Cougar--Human encounters: A search for the facts

Item

Title
Eng Cougar--Human encounters: A search for the facts
Date
1999
Creator
Eng Carnevali, Debbie D
Subject
Eng Environmental Studies
extracted text
COUGAR--HUMAN ENCOUNTERS:

A SEARCH FOR THE FACTS


by

Debbie D. Carnevali


A Thesis Submitt ed in partial fulfillment

of the requirements for the degree

Master of Environmental Studies

December 1998


This Thesis for the Master of Environmental Studies Degree

by

Debbie D. Carnevali


has been approved for

The Evergreen State College

by


d~ I!(?d;~

DR . JOHN PERKINS

Member of the Faculty


ANNA BRUCE

Wildlife Biologist

Date


ABSTRACT

Was hington State is the home of North America 's largest feline carnivo
the cougar, Felis concotor. This species is difficult to study because of its

secre tive, solitary and nocturnal nature which gives it an air of mystery and fos
fear . The cougar was thrust into the public limelight tw o years ago when

Initiative 655 was passed by Washington voters. The Initiative banned the use
hounds for hunting cougar and several other species.
Populations have increased since the bounty days before cougars were
given game status and management protection. Increases in mountain lion
sightings and encounters have raised the quest ion whet her the increase is due
solely to the two year old ban on hound hunting .

The topic of cougar-human encounters is comp lex and varied. The issu

include human population growth, habitat destruction and fragmentation, socia
and political views on carnivores, intolerance , and people' s fear due to
misinformation regarding cougars and their behavior.

Cougar biology , population dynamics and cougar management in twelv

states and two Canadian provinces (British Columbia and Alberta) were analyz
for this report. This review found that increases in cougar sightings and

encounters are greatest in areas of rapid human population growth where intru

into cougar habitat occurs. The analysis further dete rmined that cougar attack

rare and with proper safeguards and knowledge of cougar behavior people can
learn to co-exist with cougars.

INTRODUCTI ON
METHODS

2

HJ STORY

3

TAXONOMY

3

Table 1: The Family of Cats

4

5

D ISTRI B UTI ON
Figure 1: Historical and Current Range of the Cougar in the
Western Hemisphere

6

Table 2: Cougar Subspecies

7

Figure 2: Distribution of Cougar Subspecies in America

9

HABITAT

8

REPRODUCTION

10

DISPER SAL

11

Table 3: Dispersal Age, Month of Dispersal, Fate and Distance
Traveled for Nine Juvenile Cougars, 1990-92
HOl\1E RANGE

13

14

Table 4 : Average Home Ranges From Various Co ugar Stu dies
DEN SIT Y

16

17

Table 5: Cougar Population Densities Reported in Intensive
Studies In Nort h America

-111­

]8

DIET AND ENERGET ICS

]9

Table 6: Prey Items Reported Taken by Co ugars

21

Table 7: Comparison of Annua l Kill by Cougars, Derived fro m

Estimated Daily Consumption and from Frequency

of Kills

23

MANAGEMENT: BOUNTIES AND HUNTING
Table 8: Legal Status of the Cougar in the United Stat es

and Canada
COUGAR HABITAT IN WASHINGTON STATE
Tab le 9: Geographic Description of Cougar Management

Units (CMUs)

26

27

29

30

Table 10: Cougar Habitat (hectares) by Cougar Management Unit

(CMU) in Washington, 1997 (WDFW 1997a).
33
COUGAR HUNTING IN WASHlNGTON

33

Table 11: Cougar Permits Issued in Washi ngton From

1987 to 1997 .

35

Table 12: Cougar HarvestStatistics for 1997 Hunting Season

37

ADDITIONAL MORTALITY

38

Problem cougars.

38

Illegal harvest.

38

Native American Harvest.

39

Motor Vehicle Mortality.

39

-IV­

Natural Causes.

39

Cub Survival.

40

UN DERSTAND ING MYTH AND LEGEND

41

COUG AR ATT ACKS ON HUMANS IN THE UNITED STAT ES

41

Table 13: Cougar and Human Interactions Bo th Fatal and Non-Fatal

From 1890 to 1997
43
Table 14: Cougar Complaints Reported by WD FW Enforcement

Program and the Activity Cougar Were Engaging in,

1996
HUMAN POPULAnON TRENDS

50

54

Table 15: Counties with Highest Nu mber of Cougar Complaints and

Population Change
55
EXPLORING CALIFORNIA'S COUGAR HISTORY

57

WASHINGTON'S NEW COUGAR SEAS ON

58

COUGAR BEHAVlOR DURING ENCOUNTERS WITH HUMANS

AND APPROPRIATE HUMAN RESPONSES

60

Table 16: A Summary of Cougar Predatory Behavior and

Suggested Associated Human Re sponses
CONCLUSIONS AND RECOMMENDAn ONS

62

64

Research and Management

65

Ha bitat Protection

68

Education

70

BffiLIOGRAPHY

73

-v­

TABLE OF CONTENTS


APPENDIX 1:

Classification of the Felidae by W. Christopher

Wozencraft

88

APPENDIX 2:

