, San Andres Mountains, New Mexico. The males fought within a
common core area in mid-September 1991 (star), apparently over access to F15, who was
in breeding condition. M19 died (gray dot) on 27 September after moving about 19 km
north of the fight location. M19’s home range is depicted based on thirty-five locations
obtained from January to September 1991.
precipitated by the presence of a mule deer kill. It was more difficult to ascer-
tain the reason for the other associations, but we suspect some of them may have
involved an estrus female looking for a mate. Only four associations consisted
of females that we knew were non-receptive (i.e., they were pregnant or raising
young cubs). In twenty-three other cases, at least one of the pair was an adult
without cubs or her cubs had reached the age of independence. During nine of
those associations, at least one member of the pair had associated with an adult
male puma within three to fourteen days of the female-female association
(n = 6 associations), or she became pregnant within a few days of the association
(n = 3 associations). In California, direct interactions between adult females
occurred when there was an absence of a breeding male, and the interactions
coincided with the periodicity of estrus (Padley 1990). This suggests females are
more tolerant of other pumas during estrus and apparently seek them out.
MALE-FEMALE
Of 211 associations between males and females (either M-F or M-M-F), thirty-
five were between breeding pairs (the female became pregnant during the estrous
cycle in which the association was observed). Five pregnancies occurred when
females were still caring for cubs that were nine to twelve months old. Breeding
activity was probable (n = 61) or possible (n = 60) in another 121 associations.
In three other instances in which adult females were associating with subadult
males, we believed the females to be in estrus. Consequently, 75.3 percent of all
M-F associations we observed could have occurred because the female was in
estrus. Two of the associations resulted in the death of a male (described under
M-M associations).
We classified fifty-two associations (24.6 percent of all male-female associa-
tions) as non-breeding because females were either killed (n = 11), caught in a
snare (n = 1), too young to breed (n = 3), pregnant (n = 13), or raising young
cubs (n = 24) when the association occurred. Sixteen of these associations were
violent, resulting in deaths of two subadult females, nine adult females, and one
or more cubs from each of six litters. Independent females were killed when they
defended cubs apparently unrelated to the male (n = 2–3), while competing for
a food cache (n = 3–4), as a food source themselves (n = 4), and possibly during
courtship (n = 1). The skeletal remains of a thirty-month-old, apparently
untagged female (F304) indicated she was also killed by another puma (canine
punctures were evident in her braincase). We did not include her in our associ-
ation analyses because, although we suspect a male was responsible, we did not
know the status of either puma or why she was killed. (For details on these mor-
talities, see Chapter 8.) Our observations indicate no males killed females with
whom they had prior breeding associations.
Other non-breeding associations between males and females appeared ami-
cable. Pregnant females associated with resident adult males on thirteen occa-
sions for periods lasting one to two days (x– = 1.1 ± 0.3) without apparent injury.
In four of those associations, the female associated with the litter’s sire: F47 asso-
ciated with M88 once during each of three pregnancies, and F107 associated
with M88 (her father and mate) during one pregnancy. In four other instances
a pregnant female associated with a resident male known not to be the sire of
her litter. Nursing mothers also associated with resident males on nine occa-
sions; of those, four involved a mother and the sire of her cubs and lasted from
one to four days each ( x– = 2.0 ± 1.4). Five other associations involved either a
male known not to be the sire (n = 2) or who was of questionable paternity (n
= 3) and lasted from one to five days each (x– = 2.6 ± 1.8). These interludes sug-
gested that pregnant or nursing puma mothers may produce a “pseudo-estrus,”
Slide @171%
PHOTO 24. Puma F40 was killed and

eaten by territorial puma M19.
allowing them to engage males in breeding unions that may confuse paternity
or reinforce pair bonds, and consequently reduce chances of infanticide. Simi-
larly, pregnant African lion females have been observed mating with new males
at a pride takeover (Packer and Pusey 1983). Familiarity and amicable relation-
ships between resident females and territorial males should have fitness benefits
when males associate with females that are raising older cubs too. We docu-
mented thirteen instances in which mothers raising weaned cubs two to nine
months old (x– = 5.0 ± 2.1) safely associated with resident males. We knew males
were the sires of the cubs during at least three of the associations, and at least
two other associations involved males that had been with the mothers during
prior estrous cycles. Recognition of the female, and not of the cubs, is probably
what protects cubs from infanticide. M88 and M52 both killed cubs of females
they knew; however, the females were not present at the nurseries when the
males discovered the litters.
Some males were more aggressive than others, indicating individuality in
behaviors. Over the course of the study we collected data on thirty-two adult
males; at least seven or eight were known to kill females or cubs. Three males
may have been culpable for the majority of known female and cub deaths. We
knew M29 killed one adult female and one cub, but we also highly suspected
him in the deaths of three other adult females and one of those female’s cubs.
M88 was documented killing one subadult female, one adult female, and a lit-
ter of cubs. M22 killed one adult female and one subadult female. Conse-
quently, of the sixteen M-F associations that resulted in deaths, these three males
were responsible for up to 63 percent. In contrast, we observed an instance in
which resident male M38 (who was never documented in an aggressive associa-
tion with a female) usurped a deer kill made by new immigrant F240 while she
was incapacitated in a snare at the cache site. He did not harm F240, but he
dragged the deer 215 m away before feeding on it.
Our observations indicated that associations between independent pumas
involved high risk. Avoidance was the best survival tactic. In all, 207 of 269 asso-
ciations (76.9 percent) could have been instigated by an estrus female or
involved a female that was exhibiting pseudo-estrus. Mortalities occurred dur-
ing two or three of these associations (less than 2 percent of apparent breeding
associations). In contrast, mortalities occurred during eighteen or nineteen asso-
ciations not involving an estrus female (29–31 percent of non-breeding associa-
tions). Estrus and pseudo-estrus could result in recognition and tolerance dur-
ing subsequent meetings and thus be very beneficial to female survival and
reproductive success (see Chapter 8).
T h roughout their range, it appears that pumas seldom interact socially
with other pumas; when they do, it is most likely because they want to mate.
Adult pumas in Idaho associated during only 5 to 13 percent of simultane-
ous locations, and thirteen of seventeen separate interactions we re between a
single adult male and either one or two adult females (Seidensticker et al.
1973). In Colorado, independent pumas we re found in association during
only 12.3 percent of 227 simultaneous locations (Anderson et al. 1992). Of
those, 50 percent we re between single adult male and single adult female
pumas. In Florida, independent panthers were found within 0 to 500 m of
each other during only 6.5 percent of simultaneous locations (n = 3,480)
(Maehr et al. 1989). Again, associations we re most common between adult
males and adult females (87.7 percent of 227 associations where pairs we re 0
to 500 m apart ) .
Associations between Family Members

Each year an average of 74 to 76 percent of adult females within the study area
were raising cubs (see Chapter 7). Radiotelemetry data and visual observations
indicated that female pumas remained close to their nurseries when cubs were
less than eight weeks old. By monitoring the movements of twenty-seven nurs-
ing females raising forty-one different litters, we found that on any given day
during the first eight weeks after the birth of a litter, a mother was an average of
0.9 ± 0.4 km from her birth nursery (mean number of locations = 12.8 ± 4.8).
Cubs that were eighteen to thirty-five days old at capture (n = 85 cubs from 38
litters) did not have the motor skills to evade our capture. Usually we picked
them up as they attempted to crawl into protective nooks and crannies that
nurseries often provided. In contrast, cubs forty-two to fifty-nine days old at
capture (n = 38 cubs from 19 litters) quickly maneuvered over steep, rocky ter-
rain. For example, after finally locating the alternate den where F126 had left
her forty-two-day-old cubs, we had to scramble to catch them. Three researchers
were present, and all were in good physical condition; however, we managed to
capture only two of the four cubs. Six days later we saw all four siblings sleep-
ing on a ledge. Although we were within 15 to 20 m when they detected us, we
were barely able to catch one cub before the rest vanished over the rocks and
down the drainage. Such improved motor capabilities were apparent from the
movements made by family groups. By radio-tracking mothers, we were able to
determine when cubs from twenty litters were moved (or more likely moved
under their own power) at least 1.5 km from their birth nurseries. This occurred
when the cubs were an average of fifty days old. Four radio-collared cubs from
three litters made their first long-distance movements (2.8–5.5 km) from the
vicinity of their birth nurseries when they were sixty-four to seventy-one days
old. Three other tagged litters apparently moved distances of 4.5 to 7.0 km
when they were sixty-five to seventy-three days old, because each radio-collared
mother showed site affinity to the new location (she was located there consis-
tently over the next two weeks) and never returned to her birth nursery. By this
age, most cubs were probably weaned and dependent on meat. We observed
nursing in cubs as old as forty-nine days, whereas the youngest cub seen eating
meat was sixty-three days old.
By radio-collaring cubs, we obtained 454 simultaneous locations on family
groups. On average, females were located with their young (six months and
younger), radio-collared cubs during 81 percent of their telemetry locations (n
= 246), and with their older (more than six months old) radio-collared cubs dur-
ing 82 percent of their telemetry locations (n = 208). Studies in Florida and Col-
orado reported somewhat lower frequencies of interactions; females were located
within 1 km of their dependent offspring during 53 percent (n = 399) and 56
percent (n = 69) of simultaneous locations, respectively (Maehr et al. 1989,
Anderson et al. 1992). Because the majority of our locations were at apparent
day beds, it is possible females were simply returning to, and bedding with, cubs
after a night of solitary hunting. Such behavior was exhibited by female pumas
in Utah (Barnhurst and Lindzey 1989). There, females were with their cubs dur-
ing 67 percent of simultaneous locations (n = 414). However, tracks of cubs zero

to six months old were found with their mother’s tracks only 19 percent of the
time, and tracks of cubs seven to twelve months old accompanied their mother’s
tracks 43 percent of the time (Barnhurst and Lindzey 1989). Nevertheless, inter-
actions between mothers and their cubs are by far the most frequent kind of
association found between pumas and indicate that females and their depend-
ent young form the basic social unit for the species. The six adult females that
we monitored for periods exceeding forty-eight months were caring for cubs an
average of 61.8 ± 11.0 percent, and pregnant for another 17.7 ± 2.9 percent, of
their adult lives.
Communication among Pumas

In the puma, communication is vital for finding mates, maintaining family
cohesion, and avoiding competition or direct threats from other pumas. Rarely
has anyone observed direct interactions between adult pumas in the wild, so it
is practically impossible to describe postures and gestures that individuals use
during these encounters. Over the ten years of our research, we logged all of our
close encounters with pumas and tried to interpret their exhibitions toward us.
In addition, we heard their vocalizations, and described and mapped visual and
olfactory markers they left behind.
Vocalizations
The puma does not have the capability to emit the long-range calls typical of
roaring cats (i.e., African lion, tiger, African leopard, jaguar) because of the
structure of its larynx—the sound generator of voice. The vocal folds of a roar-
ing cat’s larynx are long and massive, and consequently they have a low natural
frequency and will produce high acoustical energy when vibrating. The puma’s
smaller larynx with small, sharp-edged vocal folds does not possess the necessary
equipment to similarly amplify sound into roars. Unlike those of the roaring
cats, the puma’s hyoid apparatus, which attaches to the larynx, is also completely
ossified. Consequently, the length between the puma’s oral pharynx (muscular
tube that connects the mouth and esophagus) and larynx is fixed and will not
allow for the same deepening in voice pitch (Hast 1989). Nevertheless, pumas
can emit a variety of vocalizations. From his studies on captive felids, Peters
(1978) identified several characteristic vocalizations in pumas, including the
gurgle, purr, bleat, whistle, mew, main call (i.e., the intense form of the mew),
wah-wah, and the orgasmic cry of the adult female. These sounds could be com-
bined to form more-complex vocalizations, such as the composite vocalization
of the adult female during estrus (what we term the caterwaul), or the coupling
of the gurgle with the mew. Peters did not describe vocalizations specific to ago-
nistic situations, such as growls, hisses, and spits.
Functional aspects of puma vocalizations are difficult to determine, because
an observer must be able to interpret the behaviors of both sender and receiver.
This is made even more difficult since pumas may mix different vocalizations, a
vocalization may have more than one function, and there may be more than one
vocalization for the same function. Nevertheless, four general message systems
have been outlined for carnivores (Peters and Wozencraft 1989) and applied to
vocalizations: agonistic, integrative, sexual, and neonatal.
We observed two types of agonistic messages: defensive and offensive threats.
We often heard low guttural growls, spitting, and hissing when we approached
snared pumas. Mothers also sometimes growled or hissed at us when we
approached their nurseries. Spitting has been considered a defensive threat in
felids, whereas hissing encodes a mild offensive threat (Peters and Wozencraft
1989, Wemmer and Scow 1977). The long-distance calls produced by the tiger
and leopard may also be considered agonistic, because they are used in territo-
rial advertisement (Schaller 1972, Peters and Wozencraft 1989).
We observed several types of puma vocalizations that may be integrative or
neonatal, including satisfaction, distress, solicitation, contact, maternal alarm,
and maternal assembly. Because we often observed nurseries before attempting
to capture and mark cubs, we had the opportunity to document some close-
range vocalizations between family members. Cubs and mothers sometimes
purred while cubs nursed. Continuous, pulsed, low-intensity sounds like
purring convey satisfaction. Although the functional significance of purring by
adult felids is not fully understood (Leyhausen 1979, Peters and Wozencraft
1989), we speculate that purring is probably a simple and efficient way for a
mother to communicate well-being to all of her cubs at once. Young cubs (less
than six weeks old) also emitted high-pitched, bird-like chirps and mews. The
mew sounds we heard may have been akin to the “bleating” vocalization
described by Peters (1978), which is the only call emitted during the first two
weeks of life. Since such vocalizations often occurred after the mother became
aware of our presence and ceased nursing, we suspected they were either solici-
tation or distress calls. Distress calls are made by all neonatal and juvenile carni-
vores suffering from pain, hunger, cold, or isolation from mother or siblings
(Ehret 1980). Chirps and whistles may also be contact calls. Such vocalizations
can help mothers find cubs that stray during exploratory moves around the
nursery. We also occasionally heard whistles while we were radio-tracking mem-
bers of older family groups, and suspected they were short-range contact calls
(also see Padley 1996a). Contact calls may be especially frequent when cubs
make their first excursions from the nursery and try to follow their mother
(Peters and Wozencraft 1989). One newly orphaned cub we observed from a
distance of 10 to 20 m emitted a remarkable vocalization three times. It con-
sisted of an upward-scaling, piercing whistle followed by a downward-scaling
whistle and three punctuated monotonal notes (Sweanor 1990). We believe she
was trying to contact her mother and two brothers. It is also possible we
observed ultrasonic signals between a mother and her nursing cubs that may
have conveyed more than one message. As we watched a nursery from 20 m
away, the mother detected us, opened her mouth, and apparently emitted some
kind of maternal alarm call. We heard no sound, but three of the four cubs
immediately moved about 10 m to hide in a nook between rocks within the
nursery. After about an hour the mother appeared to relax and again opened her
mouth. Although there was no audible sound, the mother apparently gave a
maternal assembly call; the cubs immediately reappeared and moved to her belly
to nurse.
Vocalizations that convey sexual messages have also been documented in the
puma. The most spectacular sound a puma emits is the caterwaul. This may be
the “scream” that is sometimes referred to in popular accounts. The caterwaul of
a puma sounds similar to that of a courting domestic cat, only much louder.
Because of its varied pitch and intensity (it is a composite vocalization with mew
and purring components; Peters 1978), and the distance and vegetative and
physiographic screening that may occur between the puma and observer, only
the higher-pitched, more-forceful sounds may be heard. Consequently, it is not
difficult to imagine how a human might perceive it as a scream. Caterwauls are
advertisement calls; they probably communicate breeding readiness and aid in
the location of a mate. In the San Andres Mountains, caterwauls were emitted
during male-female associations as well as by lone pumas. When we were able
to measure distances, we found the vocalizations could carry 400 m. Individu-
als that caterwauled were typically females we suspected were in estrus, but when
pairs were together we could not discern if one or both animals were caterwaul-
ing. During three of our visual observations of courting pairs, pumas were
within 10 m of each other when one individual caterwauled. One lone female
caterwauled by San Andrecito Camp on three occasions over a twenty-seven-day
period. We heard these caterwauls intermittently through crepuscular and noc-
turnal hours as the puma traveled through the basin surrounding our camp.
During one bout of calling, the female caterwauled thirty-one times in rapid
succession. Close-quarters vocalizations between breeding males and females
may also be somewhat distinct from the longer-distance caterwauls and may
include both advertisement and courtship messages. In one instance, we
watched a courting pair (F28 and M38) from 50 m away. F28 was emitting a
mixture of vocalizations that included louder caterwauls but also lower-ampli-
tude growls, gurgles, and mews. Before we saw the pair, we thought F28 must
be in great distress. In reality, the two cats were laying side by side. While F28
vocalized, M38 groomed his tail, appearing unaffected by his mate’s behavior.
The meanings of two other vocalizations, the yowl and the short, raspy ouch
call, were hard for us to separate or easily categorize. Both vocalizations appear
to travel a similar distance to the caterwaul. The yowl may be analogous to the
main call described by Peters (1978). Its function could include sexual identity,
dominance identity, territory advertisement, or sexual advertisement. For exam-
ple, we heard F15 emit yowls during two consecutive days before associating
with both a subadult and resident adult male. We suspect she was in estrus. In
a separate case, a lone, resident male was observed scraping during a yowling
episode; he yowled intermittently over a forty-two-minute period as he scraped
near, then traveled south past, one of our camps. The rougher ouch call was
thought by Padley (1996a) to signify frustration because he heard it after some
of his study animals unsuccessfully pursued prey. Although he did not believe it
was part of the social interactions between pumas, we could not adequately
determine this for our study animals. In the San Andres Mountains, pumas that
vocalized the ouch call would move a short distance, then call again. Usually no
more than twenty minutes elapsed between the time we heard the first and last
calls. In one case, telemetry indicated M88 was emitting ouch calls while trav-
eling toward F103. The female, who was raising one-month-old cubs, moved
away from her nursery and toward M88. Although M88 was not the father of
the litter, it is likely he was the sire of her previous litter. Individual recognition
has not been demonstrated for vocalizations of adult felids, but M88’s call may
have at least indicated his sex and possibly dominance (subordinate individuals
would probably be less likely to vocalize). By intercepting M88, F103 probably
avoided his discovery of her nursery. We did not know why M88 started calling,
but we suspect F103 exhibited a pseudo-estrus because the two pumas spent
four days together, away from the nursery. In another instance, M153 emitted
ouch calls several times as he crossed a saddle and traveled down a drainage fre-
quently used by pumas on Goat Mountain. When Bailey (1993) heard a male
leopard make similar but louder rasping calls just before entering a road com-
monly traveled by other leopards, he postulated that the leopard vocalized to
reduce the chance of a surprise encounter with a fellow traveler. It is possible
M153 vocalized for similar reasons.
Chemical Communication
Odors can be produced by urine and feces, as well as by various skin glands.
They can then be deposited in the environment as scent markers. Scent mark-
ers allow individuals to communicate indirectly with one another, since they can
remain active for long periods, even in the producer’s absence. However, to play
a role in communication, an odor must contain a message that can be decoded
by individuals that encounter it. Identity, social status, and reproductive condi-
tion may all be possible functions of scent markers. Studies on several different
mammals have demonstrated that individuals can be recognized by their odors
alone (e.g., Müller-Schwarze 1971, Gorman 1976, Kruuk et al. 1984), and
olfactory recognition of sex is common (Stoddart 1980). What functions they
may serve in puma societies needs more study.
Scrapes were the most obvious scent markers left by pumas in our study area.
A puma scraped by pushing backward with its hind feet, creating a shallow
trough about 13 to 25 cm long with a small mound of ground debris (e.g., soil,
leaves, sticks, stones) at the distal end. Other scrape measurements can be found
in the Idaho puma re s e a rch (Seidensticker et al. 1973). Scrapes we re made by
almost eve ry resident adult male on our study area, except perhaps M73. We
could not link any scrapes to M73, and we suspect his behavior was linked to
bilateral cry p t o rchidism and possibly to depressed levels of testosterone (see
Chapter 7). The mounds created by scrapes we re usually urinated on, and some
were marked with scats. Scats could contain normal or token amounts of fecal
material. Scrapes were generally placed in prominent locations and along travel
ways, such as ridgelines, channel bottoms, under large trees and ledges, and at kill
caches. Re s e a rchers in Idaho, Utah, and Wyoming found scrapes at similar sites
(Seidensticker et al. 1973, Hemker 1982, Logan and Irwin 1985, respectively).
Males in the San Andres Mountains appeared to scrape throughout their
home ranges. Some scrape sites were used by successive residents and some were
shared by up to three neighboring resident males. Typically, there were a half
dozen or so scrapes of various ages that we could see at shared sites. But a few of
the largest shared sites had more than forty scrapes. In the first three years of our
research, we identified and mapped scrapes made at 346 different scrape sites by
four adult males with overlapping home ranges. We found that 26 to 33 percent
of the scrape sites utilized by an individual male were shared with one or two
other males (Table 14-10; Plate 12). In Idaho, the only other place where puma-
scraping behavior has been described in some detail, no male puma was docu-
mented visiting the scrape site of another. However, home range overlap
between resident males in that study was minimal (Seidensticker et al. 1973).
Because many of a male’s scrape sites on our study area were shared by neigh-
boring resident males, males evidently were communicating their presence to
one another. Because we did not document frequency of scraping, we could not
determine if pumas scraped more frequently along home range boundaries or
within their core use areas. However, shared scrapes were most prevalent along
major drainages where males would presumably have the highest probability of
Table 14-10. Scrape sites (n = 346) used by four resident

adult male pumas on the San Andres Mountains from Sep-
tember 1985 to September 1988. Each puma’s annual and
cumulative home range overlapped some portion of at least
one other male’s range based on the 90 percent adaptive ker-
nel home range estimator.
Male No. of scrape sites No. of shared sites % of sites shared
1a 150 39 26.0
3 151 43 28.5
7b 52 17 32.7
22 48 14 29.2
aM1’s home range extended south of the study area boundary. Only the area
within the study area was searched for scrapes.
bOnly scrapes made in the southern one-half of M7’s home range could posi-
tively be attributed to M7. Scrapes found in the northern half of his range
were not included in the sample.
encounters (Plate 12). Male pumas in Idaho scraped more frequently in the
region of home range overlap or along home range boundaries than within the
central parts of their home ranges (Seidensticker et al. 1973). Studies on tigers,
jaguars, and leopards have reported similar results; scent marking was most fre-
quent along territorial boundaries (Rabinowitz and Nottingham 1986, Smith et
al. 1989, Bailey 1993). In tigers, marking zones also shifted as territorial bound-
aries shifted (Smith et al. 1989).
We rarely found places where adult females and subadult males scraped, a
characteristic also found for pumas in central Idaho (Seidensticker et al. 1973).
However, tracks and radiotelemetry indicated that females in the San Andres
Mountains visited scrape sites, and solitary females occasionally marked them
with urine. It was impossible to determine the frequency of such scent marking
by females, because of the lack of a corresponding visual marker. Females occa-
sionally visited scrapes, and in some instances they caterwauled in the vicinity of
well-used scrape sites, suggesting they were trying to attract mates. Although
females generally did not scrape when they defecated, they sometimes made scat
mounds. Scat mounds consisted of one or more scats buried in a pile of vegeta-
tive debris or soil. Such mounds, or toilets, were often found near kill-cache sites
and were sometimes used by more than one member of a family group. Because
the scats were hidden in a mound, their function as scent markers is question-
able. Rarely did adult male pumas make scat mounds at prey caches.
Pumas sometimes scratched the bark of cottonwood and willow trees, and
these scratch trees were usually located by scrape sites. Lamellae from puma
claws could be found in the scratches, indicating that the principal purpose was
probably claw grooming, similar to the conclusion drawn by Seidensticker et al.
(1973) in Idaho. However, scent from the feet may have also remained. Because
the same trees were recurrently inspected and marked by different individuals, it
is possible that clawing may be part of the puma’s communication system, as is
speculated for African lions and tigers (Schaller 1972, Smith et al. 1989). We
have also observed pumas in zoos rubbing their cheeks and heads on logs and
boulders, and believe that wild pumas do this too. Tigers will cheek rub, then
typically spray urine on sites that have been previously sprayed with urine
(Smith et al. 1989). Although no cheek glands have been found in cats, we
believe cheek rubbing may be used in intraspecific communication. We found
no evidence that pumas spray urine and other fluids to scent mark as do many
other big cats, including the African lion (Schaller 1972), tiger, (Smith et al.
1989), leopard (Bailey 1993), and cheetah (Caro 1994). In the tiger, lipid-rich
marking fluids are deposited at a frequency that is about fifty times higher than
that of urination (Brahmachary et al. 1992), indicating they have an important
role in communication.
Scent marking has been linked to a wide range of behaviors. Possibly two of
the most commonly reported functions may be advertisement of reproductive
status (e.g., Smith et al. 1989, Palanza et al. 1994, Converse et al. 1995), and
activities associated with the occupation of a home range or territory (e.g., Gor-
man 1984, Randall 1987, Clapperton et al. 1988, Smith et al. 1989, Lenti
Boero 1995, Brashares and Arcese 1999). In pumas, information on scent mark-
ing is primarily descriptive, and hence we must be fairly speculative about its
function. Puma scent marks seem to serve as “bulletin boards,” which males and
females visit to determine the temporal presence, reproductive status, quality,
and perhaps even the individual identity of other pumas. Research has shown
that in at least one carnivore, the dwarf mongoose, individuals can distinguish
scents based on age (Rasa 1973). If pumas can judge the age of scent at scrapes,
they may be able to determine how recently another individual has been there
and thus avoid other pumas in the vicinity. Thus, scents may facilitate spa-
tiotemporal separation between neighbors (Leyhausen and Wolff 1959, Bailey
1993). Females may also use scrapes to locate potential mates. Urine accurately
reflects the level of estrogen in the bloodstream, and increased levels of estrogen
are produced during the follicular phase of the estrous cycle (Gorman and Trow-
bridge 1989). If the olfactory message remains viable for a sufficient amount of
time, advertising at male scrape sites should enhance mating opportunities. In
tigers, females apparently scent-marked most intensively at the onset of estrus,
suggesting females were advertising their reproductive status (Smith et al. 1989).
The fact that adult male pumas advertise their presence using scrapes, whereas
females and subadult males typically do not, indicates different behavioral
strategies. Non-estrus females and subadults risk potential injury or death (to
themselves or their cubs) by revealing their location to adult males. The behav-
ior of burying feces may also reduce detection. Conversely, by advertising, an
adult male may demonstrate his dominance and thereby reduce the degree of
trespass by other males (both residents and new immigrants), increase his own
chances of breeding with resident females, and decrease the chances his mates
and offspring will be harmed. Long-term residents have the opportunity to
cover an area with scent marks, giving potential intruders ample opportunity to
retreat before there is a life-threatening encounter.
LP > > KX
1. Each year each adult male home range overlapped, sometimes completely,
the home ranges of an average of 3.3 to 4.9 females.
2. Spatial overlap between the home ranges of pairs of adult male pumas aver-
aged 18 to 28 percent each year. Overlap between core use areas was much
lower, averaging 3 to 13 percent each year. There was a tendency for over-
lap to decline between pairs of male pumas over consecutive years, but this
tendency was not as pronounced as for adult females. Each male shared its
home range with an average of 2.8 to 2.9 other adult males each year. The
total amount of area that a male puma shared with other adult males aver-
aged 50 to 68 percent each year, whereas it shared an average of 14 to 38
percent of its core area.
3. Spatial overlap between the home ranges of pairs of adult female pumas (17
to 26 percent) was similar to the amount of overlap observed between pairs
of adult males. Overlap between core areas averaged 4 to 9 percent annu-
ally. In consecutive years spatial overlap between neighboring females gen-
erally declined. Each adult female shared its home range with an average of
1.8 to 3.8 other adult females each year. The total amount of area a female
puma shared with other adult females ( x– = 43–64 percent each year) was
similar to the overlap found for adult males.
4. Spatial overlap between pairs of adult pumas of the opposite sex was greater
than for same-sex pairs, suggesting competition within each sex was greater
than competition between the sexes. The home ranges of most females were
completely engulfed by surrounding resident males. In contrast, a generous
portion of a male’s range (average of 36.6 percent) was not shared with
females.
5. Adult pumas generally occurred in random distributions, but there was a

tendency toward aggregation in females and avoidance in males. Females
became more aggregated with increasing densities.
6. Although there was often considerable spatial overlap between adult pumas
of the same sex, individuals tended to use shared areas at different times.
Spatiotemporal overlap of core areas averaged 2 percent for pairs of same-
sex pumas. Spatiotemporal overlap of female core areas was most prevalent
in closely related individuals (i.e., mother-daughter or sisters); spatiotem-
poral overlap of male core areas was stimulated by an estrus female or
resulted in fights.
7. Independent pumas associated with one another only rarely (5.1 percent of
locations). Interactions occurred most frequently among males and females
(78.4 percent of associations), and in similar, low frequencies (10.4–11.1
percent) between same-sex individuals. Most male-female interactions were
associated with courtship.
8. Fighting and intraspecific killing occurred in the population. Mortalities of
independent pumas occurred during 7.8 percent of associations. Scarring
on captured adult males indicated that almost all of them had been involved
in fights with other pumas; however, there was no evidence of fighting
between females. Males killed other males during fights over territory,
breeding females, or food caches. Males sometimes killed females and can-
nibalized young.
9. Females and their dependent young form the basic social unit for the
species. Maternal females frequently bedded with their dependent cubs
(81.5 percent of locations).
10. Scent-marking and vocalizations were believed to be important modes of
communication for maintaining spacing and finding mates.
LN N5LN5)L
1. Test for differences in home range overlap between M-M and F-F dyads
each year, using the 90 percent ADK home range estimator. Mann-Whit-
ney U test: for 1989, U = 91, P = 0.381; for 1990, U = 88, P = 0.011; for
1992, U = 352.5, P = 0.601; for 1993, U = 273.5, P = 0.871.
2. Test for differences between years (1989, 1990, 1992, 1993) in spatial over-
lap of same-sex puma dyads, using the 90 percent ADK home range esti-
mator. Kruskal-Wallis test: M-M dyads in the Treatment Area, H = 5.071,
d.f. = 3, P = 0.167; M-M dyads in the Reference Area, H = 2.038, d.f. = 3,
P = 0.565; M-M dyads in the entire study area, H = 3.046, d.f. = 3,
P = 0.385; F-F dyads in the Treatment Area, H = 3.120, d.f. = 3, P = 0.374;
F-F dyads in the Reference Area, H = 5.419, d.f. = 3, P = 0.144; F-F dyads
in the entire study area, H = 2.963, d.f. = 3, P = 0.397.
3. Test for decline in amount of spatial overlap between pairs of same-sex
pumas during consecutive years. Wilcoxon signed-rank test (1-tailed test):
for females between 1989 and 1990, n = 9, T = 11, P = 0.11; for females
between 1992 and 1993, n = 15, T = 29, P =0.04; for males between 1989
and 1990, n = 16, T = 41, P = 0.09; for males between 1992 and 1993,
n = 8, T = 10.5, P = 1.0.
4. Test for differences in spatial overlap between M-M total and F-F total each
year, using the 90 percent ADK home range estimator. Mann-Whitney
U test: for 1989, U = 52, P = 0.171; for 1990, U = 57.5, P = 0.119; for
1992, U = 133.5, P = 0.082; for 1993, U = 44, P = 0.176.
5. Testing for differences in the amount of home range overlap (90 percent
and 60 percent ADK) among M-M, F-F, and M-F dyad pairs during 1992.
Kruskal-Wallis test: for 90 percent ADK, H = 3.69, P = 0.158; for 60 per-
cent ADK, H = 6.017, P = 0.049. Sequential Mann-Whitney U tests on
pair overlaps in core areas revealed a difference between M-M and M-F
dyads: U = 341.5, P = 0.026.
6. Testing for differences between the percentage of a puma’s home range (90
percent and 60 percent ADK) that is overlapped by all other adults of the
same sex and the percentage that is overlapped by all other adults of the
opposite sex in 1992. Mann-Whitney U test: between M-M total and M-F
total, U = 41, P = 0.21 (90 percent ADK); U = 30, P < 0.001 (60 percent
ADK); between F-F total and F-M total, U = 15.5, P = 0.003 (90 percent
ADK), U = 48.5, P < 0.001 (60 percent ADK).
7. Tests for associations between Clark-Evans ratios (R) and total adult puma
density over eight six-month periods, 1989–1993. Spearman correlation
coefficient (2-tailed test): for male R and density, rs = –0.012, P > 0.2;
female R and density, rs = –0.612, P = 0.106, for male-female R and den-
sity, rs = 0.795, P = 0.025.
8. Spatiotemporal overlap indices between M-M and F-F dyads were similar
in 1989, 1990, and 1993 (P 0.44) but greater in males in 1992, based
on the 90 percent ADK home range estimator (Mann-Whitney U test,
U = 221.5, P = 0.014, nm-m = 17, nf-f = 32).
9. Test for differences in the number of associations among M-M (n1), F-F
(n2), and M- F (n 3) dyads during each of four years (1989, 1990, 1992,
1993). Kruskal-Wallis test with 2 d.f.: for 1989, H = 14.481, P = 0.001; for
1990: H = 4.575, P = 0.101; for 1992, H = 14.469, P = 0.001; for 1993,
H = 4.769, P = 0.092. Follow-up Mann-Whitney U tests indicated M-F
associations were greater than M-M or F-F associations for all years, P
0.10. M-M and F-F associations were not different in 1989, 1990, or 1993
(P > 0.382), but F-F associations were greater than M-M associations in
1992 (P = 0.096).
Chapter 15
Adaptive Significance of
Puma Social Organization
Pumas, like most felids, are solitary. Within the solitary framework, pumas are
able to find one another to breed and successfully rear young. This lifestyle has
proven very successful for pumas; historically, they may have had the largest geo-
graphic distribution of any land-dwelling mammal in the Western Hemisphere
other than humans. Even today, their social system can prove adaptive, as long
as pumas have moderate protection from human exploitation and large, inter-
connected habitat patches. In this chapter we first describe the social structure
within the desert puma population we studied. Then we examine the evolved
function of puma social organization. Is the social organization a two-strategy
approach by which male and female pumas maximize individual reproductive
success? Or did it evolve to limit the population?
The Social Structure of Desert Pumas

The puma population is composed of breeding adults, subadults, and cubs.
When cubs and subadults are successfully integrated as breeding adults into
a puma population, they represent the re p ro d u c t i ve success of their pare n t s .
Because breeding adults have the most immediate effect on fitness, and
a re consequently the ones subject to the most intense selection pre s s u re s ,
their social relationships are highly significant (Wrangham and Ru b e n s t e i n
1986).
281
Female Structure
Adult female pumas rear offspring without the aid of their mates. After inde-
pendence, many female offspring move away from their natal ranges in search
of adequate habitat to raise their own young; however, some subadults settle
close to their mothers. In areas where puma populations are unexploited by
humans and the habitat is relatively rich in resources, females may form a matri-
lineal structure whereby mothers increase their inclusive fitness by sharing habi-
tat with daughters. A matrilineal structure may be more prevalent in areas where
dispersal entails higher than average risks. Such may have been the case in the
San Andres Mountains, where females that attempted to disperse to the east or
west encountered large expanses of terrain that could not support a puma. As
matrilines develop and densities increase, females may also become more aggre-
gated. Rapid turnover in adult populations resulting from human off-take prob-
ably hinders matriline formation and may explain why they have not been well
documented in other puma populations.
In our study population, where pumas were protected from chronic human
exploitation, resident females showed strong fidelity to their home ranges over
long periods. However, established females did make occasional home range
shifts to accommodate daughters or even unrelated females. A female’s range
also tended to expand and contract as her reproductive status and food require-
ments of her family changed. As a consequence, range boundaries were not
rigid. Abandonment of an established home range appears to be extremely rare
in female pumas but might occur if a female is unable to successfully produce
offspring and she detects better opportunities elsewhere. If abandonment
occurs, we expect females to rapidly reestablish elsewhere. The puma literature
provides no evidence of transient or nomadic behavior in adult female pumas.
Still, we have no way of knowing how females might behave if the prey base they
depended on declined dramatically and then remained depressed for an
extended period. In the San Andres Mountains, we documented small increases
in female home range size but also continued site fidelity during a drought.
Unfortunately, we were unable to determine the drought’s long-term effects.
Female home ranges, which are generally much smaller than male ranges,
can also vary from being relatively exclusive to exhibiting extensive spatial over-
lap. When overlap is extensive, females usually use shared areas at different
times. Over time, and probably as neighbors became more familiar with one
another, overlap between pairs may diminish. As a result of this mutual avoid-
ance behavior, associations are infrequent and of short duration. On the occa-
sions when females do meet, they appear to behave nonaggressively.
It is unclear how females are able to avoid one another as well as they do,
especially since they typically leave no markers, such as scrapes, that humans can
CHAPTER 15. ADAPTIVE SIGNIFICANCE OF PUMA SOCIAL ORGANIZATION 283
detect. However, we still know very little about the use and effectiveness of olfac-
tory cues; these, as well as vocalizations, may play important roles in female
spacing. Because pumas are most active in dense cover and in darkness, visual
cues may be the least effective method of avoidance. However, it is possible that
females have brief encounters with one another more frequently than our
records indicated. Although we radio-tracked a few individuals continuously
over twenty-four-hour periods (Sweanor 1990), we generally located individuals
at most once per day, and the majority of these locations were at day beds. We
would easily miss most associations if they were fleeting and during the periods
when pumas were traveling.
Adult females typically share most of their home ranges with one or more
adult males, and at least one of these males sires her progeny. Although spatial
overlap is extensive, and females associate with males more than with any other
class of independent puma, associations involve great risk for the female and her
progeny. Consequently, mutual avoidance is the norm. On relatively rare occa-
sions, females associate with males to breed, reinforce pair bonds, and confuse
paternity. All these behaviors may reduce infanticide. A female may cue on
scrapes to find mates when she is in estrus, and to avoid males, especially when
she is raising cubs.
Male Structure
Within the population, resident adult males are responsible for most, if not all,
of the breeding. Male progeny disperse from their natal areas, and as subadults
they usually move long distances to avoid inbreeding and adult males. As a con-
sequence of their dispersal instincts and avoidance behavior, they often traverse
habitats that contain few, if any, pumas. When they do enter an area that sup-
ports resident pumas, they do not typically advertise their presence. Those that
do may be killed. Sometimes, as they try to avoid resident males and garner
strength for another long-distance move, subadults may localize in temporary,
transient ranges. As transients, they grow and gain experience. An increase in sex
hormones, as well as the presence of breeding females, probably causes the
young male to stop dispersing or shirking resident males. If no resident male is
there to contest him, the change from transient to resident is easy. However, in
most instances, a resident male is probably present. Now the transient must
decide whether to challenge or keep moving. A successful challenger will usurp
part, or all, of an established male’s home range, whereas an unsuccessful one
moves on—or is killed.
As do resident females, resident male pumas on our study area showed site
fidelity. Howe ve r, male fidelity had a greater tendency to decline with time.
This occurred because of home range shifts, which we re often motivated by
challenges from new male competitors or the reduction in available mates.

Responses to changes in the number of competitors and available mates result
in ranges with dynamic borders. Howe ve r, adult males probably rarely exhibit
transient or nomadic behavior. T h e re are no known cases of a resident male
abandoning his home range, but we know that seve re reductions in the num-
ber of available mates as well as competitors has resulted in large range expan-
sions. Consequently, if all females we re re m oved from a male’s range, it is plau-
sible he would begin to search farther afield for breeding opport u n i t i e s .
Whether or not he occupies a range completely disconnected from his prior
range will be largely determined by the presence of females and competing
males.
Male home ranges are typically one and one-half to three times the size of
female ranges; hence a male’s range often overlaps the ranges of several females.
Although overlap between adjacent males can be quite variable, more studies
have reported exclusivity in male ranges than in female ranges. When male over-
lap is observed within a population, it is often less than the overlap found for
resident females, but there are many exceptions (including our study in the San
Andres Mountains). Regardless of the amount of spatial overlap, resident males
appear to avoid other males. In our study area, adult males used shared space at
different times and were seldom in direct contact. As male neighbors became
more familiar with one another over time, overlap between them also declined,
but this phenomenon was not as prevalent as it was between neighboring
females. Since resident females do not have a set breeding season and only a few
are receptive at any given time, resident males must constantly patrol their home
ranges to assess the changing reproductive status and availability of females.
Consequently, a resident male must always be in breeding condition. A resident
male communicates his presence, and likely his dominance, by scraping
throughout his home range. Scrapes may be more prevalent along shared bor-
ders or travel ways in overlap zones of male home ranges. When range bound-
aries are dynamic, shared travel routes may be the best places to communicate
with neighbors. The scrapes also help females find males when they are ready to
breed. Unlike encounters between females, direct meetings between males are
occasionally violent. Fights between males can result in death, home range
shifts, or continued dispersal.
The Self-Limiting Hypothesis

The self-limiting hypothesis holds that land tenure and territoriality are mecha-
nisms by which pumas limit their own numbers, and, hence, keep a puma pop-
ulation from harming its food supply (Hornocker 1969, 1970, Seidensticker et
al. 1973). Because of the importance of land tenure and territoriality to self-lim -
itation, we examine these concepts in detail.
Land Tenure and Territoriality

Seidensticker and his colleagues introduced the concept of tenure in pumas
when they stated that pumas in their Idaho study area exhibited “a land tenure
system based on prior rights” (1973:53). As a consequence, a puma that estab-
lished a home range in a particular area was guaranteed the use of that area over
its lifetime. Neighbors as well as transients did not contest an established puma’s
rights to its home range. Instead, neighbors retreated from shared areas if they
detected the presence of another resident, and transients moved on to unoccu-
pied ranges. This was a peaceful way for pumas to space themselves over the
available habitat. Their concept of land tenure implied a degree of stability
within the puma social structure. One could envision a fixed number of home
ranges available for occupancy by resident pumas; once the vacancies were filled,
nonresidents (typically subadults looking for a place to establish residency)
moved elsewhere.
The concept of territoriality implies social dominance over a specific geo-
graphical area. It is used to describe the behavior of many carnivore species. Ter-
ritorial behavior is linked to acquisition of resources and is selected for only
when the individual’s resulting increased access to resources (and, subsequently,
fitness) outweighs the accompanying expenditure of time and energy and the
increased risk of injury and predation (Brown and Orians 1970, Kaufmann
1983, Mace et al. 1983). Territory was first used to describe avian social systems.
As the term was applied to other taxa, definitions became increasingly inclusive.
Consequently, the term is now difficult to clearly define. In their final paper on
the Idaho pumas, Seidensticker et al. (1973:54) avoided the term altogether
“because of the semantic conflicts and muddled concepts brought to mind.”
Early definitions stated simply that territory was a defended area, with the impli-
cation that defense could be exhibited through attack, threat, or advertisement
(Noble 1939, Burt 1943). Later, Pitelka (1959) introduced the concept of exclu-
sive range use. More recent authors typically define territoriality as the “area
occupied more or less exclusively by an animal or group of animals by means of
repulsion through overt defense or advertisement” (Wilson 1975:256).” How-
ever, Emlen (1957) presented a somewhat different concept that was later sup-
ported and expanded on by others (e.g., Owen-Smith 1977). He believed an
animal was territorial if, in a given area, it showed supreme dominance over
intruders. Consequently, territories need not be exclusive. Etkin (1964:24–25)
may have provided the least-restrictive definition of territoriality: “any behavior
on the part of an animal which tends to confine the movements of the animal
to a particular locality.” In this case, territories need not be defended or exclu-

sive. This was the interpretation adopted by Hornocker (1969) when he first
analyzed the winter relationships of pumas in Idaho. More recently, Kaufmann
(1983:9) synthesized the different concepts of territoriality and developed his
own definition: “a fixed portion of an individual’s or group’s range in which it
has priority of access to one or more critical resources over others, which have
priority elsewhere or at another time. This priority of access must be achieved
through social interaction.” His definition also did not require overt defense or
exclusive use.
Do Desert Pumas Exhibit Land Tenure or Territoriality?

Female pumas in the San Andres Mountains could only be considered territo-
rial under Etkin’s (1964:24–25) broad definition of the term. Females did not
exhibit the two behaviors most often cited as components of territoriality: exclu-
sive use and defense. On the contrary, spatial overlap between female ranges was
often extensive, and they did not attempt to repulse other females through fight-
ing or advertisement. However, females exhibited mutual avoidance behavior
and tended to use shared areas at different times; consequently, it might be
argued that a female using a shared area was territorial because she had “supreme
dominance over intruders” (Emlen 1957), or “priority of access” (Kaufmann
1983). Because we had no way of determining which puma retreated from a
shared area upon detecting the presence of the other, we could not adequately
assess this argument. Since dominance was not reinforced through advertise-
ment (e.g. scraping) or fighting, it is likely that individual females often made
the judgment whether to leave or stay based on the prospect of hunting success,
risks to vulnerable cubs, and possible recognition of kin or nonrelatives, and not
on which puma was there first. Consequently, we conclude that female pumas
in the San Andres Mountains were not territorial but simply exhibited mutual
avoidance behavior.
If land tenure is interpreted to mean long-term fidelity to a particular site,
then female pumas on the San Andres Mountains exhibited this behavior. How-
ever, the term implies prior rights to an area and is integral to the argument of
self-limitation in puma populations (Seidensticker et al. 1973). In this context,
there are a fixed number of home ranges that can be filled by adult females;
young adults cannot establish home ranges unless vacancies become available.
This was not the case in the San Andres Mountains. Young females often estab-
lished adult ranges in areas already occupied by long-established females. Simi-
lar behavior by female pumas was observed in Alberta, where subadult females
sometimes “established residency adjacent to their mothers, squeezing between
the home ranges of two or more older residents” (Ross and Jalkotzy 1992:424).
We also documented cases where established residents made home range shifts
to accommodate daughters and even unrelated newcomers. If females remained
neighbors for long periods, the amount of spatial overlap between them tended
to decline, but this may have been a result of mutual recognition and avoidance
and not necessarily an acknowledgment of prior rights. We did not document
any overt aggression between females that might have caused dispersal in newly
independent females. The argument that the San Andres Mountains housed a
rebuilding population, hence not all vacancies within a land tenure structure
were filled, may have been true during the early years of the study, but it was not
supported during later years. New females continued to establish adult home
ranges in areas that overlapped the ranges of well-established adults, even as the
mule deer population was rapidly declining.
In the San Andres Mountains, land tenure “based on prior rights” (Seiden-
sticker et al. 1973:53) did not accurately fit male behavior, either. Resident
males certainly encouraged young subadult males to move on, unless they killed
them first. However, males that had just reached adulthood and had no estab-
lished range sometimes contested a resident male’s rights to his home range. And
sometimes the resident lost the battle. If the system were based strictly on prior
rights, a young male would not challenge a resident but would keep moving
until he found a place that was void of resident males. Fights between residents
also resulted in a realignment of range boundaries. One resident sometimes
gained ground while the other lost it. Such give and take made for fuzzy bound-
aries and contributed to the spatial overlap we observed on our study area. In
contrast to decisions made by females, a new male’s decision to remain in an area
or move on was probably strongly affected by the presence and perceived dom-
inance of resident males. We suspect a newcomer only instigated a fight if he
believed he could successfully challenge the dominance of a resident.
We believe it is appropriate to consider male pumas in the San Andres
Mountains, and perhaps in most populations, as territorial. Males often exhibit
one or both of the behavioral components typical of territoriality: repulsion of
other males through advertisement (scraping) and fighting; and exclusiveness (in
time if not in space) of home ranges. Adult males in all puma populations prob-
ably mark their home ranges with scrapes, even though only a few studies (Sei-
densticker et al. 1973, Hemker 1982, our study) have actually provided much
detail on the behavior. Evidence of fighting between males has also been
reported across the puma’s North American range. Even in central Idaho, where
male pumas initially appeared to peacefully maintain exclusive ranges (Seiden-
sticker et al. 1973), fighting has recently been documented (Holly Akinson,
Hornocker Wildlife Institute, personal communication). At least some fights in
the San Andres Mountains occurred because males were competing for a breed-
ing female or trying to maintain or obtain territory that contained breeding

females. Although exclusive ranges have been reported for male pumas, several
studies (including ours) have also reported some degree of spatial overlap; con-
sequently, if territoriality is strictly viewed as a spatial concept, male pumas can-
not generally be considered territorial. However, we found that males in our
study area practiced mutual avoidance by using shared areas at different times.
This is similar to the way females behaved except it is more likely that males
exhibited “supreme dominance over intruders” or “priority of access,” at least
when intruders were neighboring resident males. Neighboring resident males
were probably very familiar with each other’s strengths because of prior con-
frontations. A male that entered a shared area conceivably determined whether
or not his neighbor was nearby based on the age of scent left in scrapes. A male
that remained in an area already occupied by his neighbor risked death. Death
would mean the loss of his entire territory, including a lot of area not shared
with his combatant. Consequently, it was probably in the intruder’s best inter-
est to retreat until another day.
On the San Andres Mountains, female and male pumas exhibited mutual
avoidance, and male pumas were territorial. However, we found no evidence
that these behaviors were mechanisms that limited the population, particularly
during a period of severe deer decline.
The Two-Strategies Hypothesis

This hypothesis holds that male and female pumas have evolved different strate-
gies for maximizing individual reproductive success. In this hypothesis too,
mutual avoidance and territoriality are important mechanisms. Mutual avoid-
ance enables females to successfully rear young, and territoriality enables males
to successfully compete for mates. However, these mechanisms do not function
to limit the puma population below the level set by the prey.
Female Strategy
A female puma’s reproductive success is limited by the number of offspring she
can raise to independence in her lifetime (see Chapter 7). To be successful, the
female must survive as long as possible and provide adequate nourishment and
security for her cubs. Two ways she accomplishes this is by maintaining high
home range fidelity and practicing mutual avoidance behavior. If a female puma
can enhance her progeny’s survival and opportunity for procreation after inde-
pendence, all the better. Consequently if she resides in an area that provides an
abundance of both security and food, she should not discourage daughters from
settling there, too. As long as the benefits of philopatry exceed the costs of shar-
ing resources with daughters, the mother’s inclusive fitness will increase. This
explains how female philopatry and the resulting matrilineal structure could
evolve (see Plate 4).
Unlike males, females do not need to compete directly with one another for
access to mates. Female pumas also do not aggressively compete with one
another for food resources. Instead, they avoid one another in time and space.
However, some independent female progeny that perceive competition for food
is too high in the natal area may choose to disperse and search for better habi-
tat (see Chapters 10 and 13). Dispersing females can colonize other suitable
habitats or recolonize previously occupied habitats.
Given that a female’s reproductive success is so dependent on habitat qual-
ity, habitat quality should have a great influence on female range size. Adult
female home ranges are generally much smaller than those of adult males. The
size of a resident female’s home range is probably determined most by the dis-
tribution and abundance of prey needed to support her and her cubs, as well as
cover to successfully stalk prey and ensure security. In the San Andres Moun-
tains, some of the largest female home ranges were in areas where we suspected
deer densities were lowest. An examination of puma densities, prey abundance,
and puma range sizes each year suggested that prey distribution (if not abun-
dance alone) probably had an important role in determining female range size.
The juxtaposition of potential mates may also have had some influence on
female range size, since females generally utilized larger areas when they were
solitary (and probably in breeding condition) than when they were raising cubs.
This behavior may benefit the female because she has greater opportunity to
establish positive relationships with all of the males that share or abut her home
range, and consequently confuse paternity. Our observations indicated that a
female’s subsequent encounters with these males, even when accompanied by
cubs, were generally not aggressive. Such meetings with unfamiliar males, how-
ever, sometimes proved deadly.
Overlap in female ranges should be influenced by habitat characteristics, par-
ticularly as it affects prey abundance, distribution, and stability (Brown and Ori-
ans 1970, Hixon 1980, Sandell 1989). Low to moderate prey numbers, fluctu-
ations in those numbers, and a patchy prey distribution probably contributed to
the spatial overlap we observed in resident female home ranges. Seidensticker et
al. (1973:57) suggested that the highly mobile nature of the puma’s prey made
it seem “unlikely that a female could maintain an exclusive area of sufficient size
to provide food for herself and her kittens throughout their development.” We
also observed variations in habitat characteristics throughout the San Andres
Mountains; elevation, aspect, and slope, as well as springs, had a tremendous
impact on the type of vegetation that occurred at a site, and consequently, on
the site’s suitability as deer habitat. Thus, patchiness in habitat probably corre-
sponded to variability in the spacing of the deer. In the Sierra Nevada of Cali-
fornia, pumas had overlapping home ranges where deer were more abundant
(Pierce et al. 2000b). If prey varies in space and time, the female’s range must be
larger to provide for her at all times. Consequently, the area may contain more
than enough food during part of the year, allowing more than one female to use
the same area. It is also possible that given relatively low prey densities and a
patchy distribution, no individual patch would be large enough to support a
female. In this case, she would have to visit several patches that might be well
spaced in order to survive and procreate. To maintain such an area to the exclu-
sion of all other females would require large daily movements (to patrol and
mark) and possibly physical defense. Energy expenditure alone would probably
exceed any benefits derived from exclusive use of an area. Patrolling would also
mean more time away from her vulnerable young and less time for prey acqui-
sition so she could feed those young. The added risks of injury from fighting
would be intolerable. Together, these costs would lower the female’s fitness.
Consequently, the best tactic would be avoidance of other pumas that use the
same area.
In summary, a female puma maximizes her reproductive success by main-
taining a home range in which she is familiar with vital resources, practicing
mutual avoidance, raising as many cubs as she can in her lifetime, cultivating a
matrilineal structure, and fostering amicable relationships with territorial males.
These behaviors increase a female’s own survival and her ability to raise cubs to
independence. In the San Andres Mountains, the matrilineal structure and
nonaggressive behavior of females apparently allowed female density to increase
even as the main food base—mule deer—declined. The rate of deer population
decline was strongly influenced by puma predation (see Chapter 17).
Male Strategy
A male’s reproductive success is more variable than a female’s (see Chapter 7).
Because an adult male is always in breeding condition and does not have to care
for dependent young, he has the capacity to produce many progeny in his life-
time. But first he must secure access to mates. Breeding opportunities are lim-
ited because females are dispersed in low numbers over the landscape, in estrus
at variable times, and not available for breeding during the time they are raising
young. To successfully breed with as many females as possible, the male must
traverse a large area. Yet the area cannot be too large, since the male must fre-
quently visit the different parts of his range to assess the reproductive status of
resident females and to defend against incursions from other males. The latter
could not only threaten his chances to breed but also kill the cubs he has sired.
Consequently, if he can, it is in the male’s interest to exclude other males from

an area that contains his mates. On the other hand, the male that kills another
male’s offspring can benefit his own reproductive success by causing the mother
to re-cycle and thus provide him with an opportunity to mate. Dominance in
shared areas can be obtained by repeated scent marking, direct confrontations,
and fighting. The maximum size of a male’s range is affected by the density of
adult females, his mobility, and his aggressiveness. The latter two characteristics
have probably contributed to selection for larger body size of male pumas (see
Chapter 7).
Competition from adult males and the avoidance of inbreeding have influ-
enced the behavior of the subadult male. By dispersing a long distance, he min-
imizes the chance of mating with closely related females, particularly those that
may form a matriline. Such matings could have immediate as well as long-term
repercussions on individual fitness. Long-distance moves also give young males
time to gain the body mass and experience needed to compete with other males,
and may help them locate areas with few competitors and many mates. The
added benefit of dispersal is that pumas can colonize other suitable habitats or
recolonize places from which they were previously extirpated.
It has been argued that actual fighting among highly specialized carnivores is
not advantageous to their physical well-being and surv i val (Ewer 1968,
Slide @318%
PHOTO 25. Male pumas were responsible for all of the killing of other pumas on the San
Andres Mountains.
Hornocker 1969). However, male pumas are (1) equipped with formidable
fighting machinery, (2) in a continued state of reproductive readiness and con-
sequently heightened aggression, (3) competing for a rare resource (females in
breeding condition), and (4) rarely in direct contact with other pumas, so they
may not have evolved behavioral mechanisms that prevent injuries during
intraspecific encounters (Lorenz 1971). Consequently, it is easy to understand
why fights sometimes occur. An adult male may try to dispatch a young
subadult within his territory because an inexperienced subadult is not a formi-
dable (e.g., risky) opponent; however, if ignored, he could become one. Young
adult males that are just reaching maturity may also instigate fights; they feel
aggressive but probably lack the caution of residents because they have no terri-
tory to lose. Fights between established residents should occur less frequently,
since they have become familiar with each other’s strengths and are unwilling to
risk the loss of territory (or their life) in a skirmish. Still, given the rarity of
breeding opportunities, fights between resident males may be provoked by an
estrus female.
Since male pumas are competing for a limited resource, and they fight to
protect or gain rights to breed, we might expect male ranges to be exc l u s i ve. Yet
many populations re p o rt some spatial overlap between adult males. In our pop-
ulation, spatial overlap between pairs of neighboring adult males averaged 24
p e rcent. Why was the overlap so high? In solitary carnivo res, males are affected
by the abundance, distribution, and mobility of females. Sandell (1989)
h y p o t h e s i zed that male ranges should be exc l u s i ve when females are dense and
e venly distributed, and overlapping under all other circumstances. We can eas-
ily argue that in many populations (including the San Andres Mountains),
females occur at low densities and are patchily distributed. Patchiness results
f rom an uneven distribution of re s o u rces and resulting overlapping female
ranges. Additionally, even though puma populations throughout their range
indicate a pattern of more adult females than adult males, many females are
actually unavailable for breeding because they are pregnant or raising yo u n g .
Consequently, the operational sex ratio (i.e., the ratio of adult males to females
that we re available for breeding) may approach 1:1 or even favor males. That
was the case in the San Andres Mountains. Besides being a rare re s o u rce, re c e p-
tive female pumas are also very mobile and spatially unpredictable within their
home ranges. Consequently, it is often difficult for males to monopolize a
number of fertilizable females. All of these characteristics should lead to widely
overlapping male home ranges (Lott 1991, Sandell 1989, Waser and Wi l e y
1979).
In Sandell’s (1989) analysis of the spacing behavior in solitary male carni-
vores, he believed they could adopt two different strategies to achieve matings:
stay in exclusive ranges and monopolize a number of females; or roam over large,
overlapping ranges and compete for each single female that comes into heat.
Even when females are evenly distributed, there should be a threshold density
below which the payoff is greater when the male roams in search of receptive
females than when he tries to defend an area with a depleted number of females.
Both behaviors have been observed in martens (Taylor and Abrey 1982), North
American lynxes (Bailey et al. 1986), ermines (Sandell 1986), and cheetahs
(Caro 1994). Based on Sandell’s (1989) premise that the patterns of roaming
behavior show a continuum, from extremely wide-ranging behavior to spatial
overlaps between male pairs as low as 10 percent (measured on convex poly-
gons), male pumas in the San Andres Mountains exhibited a degree of roaming
behavior. Pairs of males with overlapping ranges also directly competed for
breeding rights with resident females that were in estrus. But throughout the lit-
erature, a roaming strategy typically implies transience. For example, in chee-
tahs, roaming (or floating) males showed no consistent or predictable use of a
home range over time. Although territorial males probably obtained greater
breeding success, floaters were also successful at times (Caro 1994). In contrast,
there are no confirmed cases of a floating male having reproductive success in
any puma population. Adult male pumas on the San Andres Mountains exhib-
ited strong home range fidelity, and although they occasionally made excursions
outside their home ranges, we observed successful breeding only between resi-
dent males and the females whose home ranges they overlapped. The only other
study to examine reproductive success in pumas occurred in Yellowstone
National Park, and it reported similar results. Adult male pumas often estab-
lished overlapping ranges; these residents were responsible for all successful
breeding (Murphy et al. 1998a).
In summary, a male on the San Andres Mountains maximized his reproduc-
tive success by being territorial. Benefits of territoriality included priority in
mate access and protection of offspring and mates from other potentially canni-
balistic males. These outweighed potential benefits (i.e., possible increased
encounters with estrus females) that could have been derived from wide-ranging
or floating behavior. But given the low densities, and the very mobile and spa-
tially unpredictable nature of females, a male simply could not maintain an
exclusive territory. The male puma strategy was not a clear case of either territo-
rial defense or mate defense, but a combination of both. Although territorial
behavior sometimes caused the death or expulsion of a competitor and thus
affected male puma density, we do not believe the function of territoriality was
to limit the population below the level set by the prey (i.e., the self-limiting
hypothesis).
Pumas and Other Big Cats—Similar Strategies?

Pumas share many behavioral characteristics with other large, solitary felids,
including the leopard, tiger, and jaguar. These species inhabit similar environ-
ments: cover is dense and terrain is often rugged, thereby inhibiting movement
and providing poor visibility over long distances; prey also occur at relatively low
densities and are well dispersed. These environmental and prey characteristics
encourage a hunting style that relies on stealth and concealment. Hunts typically
consist of stalking to within a close distance of prey, followed by a short charge
(Hornocker 1970, Leyhausen 1979, Bailey 1993, Stander et al. 1997). Condi-
tions where these felids live, with the resulting high costs of sociality, may
explain why they remain solitary. Group living requires the sharing of food
resources (Packer et al. 1990); consequently groups must spend more time for-
aging or kill larger prey (Caro 1989). Larger congregations around carcasses also
increase the risk of detection and attendant competition from other carnivores
(Stander et al. 1997, Sunquist and Sunquist 1989), and heighten predation risk
to offspring (Packer and Pusey 1983).
Only the social structures of the African leopard and tiger have been studied
extensively enough to provide clear comparisons to the puma. More limited
information is available on the jaguar. Studies of all three felids indicate similar
behavioral patterns to pumas: home ranges are much larger in adult males than
in adult females, and male ranges overlap the ranges of more than one female
(Bailey 1993, Stander et al. 1997, Sunquist 1981, Rabinowitz and Nottingham
1986). Of the three species, the leopard may be the puma’s closest ecological
equivalent. It also inhabits a wide variety of habitats and had a wide, but recently
reduced, distribution (Schaller 1972, Bailey 1993). Recent radiotelemetry stud-
ies of leopards in both South Africa and Namibia, spanning two and three years,
respectively, indicate leopards behave similarly to pumas. Although both studies
found that leopards shared parts of their home ranges with adults of the same
sex (pair overlap in Namibia leopards during each of two years averaged 29 to
46 percent for males and 35 to 43 percent for females; for South African leop-
ards it averaged 19 percent for males and 18 percent or more for females), there
was temporal avoidance of shared space (Bailey 1993, Stander et al. 1997).
Direct associations between adult leopards were also rare and most probably
occurred for the purpose of breeding (Bailey 1993). Mutual avoidance was
enhanced through the use of scrapes and other olfactory cues and vocalizations;
fighting appeared to be rare. Although subadult males appeared to disperse more
slowly from their natal ranges than pumas, female leopards had a tendency to
remain philopatric, similar to female pumas on the San Andres Mountains (Bai-
ley 1993).
Tigers apparently have a greater tendency to use exclusive home ranges and
may be the most territorial of the solitary felids. Tigers in Chitwan National
Park, Nepal, have been studied for well over a decade (Sunquist 1981, Smith et
al. 1987, Smith 1993). In that population, both males and females established
territories from which other same-sex individuals were excluded. Male territorial
behavior was most pronounced when females were in estrus, whereas females
appeared to compete for food, cover, and a secure place to raise young (Sunquist
1981, Smith et al. 1987). Exclusion was obtained by scent marking, mostly
along territorial borders, and by fighting. Fights generally resulted in either one
individual (typically a subadult) leaving the area, or a shift in range boundaries.
Adult females exhibited long-term fidelity, although they sometimes made ter-
ritorial shifts to accommodate philopatric daughters. Philopatry was common in
females. Although males attempted to disperse long distances, man-induced iso-
lation of the population (similar to circumstances experienced by male Florida
panthers) sometimes forced males to return to their natal ranges (Smith 1993).
The jaguar is the only large felid that is sympatric with pumas. Only three
studies provide limited quantitative information on the movements of few adult
jaguars over an extended period of time (Crawshaw and Quigley 1991, Rabi-
nowitz and Nottingham 1986, Núñez et al. in press). Radiotelemetry research
suggests variability in home range overlap between adults of the same sex. In
populations in Brazil and Mexico, females tended to inhabit overlapping ranges
(Crawshaw and Quigley 1991, Núñez et al. in press); unfortunately, spatial rela-
tionships between males could not be determined because either no information
was provided or only one adult male was radio-collared. In Belize, the ranges of
four radio-collared adult males overlapped, but sign (tracks and visual observa-
tions) suggested that two uncollared females had exclusive ranges (Rabinowitz
and Nottingham 1986). However, there is no way to ascertain whether this
exclusivity was indicative of the population as a whole, or even whether other
females may have been present. As with the puma and leopards, jaguars appear
to practice mutual avoidance and utilize areas of overlap at different times. The
degree of marking by scrapes and feces in jaguars also varies but appears to be
most common in areas of high male overlap (Rabinowitz and Nottingham
1986). The data are sketchy, yet philopatry in female offspring may be a com-
mon trait. The four adult females studied in Brazil apparently consisted of two
mothers and their daughters (Crawshaw and Quigley 1991).
How do the solitary felids differ from the social ones? “Social behavior
evolves as an adaptation to maximize fitness, given a particular set of ecological
pressures” (Wrangham and Rubenstein 1986:452). The African lion and chee-
tah, which are the only known wild felids to exhibit sociality, generally occupy
open environments where prey occur in large, concentrated herds. In cheetahs,
females are solitary but males may form coalitions of two or three individuals
that defend a territory where females aggregate for the purpose of feeding. Coali-
tions are typically composed of littermates or of two brothers and a nonrelative
(Caro 1994). In African lions, females live in permanent prides, which consist
of one to eighteen related females, their dependent offspring, and a coalition of
one to nine adult males that are not related to the females (Schaller 1972, Packer
et al. 1988). Coalitions of male lions defend female prides rather than a specific
territory, and large coalitions can defend more than one pride simultaneously
(Packer and Pusey 1983). Smaller coalitions (two or three individuals) may be
composed of unrelated male lions, but coalitions larger than three are composed
entirely of close relatives (Packer et al. 1991). In his analysis of cheetah social
structure, Caro (1994) concluded that intense male competition resulting from
high densities and localized female distributions provides the conditions neces-
sary for the formation of male coalitions. Since most felid species (including the
puma) rarely encounter both conditions, competition is less intense, and males
remain asocial. Coalitions are typically larger in male lions than in cheetahs,
because male lion cubs, which are often born to several female pride members
simultaneously, usually disperse together (Pusey and Packer 1987). Larger male
groups appear to be further encouraged by the behavior of pride females. Dur-
ing the period when one male alliance is ousted by another, dependent offspring
are killed and females come into breeding condition. However, females also
exhibit temporary infertility. The combination of high sexual activity and tem-
porary infertility attract more coalitions, incite further competition, and typi-
cally result in a pride takeover by the largest-available coalition. This benefits the
pride females because larger coalitions are capable of retaining tenure longer
than smaller coalitions. Since it may take twenty-five months after a pride
takeover for a pride female to successfully raise a litter of offspring, longer tenure
enhances her reproductive success (Packer and Pusey 1983). The reason larger
coalitions are made up entirely of close relatives can again be measured in repro-
ductive success. Packer and colleagues assert that most members in larger coali-
tions do not get to breed, so if they are unrelated to the breeding members, they
do not realize any fitness benefits. In contrast, non-breeding relatives reap inclu-
sive fitness benefits (Packer et al. 1991).
The fundamental reason for pride or group living in female felids is not well
understood. Caro (1994) discusses two related hypotheses that suggest that the
presence of large prey at least enables females to be social. In one, the habitat
simply supports enough prey of a large-enough body mass (e.g., the same size as
the female or up to twice her body weight) for a female to feed additional mem-
bers. In an alternative hypothesis, females living at high densities improve their
inclusive fitness by sharing carcasses with kin when prey are large and highly vis-
ible. The large carcass provides enough food for more than one individual, and
the larger number of group members provides better defense of an easily
detected carcass from unrelated females as well as competitors of other species.
In their examination of why lions form groups, Packer and colleagues (1990)
agree that large prey size permits individuals to forage together, but other factors
provide the positive advantages of living in groups. These include cooperative
defense of young against infanticidal males and of territory against intruding
females. As we proposed in Chapter 7, philopatry could have been the basis for
the evolution of the pride in African lions. Resulting matrilines would have
made it easier for related females to associate in high-density prey areas and at
kills of large prey. Behavioral strategies that involved cooperative hunting and
defense of food and cubs would have enhanced the inclusive fitness of related
females that formed prides.
The social organization found in pumas as well as the other large felids indi-
cates the flexibility of these cats in the face of different environmental conditions
and the type, distribution, and density of prey that those environments support.
Other large cats have many of the same behaviors as the puma, which is not
unexpected since they share a common ancestor. As more research is conducted
on the jaguar, the African leopard, and the more-elusive species such as the snow
and clouded leopards, biologists will probably observe additional behavioral
similarities. We may also see changes in the behavior of these naturally outbred
species as their habitats become more fragmented and isolated through human
development. Changes are already occurring, since, in some populations, indi-
viduals that want to disperse are not able to do so (Beier 1995, Maehr 1997a).
In his synthesis of solitary felid behavior, Bailey (1993) speculated that fighting
might occur more frequently in isolated populations. Perhaps isolation of Chit-
wan National Park has encouraged the greater levels of aggression observed in
both the male and female tigers that live there.
In pumas, it appears that the social organization is a behavioral expression of
the life-history strategies used by individuals to maximize reproductive success.
If these behavioral patterns are adaptive because they maximize fitness, what
effect does that have on their major prey? We address that question in Part 4.
LP > > KX
1. The social organization of pumas on the San Andres Mountains apparently
enhanced individual reproductive success. Pumas employed two strate-
gies—one by females and another by males. Adult females exhibited strong
home range fidelity, practiced mutual avoidance behavior, cultivated a
matrilineal structure, and fostered amicable relationships with territorial
males. These behaviors increased a female’s ability to produce cubs and
ensured her and her cubs’ survival. Adult males were territorial. Benefits of
territoriality included priority in mate access and protection of offspring
and mates from other, potentially cannibalistic males. These outweighed
potential benefits (i.e., possible increased encounters with estrus females)
that could have been procured from floating behavior.
2. Puma social organization apparently did not function to limit the popula-
tion below the level set by the prey.
3. Pumas behave in similar ways to leopards, jaguars, and tigers, which also
live in habitat with closed vegetation and/or rugged terrain and a dispersed
prey base. Puma behavior may help biologists to better understand how
other less-studied, solitary-living felids may behave.
P a rt I V
Puma–Prey Relationships
Chapter 16
Puma Diet
Patterns of Pumas and Prey

Pumas opportunistically take advantage of the most abundant and vulnerable
prey. This generalist strategy probably enabled pumas to thrive after the late
Pleistocene extinctions that doomed their more specialized competitors and to
adapt to a wide range of habitats. In addition, prey size probably imposed selec-
tive pressures on puma body size, thus shaping the pattern in which pumas are
smallest in equatorial regions and increase in body size with latitude. The largest
pumas are in the extreme northern and southern portions of their distribution
(Kurtén 1973, Iriarte et al. 1990). Mean mass of vertebrate puma prey has been
positively correlated with puma body mass at different latitudes (Iriarte et al.
1990) and even within the same population (Murphy et al. 1998b). In other
words, larger pumas tend to kill larger animals. In an evolutionary sense, as
larger prey became more available and vulnerable farther from the equator,
larger puma variants were probably selected because they could kill larger ani-
mals and more efficiently convert acquired energy and nutrients into greater
reproductive success than could smaller pumas. By contrast, smaller pumas liv-
ing in equatorial regions more efficiently used smaller prey that were relatively
more available and vulnerable to them than larger prey. Sympatry with the
larger-bodied jaguar in those habitats may have imposed additional selective
pressure on the puma to adapt to using smaller prey (Iriarte et al. 1990) and
thereby reduce exploitation and interference competition.
In tropical regions pumas tend to prey on relatively small and taxonomically
301
302 PART IV. PUMA–PREY RELATIONSHIPS
diverse prey. In Belize, Peru, Paraguay, Argentina, and parts of Chile, pumas
thrive on prey that, on average, weigh less than 15 kg (Iriarte et al. 1990, Branch
et al. 1996, Taber et al. 1997). This includes a long list of animals in diverse
orders: pudus (the smallest deer) and peccaries, marsupials, carnivores, lago-
morphs, rodents, armadillos, anteaters, primates, and bats (Iriarte et al. 1990).
Larger ungulates become more important as their numbers and distribution
increase in puma habitat along a latitudinal scale (Iriarte et al. 1990). For exam-
ple, in southern Chile (Franklin et al. 1999), the guanaco (Lama guanicoe) was
the most important prey item by biomass (59 percent), followed by the human-
introduced European hare (Lepus capensis) (25 percent). In western Mexico,
white-tailed deer comprised the greatest prey item by biomass (66 percent). The
nine-banded armadillo (Dasypus novemcinctus) was second in importance (11
percent), with the collared peccary not far behind (9 percent) (Núñez et al.
2000).
Pumas in temperate North America prey principally upon mule deer, white-
tailed deer, and elk, with adults of these species weighing well over 15 kg. Elk,
the largest common puma prey in North America, weigh 225–350 kg, which is
easily three to ten times the mass of pumas that hunt them. Other ungulates are
used where they are locally abundant and vulnerable, such as peccary, moose,
and bighorn sheep. Smaller prey, such as rodents (e.g., porcupines, beaver [Cas -
tor canadensis]), lagomorphs, and carnivores (e.g., coyotes, striped skunk, rac-
coon [Procyon lotor]) are eaten opportunistically (Logan and Sweanor 2000, and
references therein). In localized areas, pumas also prey on livestock, usually
sheep and cattle, which may make up 0–34 percent of their diet (this study,
Cunningham et al. 1995). Pumas infrequently prey on domestic horses, and
they occasionally prey on domestic dogs and cats where residential areas invade
puma habitat (Aune 1991, Beier and Barrett 1993, Torres et al. 1996).
Puma Diet on the San Andres Mountains

Given the propensity for pumas in North America to rely on ungulate prey, we
expected the desert mule deer to be the mainstay of pumas on the San Andres
Mountains (Table 16-1). Out of 525 dead prey animals we found that showed
evidence that they had been killed and eaten by pumas, over 90 percent (n =
479) of them were deer. Of those, seventy-seven wore radio collars. We found
only ten desert bighorn sheep (1.9 percent) killed by pumas during our ten-year
study. This was partially a reflection of their very low numbers and the extremely
low chance of finding kills in the rugged terrain inhabited by the sheep. All the
sheep we found wore radio collars. Pronghorns rarely (1 percent) fell victim to
hunting pumas. Those that did were killed at springs with heavy vegetation and
CHAPTER 16. PUMA DIET 303
low visibility in the foothills and canyons at the north end of the mountains.
The pronghorns were clearly out of their usual habitat of relatively flat terrain
with short vegetation, where they ordinarily were not vulnerable to puma pre-
dation. We found only three oryx (0.6 percent) killed by pumas—all of them
calves about the size of an adult mule deer doe. This low number, again, was
probably partially a function of the very low number of oryx ranging in puma
habitat. In addition, adult male and female oryx, weighing over 200 kg and pos-
sessing spear-like horns reaching 150 cm long, are probably a formidable chal-
lenge to pumas. All other species of prey that we found consumed by pumas,
including coyote, striped skunk, badger, ringtail, jackrabbit, porcupine, and
golden eagle, each comprised less than 1 percent of the total victims. The lone
golden eagle was killed apparently as it attempted to scavenge from a puma-
killed deer. Other pumas were puma food on rare occasions (2 percent).
Although small numbers of cattle, including newborn calves, often were in
puma home ranges along the western edge of our study area, we did not find any
cattle killed by pumas during the ten years of research. Nor did ranchers in that
area report any puma-killed livestock.
Pumas sometimes killed animals but did not consume them, including four-
teen other pumas, four gray foxes, one coyote, and one long-eared owl. Biolo-
gists in Montana concluded that pumas protected food caches by killing but not
Slide @318%
PHOTO 26. Desert mule deer were the most important prey to pumas on the San Andres
Mountains.
Table 16-1. Prey killed by pumas on the San

Andres Mountains, New Mexico, 1985–1995.a,b
Species No. recorded Percent
Desert mule deer 479 91.2
Desert bighorn sheep 10 1.9
Pronghorn 5 1.0
Oryx 3 0.6
Puma 13 2.0
Coyote 3 0.6
Striped skunk 3 0.6
Badger 1 0.2
Ringtail 1 0.2
Leporids 1 0.2
Porcupine 5 1.0
Golden eagle 1 0.2
aPuma prey included puma kills and probable puma kills.
bIn addition to these animals, pumas also killed but did not
eat fourteen other pumas, four gray foxes, one coyote, and
one long-eared owl.
eating coyotes (Boyd and O’Gara 1985). Similarly, while studying resource use
by pumas, bobcats, and coyotes in the central Idaho wilderness, Koehler and
Hornocker (1991) explained that pumas defended or usurped food caches by
killing but not eating bobcats and coyotes. As we discussed previously, pumas
were killed during intraspecies competition for food and mates. We suspect that
the gray foxes and the coyote were killed as a result of interspecies competition
for food. We know from evidence of tracks and feces at caches that these carni-
vores occasionally scavenged from puma kills. We even watched a lone coyote
usurp a large mule deer buck that had just been killed by a radio-collared
subadult female puma. Another explanation is that bobcats, coyotes, and foxes
are potential predators of small puma cubs; hence, female pumas that kill those
enemies enhance their own reproductive success. We believe that bobcats and
coyotes are capable of killing puma cubs that are smaller than they are, and coy-
otes could certainly gang up on cubs (e.g., this happened to six-month-old cub
M137). Gray foxes could probably kill newborns. On two occasions we
observed gray foxes lounging on rocks within about 20 m of puma nurseries
with suckling cubs. Cubs this small could be vulnerable to fox predation when
their mother departed to forage. Killing enemies to reduce competition for food
and to protect offspring would seem to be adaptive pursuits for pumas. The
long-eared owl pillaged from its ground perch and left dead on the sand was
killed by a cub that may have simply found the bird to be an irresistible stimu-
lus to the hunt.
Prey items in puma fecal samples and stomachs we collected from the San
Andres Mountains complemented the cadaver tallies (Table 16-2). Elmer’s
(1997) analysis of 832 fecal samples and four stomachs had a more accurate rep-
resentation of puma diet because it detected a greater variety of small prey. Nat-
urally, field searches for puma-killed prey bias against small prey items because
they are often completely consumed by pumas; hence, evidence of their demise
is rarely if ever detected. Desert mule deer was the most important prey (85.6
percent), while other ungulates, including pronghorn, desert bighorn, oryx, and
javelina, each comprised less than 1 percent. Rodents were the second most
important class of food (5.5 percent), with porcupines comprising the bulk (3.2
percent). Jackrabbits and hares (i.e., Leporids) came in third (4.2 percent).
Puma hair was in 2 percent of the samples, but it was impossible to distinguish
if hair was ingested during cannibalism or grooming. Perhaps the most curious
prey item, found in one stomach, was the ornate box turtle (Terrapene ornata).
These small land turtles are most vulnerable in the peak of their activity from
mid-July through August as they take advantage of the rainy season in their
quest for food. Vegetation occurred in 20 percent of the sample and was com-
posed principally of grass but included some piñon nuts, juniper twigs, and
other woody plant parts. Although some vegetation was probably ingested acci-
dentally when it adhered to prey tissues, pumas sometimes graze sparingly on
green grass for reasons that are not known. We speculate the grass may act as an
emetic to help expel hair or as roughage that hastens expulsion of gut parasites.
Again, complementing the results of our count of prey cadavers, no remains of
livestock were found in the puma feces or stomachs.
We used Elmer’s (1997) puma diet results to estimate the relative percentage
that each prey species contributed to the total biomass consumed and the rela-
tive number of prey animals killed by pumas on the San Andres Mountains.
This helped to illuminate the importance of each prey type. To accomplish this,
we applied a correction factor developed by Ackerman (1982) in his research on
the diet and energetics of pumas in south-central Utah. While conducting feed-
ing trials on captive pumas, Ackerman found there was a positive relationship
between the number of field-collectible feces and the consumption of small prey
[i.e., feces / kg = 1.98 + 0.035 ¥ estimated prey weight (kg)]. This means that
per unit mass of the prey animal, larger prey would result in fewer collectible
feces than would small prey animals. This phenomenon occurs because large
prey are composed of proportionately more highly digestible protein-rich flesh,
Table 16-2. Prey items identified by hair and bone

remains in 832 puma fecal samples and four
stomachs collected from the San Andres Mountains,
New Mexico, during 1985–1995 (adapted from
Elmer 1997).
Species Frequency Percent frequency
Desert mule deer 716 85.6
Rodents 46 5.5
Leporids 35 4.2
Puma 17 2.0
Badger 16 1.9
Unidentified birds 6 0.7
Pronghorn 6 0.7
Skunk 6 0.7
Desert bighorn sheep 5 0.6
Oryx 3 0.4
Box turtle 1 0.1
Coyote 1 0.1
Javelina 1 0.1
Ringtail 1 0.1
Vegetation 167 20.0
but small prey are composed of relatively larger proportions of bones, hide, and
hair, which are more likely to pass through the digestive tract (Floyd et al. 1978).
Hence, Ackerman’s correction factor would give a more-accurate estimate of the
relative biomass that each prey species contributed to puma diet. We used only
those prey items that comprised more than 0.5 percent of the diet by frequency
of occurrence and applied the correction factor to those prey that weighed more
than 1 kg; for smaller prey, we used their estimated average weight (Ackerman
1982). Desert mule deer comprised 92.4 percent of the biomass consumed by
pumas (Table 16-3).
Cadavers, fecal and stomach analyses, and relative biomass and number of
each prey species consumed clearly indicated that energetic and nutritional
demands of pumas on the San Andres Mountains were strongly linked to deer.
Other ungulates were relatively minor contributors. Low availability and vul-
nerability of desert bighorn sheep, pronghorn, and oryx calves rendered these
animals as sporadic victims of puma opportunism and, consequently, of rela-
tively little importance to the overall nutritional and energy needs of pumas.
Table 16-3. Puma prey utilization based on 832 puma fecal samples and four
stomachs collected from the San Andres Mountains, New Mexico, during
1985–1995.
Correction Relative Relative number
Percent Estimated factor biomass individuals
frequency weight (kg)b (kg/feces)c consumed (%)d consumed (%)e
Preya (A) (B) (C) (D) (E)
Mule deer 85.6 44.0 3.52 92.4 41.4
Leporids 4.2 1.3 2.03 2.6 39.4
Porcupine 3.2 5.0 2.16 2.1 8.3
Badger 1.9 7.0 2.22 1.3 3.8
Squirrels 1.3 0.3 0.30 0.1 6.5
Pronghorn 0.7 40.0 3.38 0.7 0.4
Desert bighorn 0.6 53.0 3.84 0.7 0.2
aPrey that comprised more than 0.5 percent of diet by frequency of occurrence in Elmer (1997).
bEstimated weights: Mule deer—assumed 27 percent were fawns that averaged 15 kg and 73 percent
were one year or older that averaged 55 kg (masses of fawns from Anderson 1981; masses of adults
from deer captured on San Andres Mountains in this study). Leporids—assumed 50 percent were jack
rabbits and 50 percent were cottontail. Squirrels—assumed composition of 33.3 percent each of rock
squirrels, white-tailed antelope squirrels, and New Mexico ground squirrels. Masses of leporids, porcu-
pines, badgers, and ground squirrels from the Southwest Biological Museum, Albuquerque,
New Mexico. Pronghorn masses from White Sands Missile Range records. Desert bighorn sheep—
assumed 20 percent were lambs that averaged 29 kg and 80 percent were one year or older that aver-
aged 59 kg (masses from Krausman et al. 1999).
cC = 1.98 + 0.035 ¥ B (from Ackerman 1982:19).
dD = (A ¥ C) / S (A ¥ C) ¥ 100.
eE = (D 4 B) / S (D 4 B) ¥ 100.
Small prey items, rabbits and hares in particular, were probably seized oppor-
tunistically to help pumas endure periods between deer kills. Possibly young
pumas, including large cubs and independent subadults, had better success at
killing small prey relative to killing ungulates and thus may have been responsi-
ble for the bulk of those prey in feces, but we have no supporting empirical data.
We documented sixteen cases of pumas scavenging twelve mule deer, one
desert bighorn sheep, and one oryx that died of nonpredation causes. Two deer
were each used by two pumas at different times. The ungulates were scavenged
from one to twenty-nine days after death. Pumas scavenged about 9 percent of
the total number of ungulates that we found dead from nonpredation causes
(i.e., deer, sheep, oryx, and javelina that died natural deaths plus accidental
deaths resulting from unintended captures in our snares; Logan et al. 1999).
Scavenging was done by thirteen male and three female pumas. All of the males
were territorial adults. One of the female pumas was a subadult and two were
adults that were not raising cubs. In Alberta, biologists found four carcasses of
adult female moose scavenged by three adult female pumas and one adult male
puma (Ross and Jalkotzy 1996). The motivation for an individual puma to scav-
enge a particular carcass may depend upon immediate circumstances, such as
risk of harm to self or offspring from other dangerous predators (e.g., pumas,
wolves, bears, coyotes), degree of hunger, and ease of killing live prey.
Sometimes when a puma came to scavenge, cadaver tissues were fresh, such
as if the animal had been dead for less than one day. But sometimes the tissues
were putrid and laden with maggots and flies. Consumption of scavenged ani-
mals ranged from as little as 1–2 kg to almost complete consumption of edible
parts. The pumas sometimes handled the carcasses just like prey they killed—
dragging them to concealing vegetation, consuming vital organs first (i.e., heart,
lungs, liver), eviscerating stomachs and intestines, covering remains with ground
debris between meals, and scraping (males only) around feeding sites. These
handling characteristics probably increased the number of ungulate carcasses
that we included in the “probable puma kill” category. Scavenging by pumas on
wild prey is an oddity in the literature (Anderson 1983), thus contributing to
the conventional wisdom that pumas almost exclusively eat prey that they have
killed. We certainly found a higher incidence of puma scavengers than any other
studies, but we cannot reliably claim that this behavior occurred more frequently
in the desert than in other environments. Our field policy of periodically check-
ing on the status of carcasses (to be able to better characterize decomposition)
while en route to other field activities may have simply enabled us to observe
scavenging more often than did other biologists.
Clearly, pumas on the San Andres Mountains were highly dependent upon
mule deer for food. As we studied effects of puma predation on the mule deer
population, we learned that the dynamics of the puma and deer populations in
this relatively simple desert ecosystem were tightly linked.
LP > > KX
1. Pumas are opportunistic predators, capable of killing prey in a wide range
of sizes, from rabbits to moose. Prey size and competition with other pred-
ators may have influenced the evolution of puma size on a latitudinal gra-
dient (i.e., puma size increases with latitude).
2. Desert mule deer were the most important prey to pumas on the San
Andres Mountains, comprising about 91 percent of puma-killed animals we
found, about 86 percent of the diet as measured by frequency of occurrence

in feces and stomachs, and about 92 percent of total biomass consumed.
Other important prey included leporids and rodents. Other ungulates,
including desert bighorn sheep, pronghorn, and the exotic oryx, comprised
a minuscule amount of the puma’s diet.
3. Pumas, particularly males, scavenged about 9 percent of cadavers of animals
that died of nonpredation causes. Pumas handled scavenged carcasses as
they would kills.
Chapter 17
Pumas and Desert Mule Deer
Mule deer were the most numerous large prey animals for pumas on the San
Andres Mountains. Deer inhabited the mountains because food and water were
relatively abundant there. In contrast, deer were rare or locally nonexistent in the
comparatively desolate basins adjacent to the mountains. Not surprisingly,
pumas were distributed principally on the mountains where prey were most
abundant and vulnerable (recall Fig. 4-1). In that habitat, ample structural
cover—furnished by shrubs, trees, tall grass, broken terrain, rock outcrops, and
boulders—rendered advantages to the solitary, stalking, and ambush hunting
style of the puma (Logan and Irwin 1985, Laing and Lindzey 1991).
Hypotheses, Predictions, and Terms

Our research hypothesis was that puma predation was the strongest proximate
limiting factor affecting deer population growth rates. A limiting factor is any
factor that causes changes in production or loss of animals and hence con-
tributes to population limitation (i.e., the upper limit that a population can
reach in an environment; Caughley and Sinclair 1994:114). Factors such as pre-
dation, disease, and food supply are all limiting factors and can all operate on a
population simultaneously. The distinction comes in determining which of
these limiting factors most likely sets the upper limit of the prey population.
If puma predation is the primary proximate limiting factor for the deer pop-
ulation, then two predictions must hold: (1) puma predation must be the most
important proximate cause of mortality that affects deer population growth
rates, and (2) experimental removal of pumas from the Treatment Area should
311
be associated with an increase in deer survival rates, which are linked to a reduc-
tion in puma predation rates.
We tested these predictions by quantifying puma predation behavior (i.e.,
diet, predation rates) and measuring the relationships of puma density and pre-
dation rates to deer population parameters. Puma predation rates on radio-col-
lared deer were particularly informative because they came the closest to meas-
uring the total response of puma predation, which is the product of the numerical
response (i.e., changes in predator numbers with prey density) and the functional
response (i.e., variation in the number of prey consumed per predator as prey
density changes) (Holling 1959, Pech et al. 1992, Caughley and Sinclair 1994,
Messier 1995). This was our most direct way of estimating the proportion of the
breeding-age deer killed by pumas (Marshall and Boutin 1999).
Finally, we tested the self-limiting hypothesis developed by Seidensticker et
al. (1973), which postulates that pumas evolved a social system to limit their
population so that they do not overexploit their prey. The drought that struck
the San Andres Mountains during our research provided us with a natural exper-
imental setting in which to test this hypothesis. If the pumas fit the hypothesis,
we would expect them to respond to a severe deer population decline by exhibit-
ing slowed population growth or a decline in density to the extent that they did
not hasten the deer population decline.
Characteristics of Dead Deer

We found 540 dead desert mule deer during 1985–1995. Pumas killed 89 per-
cent (n = 479) while 11 percent (n = 61) died of other causes. Fawns comprised
22 percent (n = 107) of the total number of deer killed by pumas; 71 percent of
those were six months or younger (Table 17-1). When we excluded the radio-
collared deer from the total (n = 77 during 1986–1995) and included only those
deer killed by pumas that we found by chance during our field work (n = 402),
then fawns comprised 26.6 percent of the sample of deer killed by pumas.
Because pumas almost completely consumed fawns, we suspect that fawns were
somewhat underrepresented in the kill sample. Also, because fawns were almost
completely devoured, we could not ascertain the gender of sixty-seven fawns. Of
those we could identify, there were fifteen bucks and twenty-five does. The ratio
was not different from a hypothetical 1:1 sex ratio at birth [1]. The sample of
sixty-one deer that died of causes not related to puma predation (Table 17-1)
was too small to make any meaningful comparisons with puma-killed deer. Fur-
thermore, because we documented puma kills throughout the year and the sex
and age population composition counts of deer were limited to winter or fall,
we could not reliably test if pumas selected among sex and age groups of deer.
Slide @318%
PHOTO 27. A mule deer buck killed and eaten by a puma.
Nonetheless, it was clear that puma predation was the most important single
proximate cause of death in desert mule deer on the San Andres Mountains,
which supported our first prediction.
Fates of Radio-Collared Deer

Fates of radio-collared deer further emphasized the importance of puma preda-
tion as a proximate limiting factor. We monitored 175 radio-collared deer
(ninety-one bucks, eighty-four does) that were one year or older for 153,137
deer-days (74,067 buck-days, 79,070 doe-days) during biological years
1987–1988 to 1993–1994 to quantify annual survival and agent-specific mor-
tality rates. Each year we monitored twenty-eight to fifty-three bucks and
twenty-seven to fifty does. Because of the ruggedness of the mountains, we
sometimes lost radio contact with deer for days or weeks at a time. So once per
month we extended one of our weekly telemetry flights to locate radio-collared
pumas to also recontact “lost” deer. Hence, the number of days that lapsed after
a deer’s death until we could examine the carcass varied widely, averaging 19 ±
12 days (range = 3–61, median = 18). Pumas killed about sixty-two of the deer
(35.4 percent), including thirty-five bucks and twenty-seven does. Radio-col-
CHAPTER 17. PUMAS AND DESERT MULE DEER 315
lared pumas were implicated in at least thirty-nine (63 percent) of those deaths.
Twenty-nine (16.6 percent) other radio-collared deer, including eighteen bucks
and eleven does, died of causes that showed no evidence of puma predation. We
lumped all such deaths into a category called “other causes,” because we fre-
quently could not ascertain specific agents of death. Still, we could tell that none
of the deer was likely killed by pumas. In most cases, we could not determine
the proximate causes of death due to advanced tissue autolysis, but coyotes
apparently killed four of the deer (i.e., 13.8 percent of deaths from “other
causes,” 4.4 percent of all deaths). Radio-collared bucks that died averaged
6.1 ± 2.3 years of age, which was very similar to the average of 6.7 ± 2.1 years
for deceased does [2]. Because we did not sample the age distribution of the liv-
ing deer one or more years old, we could not interpret how the age distribution
of dead deer compared with survivors.
We estimated survival rates and agent-specific mortality rates for radio-col-
lared deer using Micromort software (Heisey and Fuller 1985a), the same tech-
nique that we used to compute survival rates for radio-collared pumas. Survival
rates of bucks and does one or more years old fluctuated relatively little during
biological years 1987–1988 to 1991–1992. During those years survival rates
were relatively high. Buck survival rates ranged from about 0.81 to 0.90 and doe
survival rates ranged from about 0.83 to 0.92. But survival rates for both sexes
declined substantially during the final two full years that we quantified deer sur-
vival rates (Table 17-2); survival rates for bucks and does were about 0.62 and
0.75, respectively. The years 1992–1993 to 1993–1994 included two of the
three years of severe growing-season drought on the San Andres Mountains.
During the seven-year span, bucks tended to have lower survival rates than does,
but only rates in 1990–1991 were significantly different (Table 17-2) [3]. We
projected deer survival rates for 1994–1995, even though our research ended in
March 1995, to see if the declining trend continued. We did this letting Micro -
mort compute the daily survival rates for radio-collared bucks and does (n = 14
each sex) that we monitored during the 238 days in the interval and assumed
that those rates would hold constant throughout the year (i.e., bucks =
0.99825022365, does = 0.99926552365). The resulting survival rates for bucks
(0.53) and does (0.76) suggested that survival rates for deer one or more years
old would stay low during the third year of drought.
Puma predation rates on radio-collared bucks and does fluctuated during the
seven-year span (Table 17-3). During the first five years, predation rates on
bucks ranged from about 0.06 to 0.19 and predation rates on does ranged from
about 0.04 to 0.12. Then, during the drought years, puma predation rates on
the deer increased substantially to about 0.28–0.30 on bucks and 0.23–0.25 for
does. Bucks and does were about equally susceptible to puma predation during
Table 17-2. Survival rates of radio-collared desert mule deer, San Andres
Mountains, New Mexico, 1987–1995.a
Bucks Does
Biological Yearb Deer-days Rate 95% C.L. Deer-days Rate 95% C.L.
1987–88 6748 0.90 0.77–1.0 9014 0.89 0.77–1.0
1988–89 10434 0.81 0.69–0.96 11483 0.88 0.78–1.0
1989–90 10199 0.81 0.68–0.96 14075 0.83 0.73–0.95
1990–91 13626 0.81c 0.70–0.94 12879 0.92c 0.83–1.0
1991–92 14103 0.86 0.76–0.97 12901 0.84 0.74–0.97
1992–93 11326 0.62 0.48–0.79 11243 0.75 0.62–0.90
1993–94 7631 0.62 0.46–0.83 7475 0.75 0.59–0.94
1994–95d 2286 0.53 2723 0.76
aSurvival rates and 95 percent C.L. (Confidence Limits) were computed using Micromort software
(Heisey and Fuller 1985a,b).

bBiological years spanned August–July each year. In years 1987–1988 to 1993–1994, twenty-eight to
fifty-three bucks and twenty-seven to fifty does were monitored.

cSurvival rates for bucks and does were significantly different (Z = 1.476, P = 0.07).
dThe 1994–1995 year extended only from August to March due to termination of the study. In that
period, fourteen bucks and fourteen does were monitored. Annual survival rates for bucks and does
were projected from daily survival rates computed for the 238 days the deer were monitored in the
interval; in other words, bucks = 0.99825022365 ; does = 0.99926552365.
the seven-year span; only in 1990–1991 were puma predation rates significantly
different [4]. There appeared to be a strong interaction between growing-season
(July through September) droughts and puma predation rates on deer one or
more years old. Again, we asked ourselves if the trend of increased puma preda-
tion rates would hold during 1994–1995. We used the daily survival rates and
daily puma predation rates (i.e., bucks = 0.00174978, does = 0.00073448) com-
puted using the program Micromort to estimate puma predation rates for the
entire interval (see Heisey and Fuller 1985a and Table 17-3). Results suggested
that high puma predation rates on bucks (0.47) and does (0.24) would continue
with the drought.
Death rate from “other causes” also varied for both sexes (Table 17-3). Rates
for bucks ranged from 0 to 0.13, while rates for does ranged from 0 to 0.08.
During the seven-year span, bucks appeared to be somewhat more susceptible to
“other causes” of mortality than does [5]. Rates of death from “other causes” did
not increase during the drought years, unlike puma predation rates. To the con-
trary, rates tended to decline for both sexes. In the last year of our study,
1994–1995, none of the radio-collared deer died of “other causes.”
FIGURE 17-1. Rates of survival, puma predation, and death from other causes for radio-col-
lared mule deer, San Andres Mountains, New Mexico.
Putting all these rates together—survival, puma predation, and death

f rom “other causes”—suggests that puma predation and “other causes” of
m o rtality trade off in re l a t i ve importance to survival of deer one or more
years old (Fig. 17-1). An inverse relationship emerged between rates of puma
p redation and death from “other causes,” which was stronger for does than
for bucks [6]. No t a b l y, during the drought years, puma predation on bucks
and does increased mark e d l y, while deaths from “other causes” waned. This
suggested that puma predation on deer was partially compensatory; that is,
puma predation substituted for “other causes” of mortality in some instances.
We would expect the degree of compensatory mortality via puma pre d a t i o n
to increase during drought years because more deer would likely suffer fro m
malnutrition.
Our data on condition of deer killed by pumas was too limited to test this
idea; nevertheless, it showed that puma predation is partially compensatory.