Cougar Management Units (CMUs) in Washington

90

APPENDIX 3:

Cougar Tracks

91

APPENDIX 4:

Tracks of Cougar, Bobcat, Domestic Dog and Coyote

92

-Yl­

LIST OF FIGURES

Figure I : Historical and Current Range of the Cougar in the

Western Hemisphere

6

Figure 2: Distribution of Cougar Subspecies in America

9

-VIl ­

Table 1: The Family of Cat s

4

Table 2: Cougar Subspecies

7

Tab le 3: Dispersal Age, Month of Dispers al, Fate and Distance
Traveled for Nine Juvenile Cougar s, 1990-92

13

Table 4: Average Home Ranges From Various Coug ar Studies

16

Table 5: Cougar Population Densities Repo rted in Intensive Studie s
In North America

]8

Table 6: Prey Items Reported Taken by Cougars

21

Table 7: Com pariso n of Annual Kill by Cougars , Derived from
Estimated Daily Consumption and from Frequency
of Kills

23

Table 8: Legal Status of the Cougar in the United States
and Canada

27

Table 9: Geographic Description of Cougar Management
Units (CMUs)

30

Table 10: Cougar Habitat (hectares) by Cougar Management Unit
(CMU ) in Washington, 1997 (WDF W 1997a).

33

Table 11 : Cougar Permits Issued in Washingt on From
1987 to 1997.

35

Table 12: Cougar Harvest Statistics for ]997 Hunting Season

37

Table 13: Cougar and Human Interactions Both Fatal and Non-Fatal
From 1890 to 1997

43

Table 14: Cougar Complaints Reported by WDFW Enforcement
Program and the Activity Cougar Were Engaging in,
1996

50

-vm­

Table 15: Counties with Highest Number of Cougar Complaints and

Pop ulation Change

55

Table 16: A Summary of Cougar Pre datory Behavior and Suggested

Associated Human Responses

62

-IX­

ACKN OWLEDGEMENTS

I wish to thank Dr. Oscar Sou le, Dr . John Perkins and Anna Bru

for their time, expertise and guidanc e in the development of this Thesis
A special thank you to my husband, Rick T. Carnevali, for his

patience in reading and rereadi ng my drafts and for his support, guidanc
and encouragement .

-x­

INTRODUCTION

In November 1996, Washington voters passed Initiative 655 which stat
in part:

(2) Notwithstanding RCW 77.12.240 or any other provisions o
law, it is unlawful to hunt or pursue black bear, cougar, bobcat
lynx with the aid of a dog or dogs (emphasis added) .

With the passage of this initiative, Washington became the third state to ban th

use of hounds to hunt black bears, cougars, bobcats and lynx, following Califor

in 1971 and Oregon in 1994. Conservation and animal protection groups in o
states in the West are also looking at putt ing similar initiatives before voters.
One of the most controversial and challenging conservation issues is

carnivore conservation. Large carnivores (e.g . wolves, Canis lupus, grizzly be

Ursus arctos horribilis, and the cougar, Felis concolor), have been among the
most persecuted of all North American animals (Clark et al. 1996). However,

increased public awareness and education, many people are taking a closer look

the importance of carnivores in a healthy ecos ystem . This change in attitude fr
treating carnivores as vermin to be extirpated, to important and necessary
components in a healthy ecological system, is showing up in the form of laws
protect and ensure their existence.

-1­

The passage of this initiative has thrust Washington State' s top wild lar

feline predator, the cougar, F. concotor, also known as the mountain lion, pum
panther and catamount, into the public limelight where it has become a

controversial political animal. It is important to note that Initiative 655 did no
ban the hunting of cougars; it only banned a particular method used to hunt
cougars : the use of hounds to track and tree these animals.
Although this change in the law has been in effect for only two years,
reports of cougar populations exploding and increased cougar sightings have

heightened the public's fear for its safety . This paper willlook at cougar biolo

management history , and interactions with humans in order to answer the ques
Is the increase in human-cougar encounters a direct result of the ban on using
hounds to hunt cougars? It is hoped this research wilI assist the public in

understanding cougar history , population dynamics and methods for living with
this species .

METHODS
I examined over 200 articles and boo ks in the published literature and

conducted interviews with wildlife biologists in order to evaluate cougar history
popu lation dynamics, management practices and public education programs in
twelve states and two Canadian provinces.

-2­

HISTORY
Ancestral cougars , Felis inexpe ctata, occ urred in North America from
Blancan to the Irvingtonian period during the Pleistocene epoch, three to one
million years ago, with modem cougars appearing about 100,000 years ago

(Kitchener 199 1). During the Pleistocene epoch , there were a number of coug
species, but only F. concotor survives today (Busch 1996).