During the study, we were able to examine femur marrow in 106 deer one or
more years old (fifty-six bucks, forty-six does, four undetermined sex) that were
killed by pumas 0.5–16 days ( x– = 5.1 ± 4.1, median = 4.0) before we inspected
their remains. This sample comprised 28 percent of the total number of deer
(n = 372) one or more years old that we found killed by pumas during our study.
Of the sample, seventy-four had marrow that was white and pink in color, with
a firm consistency like candlestick wax. These characteristics suggested that the
deer were probably not metabolizing fat in the long bones to meet energy
demands, and thus they could have been in good nutritional condition. Another
thirty-two deer (sixteen bucks, thirteen does, three undetermined sex) had mar-
row that was colored red, pink, yellow, amber, or orange and had the consistency
of gelatin. Characteristics of the femur marrow in this latter group indicated that
fat depletion had reached extreme levels in the long bones, the last fat reserves
available for energy metabolism. This condition suggested the animals were in
extremely poor condition and were probably dying (Mech and DelGuidice
1985). Hence, it appeared that at least 30 percent of the puma predation on this
sample of 106 deer one or more years old was compensatory mortality.
Still, survival rates appeared to respond to reduction in puma numbers on
the Treatment Area, suggesting that some puma predation was also partially
additive. An increase in buck survival in 1991–1992 corresponded with the low-
est number of pumas and is correlated with a reduction in puma predation. Sim-
ilarly, an increase in doe survival in 1990–1991 coincided with the removal of
pumas and a decline in puma predation. These dynamics satisfied our second
prediction—that removal of pumas should result in increased deer survival. It
appeared to us, as we carefully inspected the carcasses of many of the deer, that
some were in sound health prior to being killed by a puma. Pumas are large
powerful predators, perfectly capable of bringing down even the healthiest deer
if they can position themselves close to the prey for the critical final attack.
Pumas can accomplish this with an undetected stalk. Or deer, regardless of their
state of health, may place themselves in mortal danger simply by moving to a
puma waiting to ambush.
Another way we examined the effect of puma predation on deer was by pool-
ing the radio-collared bucks and does together to compute deer survival rates
and agent-specific mortality rates (Table 17-4). In doing this, we were treating
all the radio-collared deer as sentinels to puma predation (i.e., major prey items
susceptible to puma predation). We postulated that deer survival rates were
strongly related to puma predation. A linear model was unequivocal; about 90
percent of the variation in deer survival rates was explained by puma predation
rates [7]. This would further support our first prediction, because we would
Table 17-4. Survival rates and agent-specific mortality rates of radio-collared

deer (bucks and does pooled), San Andres Mountains, New Mexico,
1987–1994.a
Rates
Biological 95% Puma 95% Other 95%
Year Survival C.L. predation C.L. causes C.L.
1987–88 0.89 0.80–0.99 0.07 0.0–0.14 0.04 0.0–0.10
1988–89 0.85 0.76–0.94 0.09 0.02–0.16 0.06 0.0–0.12
1989–90 0.82 0.74–0.91 0.08 0.02–0.14 0.10 0.03–0.16
1990–91 0.86 0.79–0.94 0.12 0.04–0.19 0.03 0.0–0.06
1991–92 0.85 0.78–0.93 0.06 0.01–0.12 0.09 0.03–0.15
1992–93 0.68 0.58–0.79 0.25 0.16–0.35 0.07 0.01–0.12
1993–94 0.68 0.56–0.82 0.28 0.16–0.40 0.04 0.0–0.09
aSurvivalrates and agent-specific mortality rates with 95 percent C.L. (Confidence Limits) were com-
puted using Micromort software (Heisey and Fuller 1985b).
expect variation in survival rates of breeding-age deer to relate to deer popula-

tion growth rates.
Even if puma predation is the primary proximate limiting factor, a deer pop-
ulation can still grow if reproduction outpaces mortality. In fact, this phenome-
non has been documented. In the central Idaho wilderness, mule deer and elk
increased in numbers even though they were primary prey (70 percent aggregate
frequency in feces) for a puma population similar in density to the puma popu-
lation on the San Andres Mountains (Hornocker 1970, Seidensticker et al.
1973). As we have mentioned previously, the Idaho biologists hypothesized that
pumas in that environment evolved a social system that limited their densities
below a level set by absolute numbers of deer and elk (Seidensticker et al. 1973).
A population of mule deer in southern Utah increased even though it was the
mainstay (81 percent of biomass consumed) for a low-density puma population
that was about one-third the density of the Idaho population (Ackerman 1982,
Lindzey et al. 1994). Biologists there explained that the pumas may have been
behaving in accordance with the hypothesis offered by the Idaho researchers, or
that the deer population did not increase enough to cause a numerical response
in the puma population (Lindzey et al. 1994). And in the Northern Yellowstone
Ecosystem, where a puma population was increasing, puma “predation was nei-
ther a major source of mortality nor a significant factor limiting the numbers or
growth rates of elk and mule deer populations.” There, pumas “removed about
2–3% of elk and 3–5% of mule deer each year” (Murphy et al. 1998b:27). Biol-
ogists there explained that open, nonforested habitats used by elk and deer, and
migration of deer away from puma habitats in winter, provided those ungulates
with spatial and temporal refuge from puma predation (Murphy et al. 1998b).
Our next aim was to assess the influence of puma predation on deer population
growth on the San Andres Mountains.
Mule Deer Population Dynamics

We modeled the deer population so we could associate changes in deer popula-
tion parameters with puma population size and predation rates. The model was
a relatively simple deterministic, discrete time model that simulated the dynam-
ics of the deer population on the Treatment Area, where most data-gathering
procedures for deer population parameters were focused. By deterministic, we
mean that population parameters put into a mathematical equation determine
the estimated change in the deer population. And by discrete time, we mean the
model is applied in annual units (i.e., the biological year, August through July)
beginning with the deer birth pulse. This approach did not require us to know
what the deer population size actually was for the purpose of estimating deer
population dynamics during the study. Instead, we were able to calculate the
finite rate of increase each year (l = Nt + 1 / Nt), where l > 1 means the deer
population increased, and l < 1 means the population declined. Then by mul-
tiplying the annual ls to the projected deer population size the previous year,
starting with one thousand deer in the first year, we could simulate the trajec-
tory of the deer population.
The basic model was
Nt + 1 = B (sb ) + D (sd ) + [AD (fa ) + YD (fy ) - Fd ]
Nt = the base population of 1,000 deer one year or older at the beginning
of the biological year (August through July).
Nt + 1 = deer population size at the end of the biological year. It reflects the
change in the 1,000 deer base population (i.e., 1,000 + births – deaths).
B = estimated number of bucks (one year or older) in the base population.
D = estimated number of does (one year or older) in the base population.
AD = the number of does two years or older.
YD = the number of yearling does.
fa = fetal rate of does two years or older.
fy = fetal rate for yearling does.
sb = survival rate for bucks.
sd = survival rate for does.

Fd = the number of fawns that died.
Although most of the model parameters were gathered in surveys conducted

on the San Andres Mountains (Table 17-5), we had to rely on some estimates
from other desert mule deer populations that were studied by the New Mexico
Department of Game and Fish in other parts of southern New Mexico. Fur-
thermore, because we did not know how some parameters might vary during
drought conditions, we varied some of the parameters ourselves. Consequently,
we developed six different population simulations, or models (i.e., six models
numbered 1 to 6, Table 17-6), to simulate the dynamics of the deer population
on the Treatment Area. Model parameters, their origins and variations, and our
assumptions are described in detail in Appendix 4.
Results from the deer population model were parsimonious with our per-
ceptions of deer population dynamics through direct observation. All six mod-
els indicated that the deer population increased during the first five years
(1987–1988 to 1991–1992), relatively rapidly at first, then gradually. But dur-
ing the last three years (1992–1993 to 1994–1995) the deer population declined
rapidly (Table 17-7). Models 1 and 3 and models 5 and 6 produced the same
estimates of l each year when l was rounded off to three significant figures; this
indicated that the slight decline in yearling doe fetal rate (0.9–0.81) had virtu-
ally no effect on the model simulations. Consequently, we decided to depict the
range of deer population dynamics by using simulations from models 1, 2, 4,
and 6 (Fig. 17-2). Not surprisingly, the two parameters that affected the dynam-
ics of simulations the most were fawn production and survival rates of deer one
or more years old. If the number of surviving fawns exceeded the number of
deaths in deer one or more years old, then the population increased; if it did not,
the population declined. By extension then, the role that pumas played in these
dynamics clearly depended upon predation rates on deer.
Puma Predation and Mule Deer Population Growth

How did puma predation impact growth patterns of the deer population? Con-
sidering that deer population trends and growth rates were quite similar among
models 1, 2, 4, and 6, we chose the two models representing the most pes-
simistic (model 2) and optimistic (model 6) simulations of the deer population
to test for the influence of puma predation. Our premise—puma predation sup-
pressed deer population growth—was tested by regressing annual deer popula-
tion growth rates of models 2 and 6 (i.e., l, Table 17-7) on corresponding
annual puma predation rates on radio-collared deer (Table 17-4). Recall that we
Table 17-7. Finite rates of increase (l) estimated by modeling the desert mule
deer population on the San Andres Mountains, New Mexico, 1987–1995.
Biological Model 1 Model 2 Model 3 Model 4 Model 5 Model 6
Year la l la l lb lb
1987–88 1.13 1.13 1.13 1.17 1.17 1.17
1988–89 1.18 1.18 1.18 1.18 1.18 1.18
1989–90 1.05 1.05 1.05 1.05 1.05 1.05
1990–91 1.06 1.06 1.06 1.06 1.06 1.06
1991–92 1.06 1.06 1.06 1.06 1.06 1.06
1992–93 0.91 0.84 0.91 0.84 0.91 0.91
1993–94 0.86 0.77 0.86 0.77 0.86 0.86
1994–95 0.70 0.70 0.70 0.70 0.70 0.70
aModels 1 and 3 produced the same l each year when l was rounded off to three significant figures.
bModels 5 and 6 produced the same l each year when l was rounded off to three significant figures.
FIGURE 17-2. Desert mule deer population trend on the San Andres Mountains, New
Mexico.
FIGURE 17-3. Relationship of deer population growth rates (lambda) to puma predation
rates on radio-collared deer (bucks and does combined), San Andres Mountains, New
Mexico.
considered puma predation rates on radio-collared deer to represent the total

response of puma predation (i.e., the product of the numerical response and the
functional response). In this specific case, it was the total response on the repro-
ducing segment of the deer population. The test suggested that puma predation
rates had a negative relationship to deer population trends and explained 87 per-
cent and 79 percent of the variation in deer population growth rates for models
2 and 6, respectively [8] (Fig. 17-3). This meant that puma predation suppressed
deer population growth, thus satisfying our first prediction. The linear model
[8] suggested that the mule deer population increased when puma predation
rates were less than 0.14 and less than 0.17 for deer models 2 and 6, respectively.
Another way of looking at conditions favorable for deer population increase was
by a univariate model that used deer survival rates to account for all mortality
factors in aggregate. Our model suggested that the deer population grew when
deer survival rates exceeded 0.80 and 0.77 for deer models 2 and 6, respectively
[9]. Deer death rates from “other causes” were uninformative in explaining deer
population growth rates [10].
Deer population growth was also influenced by fawn production. A linear
model showed a strong positive relationship between ls of deer models 2 and 6
and corresponding fawn:doe ratios [11] (Fig. 17-4). This explained 80 percent
and 92 percent of the variation in deer population growth rates generated by

models 2 and 6, respectively. Furthermore, this model [11] predicted, given the
environmental conditions during our re s e a rch, that the deer population increased
when winter fawn counts exceeded about thirty-nine and thirty-seven fawns per
100 does for deer models 2 and 6, respectively. Below those ratios, the deer pop-
ulation declined.
Consequently, two key proximate factors strongly influenced growth rates of
the desert mule deer population: puma predation and fawn production. A mul-
tiple regression model using puma predation rate and fawns per 100 does as
independent variables suggests that those two factors were strong determinants
of deer population growth rates, explaining 94 percent of the variation in ls
from deer models 2 and 6 [12]. All this made sense because puma predation
directly removed animals from the deer population while fawn production
added to it. To identify the independent variable with the greatest incremental
predictive power, we sequentially examined partial correlation coefficients. The
partial correlation coefficient measures the strength of the relationship between
the dependent variable (i.e., in this case l) and a single predictor variables (e.g.,
puma predation rate) when the effects of other predictor variables (e.g., fawns /
100 does) in the model are held constant (Hair et al. 1995). Puma predation had
a stronger influence than fawn production on deer population growth rates gen-
erated by deer model 2, but for model 6, fawn production was slightly more
FIGURE 17-4. Relationship of deer population growth rates (lambda) to fawn production,
important than puma predation [12]. If puma predation also altered the num-
ber of fawns that survived, then puma predation was the strongest proximate
limiting factor affecting deer population growth rates.
Other than the proportion of fawns in the puma-kill sample (i.e., about 27
percent), we had no direct empirical data on kill rate of fawns. So, we assessed
this indirectly. We postulated that puma predation altered fawn numbers and
tested this with a measure of association between fawn counts and puma num-
bers. The test indicated a negative correlation between the ratio of fawns per 100
does counted in winter deer population surveys (Table 17-5) and adult puma
density in January in the Treatment Area (see Chapter 10, Table 10-2) [13]. The
sample of puma-killed deer had already established that pumas were eating
fawns, and now we had an indirect deduction that fawn numbers were substan-
tially altered by puma predation. Moreover, there was a negative correlation
between the ratio of fawns per 100 does and puma predation rates on radio-col-
lared deer one or more years old [14]. In other words, more fawns in the popu-
lation meant that puma predation rates on older deer would diminish; fewer
fawns meant greater predation pressure on older deer. This was a mechanism
that explained why puma predation rates in radio-collared deer did not exhibit
an increasing trend during the first puma population increase phase in the Treat-
ment Area from 1987–1988 to 1989–1990 (Fig. 17-5).
This all made sense in light of other puma research demonstrating the
FPO @ 75%
FIGURE 17-5. Relationship of adult puma numbers to predation rates on radio-collared deer
(bucks and does combined), San Andres Mountains, New Mexico.
importance of deer fawns and other juvenile hoofed prey to pumas. Biologists
in the central Idaho wilderness (Hornocker 1970) and southern Utah (Hemker
1982) found that pumas killed fawns in greater proportions than their occur-
rence within the population. In addition, female pumas selected fawns on the
east slope of California’s Sierra Nevada. There, puma mothers (the segment of
the population with the greatest energy demands, Ackerman 1982) selected
fawns less than four months old (Pierce et al. 2000a). A preference for fawns
should not be surprising; on average, they are probably considerably easier to
stalk and kill than adult deer. Probably for the same reason, pumas also selected
elk calves in central Idaho (Hornocker 1970). In Yellowstone National Park,
pumas selected elk calves above all other classes of prey (Murphy et al. 1998b).
Similarly, along the east slope of the Rocky Mountains in southwestern Alberta,
pumas selected calf moose over yearlings and adults, and male pumas appeared
to prefer moose calves even to other species of prey, including mule deer, elk,
bighorn sheep, and white-tailed deer (Ross and Jalkotzy 1996). Pumas relying
on feral horses as fodder for part of the year in California and Nevada selected
foals almost exclusively (Turner et al. 1992).
From this we inferred what seemed obvious: fawns were critical for puma
food and deer population growth. When fawn production was high, the fawn
segment of the population probably absorbed a critical level of the puma preda-
tion, in essence diverting predation pressure away from the older, reproducing
segment of the deer population. And there were enough fawns left over to out-
pace mortality in the deer population as a whole. Thus, the deer population
grew. However, during the drought years, fewer fawns were produced and sur-
vived, probably because of a combination of lower birth rates, viability, and pre-
dation. Hence, during the drought, the brunt of puma predation fell on older
deer.
Productivity and survival of mule deer fawns has been shown to be strongly
affected by habitat conditions. Mule deer does on “severely depleted” summer
range in central Utah were found to have lower ovulation rates, lower fetal rates,
and fawns with lower masses than does on “good summer range” in southern
Idaho (Julander et al. 1961). On a desert range in central Arizona, researchers
found an interaction between fawn survival, forage conditions they linked to
rainfall patterns, and predation from a host of predators, including pumas, coy-
otes, and black bears (Smith and LeCount 1979). In our Chihuahua Desert
study area, where the fawning period coincides with the summer monsoon sea-
son, mule deer does are dependent upon late-summer rains to produce high-
nutrient and high-energy foods they need for lactation. In the closely related
black-tailed deer, does in peak lactation and with single fawns increased their
food intake by 35 percent, while does with twins increased theirs by 70 percent
(Sadleir 1982). Mule deer does also have considerably higher water requirements
during late pregnancy and lactation (Short 1981); they may need to consume
more than 5 liters of free water per day (Hervert and Krausman 1986). Does try
to partially meet these requirements by seeking out habitats with better forage
characteristics and water sources (Hervert and Krausman 1986, Ordway and
Krausman 1986). We also expect the increase in herbaceous ground vegetation
(i.e., grasses, tall forbs) to more effectively hide fawns from predators (Teer et al.
1991). Once fawns are weaned, high-quality foods are needed for maintenance
and growth (Bronson 1989), and to give them the energy, strength, and agility
to better evade predators.
Drought in three consecutive growing seasons on the San Andres Mountains
must have been strenuous on deer. Ecological carrying capacity (i.e., the natural
limit of a population set by resources in the environment, Caughley and Sinclair
1994:117) for deer was low in that period of our research. Drought not only
depleted deer foods, but also caused some perennial springs to go dry. In the first
year of drought, fawns were probably most affected, followed by mothers. Fawns
are more vulnerable than adults to water deprivation because their greater rela-
tive metabolic rate and smaller body size increase their water demands (Short
1981, Hervert and Krausman 1986). Due to the strain of pregnancy and lacta-
tion, we would expect does entering winter in poor condition to have lower fer-
tility rates and to produce smaller, more-vulnerable fawns the following summer.
Lower fertility rates and lower fawn survival probably contributed to the decline
in fawn production in subsequent years. We would expect the intensity of the
effects of weather and food on deer reproduction and survival to depend on the
size of the deer population relative to ecological carrying capacity (Picton 1984).
Mule deer also abandon home ranges where traditional water sources go dry to
search for other water sources (Hervert and Krausman 1986). Not only does this
behavior impose abnormally high energy costs, but we believe it also makes deer
more vulnerable to puma predation as they explore unfamiliar surroundings. In
addition, deer normally concentrated around free water during the hot, dry part
of the year (normally April through June) on our study area. If these effects actu-
ally occurred, increasing the susceptibility of deer to physiological stress, then we
hypothesize that compensatory mortality from puma predation increased dur-
ing the drought. In other words, a larger proportion of the deer that pumas
killed would have died anyway, particularly fawns (see Bartmann et al. 1992,
Singer et al. 1997). In addition, the hot, dry ambient environment and deer
concentrations probably interacted to boost puma predation rates. Deer car-
casses decompose rapidly under those conditions—in about two or three days.
Deer concentrating around water sources could make it easier for pumas to kill
another deer, compressing time spans between kills. Dynamics such as these
could enhance food acquisition for pumas, thus increasing fertility rates and cub
survival, at least in the short term, and thereby contribute to a delay in the
decline of the puma population itself.
Some Behavioral Interactions between Deer and Pumas

Information on how prey respond behaviorally to the presence of pumas has
been limited to Hornocker’s (1970:35) observations in central Idaho. “The mere
presence of a [puma] or family of [pumas] in a locality or watershed does not
appear to alarm game animals. When a kill is made, however, the reaction is
striking. Deer and elk immediately leave the area, cross to the far side, and in
some instances leave to enter a different drainage. This behavior, observed with-
out exception, acts to distribute game animals.” On the San Andres Mountains,
we could not detect if deer abandoned locales because pumas killed prey there.
However, we made several observations indicating that some deer did not move
away from areas where pumas killed prey. As we crept upon a puma family to
count cubs, we disturbed seven deer, some from their day beds, within 100 m
of F45 and her three two-month-old cubs as they ate a buck that F45 killed that
day. Another time, we watched at least eight deer trotting through an area about
100 m from a buck that had been killed by F183 but had been usurped by a
lone coyote. The deer passed within about 170–250 m from the two predators.
In another case, we disturbed twelve deer when they were about 60 m from a
snare site where two pumas consorted. F61 was caught in a snare, and M46
stayed beside her until we approached within 50 m. F61 had killed two ten-year-
old does at the spring within a twenty-four-hour span and had cached the car-
casses about 27 m apart. One deer was cached under juniper trees where we
snared F61 and the other lay partially covered with leaves and sticks (pawed over
it by the puma) at the edge of a small spring-fed pool of water. The male puma
left the area while we chemically immobilized the female to tag her. After we fin-
ished, we left the still-groggy puma to recover in shade of juniper trees about 25
m up slope from the water. At this time the twelve deer we had disturbed earlier
walked down to the pool in single file to drink! This latter instance occurred in
July, near the peak of hot, dry weather, and suggests that water stress aggravates
vulnerability of deer to puma predation.
Observations we made in the summer drought of 1994 exhibited how pumas
exploited prey that came to water sources. In the month of June, female puma
F149 killed at least seven deer within a 60-m radius of perennial Dugout Spring,
located 1.9 airline km from the nursery where she gave birth to three male cubs
on 1 June. These deer included four does that were two to eight years old
(x– = 6 ± 2.7 years) and three young does ten to eleven months old. Consump-
tion of the carcasses varied from about 40 up to 100 percent of edible tissues.
The carcass of one of the short yearlings was abandoned after only three days,
probably because it was putrid. F149 frequently used this nearest water source
to her nursery, driven there by the high water requirements of lactation and the
extreme hot, dry ambient environment. This environment also concentrated
deer around the spring, magnifying their vulnerability to F149. But puma pre-
dation was not the only cause of death at Dugout Spring that June: two other
deer, a four-year-old buck and a seven-year-old doe, apparently lay down and
died 8–11 m from the water. Rapid tissue autolysis prevented a determination
of exact causes of death, but we surmised that the deer deaths were related to
extreme weather conditions and malnutrition. The same summer, territorial
male M36 occasionally hunted around Buckhorn windmill, which trickled
water into a small earthen impoundment. Among dense willow baccharis and
cattails, he killed five ungulates within a span of about fourteen days, including
three mule deer does that were four to six years old ( x– = 4.7 ± 1.2 years), one
fawn buck (probably an offspring of one of the does), and one yearling prong-
horn buck. He also killed a coyote that apparently came to scavenge one of his
kills. M36’s consumption of the carcasses varied from about 10 to 90 percent of
edible tissues. Rapid decomposition of the carcasses in the dry heat and the
concentrated available prey probably contributed to this extraordinarily high
kill rate.
Did Puma Predation Limit the Deer Population?

Puma predation did not—by itself—stop the deer population from increasing
on the San Andres Mountains. Deer model simulations, which we emphasize
were consistent with our field observations, indicated that the deer population
increased in the face of an increasing puma population during biological years
1987–1988 to 1990–1991 (Fig. 17-2). The deer population continued to grow
through 1991–1992 as the puma population on the Treatment Area was sup-
pressed by our experimental removal. Clearly, although predation by pumas was
slowing the rate of increase in the deer population (i.e., it was the major proxi-
mate limiting factor), it was not stopping growth. Thus, in that span of time,
puma predation did not limit the deer population. In the absence of drought,
we would have expected the deer population to continue growing until density-
dependent effects from intraspecies competition for food regulated it or until
the puma population grew to a level to limit it through predation. But in this
relatively unstable desert environment, drought drastically changed the dynam-
ics between pumas and deer. It triggered a threshold response due to a combi-
nation of lower fawn production (i.e., via lower birth and survival rates) and
substantially greater predation on breeding-age deer. From 1992–1993 through

1994–1995, the deer population declined sharply. Puma predation was the sin-
gle most important proximate cause of mortality that accelerated the decline.
However, puma predation cannot be said to have limited the deer population
(i.e., set its upper limit), because the deer population probably would have
declined during the drought even without pumas on the scene. It was the food
supply for the deer that ultimately limited the deer population, while pumas
were the major proximate cause of death driving the deer population downward.
Did Pumas Limit Their Own Density and Not Harm Their
Food Supply?
We concluded that pumas on the San Andres Mountains did not limit their
population below a level set by the prey as Seidensticker et al. (1973) hypothe-
sized for pumas in the Idaho wilderness. Instead, the large-mammal community
suffered destabilizing effects of drought and puma predation. During the deer
decline phase (i.e., 1992–1993 to 1994–1995), puma predation rates were
inversely density dependent (i.e., depensatory), meaning that pumas were killing
a larger proportion of deer as the deer population declined (Caughley and Sin-
clair 1994:15–16, Messier 1995). Furthermore, the puma population was
increasing as the deer population was declining. Thus, the puma population
exhibited a lag—a delayed numerical change in the puma population in
response to the decline of its food base (Fig. 17-6). This type of lag has been
exhibited in other tightly linked predator-prey systems such as wolves and
moose on Isle Royale National Park (McLaren and Peterson 1994) and coyotes,
lynx, and snowshoe hares in southwest Yukon (O’Donoghue et al. 1998). Not
only did puma predation expedite the deer decline on the San Andres Moun-
tains, but also we expected it to deepen and protract the low phase in deer num-
bers, at least until the puma population declined as well. In a variable environ-
ment like the desert, where the periodicity and amplitude of prey declines would
be impossible to anticipate, it is difficult for us to imagine how pumas could
evolve a social system to limit their population density below the level set by the
prey.
Independently, researchers studying pumas and mule deer in Round Valley
on the east slope of the Sierra Nevada in California also concluded that pumas
did not conform to the self-limiting hypothesis (Pierce et al. 2000b). Their
study focused on a relatively small area of about 90 km2 where deer and pumas
concentrated during the winter period extending from November through
April. In winters 1992–1997, the biologists studied the spatial and temporal dis-
tributions of ten radio-collared adult pumas along with the spatial and tempo-
FIGURE 17-6. Mule deer and puma population trends on the San Andres Mountains, New
Mexico.
ral distributions of live radio-collared mule deer and sites where 112 radio-col-
lared deer were killed by pumas. During the study, the estimated deer popula-
tion increased from about 1,344 in 1993 to 1,913 in 1997. Conversely, the
number of adult radio-collared pumas with winter distributions on the area
declined from about six in the winter season 1992–1993 to three in 1996–1997.
The biologists intimated there was a general decline in the puma population in
the area due to deaths of adults, but data on puma population dynamics and
exact mechanisms causing population decline were not stated. Such information
might have explained why the puma population did not increase along with the
deer population as would be expected in theory. Nevertheless, because the biol-
ogists found that the distribution of deer killed by individual pumas could be
explained by the distribution of live deer alone, instead of by the partitioning of
prey via territorial and mutual avoidance behavior (i.e., pumas did not sequester
prey by defending it or excluding other pumas from prey-rich areas), and
because puma distributions overlapped principally in areas where deer were
more abundant, they inferred that pumas are “most likely limited by prey avail-
ability and not territoriality” (Pierce et al. 2000b). They also suggested that the
corresponding seasonal migrations of the deer and pumas from low-elevation
winter ranges to high-elevation summer ranges “decreased the likelihood that
social interactions or a stable land-tenure system could have been regulating the
population” (Pierce et al. 2000b, also see Pierce et al. 1999).
Cases of Pumas and Other Carnivores Limiting Prey

Populations
Evidence that puma predation limits prey populations has accumulated in
recent years and suggests that pumas may not have limited their densities below
a level set by the food supply in other areas. In the Montgomery Pass Wild
Horse Territory along the California and Nevada border, biologists found that
pumas limited the feral horse population by preying on foals, particularly those
less than three months old. Over 80 percent of foal mortality was attributed to
puma predation. Mule deer were also a staple for the pumas. These deer
migrated into the wild horse territory each winter from the Sierra Nevada. Biol-
ogists believed that pumas switched seasonally between the two main prey
species, utilizing mule deer during the winter and foals during May to October
(Turner et al. 1992). This ability to switch to another prey probably magnified
the depressive effect of puma predation on the horse population because the
puma population was probably maintained at a higher level than it would have
achieved if horses were the only food source year-round. A pronghorn popula-
tion in central Arizona was limited by puma predation. The pronghorns were
vulnerable to puma predation because they inhabited rugged, brushy terrain
favored by stalking pumas (see Logan and Irwin 1985). Puma predation
accounted for about 75 percent of deaths to adult doe pronghorns. Moreover,
doe mortality from puma predation alone exceeded effective recruitment of
female progeny (Ockenfels 1994). In the Great Basin Desert of Nevada sits a
small 15-km2 basin where pumas were the main proximate mortality factor that
reduced a porcupine population from more than eighty animals to fewer than
five animals in only three years (i.e., a 94 percent reduction). In that environ-
ment, pumas depended principally upon mule deer for food. However, the mule
deer population declined precipitously over a seven-year period, 1987–1993,
apparently as a result of a persistent drought. It was during the latter two years
of that period that puma predation on porcupines increased dramatically (i.e.,
eleven of fourteen deaths the first year, eleven of sixteen deaths the second year),
driving the population to near extinction (Sweitzer et al. 1997). This case paral-
leled the effect that puma predation had on the small, isolated desert bighorn
sheep population on the San Andres Mountains (see Chapter 18).
Carnivores with which pumas evolved during the Holocene are also capable
of limiting prey populations. Wolves limited moose populations in interior
Alaska (Gasaway et al. 1983), Pukaskwa National Park, Ontario (Bergerud et al.
1983), and Isle Royale National Park Michigan (McLaren and Peterson 1994).
Also, wolf predation caused declines in caribou populations in British Colum-
bia (Bergerud and Elliot 1986, Seip 1992). Coyote predation limited a white-
tailed deer population in Texas (Teer et al. 1991). There is also evidence that
black bear predation limited a moose population in Saskatchewan (Stewart et al.
1985), and together grizzly and wolf predation limited a moose population in
Yukon (Larsen et al. 1989). In the Glacier National Park area of northern Mon-
tana and southern British Columbia, predation by a host of predators, especially
pumas and wolves, limited the white-tailed deer and elk populations (Kunkel
and Pletscher 1999). In that rugged, densely vegetated landscape pumas and
wolves mostly killed similar types of prey in similar spatial and temporal distri-
butions. The predator-caused decline in deer and elk populations, seizing of
puma kills by wolves, and direct killing of pumas by wolves suggested that those
two carnivores engaged in exploitative and interference competition for food
(Kunkel et al. 1999). Particularly in ecosystems where pumas compete with
other carnivores for the same prey, it is difficult for us to conceive that pumas
would evolve a social system to limit their own density below the level set by the
prey unless their competitors do too.
What Limits the Puma Population?

We hypothesize that the puma population on the San Andres Mountains was
ultimately limited by food. The amount of prey actually vulnerable to the stalk-
ing and ambush hunting style of pumas is influenced by habitat characteristics.
Habitat continuity, physiographic and vegetative structure, and food abundance
and distribution all affect what prey species are present and their distribution,
abundance, and vulnerability (Hornocker 1970, Seidensticker et al. 1973,
Logan and Irwin 1985, Laing and Lindzey 1991, Murphy et al. 1998b, this
study). Because of the strong link between pumas and deer, we expected the
puma population to decline after the deer decline. No alternative prey was avail-
able in high-enough biomass to buffer the effects of the deer decline. Unable to
switch to another food source, pumas could not sustain their relatively high
population density. Our research tenure did not witness a puma decline, but
there were portending signs: the adult puma population in the Treatment Area
declined slightly by January 1995, and during the drought years the rate of pop-
ulation increase in the Reference Area declined. One to two years after we left
the San Andres Mountains, two wildlife biologists—Mara Weisenberger (U.S.
Fish and Wildlife Service, San Andres National Wildlife Refuge) and Dave Hol-
derman (U.S. Army White Sands Missile Range, Environmental Services)—
supported our prediction. Their field searches suggested that the puma popula-
tion declined. Locales that traditionally held puma scrapes, or where puma
tracks were predictably found, were now barren. We speculate there was a four-
year span from the first biological year of the deer population decline
(1992–1993) to the first year that the puma population apparently declined
(1995–1996).
If puma density is ultimately limited by food, then we would expect a sig-
nificant correlation between puma density or biomass and vulnerable prey bio-
mass in areas where pumas and prey are protected from population-altering
human exploitation. So far, such data on pumas are not available to test this
hypothesis. However, support for this premise exists in literature on other large
carnivores. The ecological equivalent of the puma in the Eastern Hemisphere is
the leopard. Leopards and pumas are similar in body size and behavior, and they
use a similar wide range of habitats and prey (see Bailey 1993). While analyzing
data from African leopard populations in eleven sub-Saharan conservation areas,
Stander et al. (1997) found a significant correlation between leopard biomass
and available biomass of prey weighing 15–60 kg (r 2 = 0.72, P < 0.01). Leop-
ard densities varied widely from 0.5 to 24 cats per 100 km2. The highest
reported densities from four populations (i.e., 11–24 / 100 km2) were about two
to five times that of the highest puma densities in North America (see Chapter
10, Table 10-7). Likewise for wolves, which can compete with pumas where the
two carnivores are sympatric, Fuller (1989:20–21) examined the relationship of
wolf density to ungulate density in twenty-five areas in North America and
found that “average wolf densities are clearly correlated with biomass of ungu-
lates” (r 2 = 0.72, P < 0.001).
How Would a Puma Population Respond to a Prey Crash?

Other carnivores give insights into how pumas might respond to a severely
diminished prey base. Wolves, coyotes, bobcats, lynx, and African lions exhib-
ited lower body condition, reduced reproductive output, reduced survival, and
reduced recruitment following prey population declines (Schaller 1972, Bertram
1973, Brand et al. 1976, Mech 1977a, Todd et al. 1981, Todd and Keith 1983,
Parker et al. 1983, Knick 1990, Boertje and Stephenson 1992, Poole 1994,
Mowat et al. 1996, O’Donoghue et al. 1997). We would expect pumas, partic-
ularly cubs, to have lower survival. Cubs would be especially vulnerable because
mothers would have to hunt for longer periods, thus leaving cubs more exposed
to predation, including cannibalism. Longer time spans between feedings of
milk or meat would increase their risk of starvation, akin to African lion cubs
during periods of prey shortages (Packer et al. 1988). Pumas would need to
expend more energy to hunt increasingly scarce prey; thus, their physical condi-
tion would deteriorate. Some mothers might abandon cubs, as do cheetahs that
have difficulty securing food (Caro 1994). In addition, we expect females to
have lower pregnancy and birth rates. Some in particularly poor condition
might fail to reproduce altogether, like lynx in a snowshoe hare decline
(O’Donoghue et al. 1997). As intraspecies or interspecies competition for food
increased, we would expect greater emigration rates in subadult females. At the
extreme stage of prey shortage, pumas might increase their use of space in search
of better hunting grounds, as bobcats do during a lagomorph decline (Knick
1990). Some might even become nomadic, as lynx do during hare declines
(Ward and Krebs 1985, O’Donoghue et al. 1997). This activity might also
increase antagonistic encounters with other pumas and carnivores, which may
compete directly for prey or carrion. Pumas might also shift their diet toward
smaller prey animals that may be carriers of lethal diseases such as plague, which
killed adult female F68 in April 1995. Some pumas would simply starve to
death (Shaw 1980, T. Ruth personal communication regarding starving pumas
in northern Montana following prey decline). Starvation was the leading cause
of death among adult and subadult female leopards in South Africa. But it also
claimed some adult and subadult males (Bailey 1993). Apparently, the leopards
had difficulty capturing prey during the dry season because of the lack of stalk-
ing cover. Bailey (1993) indicated that food-stressed leopards could become
severely emaciated in one to three months. For pumas, lower birth and survival
rates should result in population decline with the potential for population col-
lapse if food shortage persisted. But before the puma population got to that
point, it could threaten small populations of prey with local extinction—the
main lesson of our next chapter.
LP > > KX
1. Pumas killed 89 percent of 540 dead deer we found; 11 percent died of
other causes. Fawns comprised about 27 percent of puma-killed deer we
found by chance, but there was probably a negative bias to finding fawns.
2. Out of 175 radio-collared deer one year and older, pumas killed about 35
percent, while about 17 percent died of other causes. Survival rates for
radio-collared deer fluctuated little during biological years 1987–1988 to
1991–1992 but declined substantially during 1992–1993 to 1994–1995
when effects of drought were prevalent. During drought years, puma pre-
dation rates also increased substantially. There appeared to be an inverse
relationship between rates of puma predation and death from other causes,
suggesting that puma predation is partially compensatory. A positive
response of deer survival rates to puma reduction suggested that puma pre-
dation is also partially additive. Deer survival rates were strongly related to
puma predation rates.
3. Results from a deterministic, discrete time model showed that the deer pop-
ulation increased during biological years 1987–1988 to 1991–1992 but
declined rapidly during 1992–1993 to 1994–1995. The deer population
dynamics were consistent with our field observations.
4. Two key proximate factors—fawn production and puma predation—
strongly influenced deer population growth.
5. Deer population limitation was determined by the interaction of puma pre-
dation, the strongest proximate cause of mortality, and deer habitat condi-
tion, as influenced by weather, which affected fawn production and survival
and adult deer health and vulnerability to predation.
6. Pumas did not adjust their population downward as the deer population
crashed. Instead, puma numbers increased or remained high as the deer
population declined, thus exhibiting a lag. Puma predation hastened the
deer decline. Consequently, we concluded that the self-limiting hypothesis
did not explain how pumas on the San Andres Mountains related to their
prey base. We hypothesized that the puma population is ultimately limited
by food.
LN N5LN5)L
1. Male:female sex ratio for fifteen buck and twenty-five doe fawns killed by
pumas did not differ from 1:1. Chi-square test: c2 = 2.50, 1 d.f., P > 0.10.
2. Ages of radio-collared bucks that died were not different from ages of radio-
collared does that died during the seven biological years 1987–1988 to
1993–1994. Two-sample t-test: t = –1.351, d.f. = 88, P = 0.18.
3. Tests for differences between buck and doe surv i val rates. Z test: 1990–1991,
Z = 1.476, P = 0.07. All other years, Z = 0.128–1.229, P = 0.11–
0.45.
4. Tests for differences in puma predation rates on bucks and does. Z test:
1990–1991, Z = 2.499, P = 0.006; all other years, Z = 0.399–0.825, P = 0.21–
0.35.
5. Tests for differences in rates of death from “other causes” for bucks and
does. Z test: years 1987–1988, 1988–1989, 1990–1991, 1992–1993,
1993–1994, Z = 1.330–2.133, P = 0.02–0.09; years 1989–1990,
1991–1992, Z = 0.282–0.802, P = 0.21–0.39.
6. Test for association between rates of puma predation and death from “other
causes” on bucks and does. Spearman correlation coefficients: bucks rs = –0.50,
P = 0.29; does rs = –0.71, P = 0.10.
7. Test for a linear relationship between survival rates of radio-collared deer
(bucks and does pooled) and puma predation rates. Linear regression: deer
survival rate = 0.9281 – 0.9129 (predation rate), P = 0.001, r2 = 0.90.
8. Test for a linear relationship between deer model 2 ls and puma predation
rates. Linear regression: l = 1.2232 – 1.5498 (predation rate), P = 0.002,
r2 = 0.87.
Test for a linear relationship between deer model 6 ls and puma predation
rates. Linear regression: l = 1.2013 – 1.1780 (predation rate), P = 0.007,
r2 = 0.79.
9. Linear regression of deer model 2 l on survival rates of radio-collared deer
(bucks and does one or more years old combined): l = – 0.3000 + 1.6323
(deer survival rate), P = 0.001, r2 = 0.89.
Linear regression of deer model 6 l on survival rates of radio-collared deer
(bucks and does one or more years old combined): l = 0.0177 + 1.2728
(deer survival rate), P = 0.003, r2 = 0.86.
10. Test for a linear relationship between deer model 2 ls and deer death rates
from “other causes.” Linear regression: l = 0.9803 + 0.5300 (death rate
from “other causes”), P = 0.84, r2 = 0.009.
Test for a linear relationship between deer model 6 ls and death rates
from “other causes.” Linear regression: l = 1.0402 + 0.0200 (death rate
from “other causes”), P = 0.99, r2 = 0.00002.
11. Test for a linear relationship between deer model 2 ls and fawns per 100 does.
Linear re g ression: l = 0.5692 + 0.0111 (fawns per 100 does), P = 0.003, r2 =
0.80.
Test for linear relationship between deer model 6 ls and fawns per 100
does. Linear regression: l = 0.6002 + 0.0109 (fawns per 100 does), P =
0.0002, r2 = 0.92.
12. Multiple regression models on ls for deer models 2 and 6: model 2 l =
0.8837 – 1.0652 (predation rate) + 0.0067 (fawns per 100 does), P = 0.004,
r2 = 0.94. Partial correlation coefficients: puma predation, r = –0.86, P =
0.03; fawns per 100 does, r = 0.71, P = 0.11.
Model 6 l = 0.8837 – 1.0652 (predation rate) + 0.0067 (fawns per 100
does), P = 0.004, r2 = 0.94. Partial correlation coefficients: puma predation,
r = –0.77, P = 0.07; fawns per 100 does, r = 0.83, P = 0.04.
13. Test for association between fawns per 100 does and adult puma density in
January in the Treatment Area, 1988–1994. Spearman correlation coeffi-
cient (1-tailed test): rs = –0.72, P = 0.05.
14. Test for association between fawns per 100 does and puma predation rates
on radio-collared deer (bucks and does pooled). Spearman correlation coef-
ficient (1-tailed test): rs = –0.68, P = 0.06.
Chapter 18
Pumas and Desert Bighorn Sheep
Pumas on the San Andres Mountains used desert bighorn sheep for food only
sparingly (i.e., 2 percent of puma kills, 0.6 percent frequency in puma fecal sam-
ples and stomachs). Nevertheless, we needed to assess the impact that puma pre-
dation had on the prospect of recovery of this endangered ungulate population.
Because this sheep population eventually went extinct, it serves as a case history
on extinction of mammal populations, and bighorn sheep populations in par-
ticular (see Berger 1990, 1999, Krausman et al. 1996, Wehausen 1999). There-
fore, we provide the historical context of desert bighorn sheep distribution in
southern New Mexico and factors known to have and suspected of having influ-
enced sheep population fluctuations.
Hypothesis and Predictions

We hypothesized that puma predation was the strongest proximate limiting fac-
tor affecting the sheep population. Hence, we predicted that (1) puma preda-
tion must be the most important proximate cause of mortality affecting sheep
population growth, and (2) experimental removal of pumas in the Treatment
Area should cause an increase in sheep survival rates that was linked to a reduc-
tion in puma predation rates.
These predictions were tested primarily by quantifying puma predation rates
on radio-collared sheep relative to other limiting factors and measuring the rela-
tionship of puma density to puma predation rates. In addition, we examined
how puma predation rates on radio-collared sheep related to puma predation
rates on radio-collared mule deer and the abundance of deer fawns.
341
Pre-history, History, and Threats

Events leading to the fate of the sheep population on the San Andres Mountains
began long before we arrived to conduct our research. Historically, there were at
least sixteen desert mountain ranges in southern New Mexico that supported
populations of desert bighorn sheep (Sandoval 1979a). Those populations, exist-
ing on mountain ranges separated by desert basins of various expanses, probably
formed a metapopulation. The San Andres Mountains, estimated to provide
about 710 km2 of suitable sheep habitat (Dunn 1994), may have had the largest
subpopulation of sheep, at least in the past one hundred years. Actually, there
were also sheep in the Organ Mountains, which were connected to the south
end of the San Andres Mountains (Sandoval 1979b). And there may have been
sheep in the Oscura Mountains, about 10 km northeast of the north end of the
San Andres Mountains. Prehistorically, it is possible that this mountain chain
supported a larger, more dispersed, and more persistent sheep population than
in historical times. Over millennia, dispersing sheep probably moved across
desert basins (i.e., matrix habitat) between mountain ranges in southern New
Mexico, albeit infrequently. Rams, especially, experience wanderlust during the
breeding season and are capable of straight-line distance travels of over 50 km
(Krausman et al. 1999). Although ewes tend to be much more sedentary, some
of them could have moved long distances during relatively cool, wet climatic
periods when food and water, and hence sheep populations, might have had
greater distribution and abundance. The most recent cooler, wetter ambient
condition in the southern New Mexico was probably the “little ice age,” which
lasted roughly from about 1600 to 1900 (Neilson 1986). Evidence of others
goes much farther back. Water levels in playa lakes, as inferred from temporal
distributions and abundances of aquatic, grassland, and desert scrub animals
found on Howell’s Ridge in southwestern New Mexico, suggest that much
longer wet periods occurred about 700 to 1,700 years ago, 2,500 to 3,300 years
ago, and from 5,000 years ago to the Pleistocene (Van Devender 1995). Sheep
immigrating into other subpopulations would have augmented populations
numerically and genetically. Immigrating sheep may have even rescued some
subpopulations from extinction and recolonized some habitat patches where
sheep had disappeared. Climate change was not the only factor that might have
affected metapopulation dynamics—so would modern human developments.
European immigrants to southern New Mexico during the 1800s and 1900s
brought new threats to the desert bighorn sheep metapopulation. Introduced
livestock, particularly sheep and goats, competed with wild sheep for food and
introduced diseases that could kill desert sheep (e.g., pneumonia, chronic sinusi-
tis, scabies; Bunch et al. 1999). Overgrazing by livestock may have transformed
CHAPTER 18. PUMAS AND DESERT BIGHORN SHEEP 343
grass-covered slopes to tree- and shrub-studded ones, diminishing habitat qual-

ity for sheep by reducing their ability to detect predators but enhancing food for
deer and cover for stalking predators (also see Berger and Wehausen 1991, Neil-
son 1986, Van Devender 1995). Settlers sometimes killed wild sheep for food
(Buechner 1960, Valdez and Krausman 1999). The construction of fences to
manage livestock movements as well as to delineate highway and railroad rights-
of-way probably eliminated the already intermittent movements of sheep
between the mountain ranges. Most patches of sheep habitat became perma-
nently isolated from one another. Thus, extant sheep populations became
increasingly vulnerable to detrimental effects that befall small isolated popula-
tions. Sheep in the Organ Mountains lingered as late as 1910. In the Guadalupe
Mountains, representing the eastern-most extent of sheep in New Mexico, sheep
disappeared about the late 1940s (Sandoval 1979b). By the time we started our
research in 1985, only four wild desert bighorn sheep populations remained in
New Mexico, all together adding up to fewer than 150 animals (Elenowitz
1987). In an environment where climate and weather fluctuations already cre-
ate a tenuous hold on life, human disruptions to the metapopulation dynamics
and changes in sheep population health and in plant and animal communities
probably heightened extinction risks to sheep populations.
Historical accounts say sheep on the San Andres Mountains were “rare” in
1902 and 1903 (J. A. Gaut, Biological Survey, cited in Sandoval 1979b). The
first official attempt to census the population was undertaken in 1942 by staff
of the newly designated San Andres National Wildlife Refuge; a minimum of
thirty-one sheep were counted. Through 1950 the population increased to at
least 140 sheep. But from 1951 to 1953 the population plummeted again to
about fifty-three sheep. According to Lang (1956) the decline was caused by a
combination of factors: severe drought, overpopulation of deer, overgrazing by
domestic livestock, and disturbances by deer hunters. During the next fourteen
years, the sheep population increased gradually, reaching an all-time estimated
high of 270 sheep in 1967. The population high justified regulated sport-hunt-
ing seasons for rams with at least three-quarter-curl horns. However, the 1967
estimate was contested by Welch (1969), who thought it too high and that an
estimate of 150–200 sheep was more realistic. Apparently, a rethinking of the
crude population estimates was being undertaken at the time, because a 1968
estimate of 250 sheep was also disputed by Augsburger (1970) and later modi-
fied by Sandoval (1979b) to about 160 sheep. The first intensive study of the
desert bighorn sheep was conducted by Sandoval (1979b). He employed a com-
bination of helicopter and ground surveys in 1975 and 1976 to estimate there
were 200 ± 18 sheep.
The sheep population again declined during the years 1978–1979. Sheep
hunting was halted. In November 1979 there were an estimated eighty sheep
(Sandoval 1980). The decline was attributed to a widespread epizootic of scabies
(Lange et al. 1980). This disease, caused by a mite (Psoroptes spp.), produces
clinical signs on the sheep that include trunk alopecia (i.e., hair loss); intense
itching; protein exudation; mucopurulent effusion from and severe folding of
the pinnae (i.e., external ears); ear canals occluded with fleshy plugs; purulent
inflammation of the outer ear; destruction of the tympanic membrane (i.e., ear
drum); ataxia; and weakness (Sandoval 1979a, Sandoval 1980, Lange et al.
1980, Lange 1982, Clark and Jessup 1990). Scabies may also cause hearing loss
(Norrix et al. 1995), hypothermia, and suppression of the immune system
(Clark and Jessup 1990). Besides scabies, some of the sheep captured in 1979
had antibodies against contagious ecthyma, and 28 percent of them had clinical
signs of this disease. Contagious ecthyma is a viral disease that causes scabby
lesions on the mouth, eyelids, forelegs, and udders. It can lead to secondary
blindness, lameness, and impaired feeding, which contribute to emaciation and
death. Lambs are particularly susceptible to starvation (Lance 1982). Sandoval
(1980) thought that scabies could have acted as a predisposing factor to conta-
gious ecthyma, and that the large-scale die-off could have been due to both dis-
eases. Physiological effects of these diseases, if not causing death directly, could
predispose sheep to death from a variety of other causes, including bacterial
infections, accidental falls from precipices, and predation.
The origin of the scabies mites in this wild sheep population has been a mys-
tery, but Sandoval (1980) suggested that domestic sheep and goats introduced
on the San Andres Mountains during the early 1900s may have passed them to
the desert bighorn sheep. He associated other scabies-caused bighorn sheep pop-
ulation declines that occurred in California, Colorado, Oregon, and Wyoming
with the introduction of domestic sheep. Furthermore, there is evidence that the
desert mule deer act as an alternate host for the mites. We found four mule deer
killed by pumas that were infested with psoroptic mites. The New Mexico
Department of Game and Fish and the U.S. Fish and Wildlife Service sampled
at least 151 deer from the San Andres Mountains during 1989–1993; twenty-
six had mites in their ears (Boyce 1994). Study of morphological characters of
mites in the sympatric sheep and deer populations on the San Andres Moun-
tains suggested that the mites are not host specific and that they may represent
a single interbreeding population (Boyce et al. 1990). Unfortunately, the extent
to which deer may transmit mites to sheep or vice versa has not been adequately
tested (Clark and Jessup 1992).
In November 1979, a sheep rescue effort was made by the U.S. Fish and
Wildlife Se rvice, the New Mexico Department of Game and Fish, and the
U.S. Army at White Sands Missile Range. Forty-nine sheep we re captured
using a helicopter, net-gun, and drug-filled darts so they could be treated

(Sandoval 1980). T h i rt y - f i veof the sheep surv i ved capture and treatments. Of
those, seven we re kept in captivity at New Mexico State Un i versity in Las
Cruces for study of the transmission and control of psoroptic mites. The
remaining twenty-eight sheep we re transported to the New Mexico De p a rt-
ment of Game and Fish desert bighorn sheep breeding facility at Red Ro c k ,
where they remained for thirteen months. T h e re, some of the sheep died of
blue-tongue viral pneumonia and contagious ecthyma. Only twe l ve surv i v i n g
sheep we re returned to the San Andres Mountains in January 1981. At that
time, the entire sheep population was estimated to be about forty animals
(Hoban 1990).
In a short span of about forty years, the sheep population on the San Andres
Mountains had fluctuated to lows less than or equal to fifty animals three times.
Biologists studying demographics of bighorn sheep in Wyoming, Montana, and
South Dakota have estimated that sheep populations with no effective immi-
gration (i.e., gene flow) need to have more than 150 total individuals to avoid
short-term (i.e., generational time) loss of genetic variability (Fitzsimmons and
Buskirk 1992). Population bottlenecks of the San Andres Mountains sheep pop-
ulation provided conditions for erosion of genetic variation through inbreeding
(i.e., mating between close relatives) and genetic drift (i.e., changes in allele fre-
quency in a small population caused by random sampling of gametes to form
zygotes). There is evidence that this may have occurred. Researchers compared
genetic variation in five major histocompatibility complex loci (tightly linked
loci involved in the immune response) and three microsatellite loci among
desert bighorn sheep from the San Andres Mountains and from eleven popula-
tions in the Mojave Desert and Peninsular Ranges in Nevada and California.
They found that San Andres sheep exhibited the lowest heterozygosity (i.e.,
genetic variation where there are different alleles at more than one locus) (Boyce
et al. 1996). Similar research, this time using ten highly variable microsatellite
loci, found that sheep from the San Andres Mountains had lower genetic varia-
tion than ten other desert bighorn sheep populations in Arizona and California
(Gutiérrez-Espeleta et al. 1998). Yet, these patterns of low genetic variation
could also result from a very small number of sheep founding the population.
Effects of lower genetic variation in the San Andres sheep are unknown and
would be very difficult to measure. Theoretically, increased expression of delete-
rious alleles could suppress individual reproductive performance, viability of
young, resistance to parasites and diseases, and the ability to adapt to a chang-
ing environment. Cumulative effects could result in lower population growth
rates, which in turn could retard the ability of the population to rebound from
population declines. A resulting small population could then be at greater risk
to extinction from unpredictable environmental events that destabilize the pop-

ulation (Lacy 1997).
Sheep Population Characteristics during Our Research

As we explained in Chapter 4, survey techniques to count sheep on the San
A n d res Mountains varied in kind and intensity. These methods pro d u c e d
minimum counts of sheep ranging from twenty-two to thirt y - s e ven (Table
18-1). Population estimates seemed to reflect survey techniques, as longer
s e a rch time and the use of a helicopter resulted in a greater number of sheep
observations. Even considering that biologists probably missed a few lone
sheep or small bands during surveys, the sheep population on the San
A n d res Mountains probably ranged from thirty to forty animals during
1986–1994.
Table 18-1. Desert bighorn sheep observed in surveys conducted on

the San Andres Mountains, New Mexico, 1986–1994.
Adults Yearlings Lambs
Unk. Unk.
Year Rams Ewes Rams Ewes Sex Rams Ewes Sex Total
1986a 9 15 3 2 4 3 36
1987a 9 13 3 3 3 5 1 37
1988a 10 11 3 5 3 2 34
1989a 10 15 3 2 3 1 34
1990b 7 9 3 2 1 22
1991c 3 10 2 1 6 22
1992d 6 13 3 5 2 2 31
1993e 7 11 5 2 5 30
1994a 12 12 2 8 34
aNumbers for 1986–1989 and 1994 were obtained from ground-based observations and
radiotelemetry data gathered year-round (San Andres National Wildlife Refuge files).
bNumbers for 1990 were from ground-based observations and radiotelemetry data gathered
during December and known mortalities that occurred during the rest of the year (San
Andres National Wildlife Refuge files).
cNumbers for 1991 were from ground-based observations and radiotelemetry data gathered
during 31 August–10 September (New Mexico Department of Game and Fish files).
dNumbers for 1992 were from ground-based observations and radiotelemetry data gathered
during May–December (San Andres National Wildlife Refuge files).

eNumbers for 1993 were from a helicopter survey conducted on 20–21 November (New
Mexico Department of Game and Fish files).

The surveys also provided data on herd composition. Based on the sheep
classified by gender during surveys, lambs apparently were born at an equal sex
ratio (17 males:16 females). This ratio held into the yearling age class (19
males:18 females). But in the adult age class, there generally were fewer males
than females, with the ratio of adult rams to adult ewes ranging from 0.30:1 to
1:1 and averaged 0.67:1 (Table 18-1). Tendency toward fewer adult rams was
probably because of proportionately greater mortality (see below). The ratio of
lambs to adult ewes ranged from 0.27:1 to 0.69:1 and averaged 0.51 ± 0.16:1.
The ratio of yearlings to adult ewes ranged from 0.17:1 to 0.73:1 and averaged
0.39 ± 0.19:1. On average, the ratio of lambs to adult ewes during the
1986–1994 surveys was quite similar to the average lamb:ewe ratio (x– = 0.50 ±
0.21) this sheep population exhibited during its initial documented growth
phase during 1941–1952 when the population increased from a low of about
twenty-seven sheep to an estimated 112 sheep. And it was similar to the average
lamb:ewe ratio ( x– = 0.54 ± 0.17) from population surveys conducted during
1965–1976 when sheep population estimates were their historical highest, rang-
ing from 182 to 270 sheep (Sandoval 1979b:13). During those times, lamb pro-
duction was high enough to offset mortality and to spur population growth. But
the difference during our research was that mortality of sheep apparently
matched or outpaced reproduction.
Fates of Radio-Collared Sheep

We and U.S. Fish and Wildlife Service biologists monitored a total of forty-three
radio-collared sheep (sixteen rams, twenty-seven ewes) from August 1985 to
March 1995 to detect puma predation and characteristics of the sheep that died.
The number of radio-collared sheep monitored each year comprised 36–83 per-
cent of the number of adults and yearlings observed during annual surveys dur-
ing 1986–1994 (Fig. 18-1). We detected twenty-six radio-collared sheep that
died. With the help of agency biologists who concentrated on the fates of these
sheep, we were able to get to sheep that died in an average of 7 ± 7 days after
death (range = 0–30, median = 5).
Pumas killed ten radio-collared sheep, including four rams (x– age = 1.7 ±
1.5, range = 0.8–four years) and six ewes (x– age = 8 ± 4.9, range = 3–16 years).
Six of these sheep had clinical scabies, with infestations ranging in severity from
relatively minor scaly lesions on the flanks to both ear canals completely
occluded with scabs. A six-year-old ewe had no scabies lesions, but she was
clearly emaciated prior to being killed by a puma. Her femur marrow was pink
and gelatinous in consistency, indicating that fat depletion had reached extreme
levels in the long bones, the last fat stores available for energy metabolism. This
FPO @ 82%
FIGURE 18-1. Number of adult and yearling desert bighorn sheep observed in annual sur-
veys and radio-collared during each year, San Andres Mountains, New Mexico.
Slide @318%
PHOTO 28. A desert bighorn ewe killed and eaten by a puma.

Slide @171%
PHOTO 29. A yearling desert bighorn
ram with ear canals occluded with
scabs. This ram was killed and eaten
by puma M3.
condition suggests that the animal was in extremely poor condition and was
probably dying (Mech and DelGuidice 1985).
We estimated that about 30 percent of puma-caused deaths in the radio-col-
lared desert bighorn sheep was compensatory mortality (i.e., the same percent-
age we found for mule deer one or more years old). Besides the emaciated ewe
mentioned previously, there were two ewes killed by male puma M23 in Janu-
ary and March 1989 that were twelve and sixteen years old, respectively.
Chances were that these two ewes would not have survived another year. They
were as old or older than a twelve-year-old ewe and two rams that were thirteen
and fourteen years old when they apparently lay down or fell in their tracks and
died from age-related ailments.
Sixteen other radio-collared sheep, including eight rams (x– age = 7.2 ± 4.0,
range = 4–14 years) and eight ewes (x– = 5.6 ± 3.4, range = 3–12 years) died from
other causes. Natural causes of death included falls from cliffs (three rams, two
ewes), aging (two rams, one ewe), scabies (two rams, one ewe), undetermined
diseases (one ewe), and breached birth (one ewe). Two ewes died of undeter-
mined natural causes. One ram died from a capture accident when he tumbled
over a cliff after getting caught in a net. One of the rams that apparently died of
scabies was scavenged by male puma M7. We also discovered an uncollared year-
ling ewe that died of undetermined causes not related to puma predation. When
we found it, a coyote was scavenging the carcass as it lay partially submerged in
a spring-fed pool.
Ten of the sixteen radio-collared sheep that died of other causes had clinical
scabies. The mildest case consisted of scabby lesions on one leg. Severest cases
apparently caused death. One six-year-old ram had hemorrhaging and suppura-
tive lesions over 40 percent of its body and was extremely emaciated. Another
six-year-old ram had scabby lesions over 75 percent of its body and suppurative
lesions over about 40 percent of its body. At the death site of the latter ram, we
found where he had been scratching himself against rocks protruding out of a
steep slope, leaving bloody scabs and hair on the rock. Furthermore, we suggest
that if scabies causes bacterial infections that impair the inner ear, which is also
vital for equilibrium, then scabies may contribute to deaths from accidental falls
(19 percent of all deaths, 29 percent of deaths from other causes). Three of the
five sheep that fell to their deaths had ear canals blocked by scabs. In all, clini-
cal scabies was found in 62 percent of the dead radio-collared sheep (i.e., puma
kills plus sheep that died of other causes), indicating that scabies was still at epi-
zootic proportions in the sheep population during our research.
Survival Rates and Agent-Specific Mortality

We used twenty-six radio-collared adult sheep (eleven rams, fifteen ewes) that
lived in the Treatment Area to quantify annual survival rates and agent-specific
mortality rates during biological years 1987–1988 to 1993–1994 to correspond
with our puma population estimates and mule deer survival and agent-specific
mortality rates in that area. We monitored those sheep for 22,240 days (7,380
ram-days, 14,860 ewe-days). These sheep lived in the Treatment Area. Two
other radio-collared adult ewes lived exclusively in the Reference Area from the
time they were originally captured in November 1989 through the end of our
study in March 1995. We did not include them in our analyses of annual sur-
vival and agent-specific mortality rates because they did not live in the Treat-
ment Area, where we experimentally removed pumas. Annual survival rates for
six to nine adult ewes monitored per year ranged from 0.63 to 1.0 (x– = 0.89 ±
0.16) (Table 18-2). We had too few rams each year (one–seven / year) to esti-
mate their survival rates, but we lumped them with ewes to boost the sample of
radio-collared sheep available each year to estimate adult sheep survival rates and
agent-specific mortality rates. With rams included, annual survival rates of adult
sheep ranged from 0.58 to 1.0 (x– = 0.84 ± 0.17). Aside from the small number
Table 18-2. Survival rates of radio-collared desert bighorn sheep, San Andres
Mountains, New Mexico, 1987–1994.a
Ewes Ewes plus ramsc
Biological Sheep-days Rate 95% C.L. Sheep-days Rate 95% C.L.
yearb
1987–88 2,192 0.85 0.62–1.0 3,656 0.90 0.74–1.0
1988–89 2,200 0.72 0.45–1.0 3,405 0.58 0.37–0.94
1989–90 2,234 1.0 1.0–1.0 3,838 1.0 1.0–1.0
1990–91 2,190 1.0 1.0–1.0 3,627 0.90 0.74–1.0
1991–92 2,196 1.0 1.0–1.0 2,928 0.88 0.69–1.0
1992–93 1,573 0.63 0.33–1.0 1,659 0.64 0.35–1.0
1993–94 2,275 1.0 1.0–1.0 3,127 1.0 1.0–1.0
aSurvival rates and 95 percent C.L. (Confidence Limits) were computed using Micromort software
(Heisey and Fuller 1985b).

bBiological years spanned August–July each year.
cIn years 1987–1988 to 1993–1994, one to seven rams and six to nine ewes were monitored.
of adult rams we could monitor, rams still showed a tendency to die at higher
rates than ewes. Of the total of thirteen adult rams we monitored during
1985–1995, 69 percent of them died, as opposed to 41 percent of the twenty-
seven radio-collared adult ewes we monitored in the same time span. This may
explain the observed tendency toward fewer adult rams than adult ewes in the
population.
Trying to estimate yearling and lamb survival rates was even more tenuous,
given their smaller numbers and variation in the surveys. We estimated survival
rates of yearling sheep by calculating a finite rate of survival (number of year-
lings surviving to two years old / number monitored) for a total of ten radio-col-
lared yearlings that we monitored from 1985 to 1994. Of those, three rams were
killed by pumas, resulting in a survival rate of 0.70. Although we could not be
certain that all yearlings counted in one year were the same individuals counted
as lambs in previous years, we calculated a rough finite rate of lamb survival by
dividing the number of yearlings observed in surveys during 1987–1994 by the
number of lambs observed in previous years from 1986 to 1993. We assumed
the difference represented the number of lambs that died. The resulting finite
lamb survival was 0.77. By multiplying these rates by the total number of lambs
observed during 1986–1994 (i.e., 56 lambs ¥ 0.77 ¥ 0.70) we estimated that
roughly thirty offspring would have survived to two years old. By recalculating
annual survival rates of adult rams and ewes (combined) for annual periods (Jan-
uary through December) 1986–1994 (using Micromort software) to correspond
with the survey data, we estimated that about thirty-four adult sheep died (i.e.,
S (1 – annual survival rate x number of adult sheep counted / year)) [1]. Hence,
in the observed sheep numbers there was roughly a net loss of four sheep dur-
ing the nine-year period. Consequently, we concluded that the desert bighorn
sheep population on the San Andres Mountains during our research was static
and vulnerable at a very low number of about thirty to forty animals. It is impor-
tant to stress that during 1987–1988 to 1991–1992, when environmental con-
ditions were favorable enough to stimulate a mule deer population increase, the
sheep population apparently did not respond similarly. During this period the
puma population throughout the entire study area was also increasing from a
biological low point; it was again experimentally reduced in the Treatment Area
in 1990–1991. Part of the reason the sheep population could not respond effec-
tively to favorable environmental conditions was the natural propensity of ewes
to produce only one lamb per year, unlike mule deer, which can produce twins.
Twinning has not yet been observed in wild desert bighorn sheep and has very
rarely been seen even in captive animals (Krausman et al. 1999). This probably
stymied the potential for reproduction to outpace realized mortality.
Puma predation and death from other causes on radio-collared adult sheep
was highly sporadic (Table 18-3). During biological years 1987–1988 to
1993–1994, four sheep were killed by pumas, including one ram and two ewes
killed by male puma M23 from 1 January to 11 March 1989. In those years, six
radio-collared adult sheep (two rams, four ewes) died of other causes. The gen-
eral pattern was that more adult sheep died from “other causes” than from puma
predation—the opposite pattern from what we documented for desert mule
deer. We removed M23 from the study area (he was translocated to northern
New Mexico) to honor our agreement with cooperators to remove any pumas
that killed more than one of the endangered sheep. He was the only puma in
our ten-year study to demonstrate an affinity for killing sheep. It is probable that
if he had stayed on the San Andres Mountains he would have killed more sheep.
After we removed M23, only one other radio-collared sheep was killed by a
puma; this occurred in August 1990 (Fig. 18-2). Puma predation rates on radio-
collared sheep (i.e., ewes and rams combined) were apparently entirely inde-
pendent of the number of adult pumas on the Treatment Area, puma predation
rates on radio-collared mule deer (i.e., bucks and does combined), and the rela-
tive abundance of highly vulnerable deer fawns (i.e., as indexed by fawns / 100
does) [2]. With so few sheep relative to the number of deer and other alternate
prey, all the pumas, except for maybe M23, apparently did not purposely hunt
them. Instead, we think pumas killed sheep that they encountered by sheer
FIGURE 18-2. Rates of surv i val, puma predation, and deaths from other causes for radio-
collared desert bighorn sheep, San Andres Mountains, New Mexico.
chance, probably while they were actually hunting the more-abundant deer that
also lived in sheep habitat. We periodically observed deer, and sometimes found
remains of deer killed by pumas, in some of the most rugged terrain used by
sheep.
Did Puma Predation Limit the Sheep Population?

We concluded that puma predation was a strong limiting factor to this remnant
desert bighorn sheep population, because puma predation was the single most
identifiable cause of death. But puma predation rates were not related to puma
density. Sporadic puma predation by itself apparently did not stop the sheep
population from growing during our research tenure. More sheep actually died
from other causes, at least half of which were probably aggravated by disease. All
mortality factors rendered the sheep population unstable, particularly in the
midst of drought and a rapidly declining mule deer population—the puma’s
chief prey. How long would the deer satisfy the energy demands of an increas-
ing puma population without escalating jeopardy to the sheep?
Finale of the Sheep Population

Dynamics between pumas and sheep changed markedly after our research
ended. Biologists Mara Weisenberger and Gary Montoya for the San Andres
National Wildlife Refuge continued monitoring the remaining, aging radio-col-

lared sheep. In 1996, seven of nine sheep (78 percent) with working radio col-
lars died. New Mexico Department of Game and Fish biologists radio-collared
a ram in April 1997, bringing the number of collared sheep available for moni-
toring to three. But in 1997, two more sheep died. Hence, in the span of only
nineteen months, from February 1996 to August 1997, nine radio-collared
adult sheep died, including two rams and seven ewes. Pumas killed one ram and
five ewes (x– age = 8.8 ± 2.0, range = 6–11 years), while one ram and two ewes
died of “other causes” (x– age = 12.0 ± 3.0, range = 9–15 years). Two of these
deaths could not be determined, but a twelve-year-old ewe fell from a precipice.
This decline seemed to have left only one eight-year-old radio-collared ewe. We
surmised that the greater puma predation rates on the radio-collared sheep were
indeed linked to the crash in the mule deer population, which by 1996 must
have been close to its nadir. Pumas probably were having to hunt their home
ranges intensively to find large prey. They could accomplish this by hunting
more often and traversing more habitat. In the process, we suspect they
increased encounter rates, and thus predation rates, on sheep and drove the
small population downward.
Biologists could not find other non-radio-collared sheep, probably indicat-
ing that attrition had occurred in them as well. This notion was supported in
December 1996 when biologists with the U.S. Fish and Wildlife Service, New
Mexico Department of Game and Fish, and White Sands Missile Range part-
nered to conduct a survey using three helicopters, each one assigned to canvass
about one-third of the San Andres Mountains. No sheep were seen in fifteen
hours of survey time. In December 1997 another helicopter survey of the moun-
tains was launched. Only one sheep was seen in five hours, the last radio-col-
lared ewe. U.S. Fish and Wildlife biologists conducted subsequent ground
searches and used cameras to monitor three springs that sheep were known to
use; no more sheep were found. With only one remaining ewe, the desert
bighorn sheep population on the San Andres Mountains was biologically
extinct. The major proximate mechanism responsible for the extinction, as indi-
cated by fates of radio-collared sheep, was puma predation.
We believe a combination of factors, both human and nonhuman, ulti-
mately brought about the biological extinction of the sheep on the San Andres
Mountains. Climatic and human-induced impacts resulting in isolation (i.e.,
disintegration of the regional metapopulation), disease (i.e., possible introduc-
tion of scabies), and the possible attendant loss of genetic variation from three
population bottlenecks resulted in a small unstable sheep population at the
brink of extinction. A naturally low reproductive rate could not offset deaths.
Finally, drought and attendant puma predation caused extinction of the popu-
lation.
Pumas and Other Sheep Populations

Puma predation is a substantial mortality factor in other bighorn sheep popula-
tions in North America. It is a major factor threatening the persistence of a fed-
erally endangered sheep population in the desert Peninsular Ranges of Southern
California. When the sheep population was listed as endangered in 1998, there
were about 330 yearling and adult animals. Biologists studying the fates of 113
(sixteen rams, ninety-seven ewes) radio-collared adult sheep during 1992–1998
found that pumas killed forty-two of sixty-one (69 percent) of the sheep that
died. On an annual basis, the puma predation rate tended to be four times the
rate from non-predation mortality. The mean annual survival rate of 0.79 was
low relative to other bighorn sheep populations in southeastern California
(greater than or equal to 0.91), northwestern Arizona (greater than or equal to
0.86), and four populations in the Mojave Desert (greater than or equal to 0.85)
(Hayes et al. 2000). It was also lower than the mean annual survival rate of 0.84
that we found for the desert bighorn sheep on the San Andres Mountains. Biol-
ogists did not assess to what extent puma predation was compensatory or addi-
tive. Still, they concluded that if high puma predation continues on adult sheep,
the population probably will not grow.
In the Mojave Desert of California, a tiny population of about a dozen
bighorn sheep experienced substantial variation in puma predation. During
1989–1992, annual survival rates for ewes averaged about 0.62, with all mor-
tality due to puma predation. However, during the following three years there
was no puma predation and the sheep population increased by 15 percent per
year (Wehausen 1996).
While studying pumas and bighorn sheep in southwestern Alberta from
1985 to 1994, biologists found that puma predation was highly variable, rang-
ing from 0 to 57 percent of winter (i.e., December through March) mortality in
a sheep population that numbered between 112 and 158 animals. Of twenty-
nine sheep killed by pumas, thirteen were lambs and sixteen were one year or
older. Pumas selected lambs. Apparently 29 percent of lambs, 44 percent of
ewes, and 50 percent of rams killed by pumas exhibited anatomical or behav-
ioral abnormalities, which may have predisposed them to predation. Not all
pumas on the study area killed sheep equally. Of five radio-collared female
pumas monitored intensively, two never killed sheep, one killed only one sheep,
one killed five sheep, and another killed seventeen sheep. This last puma killed
8.7 percent (n = 11) of the early-winter sheep population and 26.1 percent (n =
6) of the lambs in winter 1993–1994. The biologists believed that this high pre-
dation triggered a decline in the sheep population that continued into 1995 and
1996. The puma was monitored with radiotelemetry for more than ten years,
and the biologists believe she learned to kill sheep in her later life (Ross et al.
1997). This situation in Alberta, like our experience with M23, indicates that
individual pumas can cause serious mortality in small sheep populations.
LP > > KX
1. Desert bighorn sheep on the San Andres Mountains during 1985–1995
comprised a remnant population of only about thirty to forty animals. The
population probably was once part of a regional metapopulation in south-
ern New Mexico. Since the beginning of the twentieth century, the sheep
population has had a history of wide fluctuations from about fifty to two
hundred animals, including three demographic bottlenecks of fifty or fewer
animals. Human activity and development in southern New Mexico prob-
ably contributed to the decline of desert bighorn sheep regionally and to the
extinction risk for sheep on the San Andres Mountains in particular.
2. Puma predation was the single-most identifiable cause of mortality in radio-
collared sheep, claiming 23 percent of sheep monitored. Puma predation on
sheep was sporadic. Pumas seemed to kill sheep opportunistically, probably
while they were actually seeking deer. We estimated that about 30 percent
of puma predation was compensatory. There was no association between
puma predation rates on radio-collared sheep and the number of adult
pumas, puma predation rates on radio-collared deer, or fawn productivity.
More sheep died from other causes, comprising 37 percent of sheep we
monitored. Total mortality approximately matched reproduction; hence,
the sheep population was static during our research.
3. Scabies, which was responsible for the sheep population crash during
1978–1979, was still an epizootic during our research. Clinical scabies
occurred in 62 percent of all radio-collared sheep that died. Scabies proba-
bly predisposed sheep to a variety of mortality causes, including predation,
falls from cliffs, malnutrition, and disease.
4. Small population size, predation, high incidence of disease, and drought
rendered the sheep population highly unstable and vulnerable. By 1997, the
sheep population was biologically extinct due to intensified puma predation
following a crash in the deer population.
LN N5LN5)L
1. Combined annual survival rates for radio-collared adult (two years or older)
desert bighorn rams (n = 13) and ewes (n = 21) monitored for 30,188 radio-
days from 1986 to 1994: 1986 = 0.81, 1987 = 0.74, 1988 = 0.71, 1989 =
0.73, 1990 = 0.91, 1991 = 1.0, 1992 = 0.64, 1993 = 1.0, 1994 = 0.90 (x– =
0.83 ± 0.13).
2. Test for association using Spearman correlation coefficients between puma

predation rates on radio-collared adult sheep and the following parameters
in the seven-year span: numbers of adult pumas on the Treatment Area,
rs = –0.13; puma predation rates on radio-collared adult deer, rs = 0.13;
fawns/100 does, rs = 0.30. All P > 0.50.
Chapter 19
Synthesis: Pumas and Weather

Modulate Large-Mammal
Population Dynamics on the
San Andres Mountains
Puma predation was the most important proximate cause of mortality affecting
desert mule deer and desert bighorn sheep on the San Andres Mountains. The
effect that pumas had on these native ungulate populations was linked to unpre-
dictable weather events (specifically, an unusual wet period followed by drought)
and attendant effects on herbivore foods, free water, and probably security cover.
These dynamics appeared to be a classic case of biotic (i.e., puma predation,
plants, population size, disease) and abiotic (i.e., weather) forces interacting to
influence the nature of population, community, and ecosystem processes (Mat-
son and Hunter 1992, Post et al. 1999).
Both predation and weather were destabilizing forces. An unusually wet
weather pattern created an environment favoring production of an ample food
supply and cover that enabled the deer population to grow relatively rapidly,
even with puma predation slowing it down. Ample deer in the environment
probably reduced puma predation on sheep. We believe the subsequent drought,
in combination with herbivory, stressed plants and desiccated water sources.
This effectively constricted food supply and cover, thus reducing deer fertility
rates and fawn security. At the same time, the puma population was increasing.
This triggered a predation threshold response; pumas continued killing the less-
359
abundant fawns while increasing predation pressure on the now more vulnera-
ble reproduction-aged deer. Increased puma predation rates on the growth-
arrested deer population hastened the deer population decline, driving it to
lower numbers, at least in the short term, than it might have attained without
puma predation. This exposed the sheep to greater puma predation. Although
we were unable to observe the fate of puma population directly, after a lag it
apparently declined in response to its dwindling prey base. What might happen
next undoubtedly depends on whether drought persists, or if it is replaced with
a relatively normal or wet weather regime.
These dynamics pertain to an ongoing debate regarding the influence that
apex predators and plants exert on ecosystems (see Matson and Hunter 1992,
Hunter and Price 1992, Power 1992, Strong 1992, Menge 1992, McLaren and
Peterson 1994, Post et al. 1999, Terborgh et al. 1999). If predators have primacy
on system dynamics then there is top-down control. According to this scenario,
animals at the top of the food chain are food limited, and at successively lower
trophic levels animals are alternatively predator, then food limited (Power
1992). So if pumas exerted top-down control on the San Andres Mountains, the
puma population would be limited by food, chiefly deer; puma predation would
limit the deer population; and plants would be limited by food (e.g., water, soil
nutrients). But if primary producers (i.e., plants) have primacy, the system is
controlled from the bottom up. This scenario holds that animals at each trophic
level are food limited (Power 1992). Thus, food would limit the plants, which
would limit deer, which would limit pumas.
One way of sensing the relative strengths of top-down and bottom-up con-
trol is to consider what might happen if the top and bottom trophic levels were
removed from the San Andres Mountains ecosystem. Absence of pumas would
have profound effects. We would expect the deer population to grow at a faster
rate and to higher numbers before it would be limited by food. Thereafter, it
might decline at a slower rate, but in the process it might impose longer-lasting
negative impacts to food sources due to intensified and protracted overbrows-
ing. Furthermore, because deer and sheep distributions overlap substantially and
the diet of those two ungulates overlap by more than 70 percent (Sandoval
1979b), exploitation competition between deer and sheep would be expected to
increase. Hence, the ability of a small sheep population to grow would depend
upon its competitive interactions with deer and upon the quantity and quality
of food available. Puma predation, by dampening the growth rate of the deer
population, may lower exploitation competition between deer and sheep.
Hence, puma predation plays a crucial role, modulating the numbers of the
most abundant ungulate (i.e., deer) and its attendant effects on food plants and
other herbivores. We view the puma’s impacts on ecosystem dynamics as dis-
CHAPTER 19. SYNOPSIS: PUMAS AND LARGE-MAMMAL POPULATION DYNAMICS 361
Slide @318%
PHOTO 30. The puma is a keystone species in wild desert ecosystems, and it can serve as an
umbrella species in conservation efforts.
proportionately large relative to its numbers or biomass in the ecosystem, thus

qualifying the puma as a keystone species (Meffe and Carrol 1997).
Now take away the plants and what happens? The system collapses. Clearly,
it is the desert plants that have primacy in the food web, capturing most of the
energy (from the sun) needed to sustain the large-mammal community on the
San Andres Mountains. Primary productivity affects health, reproductive per-
formance, and hence population growth of ungulates and small alternate prey
(e.g., rabbits, hares, rodents), which together enable ungulate populations to
endure predation. In turn, population levels of prey determine the population
level of the puma (see Power 1992). Just during the twentieth century, severe
droughts occurred at least six times in southern New Mexico: 1908–1913,
1934–1935, 1951–1956, 1963–1965, 1971, and 1976 (Neilson 1986), with
each interposed with relatively wet periods. Clearly, weather (a subset of the
regional climate) is an unpredictable natural perturbation, whether it produces
wet cycles or drought. It can destabilize the system by acting directly on plants;
thus, it modifies relative strengths of top-down and bottom-up forces within a
relatively short time scale (i.e., within one year).
On the San Andres Mountains, top-down and bottom-up forces interacted
simultaneously and were modified by weather to shape large-mammal commu-
nity dynamics. The ecosystem is much too complicated to describe its control as
a simple dichotomy—top-down or bottom-up. Instead, it is much more impor-
tant to understand that the desert ecosystem is naturally variable and to under-
stand how biotic and abiotic factors influence its dynamics. Likely, these dynam-
ics have been operating for the life of the Chihuahuan Desert—about four
thousand years (Van Devender and Toolin 1983). Considering them in this con-
text is essential for developing management and conservation strategies for
pumas, prey, and habitat, particularly in a milieu of unprecedented human pres-
sure.
P a rt V
Pumas and People

Chapter 20
Conservation and
Management of Wild Pumas
Conservation is a confluence of our ethical obligation to protect nature and our

understanding of how nature works. Consequently, conservation is laden with
human values. Still, when people conserve species or ecosystems, then biologi-
cal diversity is perpetuated through genetic variation in species and populations,
variation in communities and ecosystems, and the interactions among life
forms. In addition, biological diversity provides opportunities for adaptation to
future environments. Humans benefit in the long term. Our security is in
healthful environments that persistently support a diversity of life, including big
mammals on the same trophic level as us. Over millions of years of evolution,
such environments nurtured and sustained humans; they ignited our spirit and
imagination and fostered a will to explore and discover.
Humans now dominate practically eve ry livable terrestrial enviro n m e n t
on Earth; thus, it is up to us to care for our celestial home. Will we re s o l ve
to conserve environments that sustain us and a resplendent array of other
large mammals, including big, free-ranging carnivo res? Or will we decide to
l i ve in a world impoverished of those beasts? In the next one hundred years,
the human legacy on Earth will be defined. Humans can have a legacy of
wisdom and self-restraint, curbing our genetic propensity to maximize our
own re p ro d u c t i ve success and attendant appetite for re s o u rces so that a myr-
iad of life forms, including big predators, may persist in intact wild ecosys-
tems. This could be one of our greatest virtues, in league with treating all
humans with respect and dignity. Or will we prove we we re not wise enough,
365
366 PART V. PUMAS AND PEOPLE
and that in humanity’s glutting of Earth we continued to drive life to extinc-

tion?
We hope, this book will help people to understand what pumas require to
thrive and what their role is in nature. In previous chapters, we established sev-
eral reasons why the puma is a model keystone species on which to design land-
scape-level conservation strategies: (1) pumas strongly influence energy flow in
ecosystems; (2) they are a potent selective force on prey animals; (3) they mod-
ulate prey populations; thus (4) they indirectly affect herbivory on plant com-
munities; (5) they influence competitive interactions between herbivores; and
(6) their persisting populations are dependent on expansive wild landscapes.
Moreover, the puma is an umbrella species, because conservation strategies ben-
efiting pumas also benefit an array of other life forms living in intact ecosystems.
The time for developing and implementing conservation plans is now, while
pumas still exist in viable populations and there still are sizable wild lands. Con-
servation strategies should consider the needs of people as well as the ecology
and evolution of pumas.
In the United States, humans have devastated big obligate carnivores. We
compete with carnivores for both living space and animals that provide us food,
fiber, and revenue, whether they be livestock or big game. Furthermore, we have
a natural fear of big carnivores. Because of their size, strength, weaponry, and
diet, they can easily overwhelm us, and even eat us. Those instances were prob-
ably more common in our prehistoric past; today, they are extraordinary. Our
tendency has been to make our surroundings safe from such beasts. Jaguars
ranged from southern Louisiana to Southern California as late as the 1800s.
Today, jaguars are rare visitors to southern Arizona and southwestern New Mex-
ico, probably having dispersed from what may be a relict population in Sonora
Mexico. Throughout their range, jaguars are listed as endangered by the U.S.
Department of Interior (USDI) (Valdez 2000). Wolves ranged the length and
breadth of the Unites States into the 1800s, but now they too are on the endan-
gered species list. Due to federal protection and recent reintroductions, their
numbers have been increasing, particularly since the 1980s. In 1995, about
1,700 to 2,000 wolves lived in northern Minnesota, but only 100 to 120 lived
in small breeding populations in Wisconsin, Michigan, Montana, and possibly
Washington (Ballard and Gipson 2000). In 1999, the Greater Yellowstone
Ecosystem had about 120 wolves, and central Idaho wild lands had about 145
wolves (U.S. Fish and Wildlife Service et al. 1999). Forested mountains and
wilderness along the borders of Arizona and New Mexico had less than twenty
Mexican wolves in early 2000 (U.S. Fish and Wildlife Service 2000). About the
same time, coastal North Carolina was home to about ninety-five wild red
wolves (Hackney 2000).
CHAPTER 20. CONSERVATION AND MANAGEMENT OF WILD PUMAS 367
Of the big flesh-eaters, only the puma exists today in viable populations
across the western United States. But that is about half of its historical range in
this country. As we discussed in Chapter 2, regulations on killing pumas in west-
ern states since 1965 allowed puma populations to recover from historical lows.
Today, self-sustaining puma populations occur in at least twelve western states—
Arizona, California, Colorado, Idaho, Montana, Nevada, New Mexico, Oregon,
Texas, Utah, Washington, and Wyoming. Because pumas live at very low densi-
ties and are highly cryptic, wildlife biologists and managers can never really
know how many pumas may live in suitable habitats within these states. Still,
wildlife agencies for seven states have volunteered educated guesses at the num-
ber of pumas, which added up to at least 21,400 animals in the late 1990s
(Logan and Sweanor 2000). Assuming that the other five states had about 2,000
pumas each, then the number rises to about 31,400 wild pumas in the western
United States. Sprinkle those pumas among the mountains and foothills of the
West and one can fathom how sparsely these animals are actually distributed. In
the eastern United States, there is reliable evidence of only one breeding popu-
lation, that of the endangered Florida panther, which numbered about eighty
animals in 1990 (Maehr et al. 1991). In Canada, pumas thrive in Alberta and
British Columbia, where their numbers have been guessed at about 700 and
2,500, respectively (Logan and Sweanor 2000). Regardless of where wild pumas
exist today, they all face the same major threats to their continued existence, and
those are human made.
Threats to Pumas
There are two major threats to self-sustaining puma populations—habitat loss
and overkill. In this section we discuss these threats, and then follow with a sec-
tion on what people can do to alleviate them.
Habitat Loss
Habitat loss due to human development is the greatest single threat to puma
conservation. Anyone who has flown in an airplane over the western United
States, or crisscrossed it by car, knows human presence is ubiquitous. Burgeon-
ing cities, towns, and industrial complexes link to one another by highways and
railways like a gigantic web of humanity laid over the land, and most sizeable
gentle areas that remain are used for agricultural fields and pasture. Anyone who
has lived in the West in the past twenty years has seen the rapidity with which
urbanization gobbles up wildlife habitat. Foothills and mountain slopes that
bore no permanent human habitation back then are now dotted with houses or
invaded by urban sprawl. Millions of us know what puma habitat looks like after
it has been usurped, perforated, and fragmented for human use. If you live in,
or have traveled through, the eastern front of the Rocky Mountains between
Colorado Springs and Fort Collins, Colorado, or the Sandia Mountains east of
Albuquerque and the Sangre de Cristo Mountains north of Santa Fe, New Mex-
ico, or the Coast and Peninsular Ranges in Southern California, you have seen
the lasting structures we have built that destroy puma habitat and disrupt puma
movements. Today, the eleven western states where pumas range (excluding
Texas) comprise the fastest-growing region in the United States. From 1990 to
1998, the human population in the region grew by about 15 percent (U.S. Cen-
sus Bureau statistics 1999). By the year 2025, there will be about 25 million
more people living in those western states, with an additional seven million peo-
ple living in Texas (U.S. Census Bureau projections 1999).
There are more people living in puma habitat today than ever before, and the
influx continues. Greater human development contributes to smaller, more frag-
mented puma populations, increasing their risk of extinction and concomitantly
increasing chances for potentially dangerous interactions between pumas and
people. Remember Fig. 9-1 in Chapter 9? Imagine an ever-expanding web of
humanity as cities, suburbs, and roads advance on the landscape. From civilized
centers, more people venture out to recreate in the outdoors. Now think of
pumas living in the mountains and those subadult pumas dispersing in random
directions. It is not hard to see that as human developments and activity increase
so will the risk that pumas will encounter humans.
Extreme human development continues to threaten puma extinction from
almost all of its historical range in the eastern United States. Natural recolo-
nization, either from the isolated Florida population or from pumas in the West,
appears to be stifled by widespread habitat loss, habitat fragmentation, and
human presence even though ample prey (e.g., white-tailed deer, wild hog,
turkey, raccoons) seems to exist. In the East, outside of Florida, there is scant evi-
dence that individual pumas have tried to wrest an unfettered existence in recent
years. In the 1990s, the Eastern Cougar Foundation (a group of naturalists, con-
servationists, and scientists dedicated to finding and conserving pumas in the
East) documented only nine confirmed cases of pumas, with one each in New
Brunswick, Ontario, Maine, Maryland, Massachusetts, Missouri, North Car-
olina, Vermont, and West Virginia. Confirmations were based on one carcass,
two video tapes, bile salts extracted from feces in one case, hair identified in feces
in two cases, DNA extracted from feces in one case, visual observation and hair
in one case, and plaster casts of tracks in one case (Eastern Cougar Foundation
2000). Origins of these pumas were unknown; they may have been wild pumas,
captive pumas that were released or escaped into the wild, or captive pumas out
on a stroll in the woods with their owners (see McBride et al. 1993).
Pumas in Florida attest to the perils of living in a heavily peopled landscape.
Collisions with vehicles were second only to intraspecies strife as a major cause
of mortality (Maehr 1997a). Isolation of the Florida panther due to habitat
destruction for urban, agricultural, and industrial developments to the north has
limited the population to only about eighty animals, making it susceptible to
extinction due to some catastrophe (e.g., global warming and attendant flood-
ing) or demographic decline due to effects of inbreeding depression or genetic
drift. Prospects of expanding the population northward by direct human inter-
vention appear bleak. In 1988, seven wild pumas caught in west Texas were
released into northern Florida to study the feasibility of translocating pumas to
help recover the species there. These animals spent only 10 to 293 days in the
wild. Two of them were killed by people and a third was found floating in a river.
Not surprisingly, one puma was grazed by a car, since the seven pumas crossed
roads about 2,612 times during their movements (i.e., approximately 2.7 cross-
ings / puma-day). In addition, when human hunters took to the woods to pur-
sue deer, hogs, and turkeys, pumas with established home ranges were dislodged,
wandered, and entered urban areas and livestock operations where they killed
goats and exotic ungulates. This necessitated the early removal of the remaining
introduced pumas from the wild (Belden and Hagedorn 1993).
A second effort was made in 1993. This time nineteen Texas pumas, com-
prising both wild-caught and captive animals, were released into northern
Florida. These pumas were in the wild for 46 to 492 days. Although fifteen of
the pumas seemed to establish home ranges, only five (26 percent) remained in
the wild through the end of the study. Five died from human causes, including
collisions with vehicles (two), illegal shooting (two), and a neck snare (one). Five
were removed for killing domestic calves and exotic Sika deer (Cervus nippon).
Four other pumas (21 percent) were removed for a variety of reasons: landowner
complaints (one), public safety (one), a cub abandoned by its mother (one), and
an extreme excursion into Georgia where the animal was out of reach of the
research team. This last animal had also previously been wounded with an arrow
and might have died if not for medical treatment. In addition, of four cubs born
in the northern Florida release area to introduced females, two died from human
causes (collision with vehicle, capture accident), and one was removed with its
mother for killing Sika deer (Belden and McCowan 1996).
At the other end of the continent, conditions can be just as grim. In the
Santa Ana Mountains in Southern California, a population of roughly twenty-
five to forty pumas living in about 2,070 km2 of habitat was practically sur-
rounded by urbanization, except for two tiny, tenuous corridors that allowed
limited puma movement to other mountain ranges. Major impediments to
movements in those corridors were eight-lane and six-lane freeways (Beier
1995). Immigration was so restricted that females in the population lacked a
breeding male for about one full year (Padley 1990, Beier and Barrett 1993).
Vehicle strikes were the single-most important cause of death, killing ten of
thirty-one pumas that died during a five-year study. Another five pumas died at
the hands of humans: one was illegally killed, one was shot by police, and three
were killed to stop depredation on livestock (Beier and Barrett 1993). The biol-
ogist that studied the population concluded that unless connectivity is main-
tained to allow pumas to move freely between habitat patches in that region, the
puma population in the Santa Ana Mountains will probably go extinct (Beier
1993, Beier 1996).
In the Southwest, most of the biologically richest habitat and linkages—the
river bottomlands—have changed forever. Look at a map of New Mexico and
find the Rio Grande—its headwaters spilling out of Colorado, flowing south
through the middle of the state, and to the western boundary of Texas. In 1583,
Spanish explorer Antonio Espejo described the bosque along the Rio Grande in
about the middle of New Mexico as being four leagues (i.e., about 19 km) wide
in places (Bolton 1916). Bottomland woods growing along the shifting, sandy,
gravelly course of the Rio Grande provided an almost-continuous corridor
shaded by cottonwood (Populus fremontii), willow (Salix spp.), and walnut
(Juglans major) trees, interspersed here and there by marshes. Compared to the
adjacent upper desert basin, the bottomland was lush with prey, water, and
cover, providing a secure link for pumas moving long distances north and south
and serving as a lucrative temporary stopover for pumas moving east and west.
The bosque probably supported its own resident puma population. Pumas
could have preyed on the same animals that Native Americans living along the
river relied upon—deer and turkeys. We suspect too that it was a vital habitat
for jaguars, linking populations in New Mexico, Texas, and Mexico. That artery
is gone now—severely fragmented or completely transformed. Most of the
bosque has been chopped down and the floodplain channeled to make room for
crops, pastures, cities, towns, and highways. Local protection and restoration
projects, such as those of the U.S. Fish and Wildlife Service at Bosque del
Apache and Sevilleta National Wildlife Refuges in the center of the state, help
to maintain east-west links for contemporary dispersing pumas and a few indi-
viduals that use strands of extant bosque and the desert uplands as their home.
Our own research documented habitat fragmentation on desert mountains.
U.S. Highway 70 divides the San Andres Mountains and the Organ Mountains
at San Augustin Pass—once a narrow, natural link of mountainous terrain about
5 km wide. This stretch became a more treacherous filter to pumas moving

north and south along the mountain chain when the highway was expanded
from four to six lanes in 1993. Prior to expansion, at least seven of our radio-
collared pumas successfully crossed the highway, including four dispersing male
subadults, one adult male, and one subadult female that frequently traveled back
and forth between our study area and the Organ Mountains, and one adult male
that occasionally explored south of the highway. After expansion, the only doc-
umented puma crossings were of one tagged adult male and one unmarked
subadult male that were both struck and killed by vehicles. None of our five
radio-collared adult pumas with home ranges abutting the widened highway
dared cross it. The wider highway and the greater volume and speed of traffic it
carried probably discouraged pumas from moving between the San Andres and
Organ Mountains and reduced survival of those that tried. If the linkage is com-
pletely severed, pumas could eventually go extinct in the Organ Mountains, par-
ticularly because female immigration from habitat patches across the vast desert
basins to the east and west would probably be extremely rare (Sweanor et al.
2000). As human development expands along the Rio Grande floodplain and
the western edge of the Sacramento Mountains, natural movements of dispers-
ing pumas across southern New Mexico will dwindle. Across the Southwest,
where human impacts affect puma movements in this way, puma subpopula-
tions, particularly the small ones, will become less resilient. If extinction of sub-
populations accelerates, entire metapopulations will unravel.
Puma Overkill
Overkilling pumas threatens their conservation because it increases risks of
extinction, especially to small populations. It can also destabilize regional popu-
lations and metapopulations. Overkill results from a variety of factors, includ-
ing sport-hunting, illegal killing, accidents, and predator control. Hunting
pumas is also associated with illegal killing and unintended mortality, such as
orphaning of cubs and infanticide. Overkilling segments of puma populations
has other risks we cannot fully comprehend because human selection is differ-
ent than natural selection.
Humans are the major cause of death in most puma populations today,
except for populations protected from sport-hunting and control and those
occupying the most remote, unpeopled habitats. In such places, the most com-
mon cause of death, and hence selection, is from other pumas (see Chapter 8).
Sport-hunters are the major known cause of death of pumas in most Western
states. In 1996 alone, hunters in Arizona, Colorado, Idaho, Montana, Nevada,
Oregon, Utah, Washington, and Wyoming reported that they killed a total of
3,003 pumas (see Logan and Sweanor 2000: Table 17-1). Hunting seasons gen-
erally span the winter months when snow cover easily exposes puma tracks to
hunters. Ordinarily, trained dogs are released to trail pumas by scent in their
tracks. In the desert Southwest where continuous snow cover is rare, hunters
depend on dogs to hunt for puma scent. Odds of dogs successfully trailing
pumas on snow are very high compared to trailing pumas on bare ground. Once
the dogs catch up to the puma, the cat uses a strategy that it probably evolved
to successfully evade predation from wolves and coyotes. It climbs a tree or
promontory to get out of reach. This strategy must have worked successfully on
wild canids; a puma need only wait for its nemesis to leave, climb down from
the perch, and be on its way. But with human hunters, pumas are captive game.
For trophy hunters, the puma’s death is practically assured.
Humans caused 50 to 100 percent of puma deaths in study populations sub-
jected to sport-hunting in Alberta, Idaho, Montana, and Wyoming (Ross and
Jalkotzy 1992, Hornocker 1970, Murphy 1983a, Logan et al. 1986, respec-
tively). Similarly, humans caused 57 to 92 percent of deaths in study popula-
tions in Arizona and along the New Mexico and Texas border where populations
were subjected to heavy killing to protect livestock (Cunningham et al. 1995,
Smith et al. 1986). Humans were also the main cause of death in populations in
Arizona, British Columbia, and Colorado that were protected from sport-hunt-
ing for research purposes. In those populations, mortalities resulted from illegal
killing inside, and legal killing outside, study area boundaries (Shaw 1977,
Spreadbury et al. 1996, Anderson et al. 1992). Even in California and Florida,
where pumas were protected from hunting statewide but where habitats were
severely fragmented by human developments, vehicle collisions were an impor-
tant cause of puma deaths (Beier and Barrett 1993, Maehr 1997a). Trends in
population size could only be ascertained for two studies: the Alberta popula-
tion increased slightly over nine years, and the Idaho population declined
slightly in eight years. In the rest of the populations, human-caused mortalities
would not have had a numerical effect on population dynamics only if all of the
deaths were compensatory (i.e., if those deaths substituted for other natural
mortality factors), which is probably not likely. Our research (see Chapter 10),
as well as puma research in Utah (Lindzey et al. 1988, 1992), indicates that
human off-take is not fully compensated for. In Alberta, a depressed puma pop-
ulation recovered after a reduction in sport-hunting pressure, suggesting that the
killing was additive mortality (Ross and Jalkotzy 1992). Generally, uncertainty
about effects of human off-take pervades puma management.
Puma-hunting management in most western states is a far cry from science.
Except in rare, intensively studied populations, wildlife managers generally do
not have reliable population estimates on which to figure allowable harvest.
Normally they depend upon bag limits, quotas, and hunting-season length to
limit off-take, and hope the harvest does not overwhelm the population’s ability
to absorb the pressure. Some western biologists have perceived general increases
in puma populations since the early 1980s because there has been an upward
trend in the number of pumas killed by hunters each year. And hunters have yet
to complain about a “shortage” of pumas. Hence, agencies have had the propen-
sity to liberalize puma hunting by extending hunting seasons and increasing kill
quotas (see Logan and Sweanor 2000). Increasing puma harvest could be a
reflection of more hunters searching for pumas each year. For example, during
thirteen hunting seasons in New Mexico (i.e., 1984–1985 to 1996–1997), the
number of puma-hunting permits issued per annum increased from 443 to 980
(121 percent), and the number of pumas reported killed per annum (reporting
is mandatory in New Mexico) increased from 79 to 168 (113 percent) (Fig. 20-
1). We found a strong linear relationship between these variables. The number
of permits issued explained 80 percent of the variation in the puma harvest [1].
Similarly in Montana, the number of puma licenses sold explained 82 percent
of the variation in the harvest from 1971 to 1990 (Aune 1991).
Puma-hunting management in most western states appears to follow what
we call a “sledgehammer approach.” In this approach, hunting off-take is pro-
gressively liberalized until population declines affect hunters to the extent that
crude population indices based on harvest data—the number, sex, and spatial
distribution of pumas killed, hunter success rates, and hunter testimony—sug-
FIGURE 20-1. Trends in number of puma-hunting permits issued and number of pumas
killed by hunters in New Mexico.
gest that the puma population has declined. When that happens, presumably
human off-take has “hammered” the population into a decline phase, the ampli-
tude of which managers cannot fully ascertain. If decline is apparent, then man-
agers may decide that people have overkilled the puma population and correct
the course by backing off on hunting pressure. The main reason for this
approach is the lack of reliable, inexpensive techniques that managers can apply
at the landscape level to track puma population dynamics relative to the man-
agement prescriptions. Because pumas live at low population densities in com-
plex habitats and are highly secretive, managers cannot fly around in airplanes
and helicopters to count pumas and estimate their numbers like they do to sur-
vey more visible big game, like deer, elk, or pronghorn. But, harvest data can be
disconnected from what may be occurring with the puma population in the
wild. The sex structure of harvested pumas does not reflect the living popula-
tion. It is biased because hunters traditionally seek big males as trophies. Spatial
distribution of hunter kills may change from year to year as hunters pursue cats
in different areas weather patterns affect hunter success, previously hunted areas
have been overharvested, or traffic from hunters is perceived to be unbearable.
In some extremely heavily hunted puma populations, hunters are in such high
competition with one another that they assail snow-covered ridges and canyons
at night with four-wheel-drive rigs and snowmobiles looking for puma tracks
with spotlights so that when legal hunting hours arrive with morning, they will
be the first ones to release their dogs on the puma’s trail. Furthermore, managers
do not comprehend how technology may enhance hunter success. Since we
began studying pumas in the West in the early 1980s, puma hunters have
increased their use of snowmobiles and all-terrain vehicles to search more habi-
tat more thoroughly, radiotelemetry to track their dogs on puma trails more effi-
ciently, and two-way radios and wireless phones to coordinate efforts between
hunters. To what extent these tools mask downtrends in puma populations is
entirely unknown. Some hunters may be reluctant to report that they may be
causing declines in puma populations because that might bring tightening
restrictions. For outfitters and guides that may earn $3,000 per puma hunt, the
incentive to exploit the puma population is high.
In addition to pumas killed legally by sport-hunters, there is also “collateral
damage” that is sometimes illegal, sometimes accidental, and which is almost
impossible for wildlife law enforcement officers and managers to quantify. Puma
hunters we have met over the years have told us disastrous stories of incidents
occurring in the bush: hunting dogs track a mother puma to a nursery and maul
kittens to death (also see Roberson 1984); a hunter mistakenly judges the sex or
size of a puma in the tree, kills it, then abandons the carcass to pursue another,
bigger trophy. Hunters sometimes kill female pumas legally and unknowingly
cause the orphaning of cubs. Cubs orphaned when they are less than nine
months old usually die, and older cubs stand less of a chance of surviving. A few
states have sanctioned “pursuit seasons” where hunters are allowed to chase
pumas with their dogs but not kill them. But this may not be the presumed
benign activity it was thought to be. Roberson (1984) discussed some of the
abuses in “pursuit seasons,” which include mauling of cubs by dogs, purposeful
wounding of pumas to restrict their movements so they could be found again
during the kill season, and illegal killing of trophy pumas. There is also the pos-
sibility that repeated harassment of individual pumas by hunters for entertain-
ment or to train their dogs may cause deleterious physiological effects (Harlow
et al. 1992). We suspect that as hunting pressure on puma populations increases,
so do “collateral damage” and the degree of uncertainty concerning the effects
of human exploitation on puma populations.
Although most states that allow puma hunting attempt to protect mothers
by outlawing the killing of females with cubs at their sides (Logan and Sweanor
2000), this does not prevent their killing. Puma mothers are susceptible to har-
vest when they are not accompanied by their cubs. Research in Wyoming
showed that during the winter hunting season, mothers were away from their
cubs about 50 percent of the time that researchers with dogs encountered them
(Logan 1983). More so, researchers studying pumas in Utah found that cub
tracks were found with their mothers only 25 percent of the time; they con-
cluded that 75 percent of mothers would not be recognized by hunters (Barn-
hurst and Lindzey 1989). In addition, we observed that distended teats and
mammary glands are not a reliable clue for hunters. They are visible in mothers
only during about the first two months after cubs are born and not necessarily
right after cubs have suckled, which empties the tissues. Since as many as three
out of four adult females may be raising cubs each year, it is possible that roughly
38–56 percent of adult females killed by hunters each year are mothers with
dependent cubs.
Further, there are at least three “beliefs” about pumas from which some west-
ern wildlife managers operate that could spawn results contradictory to man-
agement objectives. These could lead to erroneous conjecture about local puma
density and thus to overkill of pumas or their prey. One assumption, that male
pumas disperse long distances and show up in marginal habitat areas because all
the surrounding puma habitat is “saturated” with pumas, we now know is false.
Male puma dispersal is obligatory; they will even move through habitat with a
suppressed population. Long-distance dispersal in male pumas probably evolved
to avoid inbreeding and carries them through all kinds of terrain that may be
prime or marginal habitat, even nonhabitat (see Chapter 13). Another false
assumption is that male pumas fight over territory because the habitat is “satu-
rated.” Thus stories heard from hunters about fighting pumas, as well as
observed battle scars on hides from harvested cats, reinforce the notion that the
puma habitat is full. We realize now that male pumas fight to compete for access
to mates and that the rate of fighting can be independent of puma density (see
Chapter 8). Finally, some managers believe that since pumas limit their own
populations below the level set by the prey, management of human harvest of
prey populations does not necessarily have to consider effects of puma preda-
tion. Pumas will readjust their populations accordingly. However, now we
understand that pumas can limit or hasten a decline in a prey population. Thus,
puma predation along with sport-hunting could reduce prey populations. Puma
populations will likely decline after a lag period, which could be a few years after
a decline in the prey base (see Chapter 17). Furthermore, additive mortality on
prey populations imposed by sport-hunting could exacerbate declines in both
prey and puma populations.
Suppressing local puma populations may be a desired management objective.
Wildlife managers may be pressed to reduce puma numbers in areas where
encounters threaten the safety of humans and their pets (see Aune 1991, Half-
penny et al. 1991, Torres et al. 1996). Puma attacks on humans are relatively
rare. However, the rate of puma attacks has increased since 1970. Between 1890
and 1996 about fifteen people were killed and about fifty-nine people were
injured during sixty-six attack incidents in the United States and Canada com-
bined. From 1970 to 1996, ten people were killed and forty-five injured in
forty-nine attacks (Torres 1997). Two interacting reasons for the increase are
that pumas generally expanded in numbers and distribution during that period,
while numbers and distribution of humans living and recreating in puma habi-
tats exploded (Aune 1991, Torres et al. 1996). Managers may also prescribe
puma control programs to quell chronic depredation on livestock. For example,
state-sanctioned puma control to protect domestic sheep has been operating in
the Guadalupe Mountains in southeastern New Mexico since 1985. In addition,
puma control may be deemed necessary to help recover endangered ungulate
populations. For instance, in 1999 a puma bounty hunting program and
extremely high harvest quotas were prescribed for southwestern New Mexico by
the New Mexico Department of Game and Fish because puma predation was
identified as the principal proximate cause of death in the state-endangered
desert bighorn sheep. Because reduction programs create local population sinks,
wildlife managers need to monitor their region-wide effects. In addition, these
programs probably reduce the number of dispersing pumas available to augment
other subpopulations. Hence, depending upon the number, size, and distribu-
tion of population sinks, attendant overkill could threaten the resilience of a
population or metapopulation.
Another source of uncertainty is that human-caused death is a different

selection process than natural selection. Large-scale hunting of pumas by
humans, especially for sport and population control, has influenced puma pop-
ulations for only about the past two hundred years—about 0.05 percent of the
time since extant puma lineages emerged. Humans generally kill the very indi-
viduals that nature would have selected to propagate the species—big males and
females. Based on the life history and behavioral strategies that pumas appar-
ently evolved (see Parts 2 and 3), the most successful pumas should include
large, robust males and reproductively successful females. Trophy hunters seek
the biggest males; hence, they remove individuals that are likely the most fit. Big
males possess adaptive traits along with their large size, which allow them to suc-
cessfully survive in the environment, compete with other males for mates, and
father thriving young that inherit their “good genes.” Furthermore, the killing
of territorial males by hunters may increase infanticide of cubs by new immi-
grant males. This reduces reproductive success of males that won the competi-
tion to breed, and it reduces the reproductive success of females. If big males
cannot be found, then hunters may kill big females instead, the very ones that
are probably most reproductively successful in North American environments.
Larger females are probably better able to protect their offspring against
marauding male pumas and other predators and to exploit a greater variety of
large prey to more efficiently feed their young.
What is the genetic, and thus the adaptive, cost of humans changing the
selection process for pumas? We presume that killing individuals that natural
selection would have otherwise chosen to survive and reproduce will tend to
reduce fitness. Researchers investigating genetic variability in bighorn sheep, a
species with a breeding system having characteristics similar to that of pumas
(i.e., polygynous, sexual selection), have illuminated one possible mechanism for
decline in fitness. Large-horned rams, which have breeding superiority, were
more heterozygous than small-horned rams. They concluded that harvesting
large-horned rams could reduce genetic variability and thus contribute to loss in
fitness in bighorn sheep (Fitzsimmons et al. 1995). Because natural selection
produced the traits that allowed pumas to persist today, we believe that natural
selection processes should proceed at least in some large wild areas (see “Zone
Management” in this chapter). Doing so may help conserve biological diversity
and the resilience of puma populations.
Alleviating Threats
It is relatively easy to complain about the factors that threaten puma conserva-
tion. Devising solutions is much more difficult and risky. But if, as a society,
people decide to conserve self-sustaining puma populations and their wild envi-
ronments, then it is going to require an informed and involved public.
Habitat Conservation
Pumas require huge, wild, connected landscapes to support even minimum self-
sustaining populations. Flourishing habitats have sufficient persistent prey
sources and a complex terrain and vegetative structure that provides cover for
stalking, security, and nurseries (see Logan and Irwin 1985, Laing and Lindzey
1991, Logan and Sweanor 2000). The wild landscape has to be big enough to
accommodate breeding adults during their daily movements to hunt for food,
seek mates, and avoid other pumas or predators, and it must provide for any
needed seasonal habitats and movements of migrating pumas and prey. The
landscape also has to have sufficient connectivity to accommodate natural dis-
persal patterns of puma progeny. As we discussed in Chapter 10, a minimum
habitat area of 1,000–2,200 km2 is needed to sustain a nonmigratory puma
population with a greater than 98 percent probability of persisting for one hun-
dred years (Beier 1993), and that area needs to be linked to other puma-occu-
pied habitats. Puma populations in smaller habitat patches without immigration
have relatively high extinction risks. For example, twelve of one hundred simu-
lated puma populations went extinct over a one-hundred-year period for habi-
tat areas of 270 km2, the size of the Organ Mountains (Sweanor et al. 2000).
Without immigration, small populations cannot be augmented numerically and
thus are subject to greater risks of extinction from demographic and environ-
mental uncertainty and natural catastrophes. Demographic uncertainty is risk
resulting from random events on survival and reproduction of individuals. Envi-
ronmental uncertainty refers to chance events like changes in weather, food sup-
ply, competitors, predators, or disease (Meffe and Carroll 1997:217). Natural
catastrophes could be hurricanes, extensive wildfires, extensive human develop-
ments, and global warming. Furthermore, without gene flow from immigrants,
inbreeding and genetic drift may erode genetic variability, viability, and repro-
ductive vitality. Pumas could lose the ability to adapt to altered environmental
conditions (Chepko-Sade et al. 1987, Gilpin 1987, Lande 1988, Saccheri et al.
1998). Clearly, immigration is essential for numeric and genetic augmentation
in populations, especially small ones.
In North America, pumas depend primarily upon large prey such as white-
tailed deer, mule deer, elk, moose, peccary, and bighorn sheep. Yet they augment
their diet with a variety of small prey such as rabbits and rodents. In highly vari-
able environments like the desert Southwest, wildlife managers (i.e., profession-
als trained in wildlife sciences and government-appointed secretaries, directors,
commissioners, and board members responsible for overseeing wildlife manage-
ment agencies) should recognize that prey populations may fluctuate dramati-
cally with changes in weather and climate. As we have pointed out, pumas are
intrinsic participants to predator-prey dynamics in ecosystems. Their numbers
are ultimately limited by their own food supply. During stress periods such as
drought (or severe winters in more northern climes), pumas can hasten a decline
in their prey, and, after a lag, the puma population must also decline. This will
occur especially where alternate sources of prey are not available to sustain the
puma population. Resilience of the puma population is tied to the ability of the
main prey population to recover from a severe downturn.
In stress periods, managers should monitor big-game populations more
closely, because they may need to adopt more conservative harvest prescriptions
to prevent even steeper declines in the puma prey base. Remember, human
hunters are in competition with pumas for prey animals (also see Murphy et al.
1999). Yet, most human hunters today are extrinsic predators to an ecosystem.
Unlike pumas, humans generally come from outside of ecosystems in which
they hunt, and then haul their kill back out (except in subsistence situations).
Furthermore, humans are not totally dependent on hunting success to survive
and reproduce. Hunter selection may also counter natural selection, since
healthy game are typically killed for food or trophies, and high-tech weapons
(i.e., firearms, telescope sights, compound bows) make it more likely that ani-
mals are killed that might otherwise survive and reproduce. Hence, human off-
take is probably mainly additive mortality that can reduce a population further
if it is already suppressed by adverse habitat conditions (e.g., drought, severe
winters, and attendant loss of food) and predation.
Puma habitat conservation should be a combination of protecting habitat
patches and the landscape linkages that join them. In the Southwest, where
puma habitat comprises mountains separated by broad desert basins, this strat-
egy is crucial to puma conservation. Here, pumas may exist in a metapopulation
where long-term survival of puma subpopulations is dependent on the move-
ment of pumas between habitat patches (see Chapter 10; Sweanor et al. 2000).
Dispersing pumas augment subpopulations by moving through habitat and wild
land or rural matrix. In this structure, the relatively resource-barren basins (i.e.,
matrix) and even the small habitat patches, such as the Fra Cristobal and
Caballo Mountains, which are within puma dispersal distance, facilitate
numeric and gene flow to more distant and substantial habitats such as the Black
Range and Gila Wilderness. Female dispersers that do not reach the more dis-
tant patches disperse to closer patches and reproduce. Then their progeny dis-
perse to other patches. For male pumas, wild land matrix, wild land corridors,
and habitat patches are essential pathways for their much-longer dispersals.
Although it may take a few generations, an individual puma’s genes can flow
across the breadth and length of the basin and range physiography of New Mex-
ico, where habitat patches and their tenant puma populations function as step-
ping stones along the way (Sweanor et al. 2000). In huge, contiguous habitat
blocks, like Rocky Mountain masses, the same thing occurs, except dispersing
pumas are almost always moving through habitat. Throughout the first 99.95
percent of puma evolutionary history, pumas could disperse from their natal
areas in random directions with relatively little human interference, colonizing
habitats, and developing interbreeding subpopulations. They evolved into an
outbred species. It is these characteristics that made the puma such a resilient
carnivore in changing environments.
In the West, if self-sustaining, interconnected puma populations are to per-
sist as they do today, then wildlife managers need to immediately identify, map,
and conserve present puma habitat and landscape linkages. Already state and
federal wildlife agencies have baseline information to begin the mapping
process. There is statewide information on distributions of pumas, puma har-
vest, depredation incidents, and puma-human encounters to help identify occu-
pied puma range and potential landscape linkages. Wildlife agencies can also tap
into the expertise of their own local biologists, landowners, hunters, and natu-
ralists regarding puma activity. And they can rely on the intuition of puma biol-
ogists who have studied behavior, habitat use, and movement patterns to help
them identify potential links. Where critical questions arise regarding whether
or not pumas actually move through segments of wild lands, there is no better
source than the pumas themselves. Just as we learned from our research, radio-
collared pumas provide detailed information on habitats and links that they use:
subadults trace dispersal patterns that can eventually take them to other sub-
populations, and adults delineate migration routes. Computerized geographic
information systems are available for creating databases and maps from all of this
information to assist wildlife managers in formulating conservation strategies for
puma habitat at local and regional landscape scales.
If pumas are to thrive in the East, there will also need to be an increased
human tolerance toward this carnivore, and robust public support for habitat
protection, habitat restoration, and modification of many of the human-built
impediments to puma movements. There will need to be public or private funds
generated to compensate people for puma predation on privately owned ani-
mals. Wildlife agencies will need to maintain well-trained, rapid-response cap-
ture teams to react to emergency situations to safeguard humans, private prop-
erty, and pumas.
Puma habitat and wild-land linkages are a patchwork of public and private
lands. Protecting and managing them will require partnerships at all organiza-
tional levels of our society—federal, state, county, and municipal governments,
private organizations, and individuals. Public domain, which includes national

forests, Bureau of Land Management lands, national parks, national monu-
ments, national wildlife refuges, federal lands under the authority of U.S.
Department of Defense, state parks, and state wildlife management areas, are
the foundational habitats that sustain puma population in most of the West,
principally because those lands have the least human development. Rural and
undeveloped private lands (e.g., farms and ranches) furnish vital needs too, par-
ticularly those that function as movement and migration corridors for pumas
and their ungulate prey. However, in Texas, most of the puma habitat is com-
posed of privately owned land. In southern Florida, the healthiest and most pro-
ductive Florida panthers primarily inhabit private lands (Maehr 1990b). Pro-
tection of puma habitat on public domain can already be supported through
current rules and laws that govern wildlife conservation. On multiple-use pub-
lic domain lands, extraction of resources such as timber and minerals should be
planned to minimize negative impacts on pumas and their ungulate prey, par-
ticularly in critical areas where activities on adjacent unprotected lands have
destroyed or fragmented previously existing habitat or linkages. Ha b i t a t
improvements can also be made, for instance by prescribing controlled burns to
benefit ungulate prey. Outdoor recreational structures and facilities (e.g., camp-
grounds, roads, trails) could be designed to reduce risks of puma encounters and
habituation to human activities (Ruth 1991, Murphy et al. 1999). On the other
hand, local habitat modifications could be used to deter puma use of major
human activity areas and to minimize predation on endangered animals (e.g.,
cutting or prescribed fire to reduce shrub cover that attracts herbivores and pro-
vides stalking cover for pumas).
Private lands can be a problem, particularly where owners are not obligated
to protect habitat for large carnivores, either because they harbor personal ani-
mus toward predators or because other land uses, such as residential, agricul-
tural, or industrial development, yield greater monetary returns. In those cases,
land stewardship programs with alternate incentives should be developed for
landowners to compensate them for habitat protection. Government or private
organizations could purchase land or lease easements and designate them for
wildlife use. Substantial tax breaks on capital gains could be given to landown-
ers that sell land for the preservation of wildlife habitat. For current landowners
already dedicated to wildlife conservation, relief from inheritance taxes could
sustain that legacy in their families. Otherwise, heirs may opt to sell wild and
rural lands to opulent developers to offset paying high taxes. Controls will be
needed in some regions to prevent urban developments from invading critical
wildlife habitats. Support for conservation-oriented government programs
requires strong public support to push through needed legislative action or ref-
erenda. If the public at large believes that puma habitat and wild land linkages
should be protected, then the public should share the costs of protecting those
lands.
The same can be said for restoring linkages that enhance the persistence of
puma populations. An advantage is that restoration efforts aid other large animals
as well. Florida panthers benefited directly from wildlife underpasses and fencing
developed along a 64-km stretch of Interstate 75 to reduce collisions with
wildlife. Underpasses allowed resident panthers to link parts of their home ranges
and prevented the highway from becoming a population sink. Other mammals
also used the underpasses, including bobcats, raccoons, black bears, white-tailed
deer, and alligators (Alligator mississippiensis) (Foster and Humphrey 1995).
Pumas used wildlife crossing stru c t u res in Banff National Park, Canada, where
their home ranges were bisected by the Trans Canada Highway (Gloyne 1999).
Some of those same stru c t u reswere also used by elk, deer, and wolves (Clevenger
and Waltho 2000). In southern New Mexico, highway underpasses would pro b-
ably reestablish a secure link between the San Andres and Organ Mountains.
Those stru c t u res are expensive to build and thus re q u i re substantial public sup-
port to justify the expenditures. Designing them into highway development or
improvement plans from the outset would reduce the long-term costs, both in
money and to wildlife populations. In terms of dollars, it would be more expen-
sive to go back to U.S. Highway 70 at San Augustin Pass, rip out the highway at
strategic locations defined by habitat features and animal movement patterns,
and install wildlife underpasses. Still, they may be needed to restore safe travel for
mammals moving between the San Andres and Organ Mountains.
Across the West today, there are puma habitats and wild-land linkages that
have been destroyed and degraded; yet, there is still ample habitat in the public
and private domain to support thriving puma populations. Now it is up to
wildlife managers to inventory the remaining puma habitat and landscape link-
ages. It is the responsibility of the caring public to see to it that this is done.
Habitat and linkages can only become part of the working structure for puma
conservation if they are mapped and people can conceptualize their worth. Then
wildlife professionals and interested citizens can look for and quantify the
threats relative to human developments and devise strategies and enact programs
for their conservation. Pumas and a myriad of other wildlife species will benefit
and so will the quality of life for humans with strong obligations toward con-
serving wildlife, intact ecosystems, and biological diversity. In addition, habitats
and linkages that benefit pumas will enhance chances for successful restoration
of other big carnivores, such as gray wolves in the Rocky Mountains and Mexi-
can wolves and jaguars in the Southwest.
Preventing Unnecessary Overkill

Ultimately, social values determine whether or not managers and scientists are
allowed to manipulate puma populations to achieve management objectives or
to test research hypotheses. Killing individual nuisance pumas that threaten
humans and livestock is understandable, because they directly affect people’s
individual well-being, and possibly their survival. But controlling puma popu-
lations to save endangered species or to boost ungulate populations to improve
recreational hunting opportunities agitates controversy. Before proposing puma
control for those reasons, we believe wildlife managers should ask themselves at
least these basic questions: (1) Is puma predation the most important limiting
factor suppressing the ungulate population? (2) Where is the ungulate popula-
tion relative to the ecological carrying capacity of the habitat? If the population
were above carrying capacity, it would be expected to decline even with puma
control. If it were substantially below carrying capacity, then the ungulate pop-
ulation may increase, provided puma predation is mostly additive mortality. (3)
To what extent is puma predation compensatory or additive? (4) Is puma con-
trol to save a small population of animals contributing to conservation of bio-
logical diversity? (5) In the case of endangered species restoration (e.g., bighorn
sheep), to what extent would numerical augmentation of the imperiled popula-
tion offset predation? (6) Are there other options?
In addition, there are questions that managers should ask themselves about
assessing the effectiveness of control actions: (1) How much do I need to reduce
the puma population to attain the objective? (2) How will I know when I get
there? (3) How long do I need to suppress the puma population? (4) How will
the creation of a population sink affect other adjacent puma populations? Infor-
mation on effects of puma control programs is scant, but answers to some of
these questions could be attained from onsite research. That is why we strongly
urge that puma control programs be carried out as experiments that test a priori
hypotheses and include treatment and reference areas.
To know what impact human off-take is having on pumas, wildlife managers
must monitor puma populations in their habitat. Unfortunately, managers lack
monitoring techniques with known accuracy, precision, and bias. A puma pop-
ulation estimator involving sampling puma tracks on snow from a helicopter has
been tested but found to be very imprecise (Van Sickle and Lindzey 1991).
Besides, almost continuous snow cover is needed, which is rare in most of the
Southwest, and can be exceptional even in northern latitudes with chronic high
winds and variable snowfall. Counting puma track on roads in southern Utah
was found unreliable because of the poor relationship between puma density
and track-finding frequency (Van Dyke et al. 1986, Van Sickle and Lindzey
1992). Biologists studying the utility of puma track surveys in Arizona found
that track surveys probably will not detect small population fluctuations but
may be useful in detecting 30 to 50 percent changes if 30 to 190 transects, each
8 km long, are established in dry washes. They estimated that 700 transects
would be needed to detect a 10 percent decrease in a puma population and that
the expense for such efforts would be prohibitive (Beier and Cunningham
1996). Thus, it appears that puma populations would fluctuate substantially
before wildlife managers could measure the trend with any skill (i.e., the “sledge-
hammer” effect). Another method tries to identify individual pumas by meas-
urements of their tracks in the field. It has discriminated individuals in specific
research situations, but to do so the technique requires measurements of a large
number of tracks from the same individual walking on similar substrates, con-
ditions difficult to achieve in the field (Smallwood and Fitzhugh 1993, Grigione
et al. 1999). To our knowledge, this method has not been applied at the land-
scape level. Obviously, there is a need to refine all these track indices to reflect
puma abundance. Other promising, noninvasive techniques for estimating pop-
ulation size involve genotyping individual animals from tissue samples. Ernest
et al. (2000) has genotyped individual pumas by their feces in California. Sim-
ilar techniques were used to estimate population size in coyotes in the same state
(Kohn et al. 1999). Techniques recently developed for lynx (McDaniel et al.
2000) and grizzly bears (Mowat and Strobeck 2000), using DNA from hair
roots that have been snagged at bait stations in conjunction with mark-recap-
ture analysis, could be modified to estimate puma population size. These meth-
ods should someday be applied to an intensively studied puma population to
test their usefulness as indices to abundance. In the meantime, managers should
assess puma population trends with a suite of indices, including track counts,
harvest data (i.e., catch per unit effort), depredation complaint frequency (Tor-
res et al. 1996), and testimonials from reliable people that routinely contemplate
puma abundance in habitat. Until better monitoring tools are operational, we
advise that puma sport-hunting prescriptions have conservative harvests of male
pumas and protections for female pumas, regardless of whether or not they are
raising cubs.
Adaptive Management—Involving People

What can be done to provide for long-term puma conservation and the needs
of people? Earlier in this chapter, we emphasized the need to appeal to people’s
sense of responsibility, to conserve puma habitat, and to protect against puma
overkill. Dealing with those, however, will be successful only inasmuch as peo-
ple’s values toward land, animals, and other resources are acknowledged. Hence,
an adaptive, landscape-level management approach that derives from reliable
ecological information and public input is a defensible solution. Such an
approach requires that wildlife professionals have reliable management tools and
support from an informed, caring public.
ZONE MANAGEMENT
Zone management was the adaptive management approach we proposed for
pumas in New Mexico (Logan and Sweanor 1998). This involved partitioning
the state into zones with different management objectives for puma populations
in them as determined by wildlife managers and public stakeholders (Fig. 20-2).
Management in each zone should be approached as an experiment, with its own
objective(s), hypotheses, and prescriptions. But because wildlife managers can-
not afford to monitor all zones, a monitoring program should be established, at
least in select zones of each category. Managers can thus gauge their success in
meeting objectives and adjust management prescriptions accordingly. Quotas
should limit the annual puma kill in zones that allow population control and
sport-hunting. Quotas would be estimated by agency biologists as the number
necessary to achieve population objectives. Rates of puma population increase
2IAÝ iÝ Âï®
FIGURE 20-2. A hypothetical zone-management approach for pumas in New Mexico.

estimated in this study could be used as reference points for calculating first quo-
tas from initial population estimates.
We developed three zone categories: Control Zone, Hunting Zone, and
Refuge Zone. Control Zones allow for experimental puma population reduction
in localized areas where it has been deemed necessary to substantially suppress
puma numbers to protect human safety, private property (i.e., livestock, pets),
and endangered species (e.g., desert bighorn sheep). Puma control could be
accomplished by heavy sport-hunting pressure or killing by government agents.
Regardless of the quota, individual pumas involved in human safety or depre-
dation incidences could still be killed. Hunting Zones allow for sustained sport-
harvest with the objective of maintaining puma population stability (i.e., pre-
venting decline). All females should be protected with lower quotas than those
put on males, and all cubs should be protected. Light hunting could be allowed
in areas used to link Hunting Zones and Refuge Zones. Refuge Zones would not
allow human off-take, except individual pumas that threaten human safety and
private property. Refuge Zones would function as robust biological savings
accounts that contribute to puma population resilience (Weaver et al. 1996) by
countering management-related mistakes that are probably going to occur from
time to time in human-impacted populations (McCullough 1996). Refuges also
allow natural selection to act on pumas. Puma genotypes in refuges can emigrate
and become breeding members in other exploited populations. Our empirical
data on puma demographics suggests that Refuge Zones should be at least as
large as the San Andres Mountains chain, about 3,000 km2, and should be con-
nected to other habitat patches via landscape linkages (see Chapter 10, Sweanor
et al. 2000). Refuge Zones also function as reference areas against which man-
agers can compare puma population dynamics in Control and Hunting Zones.
For the state of New Mexico, we recommended that there be at least two Refuge
Zones—one in the north and one in the south (Fig. 20-2).
Applying this approach, or any other, requires that wildlife managers obtain
available information on puma numbers and distribution in the state. They
should map out potential puma subpopulations that are sources, sinks, and vul-
nerable to extinction because of small size or poor connectivity. Known and
potential landscape linkages should also be mapped. Then, with consent from
substantial stakeholders, they should proceed with developing management
objectives for zones and with mapping them.
The number and juxtaposition of Control, Hunting, and Refuge Zones
should be decided carefully to assure that management in one zone does not
jeopardize the management of an adjacent zone. Especially, boundaries of a
Control Zone should not adjoin boundaries of a Refuge Zone because pumas
killed in the Control Zone could have home ranges that overlap the Refuge
Zone. In other words, the border between the Control and Refuge Zones could
act as a sink for the Refuge Zone puma population (Woodroffe and Ginsberg
1998). This may have occurred when New Mexico wildlife managers put puma
zone management into operation in fall 1999. Most of southwestern New Mex-
ico was made a Control Zone (principally to protect desert bighorn sheep), with
the eastern boundary abutting the San Andres Mountains chain, which was
made a Refuge Zone. Managers considered the western boundary of White
Sands Missile Range (which consists of a cattle fence) to be a reasonable demar-
cation between the two zones. Yet, hunters can kill pumas on about 93 km2 of
habitat on the western edge of the San Andres chain. Thus, killing pumas on the
Control Zone side would actually exploit the Refuge Zone puma population
because, based on our research, most of the adult pumas ranging in those 93
km2 have home ranges that overlap the Refuge Zone. This problem could be
easily rectified by moving the Refuge Zone boundary west to the north-to-south
railroad line that bisects the Jornada del Muerto, a basin that clearly separates
patches of puma habitat and thus the puma subpopulations. Still, progeny dis-
persing from Refuge Zones would be expected to have substantially lower sur-
vival rates if they immediately entered a Control Zone. Likewise, Control Zones
could reduce immigration into Refuge Zones. Areas between Control and
Refuge Zones could be buffered by conservatively harvested Hunting Zones. In
the Southwest, desert basins along Refuge Zone boundaries could serve as nat-
ural buffers. Clearly, monitoring the impacts of people on puma populations in
zones will be needed to assess and modify zone boundaries and zone manage-
ment prescriptions.
Puma off-take, whether to sustain the population for sport-hunting or to
depress it for control, should be gauged relative to the rate at which the popula-
tion tends to increase (Caughley and Sinclair 1994). We suggest that the
observed rates of increase for our January population estimator (which coin-
cided with most puma hunting seasons in the West) could be used as initial ref-
erence points (see Table 10-8), provided two conditions are met: (1) wildlife
managers have a reliable estimate of puma numbers, and (2) the puma popula-
tion is increasing. We recommend using the rates of increase for adults in the
Reference Area as a guide, because they covered seven years (1989–1995) in
which puma carrying capacity fluctuated. Rates of increase in those conditions
may be reasonable averages with which to start. If the objective is to harvest a
population to provide sport-hunting opportunity, then we recommend an ini-
tial harvest rate of less than 8 percent of adult males. Female pumas should be
protected as a safety valve against overkill for two main reasons: (1) error in
puma population estimates are generally not known, and (2) population moni-
toring methods can be very imprecise and may only detect 30–50 percent
changes in puma numbers. A nominal female quota is desirable to legalize the

killing of females that are mistaken as males by hunters and individual females
encountered in the field by chance but which provide no opportunity for sex
determination before hunters kill them. Thus hunters would report those
females to be tallied by wildlife officials without fear of legal reprisal. With a
quota on males and a smaller subquota on females, a Hunting Zone would be
closed when either the male or female quota was reached. By virtue of the dis-
proportionate quotas allotted to each sex, hunters are compelled to carefully
choose males.
In Hunting Zones, we discourage harvesting a puma population at the max-
imum rate of increase for the adult population (e.g., 11 percent), because the
margin for error is small. The puma population would be harvested at the brink
of population collapse. Furthermore, environmental perturbations, such as
drought and prey declines, could cause a decline in the puma population growth
rate. This scenario is particularly realistic for the relatively variable puma habi-
tat in the Southwest. In such instances, the population may decline without har-
vest. During puma population decline, off-take, particularly at high rates, would
probably accelerate population decline.
On the other hand, if the objective is to reduce a puma population, then the
adult puma population should be harvested at a rate exceeding the rate of
increase (e.g., greater than 11 percent); both males and females should be
removed. And based on our results from the Treatment Area it may be necessary
for off-take to exceed 28 percent of the adult population to cause a decline. But
managers should be sensitive to the possibility that extended off-take at such
high rates may cause local extirpation of pumas.
The major problem wildlife managers have with developing a harvest strat-
egy is that they seldom (i.e., only in populations subjected to intensive research)
if ever (i.e., all other cases) have reliable estimates of puma numbers in any
region of interest. Zone management is an adaptive approach that addresses an
array of puma management issues—from population control to protection—
through public input, experimentation, population monitoring, and refuges
that buffer management mistakes. But zone management is not a panacea; there
are other approaches wildlife professionals and the public should pursue to aid
puma management and conservation.
RESEARCH
Research is needed in manifold areas. Whenever possible, a priori hypotheses
should be tested and empirical data gathered in a manner for inclusion into
models (e.g., population models, habitat models). Biologists should attempt to
identify key patterns to develop theory about natural phenomena, steer man-
agement and conservation strategies, and guide future research. Below, we pro-
pose urgent research needs.
• As mentioned previously, there is a need for accurate, precise, and relatively
inexpensive methods of monitoring puma population trends in habitat.
Wildlife managers require such tools to gauge impacts of population manage-
ment (i.e., sport-hunting, control) environmental perturbations (e.g.,
drought, prey fluctuations, habitat loss), and disease outbreaks.
• Puma habitat use should be studied to help wildlife managers identify habi-
tat and landscape linkages. This type of information is essential for designing
nature re s e rves intended to accommodate pumas and other large carnivo res
and for land managers assessing how human developments, resource extrac-
tion (e.g., logging, mining), and habitat modification (e.g., prescribed burn-
ing) may degrade or enhance puma habitat. It will also help to highlight are a s
of potential puma-human encounter and could be used in designing trail sys-
tems and campgrounds in parks and wildlands that minimize encounters. It
may also help managers plan habitat alterations that discourage puma use of
c e rtain areas to reduce risk to endangered species, livestock, pets, and
humans.
• Puma behavior in peopled landscapes, such as housing developments and
heavily used recreation areas, needs to be investigated to understand to what
extent pumas may avoid or tolerate human activity. Likewise, we could learn
what kinds of human behaviors or habitat modifications may attract or repel
pumas. Databases on puma-human encounters should be maintained to
divine patterns in encounters and to identify hot spots and trends.
• Long-term research should address puma population dynamics, particularly
how human off-take, prey fluctuations, and habitat loss affect them.
• The genetic structure of regional puma populations or metapopulations need
to be deciphered so that we can understand gene flow and how human mod-
ifications to habitat and landscape linkages may affect it.
• Relationships of pumas to their prey need to be studied, particularly where
puma predation may threaten endangered animals, such as desert bighorn
s h e e p. It is also important to understand to what extent puma predation lim-
its or regulates prey populations (see Caughley and Sinclair 1994:110–130)
and to determine the importance of prey switching in multi-prey systems.
These are especially important for ungulates that humans also impact
t h rough hunting and habitat modification. Such information is fundamental
for scientific management of puma and ungulate populations. Studies of
pumas and ungulates will probably have to be at least twenty years in dura-
tion to include the full range of population fluctuation (i.e., increase, peak,
decline, low) of both predator and prey. As we learned ourselves, ten ye a r s

was barely long enough to see drastic fluctuations in the deer population and
not long enough to see the numerical response of the puma population to the
deer population crash. Mo re ove r, we we re not on hand to document the bio-
logical extinction of the desert bighorn sheep. Experimental re m oval of prey
or pumas could be used to test hypotheses about how puma predation affects
p rey and vice versa.
• Additional research on translocated pumas should be done to learn more
about their behavior and survival. This could be done in concert with removal
experiments in puma-prey interaction studies. Removal of pumas from treat-
ment areas does not have to be lethal for all individuals. After all, the cats are
killing natural prey. Serious consideration and planning should be given to the
translocation of select pumas following guidelines suggested by Ruth et al.
(1998) and Belden and McCowan (1996). Those guidelines should be
improved with additional research findings.
• Competition between pumas and other carnivores needs to be studied. Cur-
rent research projects on pumas, wolves, and coyotes in the Northern Yellow-
stone Ecosystem, pumas and wolves in the central Idaho wilderness, and
pumas and jaguars in Mexico should provide vital information on effects those
carnivores have on ungulate populations as well as how they influence one
another. Once Mexican wolves are well established in the Southwest, similar
research should be done there too. Furthermore, the extent to which humans
compete with carnivores should be assessed along with our impacts on prey
populations.
• Aversive conditioning of pumas should be tested, such as using trained dogs
to drive pumas away from humans and our focal activity areas, such as homes
and campsites. Karelian bear dogs have been used in recent years to shepherd
bears from human activity areas in national parks and forests (Hunt 2000).
Perhaps an alternate breed would be the Rhodesian ridgeback, which was orig-
inally used to hunt African lions. Such guard animals may be a realistic option
for people who live or recreate in puma habitat to minimize dangerous
encounters and the need to kill pumas.
• Similarly, research should be done on protecting livestock from pumas using
guard animals. Guard dogs (e.g., Akbash, Anatolian, Komondor) reduced coy-
ote predation on domestic sheep and reliance of producers on lethal predator
control (Andelt 1992, 1999).
• Human dimensions should be studied to better understand people’s beliefs,
values, and preferences regarding pumas. For example, a 1999 study in Utah
indicated that the use of predator control actions to protect game populations
had only moderate support, and the use of sport-hunting to control black
bears and pumas, the use of dogs to hunt bears and pumas, and the use of bear
baiting were not supported by the majority of Utahans (Teel 1999). Under-
standing people’s values and perspectives toward wildlife is fundamental to a
wildlife agency’s effectiveness in responding to public expectations, developing
management prescriptions, and directing education.
EDUCATION
As an extension of research, education is basic to our understanding of nature
and to developing conservation strategies. Wildlife professionals need to be
informed about puma ecology so they can formulate realistic policies and pre-
scriptions that stand the greatest chance of success. The public needs to be
informed about pumas because knowledge helps mold their beliefs and values
and encourages realistic expectations from wildlife managers. Public workshops
and other media (e.g., printed, television, radio) are particularly important in
regions where the odds are greatest for puma-human conflicts. In recent years,
agencies in puma-range states have actively informed people about pumas, and
they need to continue these efforts.
Moreover, it should be the responsibility of people living and recreating in
puma habitat to inform themselves by gleaning information in libraries and by
tapping the knowledge of wildlife experts. People should contact their state’s
wildlife management agency if they have questions. Practically every Western
state has produced a brochure about pumas in the Living With Wildlife series
that contains a brief summary of biological information about pumas and tips
on how people should behave if they live or recreate in puma habitat. A useful
pocket book is Torres’ (1997) Mountain Lion Alert. It is filled with basic facts
and good advice about pumas. People spending time in puma habitat should
read it; parents should read it to their children. Some communities even organ-
ize their own workshops and produce informational material. A good example
is Sandia Mountain Bearwatch, composed of citizens living in the Sandia Moun-
tains east of Albuquerque, which informs people in New Mexico about wildlife
in the mountains, ways to minimize detrimental impacts to animals and chances
of dangerous encounters with bears and pumas, and how they should behave if
they meet these animals. Bearwatch accomplishes this through consulting with
wildlife professionals, researching published references, distributing their own
written materials, and organizing neighborhood workshops.
Research scientists also have an obligation to inform the public. Most often,
they are the ones who have studied certain species or ecosystems to the extent
that they have earned the status of “expert.” Scientists need to distill the often-
complicated information into a format that most people can understand. What
good is it to wildlife conservation if research results are only written in techni-
cal jargon and concepts acceptable to scientific journals, but wildlife managers
and the public cannot fathom the messages?
RESPONSIVE MANAGEMENT
As the human population expands into rural and wild landscapes the likelihood
that pumas will encounter people will increase. This is true where humans usurp
habitat from resident pumas for dwellings and supporting developments (e.g.,
roads, utilities), and where human growth intercepts dispersing pumas. To deal
with those hapless pumas, wildlife management agencies need to support indi-
viduals or teams trained in the rapid and safe capture and control of pumas.
Rapid-response agents can minimize the potential for attacks on humans and
domestic animals. At the same time, agents could save some individual pumas
by translocating them to remote puma habitat that they may have been seeking
in the first place.
Morever, one of the most important responsibilities of agency managers is to
be in touch with the public’s values and expectations regarding puma manage-
ment and conservation. A procession of twenty-two ballot measures in eleven
states during 1980–2000 to increase protection of furbearers, wolves, black
bears, and pumas (Minnis 1998) should be a strong signal to wildlife managers
that influential stakeholders want carnivores protected from methods of killing
that they believe are inhumane, unsporting, and immoral, and that they are will-
ing to circumvent wildlife agency operations to accomplish their objectives. To
protect pumas, California voters decided to ban puma sport-hunting through-
out the state in 1990, and they defeated a ballot initiative to reinstate puma
hunting in 1996. Oregon voters banned the use of dogs to hunt pumas in 1994,
and Washington voters did the same in 1996 (Logan and Sweanor 1998). By
abolishing sport-hunting or restricting methods of hunting, these measures lim-
ited options that wildlife managers could use for puma population management.
Therefore, if wildlife managers want to retain influence in the management and
conservation of pumas, they need to be responsive to stakeholders’ values. Oth-
erwise, their management authority could be frustrated.
CONSERVATION FUNDING
All wildlife conservation and management activities require funding. Hiring and
training personnel, devising and implementing management plans, holding
public meetings, conducting research, enforcing laws, monitoring populations,
producing education material, and purchasing and maintaining habitat and
equipment all require money. Conservation activities for endangered Florida
panthers receive the bulk of funding from the U.S. Fish and Wildlife Service’s
administration of the Endangered Species Act and the state of Florida’s own
nongame wildlife trust funds. However, conservation and management of the

thousands of pumas living in the West is dependent principally upon funds gen-
erated from the sale of hunting licenses and excise taxes on sporting arms and
ammunition (via the Federal Aid in Wildlife Restoration Act). In the Proceedings
of The Fifth Mountain Lion Workshop (Padley 1996b), wildlife agencies in four
Western states (Nevada, Utah, Washington, and Wyoming) reported the
amounts of money they spent on puma management. These ranged from
$30,000 to $203,000 per year and averaged $114,500. The largest amount
spent was for the state of Wyoming, which is required by law to pay for loss of
private property caused by puma predation. Three other Western states (Col-
orado, Idaho, Utah) also reimburse property owners for losses due to puma
depredation, which can easily exceed $100,000 per year. Other states did not
distinguish expenditures for puma management in their budgets, which means
puma management activities were paid for through other agency programs (e.g.,
law enforcement, game surveys, research, and information and education).
A sobering lament we have heard from wildlife managers over the years is
that their budgets do not adequately provide for scientific management of puma
populations, needed research, or habitat protection. Clearly, monies generated
from hunters alone are not enough to pay for puma management and conserva-
tion programs in the West. So all people concerned for puma conservation and
science-based management, whether they hunt or not, should be doing what
they can to support agencies charged with those responsibilities. One idea for a
revenue-generating program is for states to support a species conservation cer-
tificate that citizens can buy voluntarily, the cost of which could be commensu-
rate to the price of a hunting license or big game tag. For example, Western res-
ident puma hunters presently pay on average $20 for a license and $37 for a tag
if they kill a puma. (However, about half of the states do not require a separate
tag.) It seems reasonable that nonhunters could pay $20–$40 for something like
a Big Mammal Conservation Certificate, with all proceeds earmarked for conser-
vation activities that benefit animals such as pumas, bears, wolves, deer, elk,
javelina, and bighorn sheep. Enhanced funding for big mammals that require
huge expanses of wild and rural lands would help to conserve, along with
pumas, a diversity of other wild creatures. Hunters could buy the certificates
too. The cost would be about the price for tickets to a movie or dinner at a
restaurant for a family of four. The number of hunters purchasing puma hunt-
ing licenses in the mid-1990s numbered six hundred to fourteen hundred in
most western states. Certainly, thousands more of the nonhunting public in
each state could be encouraged to donate directly to big-mammal conservation
programs. Regardless of whether people hunt, those with the monetary means
to contribute to conservation should do so. After all, just by living—using space,
food, and fiber—all people contribute to degradation of habitat and landscape

linkages that populations of big mammals require to endure.
In addition, public support of ballot initiatives and legislation that might
generate substantial public funding for wildlife habitat and conservation activi-
ties should be explored and tailored to fit the political, economic, and cultural
characteristics of each state. For example, in 1990 Californians passed the Cali-
fornia Wildlife Protection Act, which requires that the state spend more than
$30 million dollars per year until year 2020 on wildlife habitat protection, of
which $10 million must be spent to benefit puma and deer populations.
Another idea that has successfully generated substantial funds for conservation
is the sale of specialty automobile license plates, such as Florida’s Conserve
Wildlife plate. Obviously, some states may already have programs that citizens
can finance. There are also conservation opportunities in the private sector. For
example, The Nature Conservancy protects large tracts of wildlands and works
cooperatively with rural landowners to conserve wildlife habitat.
ENGAGED CITIZENSHIP
There is nothing more important in conservation than a caring, engaged citi-
zenry. People interested in puma conservation should get involved. Learn about
pumas. Get out in their habitat and contemplate what it is like for them to make
a living there. Appreciate their wild environment. Find out how agencies
charged with wildlife and natural resource management in your state try to con-
serve self-sustaining puma populations, their habitat, and other big mammals.
If you support, are dissatisfied with, or have better ideas about how things
should be done, then become part of the decision-making processes. Use the
avenues that government agencies establish for your involvement. Contact
elected and appointed officials responsible for overseeing wildlife and resource
management agencies. Contact local, regional, and state agency personnel and
let them know what you think. Attend the game commission or wildlife board
meetings (whatever they’re called in your state) to participate in policy-making
processes. Likewise, let federal land stewards know your opinions; they are
responsible for vast puma habitat in the West. Vote for government representa-
tives that you think would do the best job in environmental and conservation
issues. Make sure that government appointees to wildlife management commis-
sions or boards have a sound understanding in biology and ecology. If there is
one thing we have learned in all our years of involvement with wildlife conser-
vation, it is that generally nothing substantial ever happens without public sup-
port. The bigger and the more organized the public participation is, the better.
Agencies crave public direction on highly volatile wildlife issues. Informed, car-
ing citizens spur conservation initiatives to benefit big carnivores such as pumas
and the myriad other species that live in their habitats and landscape linkages.
Among the many enterprises in which people can invest portions of their intel-
lect, energy, and wealth, that of conserving biological diversity through the key-
stone and umbrella services of pumas and other big mammals can be one of the
most noble.
LP > > KX
1. Major threats to pumas, and all big carnivores, are habitat loss and overkill.
2. To conserve self-sustaining puma populations indefinitely, it is essential to
conserve as much remaining puma habitat and landscape linkages as possi-
ble. In addition, degraded habitat and linkages should be restored where
possible.
3. An adaptive management approach, such as zone management, could be
developed in puma-range states to provide an array of puma management
options, including sustained sport-hunting, experimental control of popu-
lations if deemed necessary, and puma refuges. Puma population manage-
ment actions should be designed as experiments.
4. Research on pumas that would benefit their management and conservation
include population monitoring techniques, habitat use patterns, behavior,
population dynamics, genetics, relationships with prey, translocation, rela-
tionships with other carnivores, aversive conditioning, protecting livestock,
and human dimensions.
5. Educating people, especially those living in puma range, is one of the most
important activities for conservation. It needs to be pursued at both public
and private levels.
6. Rapid-response management teams need to be maintained as the human
population expands into puma habitat.
7. Additional funding needs to be developed for conservation programs.
Funds generated from sport-hunters are not enough. All citizens who care
about wildlife conservation and can afford to donate should do so.
8. Citizens who care about wildlife conservation should be active in the pub-
lic and political arenas, especially in support of conservation programs ben-
efiting keystone and umbrella species that would also benefit myriad other
wildlife species.
LN N5LN5)
1. Test for a linear relationship between number of pumas harvested per
annum and number of puma permits issued per annum in New Mexico:
Puma harvest = 36.124 + 0.104 (number of puma permits), r2 = 0.80, P <
0.0001.
Chapter 21
Epilogue
Puma concolor and Homo sapiens share a common history. As Earthlings, we have
spun around this universe for about the past 390,000 years. When technologi-
cally advanced humans arrived in North America, pumas were here. Although
they may have gone extinct with the mega-mammals in the late Pleistocene,
pumas flourished again from a small number of founders that moved north from
South America (Culver et al. 2000). Thus, the extant North American puma lin-
eage has been here about as long as humans have. Over eons we trod the same
country, hunted the same prey, and in essential instances of life, even killed and
ate each other. Our basic endeavors in life were the same: acquire food, find a
mate, keep our offspring safe.
Humans dominated. We spread throughout the puma’s range and degraded
or totally usurped their habitats. Puma distribution is now about one-half of
what it was less than two hundred years ago. Where we did not displace them,
we killed them out until the only extant populations thrived in remote rugged
mountains of the West or in the forests and swamps of south Florida. Finally, in
the mid-1960s (about three puma-lifetimes ago) we cultivated an ethical obli-
gation to protect pumas. Once again they rebounded. Each time pumas endured
the ravages of shifting climate, variable environments, and human exploitation,
they did so because they had plenty of habitat in which to take refuge and
enough genetic variability to adapt. Now as our human population expands, we
continue to threaten their existence by destroying more habitat, fragmenting
populations into ever-smaller units, and eroding their genetic ability to adapt.
As long as humans dominate the Earth, the puma’s existence is entirely in our
hands.
397
We hope this book contributes to puma conservation and all that it means—
intact wild ecosystems, landscape linkages, constrained human exploitation. We
went to the mountains to study pumas up close, spending a substantial portion
of our own lives to glean a real understanding of the animal and its environ-
ment. We canvassed the data, looked for patterns, and made comparisons to
other reliable research on pumas and other carnivores. We probed research from
extra disciplines, including paleontology and genetics, to better understand how
pumas evolved. We tested previous scientific ideas about pumas and developed
others. Scientists coming after us should test our ideas, add to the reliable ones,
discard the undependable ones, and, in the process, enhance knowledge about
pumas and their relationship to humans. Prospects for conservation should also
benefit as long as scientific interpretations are conveyed in a manner most peo-
ple can understand. We have not been entirely successfully at that here, because
this book was meant to be a scientific treatise. Yet we hope the ideas will be
understood well enough by other writers and speakers that wish to impart our
messages to interested publics—those assorted stakeholders that will most deter-
mine conservation efforts that could favor pumas and a broad diversity of life
forms.
For ourselves, some of most treasured moments of our lives will be the times
we hiked the canyons and ridges of the San Andres Mountains, experiencing
their sublime wildness, following the tracks of pumas, trying to decipher how
they made a living and how they affected that desert ecosystem. In rare dazzling
moments, we watched pumas eye to eye. Through it all we were keenly aware of
our fortune—experiencing one of the wildest large carnivores on one of the
wildest remaining landscapes on Earth. That was exhilarating—and sometimes
frightening! We can only wonder if our prehistoric ancestors were equally
enchanted by pumas, other big carnivores, and wild lands. Perhaps that is the
root of a conservation ethic today—and one that is adaptive. The wonder we
have for wildlife and wild places probably has a genetic origin connecting our
intellectual and emotional justifications for nurturing our natural environment.
Humans have prospered greatly from biologically rich, clean environments. Our
spirit to explore and survive in wild environments probably directly affected the
evolution of our supreme intelligence. We believe that humans still need expan-
sive wild places with big scary mammals that challenge us. By conserving those
life forms in their wild environments, we benefit our own survival. If we accom-
plish that, then we will prove that we have earned our self-given name—sapi -
ens—the wise.
Appendix 1
Morphological Measurements
of Pumas At Least 3.5 Months Old
on the San Andres Mountains,
New Mexico, 1985–1995
400 Appendix 1. Morphological Measurements of Pumas At Least 3.5 Months Old
Appendix 1. Morphological Measurements of Pumas At Least 3.5 Months Old 401
Appendix 2
Reproductive Chronology of
Mated Pairs of Pumas on the
San Andres Mountains,
New Mexico, 1987–1994
Estimated
Mated pairs and estimated ages a
Dates pairs Approximate gestation Litter
Female Age (mo.) Male Age (mo.) consorted b birth date c period (days) d size
F15 43 M14 71 18–19 Feb. 1987 15 May 1987 87 3

F6 49 M7 50 1 Mar. 1987 31 May 1987 92 2
F21 38 M3 61 14 Jun. 1987 10 Sep. 1987 89 4
F4 81 M22 27 3 Aug. 1987 1 Nov. 1987 91 3
F27e 38 M20 75 3–6 Jan. 1988 27 Mar. 1988 85g 2
and 1 Apr. 1988 90g
F37 62 M7 61 8 Feb. 1988 7 May 1988 90 3
F41 20 M38 45 19–21 Jun. 1988 14 Sep. 1988 88 3
or M36 46 26 Jun. 1988 81g
F15 61 M19 60 14 Aug. 1988 24 Nov. 1988 103 2
F37 78 M7 77 18 Jun. 1989 8 Sep. 1989 83 3
F57 70 M5 105 23 Jul. 1989 21 Oct. 1989 91 4
F65 21 M19 80 3 Dec. 1989 13 Mar. 1990 97 2
F41 34 M46 35 16 Dec. 1989 14 Mar. 1990 89 3
F27f 63 M53 40 11 Feb. 1990 7 Apr. 1990 56g 3
F47 38 M88 22 25–28 May 1990 24 Aug. 1990 92 3
F21 88 M3 110 5–7 Jul. 1991 4 Oct. 1991 92 4
(continues)
413
414 Appendix 2. Reproductive Chronology of Mated Pairs of Pumas
Estimated
Mated pairs and estimated ages a
Dates pairs Approximate gestation Litter
Female Age (mo.) Male Age (mo.) consorted b birth date c period (days) d size
F47 52 M3 110 25–26 Jul. 1991 25 Oct. 1991 93 4
or M88 36 27–29 Jul. 1991 91g
F28 52 M38 88 6 Jan. 1992 1 Apr. 1992 87 3
or M5 135 8 Jan. 1992 85g
F149 20 M5 141 25 Apr. 1992 25 Jul. 1992 92 4
F107 20 M88 45 25 Apr. 1992 25 Jul. 1992 92 4
F68 32 M29 56 12–21 Jun. 1992 8 Sep. 1992 89 3
or M36 94 14 Jun. 1992 87g
F47 65 M88 49 9–11 Aug. 1992 6 Nov. 1992 90 3
F45 70 M52 67 23 Aug. 1992 19 Nov. 1992 89 2
F21 106 M88 55 26 Feb. 1993 5 May 1993 89 4
F47 71 M88 55 25–28 Feb. 1993 26 May 1993 91 3
F90 32 M38 103 28 Mar. 1993 3 Jun. 1993 98 3
F126 44 M46 77 27 Jun. 1993 27 Sep. 1993 93 3
F107 38 M88 67 12 Feb. 1994 19 May 1994 97 3
F103 38 M173 41 27 Feb. 1994 31 May 1994 94 3
F149 37 M3 142 27 Feb. 1994 31 May 1994 94 3
F91 44 M46 86 12 Mar. 1994 5 Jun. 1994 86 3
F184 33 M138 48 23 Apr. 1994 28 Jul. 1994 97 4
F45 91 M151 60 22 May 1994 20 Aug. 1994 91 1h
F47 88 M193 37 24 Jul. 1994 22 Oct. 1994 91 2i
aAges for the pumas are at the time the pair consorted. We estimated ages based on dental characteristics
described in Ashman et al. (1983) and dental characteristics and appearance of known-age pumas in our
own study.
bWe recorded pumas consorting when their radiotelemetry locations overlapped and when we visually
observed them together.

cApproximate birth dates were determined from radiotelemetry data on mothers and by back-aging cubs.
dGestation periods were estimated by including the first date that mated pairs were observed and by count-
ing to the approximate birth date. Mean gestation period = 91.5 ± 4.0 days, n = 31.
eBoth cubs produced by F27 were stillborn.
fF27 aborted three fetuses on 7 April 1990, three days after she was recaptured. F27 died on 9 April 1990
due to capture-induced trauma.

gThese estimated gestation periods were not used in calculation of the mean gestation period.
hOnly one cub was visually observed with F45; it was sighted at four months and again at 6.5 months of
age. F45’s litter may have been larger, and cubs may have died during the four-month interval before we
saw the first cub.
iWe visually observed F47’s two cubs at the birth nursery on 17 November 1994, near the end of our study.
We could not subsequently capture, examine, and mark the cubs because F47’s radio collar quit working.
Appendix 3
Methods and Estimates of Annual

Home Range Size for Pumas on the
San Andres Mountains,
New Mexico
To assess if our sample size was adequate to depict a puma’s annual home range
size, we randomly chose an annual period from each of six study animals (three
males, three females), systematically deleted locations to maintain a balanced
monthly representation, and then graphed the area of each resulting home range
(based on the 90 percent minimum convex polygon and 90 percent adaptive
kernel) relative to the number of locations. The initial number of locations for
each individual ranged from forty-five to fifty-one. Based on the minimum con-
vex polygon, an average of thirty locations was needed to reach 90 percent of the
final home range size. Results were more difficult to interpret using the adaptive
kernel estimator, because home range size often fluctuated widely with each
additional location and showed a general declining trend after an initial increase.
An average of thirty-seven locations was required before the percent change
(negative or positive) in home range size decreased to less than 10 percent. The
declining trend in home range size with increasing sample size may be a result
of better performance of the adaptive kernel model as more locations are
obtained; smaller sample sizes have been found to lead to larger overestimates of
home range size (Seaman and Powell 1996).
When running CALHOME software, grid cell size was set at 50 ¥ 50. For
the adaptive kernel, we let CALHOME calculate the optimum bandwidth (or
smoothing parameter) for each data set, and then re-ran each data set using an
80 percent bandwidth. This was done because a smaller bandwidth may result
415
416 Appendix 3. Methods and Estimates of Annual Home Range Size for Pumas
in a better fit of the data (lowest least-squares cross-validation or LSCV score)

when an animal uses two or more separate core areas (Kie et al. 1994). The
bandwidth that produced the lowest LSCV score was used to determine home
range size unless the 90 percent utilization distribution was broken into a greater
number of polygons.
FIGURE A3-1. Home range sizes of pumas M5 (indicated in black) and F45 (in gray) plot-
ted against the number of locations used to determine their size. Diamonds denote the
number of locations after which home range size increased or decreased by 10 percent or
less. Solid lines represent home ranges based on the 90 percent adaptive kernel, and dotted
lines represent home ranges based on the 90 percent minimum convex polygon.
Table A3-1. Annual home range size (mean ± SD) for twenty-four adult male
and thirty adult female pumas on the San Andres Mountains, New Mexico,
1986–1994.
Home range estimate (km2)
Minimum
x– no.
Puma no. na locationsb 90% 60% 100% 90%
Males
1c 3 42.7 226.2 ± 46.2 127.5 ± 25.9 218.5 ± 16.9 155.5 ± 10.7
3 8 44.8 180.8 ± 59.5 71.3 ± 40.6 173.3 ± 51.4 117.3 ± 43.2
5 6 42.7 186.4 ± 63.3 53.2 ± 20.1 162.7 ± 48.1 118.5 ± 40.4
7 4 45.8 183.3 ± 75.9 63.9 ± 40.2 170.4 ± 94.7 121.2 ± 72.4
Appendix 3. Methods and Estimates of Annual Home Range Size for Pumas 417

Minimum
x– no.
18 4 43.8 336.4 ± 225.9 144.3 ± 126.4 367.8 ± 247.3 209.0 ± 125.7
19 3 44.7 192.4 ± 20.6 68.0 ± 2.6 185.1 ± 11.7 125.1 ± 5.3
20d 1 32.0 305.5 117.8 320.6 272.9
22 1 46.0 153.4 40.0 142.1 92.0
29 1 42.0 126.1 54.7 108.9 83.3
36 6 43.0 222.6 ± 74.0 70.6 245.5 ± 144.6 133.1 ± 32.6
38 6 43.5 248.8 ± 71.7 104.9 ± 17.3 242.9 ± 51.9 163.4 ± 36.0
46 4 46.2 229.6 ± 97.2 72.8 ± 39.7 213.5 ± 40.6 155.8 ± 38.3
49 2 46.5 113.2 ± 5.4 43.0 ± 5.3 96.1 ± 5.2 61.0 ± 4.7
52 6 40.0 104.4 ± 19.5 43.6 ± 16.8 104.7 ± 25.6 68.5 ± 17.7
53d 3 39.3 245.8 ± 115.1 59.2 ± 19.3 232.3 ± 52.3 171.7 ± 48.6
73 1 39.0 177.8 74.2 139.2 124.2
88 3 47.0 174.1 ± 40.6 67.8 ± 13.7 171.4 ± 25.2 123.1 ± 23.4
124 1 44.0 81.4 24.8 85.0 66.1
138 2 46.5 249.6 ± 93.4 79.6 ± 17.3 268.9 ± 23.5 163.2 ± 39.2
151 2 41.5 271.2 ± 77.4 71.3 ± 26.0 271.4 ± 66.3 168.8 ± 14.8
161 1 41.0 259.7 42.7 202.0 150.8
173 2 46.0 132.8 ± 8.2 61.2 ± 7.4 122.4 ± 3.5 90.2 ± 6.3
193 1 46.0 59.3 16.8 75.2 53.0
210 1 43.0 180.0 ± 70.8 70.7 194.6 128.4
x– of x– 24 43.2 193.4 ± 69.6 68.5 ± 30.3 188.1 ± 74.8 129.8 ± 50.3
Females
2 1 35.0 139.8 43.9 121.5 73.6
4 2 40.5 37.2 ± 2.1 10.4 ± 1.2 39.2 ± 3.0 22.2 ± 7.1
6 2 40.0 116.0 ± 11.4 40.4 ± 5.0 96.8 ± 0.1 60.4 ± 9.3
15 5 44.6 58.9 ± 14.2 19.0 ± 8.5 58.9 ± 14.8 37.5 ± 11.8
21 3 45.0 63.6 ± 23.1 23.3 ± 9.1 73.7 ± 9.6 46.7 ± 19.8
27 2 43.5 60.1 ± 13.7 14.6 ± 1.6 79.9 ± 1.3 33.1 ± 3.4
28 3 45.7 89.9 ± 46.5 30.0 ± 8.9 89.8 ± 47.2 59.1 ± 24.7
31 1 46.0 72.8 21.6 71.3 49.3
37 3 44.0 113.8 ± 15.2 40.6 ± 18.1 90.9 ± 2.0 59.9 ± 8.2
41 3 45.0 48.5 ± 16.0 23.0 ± 4.0 55.3 ± 17.7 38.0 ± 9.8
45 6 43.8 37.8 ± 11.4 11.4 ± 2.9 38.0 ± 8.5 27.1 ± 6.8
47 6 47.8 33.9 ± 11.0 11.4 ± 5.7 42.1 ± 13.1 24.2 ± 9.0
54 3 44.3 63.9 ± 36.5 20.1 ± 11.6 62.7 ± 32.1 40.2 ± 18.2
57 3 44.3 46.7 ± 11.2 17.9 ± 5.3 50.7 ± 4.0 33.4 ± 9.7
58 2 48.0 67.8 ± 32.6 14.0 ± 2.0 92.2 ± 32.0 41.6 ± 21.6
(continues)
418 Appendix 3. Methods and Estimates of Annual Home Range Size for Pumas
Table A3-1. Continued

Minimum
– Adaptive kernel convex polygon
x no.
65 3 41.7 61.4 ± 26.4 18.1 ± 3.3 58.8 ± 21.3 36.2 ± 11.5
68 1 47.0 30.1 8.8 25.7 16.1
86 1 44.0 92.8 28.0 117.7 57.8
87 2 41.5 107.8 ± 30.4 34.9 ± 19.1 106.8 ± 39.1 71.0 ± 19.4
89 1 43.0 45.7 16.4 43.0 28.4
90 2 44.0 64.0 ± 5.6 26.9 ± 1.1 65.6 ± 0.5 42.4 ± 2.5
91 3 44.7 45.5 ± 14.4 19.5 ± 6.8 55.4 ± 10.7 34.2 ± 8.2
103 2 49.5 54.3 ± 0.7 28.2 ± 5.0 66.6 ± 20.2 40.4 ± 2.1
107 1 47.0 67.6 19.4 72.2 52.3
126 2 41.0 80.7 ± 2.1 26.4 ± 12.7 74.2 ± 10.0 43.2 ± 12.2
128 1 44.0 47.0 21.3 46.8 33.6
130 1 44.0 66.2 21.2 78.2 43.0
147 2 39.0 150.2 ± 194.0 35.0 ± 43.5 154.5 ± 201.0 109.8 ± 143.3
149 2 43.5 61.4 ± 20.9 21.0 ± 13.4 48.0 ± 19.0 33.0 ± 18.7
184 2 45.5 71.3 ± 35.4 19.3 ± 14.0 75.8 ± 43.2 48.9 ± 30.2
x– of x– 30 43.9 69.9 ± 30.1 22.9 ± 9.1 71.7 ± 28.3 44.6 ± 18.5
an = the number of years (1 January through 31 December) for which annual home range size was
calculated. Annual home ranges were calculated only for pumas that were radio-monitored for ten
months or more of a given twelve-month period.
b The average number of locations obtained each year. Weekly aerial locations were augmented with
ground locations as long as all locations were more than three days apart.
cM1’s home range included area south of the study area (Organ Mountains) from 1986 to 1988; dur-
ing those three years 19.5, 32.1, and 55.9 percent of his locations were outside the study area.
dM1, M20, and M53 shifted their home range activities outside the study area over time. Only those
years when they spent time in the study area are reported here.
Appendix 4
A Deterministic, Discrete
Time Model That Simulated Mule
Deer Population Dynamics in
the Treatment Area, San Andres
Mountains, New Mexico,
1987–1995
See Chapter 17 for its application.

The basic deer model:
Nt + 1 = B (sb) + D (sd) + [AD (fa) + YD (fy) – Fd ]
Most model parameters were from surveys conducted on the San Andres
Mountains. However, we had to rely on some estimates from other desert mule
deer populations that were studied by the New Mexico Department of Game
and Fish in other parts of southern New Mexico. We varied some of the param-
eters ourselves to simulate variation during drought conditions. Consequently,
we developed six different population simulations, or models (i.e., six models
numbered 1 to 6), to simulate the dynamics of the deer population on the Treat-
ment Area (see Chapter 17 for details). Model parameters, their origins and vari-
ations, and our assumptions are described below.
419
420 Appendix 4. Simulated Mule Deer Population Dynamics in the Treatment Area
Nt = deer population size at the beginning of the deer biological year, which
starts with the peak of the fawning season in August (births actually occurred
from mid-July through September). Nt is the annual base population size
assumed to be a representative population of 1,000 deer one year or older.
Nt + 1 = deer population size at the end of the biological year. It reflects the
change in the base population of 1,000 deer (i.e., 1,000 + births – deaths).
B = estimated number of bucks (one year or older) in the base population. We
assumed it was proportional to the buck:doe ratio estimated from the annual
deer population composition count (Table 17-5). Models 4, 5, and 6 assume
the buck:doe ratio in year 1 was biased high, and thus they use an estimated
correction (Table 17-6).
D = estimated number of does (one year or older) in the base population. We
assumed it was proportional to the buck:doe ratio estimated from the annual
deer population composition count (Table 17-5). Models 4, 5, and 6 attempt
to correct for a potential bias in year 1 (see parameter B above).
AD = the number of does two years or older. We estimated AD by subtracting
the estimated number of yearling does from the total number of does in the
base population.
YD = the number of yearling does. We assumed they bred at 1.5 years old and
gave birth at two years old. The proportion of yearling does in the doe popu-
lation was estimated after an initial iteration of the model where all does in
year 1 were assumed to be two years or older. Then the average proportion of
yearling does (estimated from the number of fawns and does surviving to
Nt + 1) during all subsequent years was used to estimate the proportion of year-
ling does in year 1.
fa = fetal rate of does two years or older. We assumed it to be equal to the aver-
age of 1.15 fetuses per doe for twenty-seven adult does sampled on the Treat-
ment Area in April 1990, and the average of 1.10 fetuses per doe for thirty
adult does sampled on the Treatment Area in April 1991 (New Mexico
Department of Game and Fish unpublished data). Fetal rates were estimated
during the last trimester of pregnancy using the rectal abdominal palpation
method (Hulet 1972, Humphreys and Elenowitz 1988). This method tends
to underestimate the true fetal rate by 6–12 percent (Humphreys and
Elenowitz 1988); hence, fetal rates we used were probably slightly conserva-
tive. We used fetal rates of 1.15 for the first three years and 1.10 for the last
five years (Table 17-6). We assumed the sex ratio for deer births to be 1:1.
fy = fetal rate for yearling does. We assumed it to be equal to the average of 0.90
fetuses per doe for thirty-three yearlings sampled (Humphreys and Elenowitz
1988:24) in the Guadalupe Mountains, New Mexico (about 160 km east-
southeast of our Treatment Area). For models 1, 2, 4, and 5, we used 0.90 for
Appendix 4. Simulated Mule Deer Population Dynamics in the Treatment Area 421
years 1–3; additionally, for years 4–8, we assumed fy was equal to the average
of 0.81 for fifteen yearlings sampled (Humphreys and Elenowitz 1988:24) on
the MacGregor Range, New Mexico (about 75 km east-southeast of our Treat-
ment Area). For models 3 and 6, we assumed fetal rates for yearling does were
equal to 0.90 for all years (Table 17-6).
sb = survival rate for bucks. We assumed it to be equivalent to the survival rate
of radio-collared bucks (Table 17-2).
sd = survival rate for does. We assumed it to be equivalent to the survival rate of
radio-collared does (Table 17-2).
Fd = number of fawns that died. We estimated it by subtracting the number of
fawns that survived to the annual deer population composition counts from
the estimated number of fawns born [i.e., fawns born = AD (fa) + YD (fy)]
minus an additional percentage loss. Temple (1981) found that 90 percent of
the total mortality for forty radio-collared fawns on Fort Bayard, New Mexico
(about 150 km west of our Treatment Area), occurred during the first forty-
nine days of life, and 10 percent occurred during the rest of the year. For the
model, we assumed two fawn mortality schedules: (1) ninety percent of the
total fawn mortality occurred before the winter composition counts and 10
percent occurred between the composition counts and the end of the biolog-
ical year; (2) eighty percent and 20 percent of the fawn mortality occurred
during the two previously defined periods. We used schedule 2 in two of the
models (Table 17-6) to account for potential increased fawn mortality during
drought conditions (i.e., schedule 2 increased the estimated fawn mortality by
10 percent). The sex ratio of fawns that died in the population was assumed
to be 1:1.
References
Ackerman, B. B. 1982. Cougar predation and ecological energetics in southern Utah.

Master’s thesis, Utah State University.
Ackerman, B. B., F. G. Lindzey, and T. P. Hemker. 1986. Predictive energetics model of
cougars. In Cats of the world: Biology, conservation and management, edited by S. D.
Miller and D. Everett, 333–352. Washington, D.C.: National Wildlife Federation.
Ackerman, B. B., F. A. Leban, E. O. Garton, and M. D. Samuel. 1989. User’s manual for
program HOME RANGE. Contribution No. 259. Forestry, Wildlife and Range Exper-
iment Station, University of Idaho, Moscow.
Adams, D. B. 1979. The cheetah: Native American. Science 205:1155–1158.
Adaska, J. M. 1999. Peritoneal coccidioidomycosis in a mountain lion in California.
Journal of Wildlife Diseases 35:75–77.
Aiello, L. C., and P. Wheeler. 1995. The expensive tissue hypothesis: The brain and the
digestive system in human and primate evolution. Current Anthropology 36:199– 221.
Andelt, W. F. 1992. Effectiveness of livestock guarding dogs for reducing predation on
domestic sheep. Wildlife Society Bulletin 20:55–62.
———. 1999. Relative effectiveness of guarding-dog breeds to deter predation on
domestic sheep in Colorado. Wildlife Society Bulletin 27:706–714.
Anderson, A. E. 1981. Morphological and physiological characteristics. In Mule and
black-tailed deer of North America, edited by O. C. Wallmo, 27–97. Lincoln: Univer-
sity of Nebraska Press.
———. 1983. A critical review of literature on puma (Felis concolor). Colorado Division
of Wildlife Special Report No. 54.
Anderson, A. E., D. C. Bowden, and D. M. Kattner. 1992. The puma on Uncompahgre
Plateau, Colorado. Colorado Division of Wildlife Technical Publication No. 40.
Anderson, E. M. 1988. Effects of male removal on spatial distribution of bobcats. Jour -
nal of Mammology 69:637–641.
Anderson, W. D., and D. Taylor. 1983. Natural resources management plan, White Sands
Missile Range, New Mexico. Department of Army, U.S. Army, White Sands Missile
Range, New Mexico.
Armitage, K. B. 1986. Marmot polygyny revisited: Determinants of male and female
reproductive strategies. In Ecological aspects of social evolution: Birds and mammals,
edited by D. I. Rubenstein and R. W. Wrangham, 303–331. Princeton, N.J.: Prince-
ton University Press.
423
424 References
Ashman, D., G. C. Christensen, M. L. Hess, G. K. Tsukamoto, and M. S. Wickersham.

1983. The mountain lion in Nevada. Nevada Fish and Game Department, Federal Aid
in Wildlife Restoration Final Report, Proj. W-48-15.
Augsburger, J. G. 1970. Behavior of Mexican bighorn sheep in the San Andres Moun-
tains, New Mexico. Master’s thesis, New Mexico State University.
Aune, K. E. 1991. Increasing mountain lion populations and human–mountain lion
interactions in Montana. In Mountain lion-human interaction symposium and work -
shop, April 24–26, 1991, edited by C. S. Braun, 86–94. Colorado Division of Wildlife,
Denver.
Bailey, T. N. 1993. The African leopard: Ecology and behavior of a solitary felid. New York:
Columbia University Press.
Bailey, T. N., E. E. Bangs, M. F. Portner, J. C. Malloy, and R. J. McAvinchey. 1986. An
apparent overexploited lynx population on the Kenai Peninsula, Alaska. Journal of
Wildlife Management 50:279–290.
Ballard, W. B., and P. S. Gipson. 2000. Wolf. In Ecology and management of large mam -
mals in North America, edited by S. Demarais and P. R. Krausman, 321–346. Upper
Saddle River, N.J.: Prentice Hall.
Barnhurst, D. 1986. Vulnerability of cougars to hunting. Master’s thesis, Utah State Uni-
versity.
Barnhurst, D., and F. G. Lindzey. 1989. Detecting female mountain lions with kittens.
Northwest Science 63:35–37.
Barone, M. A., M. E. Roelke, J. Howard, J. L. Brown, A. E. Anderson, and D. E. Wildt.
1994. Reproductive characteristics of male Florida panthers: Comparative studies
from Florida, Texas, Colorado, Latin America, and North American zoos. Journal of
Mammalogy 75:150–162.
Bartmann, R. M., G. C. White, and L. H. Carpenter. 1992. Compensatory mortality in
a Colorado mule deer population. Wildlife Monograph No. 121.
Beasom, S. L., W. Evans, and L. Temple. 1980. The drive net for capturing western big
game. Journal of Wildlife Management 44:478–480.
Beckett, P. H. 1983. The archaic prehistory of the Tularosa Basin. In The prehistory of
Rhodes Canyon, New Mexico, edited by P. L. Eidenbach, 105–110. Tularosa, N.M.:
Human Systems Research.
Begon, M., J. L. Harper, and C. R. Townsend. 1990. Ecology, individuals, populations and
communities. Boston: Blackwell Scientific Publications.
Beier, P. 1993. Determining minimum habitat areas and habitat corridors for cougars.
Conservation Biology 7:94–108.
———. 1995. Dispersal of juvenile cougar in fragmented habitat. Journal of Wildlife
Management 59:228–237.
———. 1996. Metapopulation models, tenacious tracking, and cougar conservation. In
Metapopulations and wildlife conservation, edited by D. R. McCullough, 293–323.
Washington, D.C.: Island Press.
Beier, P., and R. H. Barrett. 1993. The cougar in the Santa Ana Mountain Range, Cali -
fornia. Orange County Cooperative Mountain Lion Study Final Report.
Beier, P., and S. C. Cunningham. 1996. Power of track surveys to detect changes in
cougar populations. Wildlife Society Bulletin 24:540–546.
Bekoff, M., and L. D. Mech. 1984. Simulation analyses of space use: Home range esti-
References 425
mates, variability, and sample size. Behavior Research Methods, Instruments and Com -
puters 16:32–37.
Belden, R. C. 1982. Florida panther recovery plan implementation: A 1982 progress report.
Federal Aid to Endangered Species Program. Florida Endangered Species Project E-1.
Belden, R. C., W. B. Frankenberger, R. T. McBride, and S. T. Schwikert. 1988. Panther
habitat use in southern Florida. Journal of Wildlife Management 52:660–663.
Belden, R. C., and B. W. Hagedorn. 1993. Feasibility of translocating panthers into
northern Florida. Journal of Wildlife Management 57:388–397.
Belden, R. C., and J. W. McCowan. 1996. Florida panther reintroduction feasibility study.
Final Report. Florida Game and Fresh Water Fish Commission, Bureau of Wildlife
Research, Tallahassee, Florida.
Berger, J. 1978. Group size, foraging, and antipredator ploys: An analysis of bighorn
sheep decisions. Behavioral Ecology and Sociobiology 4:91–99.
———. 1990. Persistence of different-sized populations: An empirical assessment of
rapid extinctions in bighorn sheep. Conservation Biology 4:91–98.
———. 1999. Intervention and persistence in small populations of bighorn sheep. Con -
servation Biology 13:432–435.
Berger, J., and J. D. Wehausen. 1991. Consequences of a mammalian predator-prey dis-
equilibrium in the Great Basin Desert. Conservation Biology 5:244–248.
Bergerud, A. T., and J. P. Elliot. 1986. Dynamics of caribou and wolves in northern
British Columbia. Canadian Journal of Zoology 64:1515–1529.
Bergerud, A. T., and R. E. Page. 1987. Displacement and dispersion of parturient cari-
bou at calving as an antipredator tactic. Canadian Journal of Zoology 65:1598–1606.
Bergerud, A. T., W. Wyett, and B. Snider. 1983. The role of wolf predation in limiting
a moose population. Journal of Wildlife Management 47:977–988.
Bertram, B. C. R. 1973. Lion population regulation. East African Wildlife Journal
11:215–225.
Blumenschine, R. J. 1991. Hominid carnivory and foraging strategies, and the socio-
economic function of early archaeological sites. Philosophical Transactions of the Royal
Society of London 334:211–221.
Boellstorff, D. E., D. H. Ownings, M. C. T. Penedo, and M. J. Hersek. 1994. Repro-
ductive behavior and multiple paternity in California ground squirrels. Animal Behav -
ior 47:1057–1064.
Boertje, R. D., and R. O. Stephenson. 1992. Effects of ungulate availability on wolf
reproductive potential in Alaska. Canadian Journal of Zoology 70:2441–2443.
Bolton, H. E. 1916. Original narratives of early American history: Spanish exploration in
the southwest, 1542–1706. New York: Charles Scribner’s Sons.
Bonney, R. C., H. D. M. Moore, and D. M. Jones. 1981. Plasma concentrations of
oestradiol-17b and progesterone, and laparoscopic observations of the ovary in the
puma (Felis concolor) during oestrus, pseudopregnancy and pregnancy. Journal of
Reproductive Fertility 63:523–531.
Boutin, S. 1992. Predation and moose population dynamics: A critique. Journal of
Boyce, M. S., and C. M. Perrins. 1987. Optimizing great tit clutch size in a fluctuating
environment. Ecology 68:142–153.
Boyce, W. M. 1994. 1993 bighorn sheep and deer health evaluations San Andres, Red Rock,
426 References
and Wheeler Peak New Mexico. Final Report prepared for the New Mexico Department
of Game and Fish, Santa Fe.
Boyce, W., M. Elliott, R. Clark, and D. Jessup. 1990. Morphometric analysis of Psoroptes
spp. mites from bighorn sheep, mule deer, cattle, and rabbits. Journal of Parasitology
76:823–828.
Boyce, W. M., P. W. Hedrick, J. E. Muggli-Crockett, S. Kalinowski, M. C. T. Penedo,
and R. R. Ramey II. 1996. Genetic variation of major histocompatibility complex and
microsatellite loci: A comparison in bighorn sheep. Genetics 145:421–433.
Boyd, D., and B. O’Gara. 1985. Cougar predation on coyotes. The Murrelet 66:17.
Boyd, D. K., and G. K. Neale. 1992. An adult cougar (Felis concolor) killed by gray
wolves (Canis lupus) in Glacier National Park, Montana. Canadian Field-Naturalist
106:524–525.
Boykin, K. G., P. L. Matusik, and B. C. Thompson. 1996. Comparative and physical
attributes of natural water sites on White Sands Missile Range. Research Completion
Report. New Mexico Cooperative Fish and Wildlife Research Unit, Las Cruces.
Brahmachary, R. L., M. P. Sarkar, and J. Dutta. 1992. Chemical signals in the tiger. In
Chemical signals in vertebrates 6, edited by R. L. Doty and D. Müller-Schwarze,
471–475. New York: Plenum Press.
Branch, L. C., M. Pessino, and D. Villarreal. 1996. Response of pumas to a population
decline of the plains vizcacha. Journal of Mammalogy 77:1132–1140.
Brand, C. J., L. B. Keith, and C. A. Fischer. 1976. Lynx responses to changing snowshoe
hare densities in central Alberta. Journal of Wildlife Management 403:416–428.
Brashares, J. S., and P. Arcese. 1999. Scent marking in a territorial African antelope: I.
The maintenance of borders between male oribi. Animal Behaviour 57:1–10.
Breternitz, C. D., and D. E. Doyel. 1983. A cultural resources overview and management
plan for the White Sands Missile Range. Phoenix, Ariz: Soil Systems.
Bronson, F. H. 1989. Mammalian reproductive biology. Chicago: University of Chicago
Press.
Brown, D. E. 1984. The wolf in the Southwest. Tucson: University of Arizona Press.
——-. 1985. The grizzly in the Southwest. Norman: University of Oklahoma Press.
Brown, D. E., F. Reichenbacher, and S. E. Franson. 1998. A classification of North Amer -
ican biotic communities. Salt Lake City: University of Utah Press.
Brown, J. L., and G. H. Orians. 1970. Spacing patterns in mobile animals. Annual
Review of Ecology Systematics 1:239–262.
Buechner, H. K. 1960. The bighorn sheep in the United States: Its past, present, and
future. Wildlife Monograph No. 44:1–174.
Bunch, T. D., W. M. Boyce, C. P. Hibler, W. R. Lance, T. R. Spraker, and E. S. Williams.
1999. Diseases of North American wild sheep. In Mountain sheep of North America,
edited by R. Valdez and P. R. Kruasman, 209–237. Tucson: University of Arizona
Press.
Burney, D. A. 1993. Recent animal extinctions: Recipes for disaster. American Scientist
81:530–541.
Burt, W. H. 1943. Territoriality and home range concepts as applied to mammals. Jour -
nal of Mammalogy 24:346–352.
Caro, T. M. 1986. The functions of stotting in Thomson’s gazelles: Some tests of the pre-
dictions. Animal Behaviour 34:663–684.
———. 1989. Determinants of asociality in felids. In Comparative socio-ecology: The
References 427
behavioral ecology of humans and other mammals, edited by V. Standen and R. A. Foley,
41–74. Special publication of the British Ecological Society, No. 8. Oxford: Blackwell
Scientific Publications.
———. 1994. Cheetahs of the Serengeti Plains: Group living in an asocial species. Chicago:
University of Chicago Press.
Caro, T. M., and C. D. Fitzgibbon. 1992. Large carnivores and their prey: The quick
and the dead. In Natural enemies: The population biology of predators, parasites and dis -
eases, edited by M. J. Crawley, 117–142. London: Blackwell Scientific Publications.
Caughley, G. 1978. Analysis of vertebrate populations. New York: John Wiley and Sons.
Caughley, G., and A. R. E. Sinclair. 1994. Wildlife ecology and management. Cambridge,
Mass.: Blackwell Science.
Chepko-Sade, B. D., and W. M. Shields with J. Berger, Z. T. Halpin, W. T. Jones, L. L.
Rogers, J. P. Rood, and A. T. Smith. 1987. The effects of dispersal and social structure
on effective population size. In Mammalian dispersal patterns: The effects of social struc -
ture on population genetics, edited by B. D. Chepko-Sade and Z. T. Halpin, 287–321.
Chicago: University of Chicago Press.
Chepko-Sade, B. D., and Z. T. Halpin, editors. 1987. Mammalian dispersal patterns: The
effects of social structure on population genetics. Chicago: University of Chicago Press.
Clapperton, B. K., E. O. Minot, and D. R. Crump. 1988. An olfactory recognition sys-
tem in the ferret Mustela furo L. (Carnivora: Mustelidae). Animal Behaviour
36:541–553.
Clark, P. J., and F. C. Evans. 1954. Distance to nearest neighbor as a measure of spatial
relationships in populations. Ecology 35:445–453.
Clark, R., and D. Jessup. 1990. Status of psoroptic scabies in and health of bighorn sheep in
the San Andres Mountains of New Mexico. Final report, International Wildlife Veteri-
nary Services, Orangevale, California, to the U.S. Fish and Wildlife Service, San
Andres National Wildlife Refuge, Las Cruces, New Mexico.
Clark, R. K., and D. A. Jessup. 1992. The health of mountain sheep in the San Andres
Mountains, New Mexico. Desert Bighorn Council Transactions 36:30–35.
Clevenger, A. P., and N. Waltho. 2000. Factors influencing the effectiveness of wildlife
underpasses in Banff National Park, Alberta, Canada. Conservation Biology 14:47–56.
Clutton-Brock, T. H. 1989. Mammalian mating systems. Proceedings of the Royal Society
of London Series B 236:339–372.
Clutton-Brock, T. H., and G. R. Iason. 1986. Sex ratio variation in mammals. The Quar -
terly Review of Biology 61:339–374.
Clutton-Brock, T. H., S. D. Albon, and F. E. Guinness. 1986. Great expectations: Dom-
inance, breeding success and offspring sex ratios in red deer. Animal Behaviour
34:460–471.
———. 1988. Reproductive success in male and female red deer. In Reproductive success:
Studies of individual variation in contrasting breeding systems, edited by T. H Clutton-
Brock, 325–343. Chicago: University of Chicago Press.
Converse, L. J., A. A. Carlson, T. E. Ziegler, and C. T. Snowdon. 1995. Communication
of ovulatory state by female pygmy marmosets Cebuella pygmaea. Animal Behaviour
49:615–621.
Coe Clough, N., and J. A. Roth. 1998. Understanding immunology. St. Louis: Mosby.
Crawshaw, P. G., Jr., and H. B. Quigley. 1991. Jaguar spacing, activity and habitat use
428 References
in a seasonally flooded environment in Brazil. Jo u rnal of Zoology, London

223:357–370.
Crow, J. F., and M. Kimura. 1970. An introduction to population genetics theory. New
York: Harper and Row.
Culver, M. 1999. Molecular genetic variation, population structure, and natural history
of free-ranging pumas (Puma concolor). Ph.D. diss., University of Maryland.
Culver, M., W. E. Johnson, J. Pecon-Slattery, and S. J. O’Brien. 2000. Genomic ances-
try of the American puma (Puma concolor). The Journal of Heredity 91:186–197.
Cunningham, S. C., L. A. Haynes, C. Gustavson, and D. D. Haywood. 1995. Evalua -
tion 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.
Daniel, W. W. 1978. Applied nonparametric statistics. Boston: Houghton Mifflin.
DeRose, M. A., and D. A. Roff. 1999. A comparison of inbreeding depression in life-
history and morphological traits in animals. Evolution 53:1288–1292.
Diamond, J. 1989. Overview of recent extinctions. In Conservation for the 21st century,
edited by D. Western and M. Pearl, 37–41. New York: Oxford University Press.
Dobson, F. S. 1982. Competition for mates and predominant juvenile male dispersal in
mammals. Animal Behaviour 30:1183–1192.
Dunn, W. C. 1994. Evaluation of desert bighorn sheep habitat in New Mexico. A Revi -
sion of the Final Report, Federal Aid in Wildlife Restoration Project W-127-R-7, Job 4.
Santa Fe: New Mexico Department of Game and Fish.
Durant, S. M. 1998. Competition refuges and coexistence: An example from Serengeti
carnivores. Journal of Animal Ecology 67:370–386.
Eastern Cougar Foundation. 2000. Eastern Cougar Foundation Newsletter. Summer.
North Spring, West Virginia.
Eaton, R. L. 1976. Why some felids copulate so much. World’s Cats 3:73–94.
Ehret, G. 1980. Development of sound communication in mammals. In Advances in the
study of behavior, edited by J. S. Rosenblatt, R. A. Hinde, C. Beer, and M. C. Busnel,
11:179–225. New York: Academic Press.
Eidenbach, P. L., editor. 1983. The prehistory of Rhodes Canyon, New Mexico. Tularosa,
N.M.: Human Systems Research.
Eidenbach, P. L., 1989. The West that was forgotten: Historic ranches of the northern San
Andreas Mountains, White Sands Missile Range, New Mexico. Tularosa, N.M.: Human
Systems Research.
Eidenbach, P. L., and B. Morgan, editors. 1994. Homes on the range: Oral recollections of
early ranch life on the U.S. Army White Sands Missile Range, New Mexico. Mesilla,
Eisenberg, J. F. 1981. The mammalian radiations: An analysis of trends in evolution, adap -
tation, and behavior. Chicago: University of Chicago Press.
Elenowitz, A. S. 1987. Status of bighorn sheep in New Mexico, 1986. Desert Bighorn
Council 1987 Transactions 31:30–32.
Elmer, M. 1997. Cougar food habit dynamics in the San Andres Mountains, New Mex-
ico. Master’s thesis, University of Idaho.
Emlen, J. T. Jr. 1957. Defended area? A critique of the territory concept and of conven-
tional thinking. Ibis 99:352.
References 429
Emlen, S. T., and L. W. Oring. 1977. Ecology, sexual selection, and the evolution of
mating systems. Science 197:215–223.
Erlinge, S., and M. Sandell. 1986. Seasonal changes in the social organization of male
stoats, Mustela ermined: An effect of shifts between two decisive resources. Oikos
47:57–62.
Ernest, H. B., M. C. T. Penedo, B. P. May, M. Syvanen, and W. M. Boyce. 2000. Mol-
ecular tracking of mountain lions in the Yosemite Valley region in California: Genetic
analysis using microsatellites and faecal DNA. Molecular Ecology 9:433–441.
Erwin, D. H. 1998. The end and the beginning: Recoveries from mass extinctions.
Trends in Ecology and Evolution 13:344–349.
Etkin, W. 1964. Cooperation and competition in social behavior. In Social behavior and
organization among vertebrates, edited by W Etkin, 1–34. Chicago: University of
Chicago Press.
Evans, W. 1983. The cougar in New Mexico: Biology, status, depredation of livestock, and
management recommendations. Response to House Memorial 42. New Mexico Game
and Fish Department, Santa Fe.
Ewer, R. F. 1968. Ethology of mammals. London: Elek Science.
Ewer, R. F. 1973. The carnivores. Ithaca, N.Y.: Cornell University Press.
Findley, J. S., A. H. Harris, D. E. Wilson, and C. Jones. 1975. Mammals of New Mex -
ico. Albuquerque: University of New Mexico Press.
Fitter, R. 1986. Wildlife for man. London: Collins.
Fitzsimmons, N. N., and S. W. Buskirk. 1992. Effective population sizes for bighorn
sheep. Biennial Symposium of the Northern Wild Sheep and Goat Council 8:1–7.
Fitzsimmons, N. N., S. W. Buskirk, and M. H. Smith. 1995. Population history, genetic
variability, and horn growth in bighorn sheep. Conservation Biology 9:314–323.
Floyd, T. J., L. D. Mech, and P. A. Jordan. 1978. Relating wolf scat content to prey con-
sumed. Journal of Wildlife Management 43:528–532.
Foster, M. L., and S. R. Humphrey. 1995. Use of highway underpasses by Florida pan-
thers and other wildlife. Wildlife Society Bulletin 23:95–100.
Frank, D. H., and E. J. Heske. 1992. Seasonal changes in space use patterns in the south-
ern grass-hopper mouse, On ychomys torridus torridus. Journal of Ma m m a l o gy
73:292–298.
Franklin, W. L., W. E. Johnson, R. J. Sarno, and J. A. Iriarte. 1999. Ecology of the Patag-
onia puma Felis concolor patagonica in southern Chile. Biological Conservation
90:33–40.
Freeman, S., and J. C. Herron. 1998. Evolutionary analysis. Upper Saddle River, N.J.:
Prentice Hall.
Fuller, T. K. 1989. Population dynamics of wolves in north-central Minnesota. Wildlife
Monograph No. 105.
Garrott, R. A., G. C. White, R. M. Bartmann, L. H. Carpenter, and A. W. Alldredge.
1987. Movements of female mule deer in northwest Colorado. Journal of Wildlife
Gasaway, W. C., R. O. Stephenson, J. L. Davis, P. E. K. Shepherd, and O. E. Burris.
1983. Interrelationships of wolves, prey, and man in interior Alaska. Wildlife Mono-
graph No. 84.
Gasaway, W. C., R. D. Boertje, D. V. Grangaard, D. G. Kelleyhouse, R. O. Stephenson,
and D. G. Larsen. 1992. The role of predation in limiting moose at low densities in
Alaska and Yukon and implications for conservation. Wildlife Monograph No. 120.
430 References
Gashwiler, J. S., and W. L. Robinette. 1957. Accidental fatalities of the Utah cougar.
Journal of Mammalogy 38:123–126.
Gay, S. W., and T. L. Best. 1995. Geographic variation in sexual dimorphism of the
puma (Puma concolor) in North and South America. The Southwestern Naturalist
40:148–159.
Geist, V. 1966. Validity of horn segment counts in aging bighorn sheep. Journal of
———. 1985. On Pleistocene bighorn sheep: Some problems on adaptation, and rele-
vance to today’s American megafauna. Wildlife Society Bulletin 13:351–359.
———. 1998. Deer of the world: Their evolution, behavior, and ecology. Mechanicsburg,
Penn.: Stackpole Books.
Gilpin, M. E. 1987. Spatial stru c t u re and population vulnerability. In Viable populations for
c o n s e rva t i o n, edited by M. E. Soulé, 125–139. New York: Cambridge University Press.
Ginsberg, J. R., and T. P. Young. 1992. Measuring association between individuals or
groups in behavioural studies. Animal Behaviour 44:377–379.
Gittleman, J. L., and P. H. Harvey. 1982. Carnivore home-range size, metabolic needs
and ecology. Behavioral Ecology and Sociobiology 10:57–63.
Gloyne, C. 1999. Cougars and roads: Their use of wildlife crossing structures on the
Trans Canada Highway in Banff National Park, Canada, through analysis of their
tracks. Master’s thesis, University of East Anglia, Norwich, United Kingdom.
Goldman, E. A. 1946. Classification of the races of the puma, Part 2. In The puma: Mys -
terious American cat, edited by S. P. Young and E. A. Goldman, 177–302. Washing-
ton, D.C.: The American Wildlife Institute.
Gonyea, W. J. 1976. Adaptive differences in the body proportions of large felids. Acta
Anatomica 96:81–96.
Goodrich, J. M., and S. W. Buskirk. 1998. Spacing and ecology of North American
badgers (Taxidea taxus) in a prairie-dog (Cynomys leucurus) complex. Journal of Mam -
malogy 79:171–179.
Gorman, M. L. 1976. A mechanism for individual recognition by odour in Herpestes
auropunetatus (Carnivora: Viverridae). Animal Behaviour 24:141–145.
———. 1984. Scent marking and territoriality. Acta Zoologica 171:49–53.
Gorman, M. L., and B. J. Trowbridge. 1989. The role of odor in the social lives of car-
nivores. In Carnivore behavior, ecology and evolution, edited by J. L. Gittleman, 57–88.
Ithaca, N.Y: Cornell University Press.
Grafen, A. 1988. On the uses of data on lifetime reproductive success. In Reproductive
success: Studies of individual variation in contrasting breeding systems, edited by T. H.
Clutton-Brock, 454–471. Chicago: University of Chicago Press.
Greenwood, P. J. 1980. Mating systems, philopatry and dispersal in birds and mammals.
Animal Behaviour 28:1140–1162.
Gregg, C. T. 1985. Plague: An ancient disease in the twentieth century, rev. ed. Albu-
querque: University of New Mexico Press.
Grigione, M. M., P. Burman, V. C. Bleich, and B. M. Pierce. 1999. Identifying individ-
ual mountain lions Felis concolor by their tracks: Refinement of an innovative tech-
nique. Biological Conservation 88:25–32.
Gutiérrez-Espeleta, G. A., S. T. Kalinowski, W. M. Boyce, and P. W. Hedrick. 1998.
Genetic variation in desert bighorn sheep. Desert Bighorn Council Transactions
42:1–10.
References 431
Hackney, R. 2000. Northeastern North Carolina and the red wolf: A cautious optimism.
International Wolf 10:18–19.
Hailey, M. P., C. J. Deutsch, and B. J. Le Boeuf. 1994. Size, dominance and copulatory
success in male northern elephant seals, Mirounga angustirostris. Animal Behaviour
48:1249–1260.
Hair, J. F., R. E. Anderson, R. L. Tatham, and W. C. Black. 1995. Multivariate data
analysis: With readings. Upper Saddle River, N.J.: Prentice Hall.
Halfpenny, J. C., M. R. Sanders, and K. A. McGrath. 1991. Human-lion interactions in
Boulder County, Colorado: Past, present, and future. In Mountain lion–human inter -
action symposium and workshop, April 24–26, 1991, edited by C. S. Braun, 10–16.
Colorado Division of Wildlife, Denver.
Halloran, A. F. 1946. The carnivores of the San Andres Mountains, New Mexico. Jour -
nal of Mammalogy 27:154–161.
Harlow, H. J., F. G. Lindzey, W. D. Van Sickle, and W. A. Gern. 1992. Stress response
of cougars to nonlethal pursuit by hunters. Canadian Journal of Zoology 70:136–139.
Harris, A. H. 1987. Reconstruction of mid-Wisconsin environments in southern New
Mexico. National Geographic Research 3:142–151.
Harvey, M. J., and R. W. Barbour. 1965. Home range of Microtus ochrogaster as deter-
mined by the modified minimum area method. Journal of Mammalogy 46:398–402.
Hast, M. H. 1989. The larynx of roaring and non-roaring cats. Journal of Anatomy
163:117–121.
Hawley, J. W. 1983. Quaternary geology of the Rhodes Canyon (RATSCAT) site. In The
prehistory of Rhodes Canyon, New Mexico, edited by P. L. Eidenbach, 17–32. Tularosa,
Hawley, J. W., G. O. Bachman, and K. Manley. 1976. Quaternary stratigraphy of basin
and range and Great Plains provinces. New Mexico and western Texas. In Quaternary
stratigraphy of North America, edited by W. C. Mahaney, 235–274. Stroudsburg, Penn.:
Stroudsburg, Dowden, Hutchinson and Ross.
Hayes, C. L., E. S. Rubin, M. C. Jorgensen, R. A. Botta, and W. M. Boyce. 2000.
Mountain lion predation of bighorn sheep in the Peninsular Ranges, California. Jour -
nal of Wildlife Management 64:954–959.
Hayne, D. W. 1949. Calculation of size of home range. Journal of Mammalogy 30:1–18.
Hedrick, P. W. 1995. Gene flow and genetic restoration: The Florida panther as a case
study. Conservation Biology 9:996–1007.
Heisey, D. M., and T. K. Fuller. 1985a. Evaluation of survival and cause-specific mor-
tality rates using telemetry data. Journal of Wildlife Management 49:668–674.
———. 1985b. Micromort user’s guide. St. Paul, Minn.: NH Analytical Software.
Hemker, T. P. 1982. Population characteristics and movement patterns of cougars in
southern Utah. Master’s thesis, Utah State University.
Hemker, T. P., F. G. Lindzey, and B. B. Ackerman. 1984. Population characteristics and
movement patterns of cougars in southern Utah. Journal of Wildlife Management
48:1275–1284.
Hemker, T. P., F. G. Lindzey, B. B. Ackerman, and A. J. Button. 1986. Survival of cougar
cubs in a non-hunted population. In Cats of the world: Biology, conservation, and man -
agement, edited by S. D. Miller and D. D. Everett, 327–332. Washington, D.C.:
National Wildlife Federation.
432 References
Hemmer, H. 1978. The evolutionary systematics of living Felidae: Present status and
current problems. Carnivore 1:71–79.
Hemming, J. E. 1969. Cemental deposition, tooth succession, and horn development as
criteria of age in Dall sheep. Journal of Wildlife Management 33:552–558.
Hervert, J. J., and P. R. Krausman. 1986. Desert mule deer use of water developments
in Arizona. Journal of Wildlife Management 50:670–676.
Hiraiwa-Hasegawa, M. 1993. Skewed birth sex ratios in primates: Should high-ranking
mothers have daughters or sons? Trends in Ecology and Evolution 8:395–400.
Hixon, M. A. 1980. Food production and competitor density as the determinants of
feeding territory size. The American Naturalist 4:510–530.
Hoban, P. A. 1986. San Andres Mountains National Wildlife Refuge: Annual narrative
report. U.S. Department of Interior, Fish and Wildlife Service.
———. 1990. A review of desert bighorn sheep in the San Andres Mountains, New
Mexico. Desert Bighorn Council Transactions 34:14–22.
Hoffmeister, D. F. 1986. Mammals of Arizona. Tucson: University of Arizona Press.
Holling, C. S. 1959. The components of predation as revealed by a study of small mam-
mal predation of the European sawfly. Canadian Entomologist 91:293–320.
Hopkins, R. A. 1989. Ecology of the puma in the Diablo Range, California. Ph.D. diss.,
University of California, Berkeley.
Hopkins, R. A., M. J. Kutilek, and G. L. Shreve. 1986. The density and home range
characteristics of mountain lions in the Diablo Range, California. In Cats of the world:
Biology, conservation, and management, edited by S. D. Miller and D. D. Everett,
223–235. Washington, D.C.: National Wildlife Federation.
Hornocker, M. G. 1969. Winter territoriality in mountain lions. Journal of Wildlife
———. 1970. An analysis of mountain lion predation upon mule deer and elk in the
Idaho Primitive Area. Wildlife Monograph No. 21.
Hornocker, M. G., and T. Bailey. 1986. Natural regulation in three species of felids. In
Cats of the world: Biology, conservation, and management, edited by S. D. Miller and D.
D. Everett, 211–220. Washington, D.C.: National Wildlife Federation.
Hulet, C. V. 1972. A rectal abdominal palpation technique for diagnosing pregnancy in
the ewe. Journal of Animal Science 35:192.
Hummel, M., S. Pettigrew, and J. Murray. 1991. Wild hunters: Predators in peril. Niwot,
Colo.: Roberts Rinehart.
Humphreys, D., and A. Elenowitz. 1988. New Mexico’s mule deer population/environ -
ment/hunt computer model. Final Report, Federal Aid in Wildlife Restoration, Project
W-124-R-11, Work Plan 5, Job 1–4. New Mexico Department of Game and Fish,
Santa Fe.
Hunt, C. 2000. Bear shepherding to reduce human-bear conflict—the partners in life
program. International Bear News 9:14–17.
Hunt, R. M. 1996. Biogeography of the order Carnivora. In Carnivore behavior, ecology,
and evolution, Vol. 2, edited by J. L. Gittleman, 485–541. Ithaca, N.Y.: Cornell Uni-
versity Press.
Hunter, L. T. B., and J. D. Skinner. 1998. Vigilance behaviour in African ungulates: The
role of predation pressure. Behaviour 135:195–211.
Hunter, M. D., and P. W. Price. 1992. Playing chutes and ladders: Heterogeneity and
References 433
the relative roles of bottom-up and top-down forces in natural communities. Ecology
73:724–732.
Iriarte, J. A., W. L. Franklin, W. E. Johnson, and K. H. Redford. 1990. Biogeographic
variation of food habits and body size of the American puma. Oecologia 85:185–190.
James, W. H. 1996. Evidence that mammalian sex ratios at birth are partially controlled
by parental hormone levels at the time of conception. Journal of Theoretical Biology
180:271–286.
———. 2000. The hypothesized hormonal control of offspring sex ratio: Evidence from
families ascertained by schizophrenia and epilepsy. Journal of Theoretical Biology
206:445–447.
Jennrich, R. I., and F. B. Turner. 1969. Measurement of non-circular home range. Jour -
nal of Theoretical Biology 22:227–237.
Jiménez, J. A., K. A. Hughes, G. Alaks, L. Graham, and R. C. Lacey. 1994. An experi-
mental study of inbreeding depression in a natural habitat. Science 266:271–273.
Johnson, N. 1984. Idaho. In Proceedings of the second mountain lion workshop, Zion
National Park, 27–29, November1984, edited by J. Roberson and F. Lindzey, 37–38.
Utah Division of Wildlife Resources, Salt Lake City.
Johnson, W. E., and S. J. O’Brien. 1997. Phylogenetic reconstruction of the felidae using
16S rRNA and NADH-5 mitochondrial genes. Journal of Molecular Evolution 44
(Supplement 1):S98–S116.
Julander, O., W. L. Robinette, and D. A. Jones. 1961. Relation of summer range condi-
tion to mule deer herd productivity. Journal of Wildlife Management 25:54–60.
Kaufmann, J. H. 1983. On the definitions and functions of dominance and territorial-
ity. Biological Review 58:1–20.
Kie, J. G., J. A. Baldwin, and C. J. Evans. 1994. CALHOME home range analysis pro -
gram electronic user’s manual. U.S. Department of Agriculture, Forest Service, Pacific
Southwest Forest Experiment Station.
Knick, S. T. 1990. Ecology of bobcats relative to exploitation and prey decline in south-
eastern Idaho. Wildlife Monograph No. 108.
Koehler, G. M., and M. G. Hornocker. 1991. Seasonal resource use among mountain
lions, bobcats, and coyotes. Journal of Mammalogy 72:391–396.
Kohn, M. H., E. C. York, D. A. Kamradt, G. Haught, R. M. Sauvajot, and R. K. Wayne.
1999. Estimating population size by genotyping faeces. Proceedings of the Royal Society
of London B. 266:657–663.
Kottlowski, F. E., R. H. Flower, M. L. Thompson, and R. W. Foster. 1956. Stratigraphic
studies of the San Andres Mountains, New Mexico. New Mexico Bureau of Mines and
Mineral Research Memoir 1, Socorro, New Mexico.
Krackow, S. 1995. Potential mechanisms for sex ratio adjustment in mammals and birds.
Biological Reviews 70:225–241.
Krausman, P. R., R. C. Etchberger, and R. M. Lee. 1996. Persistence of mountain sheep
populations in Arizona. Southwestern Naturalist 41:399–402.
Krausman, P. R., A. V. Sandoval, and R. C. Etchberger. 1999. Natural history of desert
bighorn sheep. In Mountain sheep of North America, edited by R. Valdez and P. R.
Krausman, 139–191. Tucson: University of Arizona Press.
Krebs, C. J. 1999. Ecological methodology. Menlo Park, Calif.: Benjamin/Cummings.
Krebs, C. J., S. Boutin, R. Boonstra, A. R. E. Sinclair, J. N. M. Smith, M. R. T. Dale,
434 References
K. Martin, and R. Turkington. 1995. Impact of food and predation on the snowshoe
hare cycle. Science 269:1112–1115.
Kruuk, H., M. L. Gorman, and A. Leitch. 1984. Scent marking with the subcaudal
gland by the European badger, Meles meles L. Animal Behavior 32: 899–907.
Kruuk, L. E. B., T. H. Clutton-Brock, S. D. Albon, J. M. Pemberton, and F. E. Guiness.
1999. Population density affects sex ratio variation in red deer. Nature 399:459–461.
Kucera, T. E. 1991. Adaptive variation in sex ratios of offspring in nutritionally stressed
mule deer. Journal of Mammalogy 72:745–749.
Kunkel, K., and D. H. Pletscher. 1999. Species-specific population dynamics of cervids
in a multipredator ecosystem. Journal of Wildlife Management 63:1082–1093.
Kunkel, K. E., T. K. Ruth, D. H. Pletscher, and M. G. Hornocker. 1999. Winter prey
selection by wolves and cougars in and near Glacier National Park, Montana. Journal
of Wildlife Management 63:901–910.
Kurtén, B. 1973. Geographic variation in size in the puma (Felis concolor). Commenta -
tiones Biologicae 63:1–8.
———. 1976. Fossil puma (Mammalia: Felidae) in North America. Netherlands Journal
of Zoology 26:502–534.
Kurtén, B., and E. Anderson. 1980. Pleistocene mammals of North America. New York:
Columbia University Press.
Lack, D. 1947. The significance of clutch size. Ibis 89:302–352.
Lacy, R. C. 1987. Loss of genetic diversity from managed populations: Interacting effects
of drift, mutation, immigration, selection, and population subdivision. Conservation
Biology 1:143–158.
———. 1997. Importance of genetic variation to the viability of mammalian popula-
tions. Journal of Mammalogy 78:320–335.
Laing, S. P., and F. G. Lindzey. 1991. Cougar habitat selection in south-central Utah. In
Mountain lion–human interaction symposium and workshop, April 24–26, 1991, edited
by C. E. Braun, 27–37. Colorado Division of Wildlife, Denver.
———. 1993. Patterns of replacement of resident cougars in southern Utah. Journal of
Mammalogy 74:1056–1058.
Lair, H. 1987. Estimating the location of the focal center in red squirrel home ranges.
Ecology 68:1092–1101.
Lance, W. R. 1982. Contagious ecthyma. In Diseases of wildlife in Wyoming, edited by E.
T. Thorne, N. Kingston, W. R. Jolley, and R. C. Bergstrom, 2–5. Wyoming Game and
Fish Department, Cheyenne.
Lande, R. 1988. Genetics and demography in biological conservation. Science
241:1455–1460.
Lang, E. M. 1956. Sheep survey. Federal Aid in Wildlife Restoration Project W-75-R-3,
Job 8. New Mexico Department of Game and Fish, Santa Fe.
Lange, R. E. 1982. Psoroptic scabies. In Diseases of wildlife in Wyoming, edited by E. T.
Thorne, N. Kingston, W. R. Jolley, and R. C. Bergstrom, 244–247. Wyoming Game
and Fish Department, Cheyenne.
Lange, R. E., A. V. Sandoval, and W. P. Meleney. 1980. Psoroptic scabies in bighorn
sheep (Ovis canadensis mexicana) in New Me x i c o. Journal of Wildlife Diseases
16:77–82.
Larsen, D. G., D. A. Gauthier, and R. L. Markel. 1989. Causes and rate of moose mor-
tality in the southwest Yukon. Journal of Wildlife Management 53:548–557.
References 435
Le Boeuf, B. J., and J. Reiter. 1988. Lifetime reproductive success in northern elephant
seal. In Reproductive success: Studies of individual variation in contrasting breeding sys -
tems, edited by T. H. Clutton-Brock, 344–362. Chicago: University of Chicago Press.
Lenti Boero, D. 1995. Scent-deposition behaviour in alpine marmots (Marmota mar -
mota): Its role in territory defense and social communication. Ethology 100:26–38.
Leyhausen, P. 1965. The communal organization of solitary mammals. Symposia of the
Zoological Society of London 14:249–263.
———. 1979. Cat behaviour: The predatory and social behaviour of domestic and wild
cats. Translated by B. A. Tonkin. New York: Garland Press.
Leyhausen, P., and R. Wolff. 1959. Das Revier einer Hauskatze. Zeitschrift für Tierpsy -
chologie 16:666–670.
Lilliefors, H. W. 1967. On the Kolmogorov-Smirnov test for normality with mean and
variance unknown. Journal of the American Statistical Association 64:399–402.
Lindstedt, S. L., B. J. Miller, and S. W. Buskirk. 1986. Home range, time and body size
in mammals. Ecology 67:413–418.
Lindzey, F. G., B. B. Ackerman, D. Barnhurst, and T. P. Hemker. 1988. Survival rates of
mountain lions in southern Utah. Journal of Wildlife Management 52:664–667.
Lindzey, F. G., B. B. Ackerman, D. Barnhurst, T. Becker, T. P. Hemker, S. P. Laing, C.
Mecham, and W. D. Van Sickle. 1989. Boulder-Escalante cougar project. Final Report.
Utah Division of Wildlife Resources, Salt Lake City.
Lindzey, F. G., W. D. Van Sickle, S. P. Laing, and C. S. Mecham. 1992. Cougar popu-
lation response to manipulation in southern Utah. Wildlife Society Bu l l e t i n
20:224–227.
Lindzey, F. G., W. D. Van Sickle, B. B. Ackerman, D. Barnhurst, T. P. Hemker, and S.
P. Laing. 1994. Cougar population dynamics in southern Utah. Journal of Wildlife
Litvaitis, J. A., J. A. Sherburne, and J. A. Bissonette. 1986. Bobcat habitat use and home
range size in relation to prey density. Journal of Wildlife Management 50:110–117.
Logan, K. A. 1983. Mountain lion population and habitat characteristics in the Big
Horn Mountains of Wyoming. Master’s thesis, University of Wyoming.
Logan, K. A., and L. L. Irwin. 1985. Mountain lion habitats in the Big Horn Moun-
tains, Wyoming. Wildlife Society Bulletin 13:257–262.
Logan, K. A., and L. L. Sweanor. 1998. Cougar management in the West: New Mexico
as a template. In Proceedings of the western association of fish and wildlife agencies, June
26–July 2, 1998, Jackson, Wyoming, 101–111. Wyoming Game and Fish Department,
Cheyenne.
———. 2000. Puma. In Ecology and management of large mammals in North America,
edited by S. Demarias and P. R. Krausman, 347–377. Upper Saddle River, N.J.: Pren-
tice Hall.
Logan, K. A., L. L. Irwin, and R. Skinner. 1986. Characteristics of a hunted mountain
lion population in Wyoming. Journal of Wildlife Management 50:648–654.
Logan, K. A., E. T. Thorne, L. L. Irwin, and R. Skinner. 1986. Immobilizing wild moun-
tain lions (Felis concolor) with ketamine hydrochloride and xylazine hydrochloride.
Journal of Wildlife Diseases 22:97–103.
Logan, K. A., L. L. Sweanor, J. Frank Smith, and Maurice G. Hornocker. 1999. Cap-
turing pumas with foot-hold snares. Wildlife Society Bulletin 27:201–208.
Lopez-Gonzalez, C. A. 1999. Implicaciones para la Conservacion y el Manejo de Pumas
436 References
(Puma concolor) Utilizando como Modelo una Poblacion Sujeta a Caceria Deportiva.
Ph.D. Tesis, Universidad Nacional Autonoma de Mexico.
Lorenz, K. 1971. Studies in animal and human behaviour, Volume 2. Translated by Robert
Martin. London: Methuen and Company.
Lott, D. F. 1991. Intraspecific variation in the social systems of wild vertebrates. Cambridge:
Cambridge University Press.
Macdonald, D. W. 1983. The ecology of carnivo re social behaviour. Na t u re
301:379–384.
Mace, G. M., P. H. Harvey, and T. H. Clutton-Brock. 1983. Vertebrate home-range size
and energetic requirements. In The ecology of animal movement, edited by I. Swingland
and P. J. Greenwood, 32–53. Oxford: Clarendon Press.
Maehr, D. 1990a. Florida panther movements, social organization, and habitat utilization.
Final Report. Bureau of Wildlife Research, Florida Game and Fresh Water Fish Com-
mission, Tallahassee.
Maehr, D. S. 1990b. The Florida panther and private lands. Conservation Biology
4:167–170.
———. 1996. Comparative ecology of bobcat, black bear, and Florida panther in south
Florida. Ph.D. diss., University of Florida, Gainesville.
Maehr, D. 1997a. The comparative ecology of bobcat, black bear, and Florida panther
in south Florida. Bulletin of the Florida Museum of Natural History, 40.
Maehr, D. S. 1997b. The Florida panther: Life and death of a vanishing carnivore. Wash-
ington, D.C.: Island Press.
Maehr, D. S., and G. B. Caddick. 1995. Demographics and genetic introgression in the
Florida panther. Conservation Biology 9:1295–1298.
Maehr, D. S., and J. A. Cox. 1995. Landscape features and panthers in Florida. Conser -
vation Biology 9:1008–1019.
Maehr, D. S., and C. T. Moore. 1992. Models of mass growth for three North Ameri-
can cougar populations. Journal of Wildlife Management 56:700–707.
Maehr, D., J. C. Roof, E. D. Land, and J. W. McCowan. 1989. First reproduction of a
panther (Felis concolor coryi) in southwestern Florida. U.S.A. Mammalia 53:129–131.
Maehr, D., E. D. Land, and J. C. Roof. 1991. Social ecology of Florida panthers.
National Geographic Research and Exploration 7:414–431.
Marshall, J. P., and S. Boutin. 1999. Power analysis of wolf–moose functional responses.
Journal of Wildlife Management 63:396–402.
Marshall, L. G., S. D. Webb, J. J. Sepkoski, Jr., and D. M. Raup. 1982. Mammalian evo-
lution and the great American interchange. Science 215:1351–1357.
Martin, L. D. 1980. Functional morphology and the evolution of cats. Transactions of
the Nebraska Academy of Sciences 8:141–154.
Matson, P. A., and M. D. Hunter. 1992. The relative contributions of top-down and
bottom-up forces in population and community ecology. Ecology 73:723.
Mayr, E. 1996. The modern evolutionary theory. Journal of Mammalogy 77:1–7.
McBride, R. T. 1976. The status and ecology of the mountain lion Felis concolor stanleyana
of the Texas–Mexico border. Master’s thesis, Sul Ross State University, Alpine, Texas.
McBride, R. T., R. M. McBride, J. L. Cashman, and D. S. Maehr. 1993. Do mountain
lions exist in Arkansas? Proceedings of the Annual Conference of Southeastern Fish and
Wildlife Agencies 47:394–402.
McCullough, D. R. 1996. Spatially structured populations and harvest theory. Journal of
References 437
McDaniel, G. W., K. S. McKelvey, J. R. Squires, and L. F. Ruggiero. 2000. Efficacy of

lures and hair snares to detect lynx. Wildlife Society Bulletin 28:119–123.
McLaren, B. E., and R. O. Peterson. 1994. Wolves, moose, and tree rings on Isle Royale.
Science 266:1555–1558.
Mech, L. D. 1977a. Productivity, mortality, and population trends of wolves in north-
eastern Minnesota. Journal of Mammalogy 58:559–574.
———. 1977b. Wolf-pack buffer zones as prey reservoirs. Science 198:320–321.
Mech, L. D., and G. D. DelGuidice. 1985. Limitations of the marrow fat technique as
an indicator of body condition. Wildlife Society Bulletin 13:204–206.
Meffe, G. K., and C. R. Carroll. 1997. Principles of conservation biology. 2d ed. Sunder-
land, Mass: Sinauer Associates.
Mehrer, C. F. 1975. Some aspects of reproduction in captive mountain lions Felis con -
color, bobcats Lynx rufus, and lynx Lynx canadensis. Ph.D. diss., University of North
Dakota, Grand Forks.
Menge, B. A. 1992. Community regulation: Under what conditions are bottom-up fac-
tors important on rocky shores? Ecology 73:755–765.
Messier, F. 1991. The significance of limiting and regulating factors on the demography
of moose and white-tailed deer. Journal of Animal Ecology 60:377–393.
———. 1995. On the functional and numerical responses of wolves to changing prey
density. In Ecology and conservation of wolves in a changing world, edited by L. N. Car-
byn, S. H. Fritts, and D. R. Seip, 187–197. Canadian Circumpolar Institute, Occa-
sional Publication 35.
Mills, M. G. L. 1990. Kalahari hyaenas. London: Unwin Hyman.
Minnis, D. L. 1998. Wildlife policy-making by the electorate: An overview of citizen-
s p o n s o red ballot measures on hunting and trapping. Wildlife Society Bu l l e t i n
26:75–83.
Minta, S. 1993. Use of spatio-temporal interaction statistics on badgers to test predic-
tions of territorial polygyny theory. Oecologia 96:402–409.
Mitani, J. C., J. Gros-Louis, and A. F. Richards. 1996. Sexual dimorphism, the opera-
tional sex ratio, and the intensity of male competition in polygynous primates. Amer -
ican Naturalist 147:966–980.
Modig, A. O. 1996. Effects of body size and harem size on male reproductive behaviour
in the southern elephant seal. Animal Behaviour 51:1295–1306.
Mohr, C. O. 1947. Table of equivalent populations of North America small mammals.
American Midland Naturalist 37:223–249.
Moore, J., and R. L. Ali. 1984. Are dispersal and inbreeding avoidance related? Animal
Behaviour 32:94–112.
Morris, D. W. 1982. Age-specific dispersal strategies in iteroparous species: Who leaves
when? Evolutionary Theory 6:53–65.
Mowat, G., B. G. Slough, and S. Boutin. 1996. Lynx recruitment during a snowshoe
hare population peak and decline in southwest Yukon. Journal of Wildlife Management
60:441–452.
Mowat, G., and C. Strobeck. 2000. Estimating population size of grizzly bears using hair
capture, DNA profiling, and mark-recapture analysis. Journal of Wildlife Management
64:183–193.
Müller-Schwarze, D. 1971. Pheromones in black-tailed deer (Odocoileus hemionus
columbianus). Animal Behaviour 19:141–152.
438 References
Muñoz, D. 1983. San Andres National Wildlife Refuge: Annual narrative report. U.S.
Department of Interior Fish and Wildlife Service.
Murphy, K. M. 1983a. Relationships between a mountain lion population and hunting
pressure in western Montana. Master’s thesis, University of Montana.
———. 1983b. Relationships between a mountain lion population and hunting pressure in
western Montana. Final Report. Federal Aid in Wildlife Restoration Project W-120-R-
13 and 14, Job 1. Montana Department of Fish, Wildlife and Parks, Helena.
———. 1998. The ecology of the cougar (Puma concolor) in the northern Yellowstone
ecosystem: Interactions with prey, bears, and humans. Ph.D. diss., University of Idaho,
Moscow.
Murphy, K. M., M. Culver, M. Menotti-Raymond, V. David, M. G. Hornocker, and S.
J. O’Brien. 1998a. Cougar reproductive success in the Northern Yellowstone Ecosys-
tem. Chapter 3 in The ecology of the cougar (Puma concolor) in the northern Yellow-
stone ecosystem: Interactions with prey, bears, and humans, K. M. Murphy, 78–112.
Ph.D. diss., University of Idaho, Moscow.
Murphy, K. M., G. S. Felzien, M. G. Hornocker, and T. Lemke. 1998b. Cougar food
habits, prey selection, and predation rates in the northern Yellowstone ecosystem.
Chapter 1 in The ecology of the cougar (Puma concolor) in the northern Yellowstone
ecosystem: Interactions with prey, bears, and humans. K. M. Murphy, 2–62. Ph.D.
diss., University of Idaho, Moscow.
Murphy, K. M., P. I. Ross, and M. G. Hornocker. 1999. The ecology of anthropogenic
influences on cougars. In Carnivores in ecosystems: The Yellowstone experience, edited by
T. W. Clark, A. P. Curlee, S. C. Minta, and P. M. Kareiva. New Haven, Conn.: Yale
University Press.
Musgrave, M. E. 1926. Some habits of mountain lions in Arizona. Journal of Mammal -
ogy 7:282–285.
Nash, W. G., and S. J. O’Brien. 1982. Conserved regions of homologous G-banded
chromosomes between orders in mammalian evolution: Carnivores and primates. Pro -
ceedings of the National Academy of Sciences 79:6631–6635.
Neal, D. L., G. N. Steger, and R. C. Bertram. 1987. Mountain lions: Preliminary find -
ings on home-range use and density in the central Sierra Nevada. Research Note PSW-
392. Pacific Southwest Forest and Range Experiment Station, Forest Service, U.S.
Department of Agriculture, Berkeley, California.
Neilson, R. P. 1986. High-resolution climatic analysis and Southwest biogeography. Sci -
ence 232:27–33.
Noble, G. K. 1939. Dominance in the life of birds. Auk 56:263–273.
Norrix, L. W., D. W. DeYoung, P. R. Krausman, R. C. Etchenberger, and T. J. Glattke.
1995. Conductive hearing loss in bighorn sheep. Jo u rnal of Wildlife Diseases
31:223–227.
Noss, R. F., H. B. Quigley, M. G. Hornocker, T. Merrill, and P. C. Paquet. 1996. Con-
servation biology and carnivore conservation in the Rocky Mountains. Conservation
Biology 10:949–963.
Nowak, R. M. 1976. The cougar in the United States and Canada. U.S. Department of
Interior Fish and Wildlife Service, Washington, D.C., and New York Zoological Soci-
ety, New York.
———. 1991. Walker’s mammals of the world. Vol. 2. Baltimore: John Hopkins Univer-
sity Press.
Nowell, K., and P. Jackson, editors. 1996. Wild cats: Status and conservation action plan.
References 439
The World Conservation Union, Species Survival Commission, Cat Specialist Group,
Gland, Switzerland.
Núñez, R. B. 1999. Hábitos Alimentarios del jaguar (Panthera onca, Linnaeus 1778) y del
puma (Puma concolor, Linnaeus 1771) en la Reserva de la Biósfera Chamela-Cuixmala,
Jalisco, Mexico. Master’s Tesis, Licenciatura, Universidad Michoacana de San Nicolas
de Hidalgo, Morelia, Michoacán.
Núñez, R., B. Miller, and F. Lindzey. 2000a. Food habits of jaguars and pumas in Jalisco,
Mexico. Journal of Zoology, London 252:373–379.
Núñez, R., B. Miller, and F. Lindzey. 2000b. Ecology of jaguars in the Chamela-Cuix-
male Biosphere Reserve, Jalisco, Mexico. In El jaguar en el nuevo milenio. Una evalua -
cion de su estado, deteccion deprioridades y recomendaciones para la conservacion de los
jaguares en America, edited by R. A. Medellin, C. Chetkiewicz, A. Rabinowitz, K. H.
Redford, J. G. Robinson, E. Sanderson, and A. Taber. Mexico, D.F.: Universidad
Nacional Autonoma de Mexico.
Nunn, C. L., and M. E. Pereira. 2000. Group histories and offspring sex ratios in ring-
tailed lemurs (Lemur catta). Behavioral Ecology and Sociobiology 48:18–28.
Nunney, L., and D. R. Elam. 1994. Estimating the effective population size of conserved
populations. Conservation Biology 8:175–184.
O’Brien, S. J. 1996. Molecular genetics and phylogenetics of the Felidae. In Wild cats:
Status and conservation action plan, edited by K. Nowell and P. Jackson, xxiii–xxiv. The
World Conservation Union, Species Survival Commission, Cat Specialist Group,
Gland, Switzerland.
———. 1997. The human-cat connection. National Geographic 191:77–85.
O’Brien, S. J., J. S. Martenson, C. Packer, L. Herbst, V. Vos, P. Joslin, J. Ott-Joslin, D.
E. Wildt, and M. Bush. 1987. Biochemical genetic variation in geographic isolates of
African and Asiatic lions. National Geographic Research 3:114–124.
O’Brien, S. J., M. E. Roelke, N. Yuhi, K. W. Richards, W. E. Johnson, W. L. Franklin,
A. E. Anderson, O. L. Bass, Jr., R. C. Belden, and J. S. Martenson. 1990. Genetic
introgression within the Florida panther, Felis concolor coryi. National Geographic
Research 6:485–494.
Ockenfels, R. A. 1994. Factors affecting adult pronghorn mortality rates in central Ari-
zona. Arizona Game and Fish Department, Wildlife Digest 16:1–11.
O’Donoghue, M., S. Boutin, C. J. Krebs, and E. J. Hofer. 1997. Numerical responses of
coyotes and lynx to the snowshoe hare cycle. Oikos 80:150–162.
O’Donoghue, M., S. Boutin, C. J. Krebs, G. Zuleta, D. L. Murray, and E. J. Hofer.
1998. Functional responses of coyotes and lynx to the snowshoe hare cycle. Ecology
79:1193–1208.
Ordway, L. L., and P. R. Krausman. 1986. Habitat use by desert mule deer. Journal of
Ott, L. 1988. An introduction to statistical methods and data analysis. Boston: PWS-Kent
Publishing.
Owen-Smith, N. 1977. On territoriality in ungulates and an evolutionary model. Quar -
terly Reviews of Biology 52:1–38.
———. 1987. Pleistocene extinctions: The pivotal role of megaherbivores. Paleobiology
13:351–362.
———. 1989. Megafaunal extinctions: The conservation message from 11,000 years
B.P. Conservation Biology 3:405–412.
Packer, C. 1986. The ecology of sociality in felids. In Ecological aspects of social evolution,
440 References
edited by D. I. Rubenstein and R. W. Wrangham, 429–451. Princeton, N.J.: Prince-

ton University Press.
Packer, C., and A. E. Pusey. 1982. Cooperation and competition within coalitions of
male lions: Kin selection or game theory? Nature 296:740–742.
———. 1983. Adaptations of female lions to infanticide by incoming males. American
Naturalist 121:716–728.
Packer, C., D. A. Gilbert, A. E. Pusey, and S. J. O’Brien. 1991. A molecular genetic
analysis of kinship and cooperation in African lions. Nature 351:562–565.
Packer, C., A. E. Pusey, H. Rowley, D. A. Gilbert, J. Martenson, and S. J. O’Brien. 1991.
Case study of a population bottleneck: Lions of the Ngorongoro Crater. Conservation
Biology 5:219–230.
Packer, C., D. Scheel, and A. E. Pusey. 1990. Why lions form groups: Food is not
enough. American Naturalist 136:1–19.
Packer, C., L. Herbst, A. E. Pusey, J. D. Bygott, J. P. Hanby, S. J. Cairns, and M. B. Mul-
der. 1988. Reproductive success of lions. In Reproductive success: Studies of individual
variation in contrasting breeding systems, edited by T. H. Clutton-Brock, 363–383.
Chicago: University of Chicago Press.
Padley, W. D. 1990. Home ranges and social interaction of mountain lions (Felis con -
color) in the Santa Ana Mountains, California. Master’s thesis, California Polytechnic
University, Pomona.
Padley, W. D. 1996a. Mountain lion (Felis concolor) vocalizations in the Santa Ana
Mountains, California. Paper presented at the Fifth Mountain Lion Workshop, 27
February – 1 March 1996, San Diego, California.
Padley, W. D., editor. 1996b. Proceedings of the fifth mountain lion workshop, 27 Febru -
ary–1 March 1996, San Diego, California. The Southern California Chapter of the
Wildlife Society.
Palanza, P., S. Parmigiana, and F. S. vom Saal. 1994. Male urinary cues stimulate intra-
sexual aggression and urine-marking in wild female mice, Mus musculus domesticus.
Animal Behaviour 48:245–247.
Parker, G. R., J. W. Maxwell, L. D. Morton, and G. E. Smith. 1983. The ecology of the
lynx Lynx canadensis on Cape Breton Island Canada. Canadian Journal of Zoology
61:770–786.
Pech, R. P., A. R. E. Sinclair, A. E. Newsome, and P. C. Catling. 1992. Limits to preda-
tor regulation of rabbits in Australia: Evidence from predator-removal experiments.
Oecologia 89:102–112.
Pecon-Slattery, J., and S. J. O’Brien. 1998. Patterns of Y and X Chromosome DNA
sequence divergence during the felidae radiation. Genetics 148:1245–1255.
Pederson, J. 1996. The new bighorn sheep of the Fra Cristobal Range. New Mexico
Wildlife 41:14–18.
Peters, G. 1978. Vergleichende untersuchung zur lautgeburg einiger feliden (Mammalia,
Felidae). Spixiana, Supplement 1.
Peters, G., and W. C. Wozencraft. 1989. Acoustic communication by fissiped carnivores.
In Carnivore behavior, ecology and evolution, edited by G. L. Gittleman, 14–56. Ithaca,
N.Y.: Cornell University Press.
Phillips, D. M., D. J. Harrison, and D. C. Payer. 1998. Seasonal changes in home range
area and fidelity in martens. Journal of Mammalogy 79:180–190.
References 441
Picton, H. D. 1984. Climate and the prediction of reproduction of three ungulate

species. Journal of Applied Ecology 21:869–879.
Pierce, B. M., V. C. Bleich, and R. T. Bowyer. 2000a. Prey selection by mountain lions
and coyotes on mule deer: Effects of hunting style, body size, and reproductive state.
Journal of Mammalogy 81:462–472.
———. 2000b. Social organization of mountain lions: Does a land-tenure system reg-
ulate population size? Ecology 81:1533–1543.
Pierce, B. M., V. C. Bleich, J. D. Wehausen, and R. T. Bowyer. 1999. Migratory patterns
of mountain lions: Implications for social regulation and conservation. Journal of
Mammalogy 80:986–992.
Pitelka, F. A. 1959. Numbers, breeding schedule and territoriality in pectoral sandpipers
of northern Alaska. Condor 61:233–264.
Pollock, K. H., S. R. Winterstein, C. M. Bunck, and P. D. Curtis. 1989. Su rv i val analysis
in telemetry studies: The staggered entry design. Jo u rnal of Wildlife Management 53:7–15.
Poole, K. G. 1994. Characteristics of an unharvested lynx population during a snowshoe
hare decline. Journal of Wildlife Management 58:608–618.
Post, E., R. O. Peterson, N. C. Stenseth, and B. E. McLaren. 1999. Ecosystem conse-
quences of wolf behavioural response to climate. Nature 401:905–907.
Power, M. E. 1992. Top-down and bottom-up forces in food webs: Do plants have pri-
macy? Ecology 73:733–746.
Pusey, A. E., and C. Packer. 1987. The evolution of sex-biased dispersal in lions. Behav -
iour 101:275–310.
Pusey, A., and M. Wolf. 1996. Inbreeding avoidance in animals. Trends in Ecology and
Evolution 11:201–206.
Putman, R. 1988. The natural history of deer. Comstock Publishing Associates, Ithaca,
Quigley, H. B., G. M. Koehler, and M. G. Hornocker. 1989. Dynamics of a mountain
lion population in central Idaho over a 20-year period. Paper presented at the Third
Mountain Lion Workshop, 6–8 December 1988, Prescott, Arizona.
Rabb, G. B. 1959. Reproductive and vocal behavior in captive pumas. Journal of Mam -
malogy 40:616–617.
Rabinowitz, A. R., and B. G. Nottingham, Jr. 1986. Ecology and behaviour of the jaguar
(Panthera onca) in Belize, Central America. Journal of Zoology, London 210:149–159.
Rail, C. D. 1985. Plague ecotoxicology. Springfield, Ill.: Charles C. Thomas.
Randall, J. A. 1987. Sandbathing as a territorial scent-mark in the banner tail kangaroo
rat, Dipodomys spectabilis. Animal Behaviour 35:426–434.
Rasa, O. A. E. 1973. Marking behaviour and significance in the African dwarf mon-
goose, Helogale undulata rufula. Zeitschrift für Tierpsychologie 32:449–488.
Read, A. F., and P. H. Harvey. 1989. Life history differences among the eutherian radi-
ations. Journal of Zoology, London 219:329–353.
Redford, K. H. 1992. The empty forest. BioScience 42:412–422.
Rettenberger, G., C. Klett, U. Zechner, J. Bruch, W. Just, W. Vogel, and H. Hameister.
1995. ZOO-FISH analysis: Cat and human karyotypes closely resemble the putative
ancestral mammalian karyotype. Chromosome Research 3:479–486.
Ricklefs, R. E. 1990. Ecology. 3d ed. New York: W. H. Freeman.
Roberson, J. 1984. Pursuit seasons. In Proceedings of the second mountain lion workshop,
442 References
Zion National Park, 27–29 November 1984, edited by J. Roberson and F. Lindzey,
191–203. Utah Division of Wildlife Resources, Salt Lake City.
Robinette, W. L., D. A. Jones, G. Rogers, and J. S. Gashwiler. 1957. Notes on tooth
development and wear for Rocky Mountain mule deer. Journal of Wildlife Management
21:134–153.
Robinette, W. L., J. S. Gashwiler, and O. W. Morris. 1961. Notes on cougar productiv-
ity and life history. Journal of Mammalogy 42:204–217.
Roelke, M. E., D. J. Forrester, E. R. Jacobson, G. V. Kollias, F. W. Scott, M. C. Barr, J.
F. Everman, and E. C. Pirtle. 1993a. Seroprevalence and infectious disease agents in
free-ranging Florida panthers (Felis concolor cory i). Journal of Wildlife Diseases
29:36–49.
Roelke, M. E., J. S. Martenson, and S. J. O’Brien. 1993b. The consequences of demo-
graphic 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. Journal of Wildlife Management 56:417–426.
———. 1996. Cougar predation on moose in southwestern Alberta. Alces 32:1–8.
Ross, P. I., M. G. Jalkotzy, and P. Daoust. 1995. Fatal trauma sustained by cougars, Felis
c o n c o l o r, while attacking prey in southern Alberta. Canadian Fi e l d - Na t u ralist
109:261–263.
Ross, P. I., M. G. Jalkotzy, and M. Festa-Bianchet. 1997. Cougar predation on bighorn
sheep in southwestern Alberta during winter. Canadian Journal of Zoology
74:771–775.
Rutberg, A. T. 1987. Adaptive hypotheses of birth-synchrony in ruminants: An inter-
specific test. American Naturalist 130:692–710.
Ruth, T. K. 1991. Mountain lion use of an area of high recreational development in Big
Bend National Park, Texas. Master’s thesis, Texas A&M University.
Ruth, T. K., K. A. Logan, L. L. Sweanor, M. G. Hornocker, and L. J. Temple. 1998.
Evaluating cougar translocation in New Mexico. Journal of Wildlife Management
62:1264–1275.
Saccheri, I., M. Kuussaari, M. Kankare, P. Vikman, W. Fortelius, and I. Hanski. 1998.
Inbreeding and extinction in a butterfly metapopulation. Nature 392:491–494.
Sadleir, R. M. F. S. 1982. Energy consumption and subsequent partitioning in lactating
black-tailed deer. Canadian Journal of Zoology 60:382–386.
Saiz, R. B. 1975. Ecology and behavior of the gemsbok on the White Sands Missile
Range, New Mexico. Master’s thesis, Colorado State University.
Sale, M., and K. Laumbach. 1989. Reconnaissance in the Upper Jornada del Muerto and
Hembrillo Canyon and Other Special Projects, White Sands Missile Range, New Mexico.
Tularosa, N.M.: Human Systems Research.
Saltz, D., and P. U. Alkon. 1985. A simple computer-aided method for estimating radio-
location error. Journal of Wildlife Management 49:664–668.
Sandell, M. 1986. Movement patterns of male stoats Mustela erminea during the mating
season: Differences in relation to social status. Oikos 47:63–70.
———. 1989. The mating tactics and spacing patterns of solitary carnivores. In Carni -
vore behavior, ecology and evolution, edited by G. L. Gittleman, 164–182. Ithaca, N.Y.:
Cornell University Press.
Sandoval, A. V. 1979a. Bighorn sheep status report from New Mexico. Desert Bighorn
Council Transactions 23:82–87.
References 443
———. 1979b. Preferred habitat of desert bighorn sheep in the San Andres Mountains,
New Mexico. Master’s thesis, Colorado State University.
———. 1980. Management of a psoroptic scabies epizootic in bighorn sheep (Ovis
canadensis mexicana) in New Mexico. Desert Bighorn Council Transactions 24:21–28.
Saunders, G. C. 1970. Development of the immune response. In Biology of the immune
response, edited by P. Abramoff and M. F. La Via, 93–135. New York: McGraw-Hill.
Savage, D. E., and D. E. Russell. 1983. Mammalian Paleofauna of the World. London:
Addison- Wesley.
Schaller, G. B. 1972. The Serengeti lion. Chicago: University of Chicago Press.
———. 1993. The last panda. Chicago: University of Chicago Press.
Schaller, G. B., and P. G. Crawshaw. 1980. Movement patterns of jaguars. Biotropica
12:161–168.
Schenk, A., and K. M. Kovacs. 1995. Multiple mating between black bears revealed by
DNA fingerprinting. Animal Behavior 50:1483–1490.
Schmidt, J. L. 1978. Early management: Intentional and otherwise. In Big game of North
America: Ecology and management, edited by J. L. Schmidt and D. L. Gilbert,
257–270. Harrisburg, Penn.: Stackpole Books.
Schmidt, K., W. Jedrzejewski, and H. Okarma. 1997. Spatial organization and social
relations in the Eurasian lynx population in Bialowieza Primeval Forest, Poland. Acta
Theriologica 42:289–312.
Schoener, T. W. 1981. An empirically based estimate of home range. Theoretical Popula -
tion Biology 20:281–325.
Seaman, D. E., and R. A. Powell. 1996. An evaluation of the accuracy of kernel density
estimators for home range analysis. Ecology 77:2075–2085.
Seidensticker, J. 1997. Saving the tiger. Wildlife Society Bulletin 25:6–17.
Seidensticker, J. C., M. G. Hornocker, W. V. Wiles, and J. P. Messick. 1973. Mountain
lion social organization in the Idaho Primitive Area. Wildlife Monograph No. 35.
Seip, D. R. 1992. Factors limiting woodland caribou populations and their interrela-
tionships with wolves and moose in southeastern British Columbia. Canadian Journal
of Zoology 70:1494–1503.
Shaffer, M. 1987. Minimum viable populations: Coping with uncertainty. In Viable pop -
ulations for conservation, edited by M. E. Soulé, 69–86. New York: Cambridge Uni-
versity Press.
Shaw, H. G. 1977. Impact of mountain lion on mule deer and cattle in northwestern
Arizona. In Proceedings of the 1975 predator symposium, June 16–19, 1975, edited by
R. L. Phillips and C. Jonkel, 17–32. University of Montana, Missoula.
———. 1980. Ecology of the mountain lion in Arizona. Final Report, Federal Aid in
Wildlife Restoration Project W-78-R, Work Plan 2, Job 13. Arizona Game and Fish
Department, Phoenix.
Shields, W. M. 1982. Philopatry, inbreeding and the evolution of sex. Albany: State Uni-
versity of New York Press.
———. 1987. Dispersal and mating: Investigating their causal connections. In Mam -
malian dispersal patterns: The effects of social structure on population genetics, edited by
B. D. Chepko-Sade and Z. T. Halpin, 3–24. Chicago: University of Chicago Press.
Short, H. L. 1981. Nutrition and metabolism. In Mule and black-tailed deer of North
America, edited by O. C. Wallmo, 99–127. Washington, D.C.: The Wildlife Manage-
ment Institute.
Sibley, R. M., and P. Calow. 1986. Physiological ecology of animals: An evolutionary
approach. Palo Alto, Calif.: Blackwell Scientific Publications.
444 References
Sinclair, A. R. E. 1992. Do large mammals disperse like small mammals? In Animal dis -
persal: Small mammals as a model, edited by N. C. Stenseth and W. Z. Lidicker, Jr.,
229–242. London: Chapman and Hall.
Sinclair, A. R. E., S. A. R. Mduma, and P. Arcese. 2000. What determines phenology
and synchrony of ungulate breeding in Serengeti? Ecology 81:2100–2111.
Sinclair, D. F. 1985. On tests of spatial randomness using mean nearest neighbor dis-
tance. Ecology 66:1084–1085.
Singer, F. J., A. Harting, K. K. Symonds, and M. B. Coughenour. 1997. Density
dependence, compensation, and environmental effects on elk calf mortality in Yellow-
stone National Park. Journal of Wildlife Management 61:12–25.
Sitton, L. W., and S. Wallen. 1976. California mountain lion study. Sacramento: Califor-
nia Department of Fish and Game.
Smallwood, K. S., and E. L. Fitzhugh. 1993. A rigorous technique for identifying indi-
vidual mountain lions Felis concolor by their tracks. Biological Conservation 65:51–59.
Smith, C., R. Valdez, J. L. Holechek, P. J. Zwank, and M. Cardenas. 1998. Diets of
native and non-native ungulates in southcentral New Mexico. Southwestern Naturalist
43:163–169.
Smith, J. L. D. 1993. The role of dispersal in structuring the Chitwan tiger population.
Behaviour 124:165–195.
Smith, J. L. D., and C. McDougal. 1991. The contribution of variance in lifetime repro-
duction to effective population size in tigers. Conservation Biology 5:484–490.
Smith, J. L. D., C. W. McDougal, and M. E. Sunquist. 1987. Female land tenure sys-
tem in tigers. In Tigers of the world: The biology, biopolitics, management and conserva -
tion of an endangered species, edited by R. L. Tilson and U.S. Seal, 97–109. Park Ridge,
N.Y.: Noyes Press.
Smith, J. L. D., C. McDougal, and D. Miquelle. 1989. Scent marking in free-ranging
tigers, Panthera tigris. Animal Behaviour 37:1–10.
Smith, R. H., and A. LeCount. 1979. Some factors affecting survival of desert mule deer
fawns. Journal of Wildlife Management 43:657–665.
Smith, R. L. 1984. Human sperm competition. In Sperm competition and the evolution
of animal mating systems, edited by R. L. Smith, 601–659. Orlando, Fl.: Academic
Press.
Smith, T. E., R. R. Duke, M. J. Kutilek, and H. T. Harvey. 1986. Mountain lions (Felis
concolor) in the vicinity of Carlsbad Caverns National Park, New Mexico, and
Guadalupe Mountains National Park, Texas. Final Report. U.S. Department of Interior,
National Park Service, Santa Fe.
Spreadbury, B. 1989. Cougar ecology and related management implications and strate-
gies in southeastern British Columbia. Master’s thesis, University of Calgary, Alberta.
Spreadbury, B. R., K. Musil, J. Musil, C. Kaisner, and J. Kovak. 1996. Cougar popula-
tion characteristics in southeastern British Columbia. Journal of Wildlife Management
60:962–969.
Stander, P. E., P. J. Haden, Kaqece, and Ghau. 1997. The ecology of asociality in Namib-
ian leopards. Journal of Zoology, London 242:343–364.
Stearns, S. C. 1992. The evolution of life histories. Oxford: Oxford University Press.
Stenseth, N. C., and W. Z. Lidicker, Jr., editors. 1992. Animal dispersal: Small mammals
as a model. London: Chapman and Hall.
Stewart, R. R., E. H. Kowal, R. Beaulieu, and T. W. Rock. 1985. The impact of black
bear removal on moose calf survival in east-central Saskatchewan. Alces 21:403–418.
References 445
Stoddart, D. M. 1980. The ecology of vertebrate olfaction. London: Chapman and Hall.
Strong, D. R. 1992. Are trophic cascades all wet? Differentiation and donor-control in
speciose ecosystems. Nature 73:747–754.
Stuart, A. J. 1991. Mammalian extinctions in the late Pleistocene of Northern Eurasia
and North America. Biological Review 66:453–562.
Sunquist, M. E. 1981. The social organization of tigers (Panthera tigris) in Royal Chitawan
National Park, Nepal. Smithsonian Contributions to Zoology No. 336. Washington,
D.C.: Smithsonian Institution Press.
Sunquist, M. E., and F. C. Sunquist. 1989. Ecological constraints on predation by large
felids. In Carnivore behavior, ecology and evolution, edited by J. L. Gittleman, 283–301.
Ithaca, N.Y.: Cornell University Press.
Sweanor, L. L. 1990. Mountain lion social organization in a desert environment. Mas-
ter’s thesis, University of Idaho.
Sweanor, L. L., K. A. Logan, and M. G. Hornocker. 2000. Cougar dispersal patterns,
metapopulation dynamics, and conservation. Conservation Biology 14:798–808.
Sweitzer, R. A., S. H. Jenkins, and J. Berger. 1997. Near-extinction of porcupines by
mountain lions and consequences of ecosystem change in the Great Basin Desert. Con -
servation Biology 11:1407–1417.
Swihart, R. K., and N. A. Slade. 1985. Influence of sampling interval on estimates of
home-range size. Journal of Wildlife Management 49:1019–1025.
———. 1986. The importance of statistical power when testing for independence in
animal movements. Ecology 67:255–258.
Taber, A. B., A. J. Novaro, N. Neris, and F. H. Colman. 1997. The food habits of sym-
patric jaguar and puma in the Paraguayan Chaco. Biotropica 29:204–213.
Taylor, M. E., and N. Abrey. 1982. Marten, Martes americana, movements and habitat
use in Algonquin Provincial Park, Ontario. Canadian Field-Naturalist 96:439–447.
Teel, T. L. 1999. Utah stakeholders’ attitudes toward wildlife-related issues. Master’s the-
sis, Utah State University.
Teer, J. G., D. L. Drawe, T. L. Blankenship, W. F. Andelt, R. S. Cook, J. G. Kie, F. F.
Knowlton, and M. White. 1991. Deer and coyotes: The Welder experiment. Transac -
tions of the 56th wildlife and natural resources conference, March 17–22, 1991, Edmon -
ton, Alberta, 550–560. The Wildlife Management Institute, Washington, D.C.
Temple, L. 1981. Big game research. Final Report. Federal Aid in Wildlife Restoration,
Project W-124-R-5. New Mexico Department of Game and Fish, Santa Fe.
Templeton, A. R. 1987. Inferences on natural population structure from genetic studies
on captive mammalian populations. In Mammalian dispersal patterns: The effects of
social structure on population genetics, edited by B. D. Chepko-Sade and Z. T. Halpin,
257–272. Chicago: University of Chicago Press.
Terborgh, J., J. A. Estes, P. Paquet, K. Ralls, D. Boyd - Hager, B. J. Miller, and R. F. Noss.
1999. The role of top carnivo res in regulating terrestrial ecosystems. In Continental con -
s e rva t i o n, edited by M. E. Soulé and J. Terborgh, 39–64. Washington, D.C.: Island Press.
Thomas, C. S., and J. C. Coulson. 1988. Reproductive success of kittiwake gulls, Rissa
tridactyla. In Reproductive success: Studies of individual variation in contrasting breeding
systems, edited by T. H. Clutton-Brock, 251–262. Chicago: University of Chicago
Press.
Thorne, E. T., N. Kingston, W. R. Jolley, and R. C. Bergstrom, editors. 1982. Diseases
of wildlife in Wyoming, 2d ed. Wyoming Game and Fish Department, Cheyenne.
446 References
Thrapp, D. L. 1980. Victorio and the Mimbres Apaches. Norman: University of Okla-
homa Press.
Timossi, I. C., and R. H. Ba r rett. 1985. Mi c ro - d i xon user’s manual, version 10-1-85.
De p a rtment Forestry and Resource Management, University of California, Berke-
ley.
Todd, A. W., and L. B. Keith. 1983. Coyote demography during a snowshoe hare decline
in Alberta. Journal of Wildlife Management 47:394–404.
Todd, A. W., L. B. Keith, and C. A. Fischer. 1981. Population ecology of coyotes dur-
ing a fluctuation of snowshoe hares. Journal of Wildlife Management 45:629–640.
Torres, S. G. 1997. Mountain lion alert. Helena, Mont.: Falcon Publishing.
Torres, S. G., T. M. Mansfield, J. E. Folley, T. Lupo, and A. Brinkhaus. 1996. Mountain
lion and human activity in California: Testing speculations. Wildlife Society Bulletin
24:451–460.
Trivers, R. 1985. Social Evolution. Menlo Park, Calif.: Benjamin/Cummings.
Trivers, R. L. 1972. Parental investment and sexual selection. In Sexual selection and the
descent of man, edited by B. Campbell, 136–179. Chicago: Aldine Press.
Trivers, R. L., and D. E. Willard. 1973. Natural selection of parental ability to vary the
sex ratio of offspring. Science 179:90–92.
Turner, J. W., M. L. Wolfe, and J. F. Kirkpatrick. 1992. Seasonal mountain lion preda-
tion on a feral horse population. Canadian Journal of Zoology 70:929–934.
U.S. Census Bureau. 1999. Statistical abstract of the United States. Washington, D.C.
U.S. Fish and Wildlife Service. 2000. Mexican wolf recovery program project update, July
16, 1999–February 20, 2000. Albuquerque, New Mexico.
U.S. Fish and Wildlife Service, Nez Perce Tribe, National Park Service, and USDA
Wildlife Services. 1999. Rocky Mountain wolf recovery 1999 annual report. U.S. Fish
and Wildlife Service, Helena, Montana.
Valdez, R. 2000. Jaguar. In Ecology and management of large mammals in North America,
edited by S. Demarais and P. R. Krausman, 378–388. Upper Saddle River, N.J.: Pren-
tice Hall.
Valdez, R., and P. R. Krausman. 1999. Description, distribution, and abundance of
mountain sheep in North America. In Mountain sheep of North America, edited by R.
Valdez and P. R. Krausman, 3–22. Tucson: University of Arizona Press.
Van Ballenberghe, V. 1983. Rate of increase of white-tailed deer on the George Reserve:
A re-evaluation. Journal of Wildlife Management 47:1245–1247.
Van Devender, T. R. 1995. Desert grassland history: Changing climates, evolution, bio-
geography, and community dynamics. In The desert grassland, edited by M. P.
McClaren and T. R. Van Devender, 68–99. Tucson: University of Arizona Press.
Van Devender, T. R., and L. J. Toolin. 1983. Late Quaternary vegetation of the San
Andres Mountains, Sierra County, New Mexico. In The prehistory of Rhodes Canyon,
edited by P. L. Eidenbach, 33–55. Tularosa, N.M.: Human Systems Research.
Van Dyke, F. G., R. H. Brocke, and H. G. Shaw. 1986. Use of road track counts as
indices of mountain lion presence. Journal of Wildlife Management 50:102–109.
Van Orsdol, K. G., J. P. Hanby, and J. D Bygott. 1985. Ecological correlates of lion
social organization (Panthera leo). Journal of Zoology, London 206:97–112.
Van Sickle, W. D., and F. Lindzey. 1991. Evaluation of a cougar population estimator
based on probability sampling. Journal of Wildlife Management 55:738–743.
———. 1992. Evaluation of road track surveys for cougars (Felis concolor). Great Basin
Naturalist 52:232–236.
References 447
Van Valkenburgh, B., and F. Hertel. 1993. Tough times at La Brea: Tooth breakage in
large carnivores of the late Pleistocene. Science 261:456–459.
Van Valkenburgh, B., F. Grady, and B. Kurtén. 1990. The plio-pleistocene cheetah-like
Miracinonyx inexpectatus of North America. Journal of Ve rt e b rate Paleontology
10:434–454.
Vaughan, T. A. 1997. Mammalogy. Philadelphia: Saunders College.
Waid, D. D. 1990. Movements, food, habits, and helminth parasites of mountain lions
in southwestern Texas. Ph.D. diss., Texas Tech University, Lubbock.
Ward, R. M. P., and C. J. Krebs. 1985. Behavioural responses of lynx to declining snow-
shoe hare abundance. Canadian Journal of Zoology 63:2817–2824.
Waser, P. M., and R. H. Wiley. 1979. Mechanisms and evolution of spacing in animals.
In Handbook of behavioral neurobiology, social behavior and communication, edited by
P. Marker and J. G. Vandenbergh, 159–223. New York: Plenum Press.
Watson, A., and R. Moss. 1970. Dominance, spacing behavior and aggression in relation
to population limitation in vertebrates. In Animal populations in relation to their food
resources, edited by A. Watson, 167–220. Oxford: Blackwell Scientific Publications.
Wayne, R. K., and K. Koepfli. 1996. Demographic and historical effects on genetic vari-
ation in carnivores. In Carnivore behavior, ecology, and evolution, Vol. 2, edited by J. L.
Gittleman, 453–484. Ithaca, N.Y.: Cornell University Press.
Weaver, J. L., P. C. Paquet, and L. F. Ruggiero. 1996. Resilience and conservation of
large carnivores in the Rocky Mountains. Conservation Biology 10:964–976.
Webb, S. D. 1976. Mammalian faunal dynamics of the great American interchange.
Paleobiology 2:220–234.
Weber, W., and A. Rabinowitz. 1996. A global perspective on large carnivore conserva-
tion. Conservation Biology 10:1046–1054.
Weckerly, F. W. 1998. Sexual-size dimorphism: Influence of mass and mating systems in
the most dimorphic mammals. Journal of Mammalogy 79:33–52.
Wehausen, J. D. 1996. Effects of mountain lion predation on bighorn sheep in the Sierra
Nevada and Granite Mountains of California. Wildlife Society Bulletin 24:471–479.
———. 1999. Rapid extinction of mountain sheep populations revisited. Conservation
Biology 13:378–384.
Welch, R. D. 1969. Behavioral patterns of desert bighorn sheep in south-central New
Mexico. Desert Bighorn Council Transactions 13:114–129.
Wemmer, C., and K. Scow. 1977. Communication in the Felidae with emphasis on scent
marking and contact patterns. In How animals communicate, edited by T. A. Sebeok,
749–766. Bloomington: Indiana University Press.
Werdelin, L. 1996. Carnivoran ecomorphology: A phylogenetic perspective. In Carni -
vore behavior, ecology, and evolution, Vol. 2, edited by J. L. Gittleman, 582–624. Ithaca,
White, G. C., and R. A. Garrott. 1990. Analysis of wildlife radio-tracking data. San
Diego: Academic Press.
White, P. A., and D. K. Boyd. 1989. A cougar, Felis concolor, killed and eaten by gray
wolves, Canis lupus, in Glacier National Park, Montana. Canadian Field-Naturalist
103:408–409.
White Sands Missile Range. 1994. Integrated natural re s o u rces management plan
1994–1998. White Sands Missile Range, New Mexico. Environmental Services Division,
White Sands Missile Range, New Mexico.
Wienberg, J., R. Stanyon, W. G. Nash, P. C. M. O’Brien, F. Yang, S. J. O’Brien, and M.
448 References
A. Ferguson-Smith. 1997. Conservation of human vs. feline genome organization

re vealed by re c i p rocal chromosome painting. Cytogenetics and Cell Genetics
77:211–217.
Wildt, D. E., M. Bush, K. L. Goodrowe, C. Packer, A. E. Pusey, J. L. Brown, P. Joslin,
and S. J. O’Brien. 1987. Reproductive and genetic consequences of founding isolated
lion populations. Nature 329:328–331.
Williams, J. S. 1992. Population characteristics and habitat use of mountain lions in the
Sun River area of northern Montana. Master’s thesis, Montana State University.
Wilson, E. O. 1975. Sociobiology: The new synthesis. Cambridge: Belknap Press of Har-
vard University Press.
Wilson, P. J. 1980. Man, the promising primate: The conditions of human evolution. New
Haven, Conn.: Yale University Press.
Woodroffe, R., and J. R. Ginsberg. 1998. Edge effects and the extinction of populations
inside protected areas. Science 280:2126–2128.
Woodroffe, R., and A. Vincent. 1994. Mother’s little helpers: Patterns of male care in
mammals. Trends in Ecology and Evolution 9:294–297.
Worton, B. J. 1987. A review of models of home range for animal movement. Ecological
Modeling 38:277–298.
———. 1989. Kernal methods for estimating the utilization distribution in home range
studies. Ecology 70:164–168.
Wozencraft, W. C. 1993. Order Carnivora. In Mammal species of the world: A taxonomic
and geographic reference, 2d ed., edited by D. E. Wilson and D. M. Reeder, 286–346.
Washington, D.C.: Smithsonian Institution Press.
Wrangham, R. W. 1980. An ecological model of female-bonded primate groups. Behav -
iour 75:262–300.
Wrangham, R. W., and D. I. Rubenstein. 1986. Social evolution in birds and mammals.
In Ecological aspects of social evolution, edited by D. I. Rubenstein and R. W. Wrang-
ham, 452–470. Princeton N.J.: Princeton University Press.
Young, S. P. 1946. History, life habits, economic status, and control, part 1. In The
puma: Mysterious American cat, edited by S. P. Young and E. A. Goldman, 1–173.
Washington, D.C.: The American Wildlife Institute.
Zar, J. H. 1984. Biostatistical analysis. 2d ed. Englewood Cliffs, N.J.: Prentice Hall.
About the Authors
KENNETH LOGAN and LINDA SWEANOR met on a bighorn sheep study in the
North Absaroka Wilderness of Wyoming in 1983. They married in 1985, and
have been studying pumas ever since. Kenneth has a B.S. in Range and Wildlife
Management from Texas A&I University, an M.S. in Zoology and Physiology
from the University of Wyoming, and a Ph.D. in Wildlife Sciences from the
University of Idaho. Linda earned a B.S. in Wildlife Biology and Zoology from
Colorado State University and an M.S. in Wildlife Sciences from the University
of Idaho. As ecologists with the Hornocker Wildlife Institute, they directed field
research on pumas, mule deer, and desert bighorn sheep in the Chihuahua
Desert from 1985 to 1995 and thereafter developed New Mexico’s puma man-
agement plan by working with the public and other wildlife professionals.
Presently, Kenneth and Linda are scientists with the University of California at
Davis, School of Veterinary Medicine, Wildlife Health Center. They are study-
ing interactions of pumas with mule deer, bighorn sheep, and humans in south-
ern California. Their passion is raising their son Oren.
Index
Abrey, N., 293 Andelt, W. F., 390

Accidents as cause of puma deaths, 129, Anderson, Allen E., 7, 8, 18, 126, 128,
137–38 136, 149, 152, 153, 223, 267, 269,
Ackerman, B. B., 89, 138, 173, 189, 196, 274, 372
198, 305, 306, 320, 329 Anderson, E. M., 10, 12, 13, 90, 101, 219
Adams, D. B., 10 Anderson, W. D., 20, 36, 42, 80, 308
Adaptability of puma, 15 Apache, 34
Adaptive kernel method to estimate home Arcese, P., 275
range, 189, 190, 249 Armadillo, 302
Adaptive management, 384–95, 395 Armitrage, K. B., 102
conservation funding, 392–94 Arvas, Thomas, xxiv
education, 391–92 Ashman, David, 7, 8, 48, 70, 90
engaged citizenship, 394–95 Augsburger, J. G., 343
research, 388–91 Aune, K. E., 302, 373, 376
responsive management, 392 Avoidance of other pumas, 267, 282–83,
zone management, 385–88 286, 288, 290
Adult pumas: by females to reduce infanticide, 140,
age structure, 77–80 141, 283
defined, 64
home range, see Home range for adult Badger, 303
pumas Bailey, Ted, xxiv, 102, 140, 198, 219, 272,
mortality and survival of: 274, 275, 293, 294, 297, 337, 338
human-caused, 115–16 Ballard, W. B., 366
natural, 127–39, 143 Barnhurst, D., 42, 269, 375
sex ratios of, 73–75, 292 Barone, M. A., 103, 109, 243
Aerial telemetry, 190 Barrett, R. H., 74, 119, 128, 136, 240,
African Leopard, The (Bailey), xxiv 275, 302, 370, 372
Age of pumas: Bartmann, R. M., 330
dental characteristics to determine, Beasom, S. L., 52
47–49 Beckett, P. H., 33, 34
eye color and, 50 Beier, P., 74, 119, 128, 136, 147, 149, 151,
Age structure of puma population, 75–81 178, 239, 240, 244, 275, 297, 302,
Akinson, Holly, 287 370, 372, 378, 384
Alberta, Canada, puma study in, see Belden, R. C., 8, 168, 197, 369, 390
Jalkotzy, Martin; Ross, Ian Berger, J., 343
Alkon, P. U., 55 Bergerud, A. T., 335
451
452 Index
Bertram, C. R., 337 response to prey crash, 338

Biological diversity, 365 social organization of, 183
Birth-interval home range, 192–95 transient behavior in, 244
Birth intervals of pumas, 87, 92–93, 110 Chemical communication, 273–76, 277,
Black bears, 14, 32–33 283, 284
Bobcats, 202, 219, 304, 337, 338 Chepko-Sade, B. D., 153, 178, 378
Boellstorff, D. E., 105 Chihuahua Desert, 14, 29
Boertje, R. D., 337 Clapperton, B. K., 275
Bolton, H. E., 370 Clark, P. J., 255, 256
Bonney, R. C., 89 Clark, R., 344
Boutin, S., 313 Clevenger, A. P., 382
Boyce, W. M., 84, 136, 344, 345 Climate of research area, 27–29, 168
Boyd, D., 304 large-mammal population dynamics and,
Boykin, K. G., 27 359–62, 379
Brahmachary, R. L., 275 Clouded leopards, 297
Branch, L. C., 302 Clutton-Brock, T. H., 72, 104, 110, 184
Brand, C. J., 337 Coccidiodmycosis, 245
Brashares, J. S., 275 Coe Clough, N., 118
Breeding, see Reproduction of pumas Colorado puma study, see Anderson, Allen
Breternitz, C. D., 33, 34, 35 E.
British Columbia puma study, see Communication among pumas, 269–76,
Spreadbury, B. R. 277
Bronson, F. H., 89, 90, 95, 96, 109, 330 chemical, 273–76, 277, 283, 284
Brown, D. E., 16, 27 vocalizations, 269–72, 277
Brown, J. L., 285 Conservation and management of wild
Buechner, H. K., 343 pumas, 365–98
Bunch, T. D., 342 alleviating threats, 377–95
Bureau of Land Management (BLM), 23 adaptive management, 384–95
Burney, D. A., 12, 13 habitat conservation, 378–82
Burt, W. H., 189, 285 preventing unnecessary overklil,
Buskirk, S. W., 198, 345 383–84
funding for conservation, 392–94
Caddick, G. B., 86, 103, 120 threats to pumas:
CALHOME software, 189, 413 habitat loss, 367–71
California Wildlife Protection Act, 394 puma overkill, 371–77
Calow, P., 71 Conservation Biology, 145
Canadian lynx, 202, 219 Contagious ecthyma, 344, 345
Cannibalism, 120, 222, 305 Converse, L. J., 275
Caro, T. M., 183, 244, 245, 275, 294, 296, Coulson, J. C., 97
338 Cox, J. A., 8
Carroll, C. R., 176, 361, 378 Coyotes, 32, 121, 302, 303, 304, 333, 337
Catching pumas, 42–51 Crawshaw, P. G., 198, 295
Caterwaul, 271–72 Crow, J. F., 178
Caughley, G., 85, 169, 311, 313, 330, 333, Cryptorchidism, 108–109, 134, 243
386, 389 Cubs, puma:
Cheek rubbing as form of chemical age structure of the population, 75–77
communication, 275 birth weights, 20
Cheetahs, 9, 293 capturing nursling, 47–48, 49
chemical communication among, 275 color of coats, 20
coalitions formed by, 295–97 defined, 64
Index 453
infanticide, see Infanticide home range size of adult pumas and,

mass, gender and, 71–72 204–10
mortality and survival of: female response to changes in puma
human-caused, 115–16 density, 206–208
of matrilineal mothers, 101, male response to changes in puma
125–26 density, 208–10
natural, 117–22 self-limiting hypothesis and,
mother’s defense of, 141 210–11
orphaned, 121–22, 124, 371, 375 prey density and, 168
parental investment, 96, 111, 268–69 Dental characteristics, 21, 48–50
sex structure of, 69–72 DeRose, M. A., 243
vocalizations, 270–71 Description of puma, 18–21
Culver, M., 10, 11, 17, 179, 242, 243 body size, 18–19, 21
Cunningham, S. C., 129, 136, 176, 302, claws and feet, 19
372, 384 color, 19–20
sex organs, 21
Daniel, W. W., 59 teeth, 21
Definitions: Desert bighorn sheep, 14, 15, 30, 35, 302
adults, 64 diseases affecting, 344–45, 347, 349,
cubs, 64 350, 355, 357
deterministic, 321 as endangered species, 31
discrete time, 321 extinction on San Andres Mountains,
dispersers, 64, 145 354–55, 357
duration of transient behavior, 231 history of New Mexico, prior to San
emigrants, 65, 145 Andres Mountains puma study,
fidelity to home range, 211 342–46, 357
functional response, 313 monitoring radio-collared, 51, 53–54,
home range, 64, 185, 189 57
immigrants, 65, 145 fates of, 347–50
independent home range, 231 mortality of, 58, 347–50, 357
limiting factor, 311 survival rates and agent-specific
natal home range, 231 mortality, 350–54
numerical response, 313 pumas and, 302, 305, 307, 341–62
philopatric, 65, 145, 231 hypothesis and predictions, 341
recruits, 65, 145 other populations outside New
residents, 64 Mexico, 356–57
site attachment, 231 predation, 341, 347, 349, 352–54,
spatiotemporal relationships, 257 355, 357, 359–62, 378
subadult home range, 231 puma control and survival of,
subadults, 64 36–37, 341
total response, 313 statistics, 357–58
transient home range, 231, 244 summary, 357
DelGuidice, G. D., 319, 349 surveys of population characteristics,
Density of puma population, 160–69, 346–47
179–80 weather and population dynamics of,
comparison of San Andres Mountain 359–62
pumas with other populations, Desert mule deer, 14, 15, 30, 31, 35, 88,
166–68 302, 307, 311–40, 378
habitat and, 168–69 characteristics of dead, 313–14
definitions, 311–13
454 Index
Desert mule deer (continued) desert bighorn sheep and, 344–45, 347,
deterministic, discrete time model that 349, 350, 355, 357
simulated population dynamics of, on puma mortality and, 120–21, 129,
Treatment Area, 419–21 136–37
hunting by humans, 31 Dispersers, puma, 148–53, 154–55, 375
mortality and survival rates, 58, 314–21, defined, 64, 145
339 population density and, 152–53
pumas and, 311–40, 355 subadult:
behavioral interactions, 331–32 females, 233–36, 240–43, 245–46
limit on the deer population, frustrated, 239–40
332–33 hypotheses to explain, 240–43, 246
mule deer population dynamics, males, 236–39, 240–43, 246, 283
321–31, 339, 359–62 timing of dispersal, 148–49
partially compensatory nature of Dobson, F. S., 153, 240
puma predation of, 318–19 Dogs used to catch puma, 42
predation and mule deer Doyel, D. E., 33, 34, 35
population growth, 322–31, 339 Drugs, immobilizing, 47, 48
as primary source of food, 176, Dunn, W. C., 342
302, 303, 305, 306, 308–309 Duration of transient behavior defined, 231
research hypotheses and Dwarf mongoose, 275
predictions, 311–13, 319–20
self-limiting hypothesis and, Ear tags, 50
333–35, 339 Eastern Cougar Foundation, 368
statistics, 339–40, 419–21 Eaton, R. L., 72
summary, 338–39 Education as conservation strategy, 391–92
radio-collared, 49, 51–53, 57 Ehret, G., 270
fates of, 314–21, 338 Eidenbach, P. L., 26, 34, 35, 36
weather and population dynamics of, Eisenberg, J. F., 14
359–62 Elam, D. R., 178
Diet of pumas, 301–309 Elenowitz, A., 52, 343, 420
patterns of pumas and prey, 301–302 Elk, 302, 335, 378
prey size and puma body size, 301–302, Elliot, J. P., 335
308 Elmer, Mike, 58, 305
on the San Andres Mountains, 302–308 Emigration, 153, 155, 176
scavenging, 307–308, 309 defined, 65, 145
summary, 308–309 Emlen, J. T., Jr., 184, 285, 286
see also Prey of pumas Endangered species, 366
Direct interactions between pumas, 259–69 Endangered Species Act of 1973, 15, 392
for breeding activity, 259–60, 265, 267 Engaged citizenship, puma conservation
between family members, 268–69 and, 394–95
independent pumas, associations Erlinge, S., 184
between: Ermines, 293
female-female, 264–65 Erwin, D. H., 14
male-female, 265–67 Etkin, W., 285–86
male-male, 260–64 Evans, Dr. Wain, xxiii, xxiv, 36
for intraspecies strife, 260–63, 265, 267 Evans, F. C., 255, 256
statistics, 278–79 Evolutionary ecology, 6
summary, 277 Ewer, R. F., 17, 124
Disease: Eye color, age and, 50
Index 455
Fecundity rates of pumas, 85–86, 110 Gilpin, M. E., 178, 378

Fidelity, adult puma home range, see Home Ginsberg, J. R., 212
range for adult pumas, fidelity Gipson, P. S., 366
Fidelity Index, 212, 213–14, 215 Gittleman, J. L., 198
Findley, J. S., 16 Gloyne, C., 382
Fisher, Amy, 49 Golden eagle, 303
Fitness of adult female pumas, factors Goldman, E. A., 17
associated with, 140–41 Gonyea, W. J., 19
Fitness of adult male pumas, factors Goodrich, J. M., 198
associated with, 141–42 Gorman, M. L., 273, 275, 276
Fitzhugh, E. L., 384 Grafen, A., 97
Fitzsimmons, N. N., 345, 377 Gray fox, 32, 304
Floater, see Transient home range Gray wolves, 14, 382
Florida panthers, 7, 295, 369, 372 Gregg, C. T., 136
cryptorchidism among, 109 Grigione, M. M., 384
direct interaction between, 267 Grizzly bears, 14, 16, 33, 384
dispersal by, 149 Guanaco, 302
as endangered species, 15, 367, 392–93 Gutiérrez-Espeleta, G. A., 345
habitat loss, 369, 372
habitat preservation, 382 Habitat, puma, 336, 389
home range shifts, 219, 224 conservation of, 378–82
inbreeding in, 103, 243 home range size and, 197–98
intraspecies strife among, 136, 369 loss of, as threat, 367–71
litter size, 86 population density and, 168–69
longevity of, 80 Hackney, R., 366
male philopatry, 239 Hagedorn, B. W., 369
mortality/survival rates: Hailey, M. P., 110
of cubs, 119–20 Hair, J. F., 327
reproduction of, 90–91, 95 Halfpenny, J. C., 376
transient behavior in, 244 Halpin, Z. T., 153
Floyd, T. J., 306 Hand capture of cubs, 48, 51
Foster, M. L., 382 Hares, 302, 306, 333, 338
Frank, D. H., 198 Harlow, H. J., 375
Franklin, W. L., 302 Harris, A. H., 13
Freeman, S., 6, 10, 84, 96, 97 Harvey, P. H., 14, 198
Fuller, T. K., 127, 128, 168, 315, 316, 317, Hast, M. H., 269
320, 337 Hawley, J. W., 26
Fulwyler, Major General Niles, xxiv Hayne, D. W., 189
Functional response, defined, 313 Hedrick, P. W., 103, 243
Funding for puma conservation, 392–94 Heisey, D. M., 127, 128, 315, 316, 317,
320
Garrott, R. A., 189, 213 Hembrillo Base Camp, 40
Gasaway, W. C., 335 Hemker, T. P., 98, 119, 149, 168, 195,
Gashwiler, J. S., 138 201, 204, 273, 287, 329
Gaut, J. A., 343 Hemmer, H., 17
Geist, V., 14, 53 Hemming, J. E., 53
Geographic distribution and status of Herron, J. C., 6, 10, 84, 96, 97
puma, 15–16 Hertel, F., 12
Gestation period, puma, 91, 110 Hervert, J. J., 330
456 Index
Heske, E. J., 198 definitions, 231

Hiraiwa-Hasegawa, M., 72 dispersal:
Hoban, P. A., 35, 345 by females, 233–36, 240–43,
Hoffmeister, D. F., 16 245–46
Holderman, Dave, 336 frustrated, 239–40
Holling, C. S., 313 hypotheses to explain, 240–43, 246
Holocene epoch, 12, 29 by males, 236–39, 240–43, 246
Home range: philopatry in females, 232–33, 235–36,
for adult pumas, see Home range for 245
adult pumas statistics, 246
defined, 64, 185, 189 summary, 245–46
methods and estimates of annual size of, transient behavior, 244–245, 246, 283
on San Andres Mountains, 413–16 Hopkins, R. A., 74, 80, 201, 219, 221–22,
scent markings and, 275 224, 274, 275
for subadults, see Home range for Hornocker, Maurice, xxiii–xxv, 6–7, 21, 42,
subadult pumas 63, 86, 136, 138, 152, 175, 183,
Home range for adult pumas, 189–229, 245, 286, 294, 304, 320, 329, 336,
282, 284 372
annual, 191–92 self-limiting hypothesis and, 184–85,
birth-interval, 192–95 284
comparison of San Andres Mountain Hornocker Wildlife Institute, xxiii–xxiv
pumas to other populations, 201 Horses, feral, 335
delineating the, 189–91 Humphrey, S. R., 382
energy demands and, 195–98 Humphreys, D., 52, 420
factors influencing size of, 195–201 Hunt, C., 390
energy demands, 195–98 Hunt, R. M., 9
gender, 198–201, 282 Hunter, M. D., 359, 360
prey abundance, and puma density, Hunting and predator control, 115–16,
201–11 164–65, 176–77
fidelity, 211–25, 283–84 government sanctioned, 16, 36–37, 376
benefits of fidelity and the two- management of puma hunting, problems
strategies hypothesis, 223–25 with, 373–76
defined, 211 natural selection process, effects on, 377,
distances between mean locations 379
method, 213, 214–16 overkill as threat to pumas, 371–77
Fidelity Index, 212, 213–14, 215 preventing unnecessary overkill, 383–84
home range shifts in pumas, prohibition of, 16, 37, 176, 367
216–22 tenure of territorial males affected by,
homing by translocated pumas, 142
222–23
methods of determining, 212–16 Iason, G. R., 72
self-limiting hypothesis and, 225 Idaho puma study, see Seidensticker, J. C.
lifetime, 195 Immigrants, puma, 153, 154, 155, 159
old males, 245 defined, 65, 145
range size, prey abundance, and puma Inbreeding, 102–103
density, 201–11 avoidance of, 240, 243, 246, 283
seasonal, 191–92 Independence of puma progeny, 146–48,
statistics, 227–29 154
summary, 225–27 timing of, resumption of breeding
Home range for subadult pumas, 231–46 behavior in mothers and, 147–48
Index 457
Independent home range defined, 231 Koepfli, K., 177

Infanticide, 120, 140, 142, 266–67, 371 Kohn, M. H., 384
avoidance of other pumas by mothers to Kottlowski, F. E., 26, 27
reduce threat of, 140, 141 Kovacs, K. M., 105
Interactions between pumas: Krackow, S., 72
communication, see Communication Krausman P. R., 330, 342, 343, 352
among pumas Krebs, C. J., 65, 202, 338
direct, see Direct interactions between Kruuk, H., 273
pumas Kruuk, L. E. B., 72
spacial relationships, see Spatial Kunkel, K., 335
relationships Kurtén, B., 10, 12, 13, 18, 301
statistics, 277–79
summary, 276–77 Lack, D., 84, 378
Intraspecies strife, 287–88, 375–76 Lacy, R. C., 178, 346
as cause of adult puma mortality, Laing, S. P., 101, 159, 168, 197, 222, 336,
129–36, 139–42, 143, 239 378
direct interactions for, 260–63, 265, 267 Lair, H., 191
Iriarte, J. A., 21, 301, 302 Lance, W. R., 344
Irwin, L. L., 168, 197, 273, 335, 336, 378 Lande, R., 378
Land tenure system, 185, 186, 285, 286, 287
Jackrabbits, 303, 306 Lang, E. M., 343
Jackson, P., 16, 17 Lange, R. E., 344
Jaguars, 14, 16, 32, 102, 365, 370, 382 Larsen, D. G., 335
comparison of social structures of pumas Laumbach, K., 34
and, 294, 295, 297, 298 Le Boeuf, B. J., 104, 110
home range size for, 198, 295 LeCount, 329
scent markings, 274, 295 Lenti Boero, D., 275
size of, 18 Leopards, 102, 338
Jalkotzy, Martin, 7, 63, 70, 77, 86, 90, 95, chemical communications among, 274,
98, 101, 119, 136, 138, 142, 147, 275
149, 153, 166, 167, 168, 174, 185, comparison of social structure of pumas
195, 204, 224, 242, 286, 308, 329, and, 294, 297, 298
372 home ranges, 198, 219, 294
James, W. H., 72 prey biomass and population densities,
Javelina, 30, 305, 307 168, 337
Jennrich, R. I., 189 vocalizations, 272
Jessup, D., 344 Leyhausen, P., 17, 183, 270, 275, 294
Jiménez, J. A., 243 Lidicker, W. Z., 153
Johnson, W. E., 9, 10 Limiting factors:
Jornada Experimental Range, 28, 35 defined, 311
Julander, O., 329 puma predation as primary proximate,
for desert mule deer population,
Kaufmann, J. H., 286 311–13, 314
Keith, L. B., 337 of San Andres Mountains puma
Keystone species, pumas as, 365 population, 336–37
Kie, J. G., 189, 190 Lindstedt, S. L., 198
Kimura, M., 178 Lindzey, Fred, 7, 8, 63, 74, 86, 90, 95,
Knick, S. T., 337, 338 101, 128, 138, 159, 167, 168, 174,
Koch, James, xxiv 175, 186, 197, 222, 269, 320, 336,
Koehler, G. M., 304 372, 375, 378, 383
458 Index
Linneaus, 16 survival rate of cubs with matrilineal

Lions, African, 102, 109, 337 mothers, 101, 125–26
chemical communication among, 275 Matson, P. A., 359, 360
coalitions formed by, 295–97 Mayr, E., 6
home range size and prey abundance, Mech, L. D., 319, 337, 349
202 Meffe, G. K., 176, 361, 378
inbreeding, 243 Mehrer, C. F., 124
social organization of, 183 Menge, B. A., 360
transient behavior in, 244 Messier, F., 168, 313, 333
Litter size, 83–84, 86–87, 110 Metapopulation dynamics, 175–79,
Litvaitis, J. A., 202 180
Logan, Kenneth A., xxiv, xxvii–xxxi, 15, Micromort survival analysis software, 127,
42, 43, 45, 47, 65, 70, 74, 86, 90, 128–29, 162, 316, 352
98, 106, 149, 152, 167, 168, 197, Minimum convex polygon (MCP) method
245, 273, 302, 308, 335, 336, to estimate home range, 189–90,
367, 371, 372, 373, 375, 378, 385, 231, 249
392 Minnis, D. L., 392
Long-eared owl, 305 Minta, S., 198
Lopez-Gonzalez, C. A., 16, 70, 101, 119 Mitani, J. C., 110
Lorenz, K., 292 Montoya, Gary, 354–55
Lott, D. F., 198, 292 Moore, C. T., 72
Lynxes, American, 293, 333, 337, 384 Moose, 302, 333, 378
response to prey crash, 338 Morgan, B., 36
Morphological measurements of puma 3.5
McBride, R. T., 136 months or older on San Andres
McCowan, J. W., 390 Mountains, 399–412
McCullough, D. R., 385 Morris, D. W., 240
McDaniel, G. W., 384 Mortality of desert bighorn sheep, see
Macdonald, D. W., 184, 201 Desert bighorn sheep, mortality of
McDougal, C. W., 87, 109, 142 Mortality of desert mule deer, see Desert
Mace, G. M., 285 mule deer, mortality and survival
McLaren, B. E., 333, 335, 360 rates
Maehr, D., 8, 80, 91, 95, 120, 149, 219, Mortality/survival of puma, 115–44
222, 224, 244, 267, 269, 297, 367, human-caused, 115–16, 142
372, 381 investigation methods, 58–59
Maehr, D. S., 72, 74, 86, 103, 136 male/female, 120, 122, 126, 127–28,
Marshall, L. G., 11, 313 143
Martin, L. D., 14 natural, 117–45
Mass, puma: of adults, 127–39, 143
of adult male, 109–10, 142 of cubs, 117–22, 142–43
home range size and, 198 of subadults, 122–27, 143
of cubs, gender and, 71–72 reasons pumas kill other pumas, 139–42,
morphological measurements of puma 143
3.5 months or older on San Andres statistics, 143–44
Mountains, 399–412 summary, 142–43
prey size and puma body size, 301–302, Mowat, G., 337, 384
308 Mule deer, desert, see Desert mule deer
Mating, see Reproduction of pumas Müller-Schwarze, D., 273
Matrilines, puma, 99–101, 233, 282, 290, Muñoz, D., 36
297 Murphy, Ke r ry, 7, 97, 104, 105, 107, 109,
longevity of, 102 293, 301, 321, 336, 372, 379, 381
Index 459
Museum of Southwestern Biology, 37 Padley, W. D., 265, 272, 393

Palanza, P. S., 275
Natal home range, defined, 231 Parental investment in offspring, 96,
National Cancer Institute,Laboratory of 111
Genetic Diversity, 50 Parker, G. R., 337
Neal, D. L., 166, 201, 274 Paw petroglyph, 33
Nearest-neighbor analysis, 254–56 Peccary, 14, 302, 378
Neilson, R. P., 342, 343, 361 Pech, R. P., 313
Nevada Department of Wildlife, 7 Pecon-Slattery, J., 9, 10
Nevada puma study, see Ashman, David Pederson, J., 31
New Mexico Department of Agriculture, Pereira, M. E., 72
Veterinary Diagnostic Service, 58, Perrins, C. M., 84
136 Peters, G., 269–70, 271, 272
New Mexico Department of Game and Peterson, R. O., 333, 335, 360
Fish, xxiv, 16, 31, 32, 36, 37, 49, Peterson Method, 65
52, 53, 232, 322, 344, 345, 355, Philopatric lions, 297
376, 420 Philopatric pumas, 148–53, 154
Noble, G. K., 285 defined, 65, 145, 231
Nomadic behavior, see Transient home subadult females, 232–33, 235–36
range Phylogeny of pumas, 9–15
Norrix, L. W., 344 Physiography and geology of study area,
Northern Yellowstone Ecosystem, study of 26–27
pumas in, see Murphy, Kerry Picton, H. D., 330
Noss, R. F., 178 Pierce, B. M., 88, 168, 186, 211, 223, 290,
Nottingham, B. G., 274, 294, 295 329, 333, 334, 335
Nowak, R. M., 15, 16, 17 Pitelka, F. A., 285
Nowell, K., 16, 17 Pleistocene, 12–14
Numerical response, defined, 313 mass extinction of mammals during,
Nuñez, R. B., 295 11–12, 13–14, 301
Nunn, C. L., 72 Pollock, K. H., 126, 128
Nunney, L., 178 Poole, K. G., 337
Population of pumas, study area:
O’Brien, S. J., 8, 9, 10, 103, 243 age structure, 75–81
Ockenfels, R. A., 335 census of, methods for, 65–68, 80
O’Donoghue, M., 333, 337, 338 density of, see Density of puma
Odors, chemical communication through, population
see Chemical communication experimentally removing pumas,
O’Gara, B., 304 157–60, 179, 222–23, 390
Ordway, L. L., 330 growth rates, 169–75, 180
Organ Mountains, 26, 27, 37, 178, limiting factor for, 336–37
370–71, 378, 382 metapopulation dynamics, 175–79, 180
desert bighorn sheep in, 342, 343 observed, 51
Orians, G. H., 285 self-limiting hypothesis, see Self-limiting
Ornate box turtle, 305 hypothesis
Orphaned cubs, 121–22, 124, 371, 375 sex structure, 69–75, 80–81, 292
Oryx, 31–32, 303, 305, 307 structure of, 69–81, 159
Oscura Mountains, 36–37, 342 Porcupine, 303
Owen-Smith, N., 13, 285 Post, E., 360
Powell, R. A., 190
Packer, C., 21, 104, 109, 140, 243, 266, Power, M. E., 360, 361
294, 296, 297, 337 Precipitation in study area, 26–27
460 Index
Prey biomass (prey density), 168 birth intervals, 87, 92–93, 110
dispersal and, 241–43 chemical communication and, 275–76,
home range size and, 202–204 284
Prey of pumas: direct interactions for breeding, 259–60,
animals killed but not consumed, 265, 267
303–304 dispersal and competition for mates,
cases of puma and other carnivores 240–41
limiting prey populations, 335–36 fecundity rates, 85–86, 110
densities of, see Prey biomass (prey fidelity, 105
density) gestation, 91, 110
diet, 301–309 litter size, 83–84, 86–87, 110
patterns of pumas and prey, mating, 91–92
301–302 natality, 83–87, 110
prey size and puma body size, parental investment, 96, 111, 268–69
301–302, 308 as polyestrous, 89, 90
on the San Andres Mountains, polygynous, 74, 105
302–308 promiscuity, 74, 105, 140–41
scavenging, 307–308, 309 puberty and first litters, 93–96, 110–11
summary, 308–309 reproductive chronology of mated pairs
method of attack, 21 on San Andres Mountains, 413–14
response to prey crash, 337–38 reproductive success, 96–110, 111
see also Desert mule deer; specific prey, females, 98–103, 111
e.g. Desert bighorn sheep males, 103–10, 111
Price, P. W., 360 scent marking and, 275
Prides, forming of, 296–97 statistics, 111–13
Pronghorns, 30, 302–303, 305 summary, 110–11
Protection of puma, legal, 15, 16 timing of births, 88–91, 110
Puberty of pumas, first litters and, 93–96, two-strategies hypothesis, see Two-
110–11 strategies hypothesis
Puma concolor, 10 Reproductive success of pumas, 96–110,
Pusey, A. E., 140, 243, 266, 111
294, 296 females, 98–103, 111
males, 103–10, 111
Quigley, H. B., 295 Research area, see San Andres Mountains,
New Mexico, as puma study area
Rabinowitz, A. R., 274, 294, 295 Research hypotheses and predictions,
Raccoon, 302 63–64
Radiotelemetry, 45, 50 Research on pumas, future needs for,
study aided by, 51, 54–57 388–91
Rail, C. D., 136, 137 Research team, 23
Randall, J. A., 275 Residents defined, 64
Ranging behavior, see Home range Responsive management, 392
Rasas, O. A. E., 275 Ricklefs, R. E., 69, 160
Rattlesnake bites, 138 Ringtail, 303
Read, A. F., 14 Roamer, see Transient home range
Recruits, puma, 154, 155 Roberson, J., 374, 375
after experimentally removing pumas Robinette, W. L., 53, 70, 72, 138
from Treatment Area, 159 Rodents, 302, 378
defined, 65, 145 Roelke, M. E., 8, 103, 243
Reiter, J., 104, 110 Roff, D. A., 243
Reproduction of pumas, 83–113, 281 Ross, Ian, 7, 15, 63, 70, 77, 80, 86, 90, 95,
Index 461
98, 101, 119, 136, 138, 142, 147, Schaller, G. B., 183, 198, 244, 245, 270,
149, 153, 166, 167, 168, 174, 185, 275, 294, 296, 337
195, 204, 224, 242, 286, 308, 329, Schenk, A., 105
357, 372 Schmidt, J. L., 219
Roth, J. A., 118 Schoener, T. W., 191
Rubenstein, D. I., 281 Scow, K., 270
Russell, D. E., 10 Scrapes as form of chemical
Ruth, T. K., 47, 98, 106, 107, 139, 141, communication, 273–76, 284
157, 338, 381, 390 Seaman, D. E., 190
Seidensticker, J. C., 7, 63, 64, 70, 74, 104,
Saccheri, I., 178, 243, 378 146–47, 152, 159, 167, 168, 175,
Sadleir, R. M., 330 183, 186, 201, 204, 219, 245, 267,
Saiz, R. B., 31–32 273, 274, 275, 289, 320, 336
Sale, M., 34 home range fidelity and, 223
Salman, David, xxiv self-limiting hypothesis and, 184–85,
Saltz, D., 55 210, 211, 284–85, 286, 287, 313,
San Andrecito Camp, 39–40, 41 333
San Andres Mountain National Wildlife Seip, D. R., 335
Refuge, 35 Self-limiting hypothesis, 183, 184–87,
San Andres Mountains, New Mexico, as 284–88, 333–35, 376
puma study area, 23–26 land tenure and, 285
catching wild pumas, 42–51 San Andres study and, 186–87, 210–11,
climate, 27–29, 168, 359–62, 379 225, 286–88, 293, 298, 313, 333,
counting pumas, 65–68 339
fauna of, 30–33 territoriality and, 285–86
flora of, 29–30 Septicemic plague (Yersinia pestis), 136–37
human use, history of, 33–36 Sex organs of pumas, 21
life afield, 39–42 Sex structure:
map, 24 of desert bighorn sheep, 347
physiography and geology of, 26–27 of puma population, 69–75, 80–81, 292
puma exploitation, history of, 36–37 Sexual selection, 109
reasons for choosing, 23–26 Shaffer, M., 178
Reference Area (RA), 26 Shaw, H. G., 42, 74, 338, 372
research hypotheses and predictions, Shields, W. M., 177, 243
63–64 Short, H. L., 330
terms for pumas, 64–65 Sibly, R. M., 71
Treatment Area (TA), 26 Sierra Oscura Mountains, 26
experimentally removing pumas Sinclair, A. R. E., 240, 311, 313, 330, 333,
from, 157–60, 179, 222–23, 390 386, 389
see also specific aspects of puma behavior Sinclair, D. F., 88, 255
San Andres Wildlife Refuge, 343, Singer, F. J., 330
354–55 Site attachment defined, 231
Sandell, M., 183, 184, 198, 201, 202, 203, Sitton, L. W., 274
244, 292–93, 293 Slade, N. A., 191
Sandoval, A. V., 35, 36, 342, 343, 344, Smallwood, K. S., 384
345, 347 Smith, C. R., 31–32
SAS Institute, Inc., 59 Smith, Frank, xxiv, 42, 57, 149
Saunders, G. C., 118 Smith, J. L. D., 87, 102, 109, 142, 198,
Savage, D. E., 10 222, 224, 274, 275, 276, 295
Scabies, 344, 347, 349, 350, 355, 357 Smith, R. H., 329
Scent markers, 273–76, 284, 295 Smith, R. L., 105
462 Index
Smith, T. E., 274, 372 overview of analytical and statistical

Snares, foothold, 43–47, 50–51, 115 methods, 59
vocalization of snared pumas, 270 sex structure, 81
Snow leopards, 102, 297 Stearns, S. C., 6, 71, 96
Snowshoe hares, 333, 338 Stenseth, N. C., 153
Social organization of pumas, 7 Stephenson, R. O., 337
adaptive significance of, 281–98 Stoddart, D. M., 273
communication, see Communication Striped skunk, 302, 303
among pumas Strobeck, C., 384
comparison with other big cats, 294–97, Strong, D. R., 360
298 Stuart, A. J., 12, 13
direct interactions between pumas, see Studies of pumas, synopsis of previous, 6–8
Direct interactions between pumas Subadults, puma:
self-limiting hypothesis and, see Self- age structure of the population,
limiting hypothesis 75–77
social structure of desert pumas, defined, 64
281–84 home range of, see Home range for
female structure, 282–83 subadult pumas
male structure, 283–84 mortality and survival of:
spatial relationships, see Spatial human-caused, 115–16
relationships natural, 122–27, 143
two-strategies hypothesis and, see Two- sex ratio of, 73
strategies hypothesis Sunquist, M. E., 198, 222, 224, 294, 295
Source-sink metapopulation structure, Survival of puma, see Mortality/survival of
177–78, 180 puma
Spatial relationships of pumas, 247–56, Sweanor, Linda L., xxvii–xxxi, 15, 44, 56,
284, 286, 292 145, 152, 153, 175, 178, 191, 192,
home range overlap indices, 249–54 212, 219, 240, 271, 283, 302, 367,
nearest-neighbor analysis, 254–56 371, 373, 378, 379, 380, 385, 392
spatiotemporal relationships, 257–59, Sweitzer, R. A., 335
275 Swihart, R. K., 191
statistics, 277–78
summary, 276–77 Taber, 302
Spreadbury, B. R., 70, 86, 90, 136, 167, Tagging, 50
185, 204, 222, 234, 245, 274, 372 Tattooing, 49
SSPS, Inc., 59 Taxonomy, puma, 16–17
Stander, P. E., 168, 198, 203, 294, 337 Taylor, D., 36
Starvation, 120–21 Taylor, M. E., 293
Statistics, study, 395 Teel, T. L., 391
age structure, 81 Teer, J. G., 330, 335
body mass, 21 Teeth of pumas, 21, 48–50
desert bighorn sheep, 357–58 Temperature in study area, 28–29
desert mule deer, 339–40, 419–21 Templeton, A. R., 177
home range for adult pumas, 227–29, Terborgh, J., 360
413–16 Territoriality, 285–86, 287, 293
home range for subadult pumas, 246 Thomas, C. S., 97
interactions between pumas, 277–79 Thorne, E. T., 136, 137
morphological measurements, 399–412 Thrapp, D. L., 34
mortality/survival, 143–44 Tigers, 109, 142
Index 463
chemical communication among, 274, cubs, 270–71

275 yowl and raspy ouch, 272
comparison of social structure of pumas
and, 294, 297, 298 Waid, D. D., 200
home range, 295 Wallen, S., 274
fidelity, 224, 295 Waltho, N., 382
size, 198 Ward, R. M. B., 202, 338
scent markings, 274, 295 Waser, P. M., 292
Todd, A. W., 337 Wayne, R. K., 177
Toolin, L. J., 362 Weather, see Climate of research area
Torres, S. G., 302, 376, 384 Webb, S. D., 10
Total response defined, 313 Weckerly, F. W., 109
Transient home range, 244–45, 246, 282, Wehausen, J. D., 343, 356
283 Weisenberger, Mara, 336, 354–55
defined, 231, 244 Welch, R. D., 343
Translocation of pumas, experimental, Wemmer, C., 270
157–60, 179, 390 Werdelin, L., 9
homing by translocated pumas, White, G. C., 189, 213
222–23 White Sands Missile Range, xxiv, 23, 24,
Trapping, 42–43, 116 31, 32, 35, 37, 344, 355
Trivers, R., 6, 70, 72, 96, 109, 183, 198 White Sands National Monument, 35
Trowbridge, B. J., 276 White-tailed deer, 14, 302, 335, 378
Turner, F. B., 189, 329 Wildt, D. E., 243
Turner, J. W., 335 Wiley, R. H., 292
Two-strategies hypothesis, 183–84, Willard, D. E., 72
288–93, 297–98 Williams, J. S., 136
female strategy, 288–90 Wilson, E. O., 285
home range fidelity and, 223–25 Windy season in study area, 29
male strategy, 290–93 Wolf, M., 243
Wolff, R., 275
U.S. Census Bureau, 368 Wolves, 16, 32, 33, 333, 335, 366,
U.S. Department of Interior, 366 382
U.S. Fish and Wildlife Service, 36, 52, 53, prey biomass and population densities
54, 344, 355, 366, 370, of, 168, 337
392 Wood, J. P., Jr., 33, 36
U.S. Forest Service, 35 Woodroffe, R., 96
Utah puma study, see Lindzey, Fred Worton, B. J., 189, 190
Wozencraft, W. C., 16, 17, 270, 271
Valdez, R, 343, 366 Wrangham, R. W., 183, 184, 281,
Van Ballenberghe, V., 169 295
Van Devender, T. R., 342, 343, 362 Wyoming puma study, see Logan, Kenneth
Van Orsdol, K. G., 202 A.
Van Sickle, W. D., 383 Yersinia pestis, 136–37
Van Valkenburgh, B., 10, 11, 12 Young, S. P., 15, 16, 20, 179
Vaughan, T. A., 9 Young, T. P., 212
Vincent, A., 96
Vocalizations, puma, 269–72, 277, 283 Zar, J. H., 59
caterwaul, 271–72 Zone management, 385–88
Island Press Board of Directors
Chair, HENRY REATH CAROLYN PEACHEY

President, Collector’s Reprints, Inc. Campbell Peachey & Associates
Vice-Chair, VICTOR M. SHER WILL ROGERS

Environmental Lawyer Trust for Public Lands
Secretary, DANE NICHOLS CHARLES C. SAVITT

Chairman, The Natural Step, U.S. President, Center for Resource
Economics/Island Press
Treasurer, DRUMMOND PIKE
President, The Tides Foundation SUSAN E. SECHLER
Director of Global Programs,
WILLIAM M. BACKER The Rockefeller Foundation
Backer, Spielvogel, Bates (ret.)
PETER R. STEIN
ROBERT E. BAENSCH Managing Partner,
Professor of Publishing, The Lyme Timber Company
New York University
RICHARD TRUDELL
MABEL H. CABOT Executive Director,
American Indian Resources Institute
DAVID C. COLE
Sunnyside Farms, LLC WREN WIRTH
President,
CATHERINE M. CONOVER The Winslow Foundation
GENE E. LIKENS
Director, The Institute of
Ecosystem Studies

Desert Puma - Evolutionary Ecology and Conservation

Hochgeladen von

Dokumentinformationen

Copyright

Verfügbare Formate

Dieses Dokument teilen

Dokument teilen oder einbetten

Freigabeoptionen

Stufen Sie dieses Dokument als nützlich ein?

Sind diese Inhalte unangemessen?

Copyright:

Verfügbare Formate

Desert Puma - Evolutionary Ecology and Conservation

Hochgeladen von

Copyright:

Verfügbare Formate

About Island Press

About the Hornocker Wildlife Institute

HORNOCKER WILDLIFE INSTITUTE,

Washington • Covelo • London

All rights reserved under International and Pan-American Copyright Conventions. No

ISLAND PRESS is a trademark of The Center for Resource Economics.

Library of Congress Cataloging-in-Publication Data

British Cataloguing-in-Publication Data available.

Printed on recycled, acid-free paper

Manufactured in the United States of America

List of Figures, Tables, and Photos xv

Part 1. Setting the Stage

Chapter 4. Studying Wild Pumas 39

Part 2. Puma Life History Strategies and

Chapter 8. Mortality and Survival 115

Part 3. Puma Behavior and Social Organization

Lifetime Home Range 195

Part 4. Puma–Prey Relationships

How Would a Puma Population Respond to a Prey Crash? 337

Part 5. Pumas and People

Appendix 1. Morphological Measurements of Pumas At Least 3.5 Months Old on

Appendix 2. Reproductive Chronology of Mated Pairs of Pumas on the San

10-1 Observed exponential rates of increase for adult pumas in the

radio-collared desert mule deer, San Andres Mountains, New Mexico,

14 Six-week-old cub F162 with eartag, tattoo, and radio collar. 49

public. Research focuses on scholarly, creative efforts designed to

The project in New Mexico that we envisioned would become a flagship

Department operations. Amy Fisher, Darrel Weybright, and Doug Humphreys

Davis), Paul Krausman (University of Arizona), Brian Miller (Denver Zoologi-

—KENNETH A. LOGAN and LINDA L. SWEANOR

Setting the Stage

Rationale and Approach

When we started this project, we made a significant long-term commitment.

Synopsis of Previous Studies

Pumas Past and Present

geographic distributions of individuals in a species) on pumas across the West-

involving a small number of individuals who migrated ‘out of South America’

bivore fauna. In the absence of megaherbivores, grasslands turned to woodlands,

Puma Distribution and Status

Pumas in the Southwest

PHOTO 2. Puma F47’s four-week-old cubs at the birth nursery.

Genders of adult pumas can be distinguished by their external sex organs.

Our Outdoor Laboratory

PHOTO 4. Upper Sulphur Canyon.

Physiography and Geology

Department of Agriculture’s Jornada Experimental Range (Fig. 3-2). During our

tures averaged 3.4 degrees Celsius. Extreme temperatures in summer reached

(Mimosa biuncifera), and southwestern condalia (Condalia lycioides) typify this

History of Human Use

PHOTO 5. Puma paw petroglyph near Three Rivers, New Mexico.

History of Puma Exploitation

Studying Wild Pumas

on a promontory, overlooked extraordinary puma habitat at the confluence of

PHOTO 6. Hembrillo Basin Camp.

PHOTO 7. San Andrecito Camp.

PHOTO 8. Puma tracks in the sand.

PHOTO 9. Puma M7 caught in a snare by his right forefoot.

PHOTO 10. Linda Sweanor using a

PHOTO 11. Ken Logan hauling puma

PHOTO 12. Puma F27 at her birth nursery.

and from January to February in 1995.

Mountains population and immigrant recruits from other puma populations.

PHOTO 14. Six-week-old cub F162

descriptions of dental characteristics. As the study progressed, we noted dental