TAXONOMY

One of the first classifications of the family Felidae was done by Carolu

Linneaus, the father of taxonomy. It was Linneaus who first named the cougar

placing it in the genus Felis. There have been disagreements between scientists

over how to classify the 37 species of cats that exist today. The main controve
involves the division of the family Felidae into genera and subgenera (Nowak

1991) . Each taxonomist had his/her own criteria for classifying the cats (Nowa
199 1). Many scientists today divide the cat family into four genera: Panthera

large roaring cats ; Felis, the smaller purring cats; Neofe lis, the Clouded Leop
and Acinonyx, the Cheetah (Table 1):

-3­

Tiger iPanthera tigris)
Lion (Panthera leo )
Panthera ----­ -------­ Leopard (Panthera pardus )
Jagu ar (panthera onca )
Snow Leopard (Panthera uncia)

Cou gar (Felis concolor)
Felis ------------------ Ocelo t (Felis pardalisi
Caracal (Felis caracal)
and 27 other species of Felis

Neofelis -------------­ Clouded leopard (Neofelis nebulo

Acinonyx ----­ ------­ Cheetah (Acinonyx jubatus)
(Hansen 1992).

F. concotor is the most widely used scientific name for the cougar.
However, a recent reclassification occurred in 1993 in W. Christopher

Wozencraft's, "Classification of the Felidae" (Appendix 1). Under Wozencraft

classification, the cougar is known as Puma concolor. This classification is also
widely used and has been adopted by the Convention on International Trade in
Endangered Species of Wild Fauna and Flora (CITES) and the World

-4­

Conservation Mo nitoring Centre (WCMC).

This paper will use the classificat

by Carolu s Linneaus, F. concotor .

DISTRIB UTION
Cougars once ranged from British Columbia to the Straits of Magellan

South America , and coast to coast from the northern United States and souther

Canada southward (Russell 1978) . This extensive distribution indicates a plasti

for living in many habitat types and for utilizing a variety of prey species (Hopk
1984).

The cougar is widely distributed thro ughout its current range acro ss No

and South America (Figure 1). They have been extirpated from most of eastern
North America and population figures for South America are incomplete. It is
estimated that there are 27-30 subspecies scattered across North and South
America, depending on which taxonomic list is used (Table 2).

-5­

II

i
I
I

I
II;.

C li RRE!': T RANGE
j!

I! ~
I,

" r--- ,-- _

FIG URE 1: HISTORICAL AND CURRE NT RA NGE OF
COU GAR IN THE WESTERN HEMISPHER
(Status of Cougar in Mexico, Central and Sou
America is unknown) (Hansen 199 2)
-6­

TABLE 2: Cougar Subspecies in North and South America:
Cougar (Felis con c%r) Subspecies
North America

F. c. cOlIgllar--Eastern cougar

F. c. browni--Yuma puma

F. c. schogeri--Wisconsin cougar

F c. improcera--Baja California couga

F c. missoll/ensis--Missoula cougar

F. c. azteca--Mexican cougar

F. c. hippo/estes--Colorado cougar

F c. stanleyana-» Texas cougar

F. c. oregonensis--Oregon cougar

F. c. cOlyi--Florida panther

F. c. vancollverensis-- Vancouver

F. c. mayensis--Mayan cougar

Island cougar
F. c. ca/ifomica--California cougar

F. c. costaricensis--Costa Rican puma

F c. kaibabensis--Kaibab cougar

South America

F. c. conc%r--Brazilian cougar

F. c. acrocodia--Mato Grosso cougar

F c. bangsi--Colombian cougar

F. c. plIma--Chilean puma

F. c. soderstromi--Ecuador cougar

F. c. cabrerae--Argentine puma

F. c. disc%r--Arnazon cougar

F. c. pearsoni--Pearson's puma

F. c. incarum--Incan cougar

F. c. patagonica--Patagonia puma

F c. osgoodi--Bolivian cougar

F c. arancanus-Asvies puma

(Busch 1996).
In a review of the literature, Washington's cougar population would be

Felis concotor oregonensis. However, E. A. Goldman (1946), in his

-7­

Washington: F. c. oregonensis and F. c. olympus, a subspecies found on the
Olympic Peninsula (Figure 2).

HABI TAT

The cougar has one of the greatest natural distributions of any mammal

the Weste rn Hemisphere (Nowak 1991). It can thrive in montane co niferous a

lowland tropical forests, swamps, grassland, dry brush country or any other are
which provides adequate food and cover. In studies conducted in the remote
wilderness area in the Idaho Primitive Area, (now known as the Frank Church

River of No Return Wilderness Area), researchers found that cougars preferred

steep, rocky areas covered with dense stands of Douglas fir and ponderosa pine
with sagebrush and grasslands mixed among the bluffs and talus slopes (Horno
1970; Seidensticker et al. 1973). In the Bighorn Mountains of northern
Wyoming , researchers found similar results . Cougars frequented canyonland

habitats with steep, rugged slopes (greater than 45 degrees) containing a mix o

conifer and brushy mountain mahogany cover. Grasslands and sagebrush areas

with gentle slopes (less than 20 degrees) were generally avoided (Hansen 1992
Cougars normally avoid large open spaces where there is insufficient cover .

Suitable cougar habitat is a combination of veget ation, top ography , prey numbe
and prey vulnerability (which depends on both stalking and escape cover).

Persecution, habitat loss and reduction in prey species have brought bot
the Eastern cougar, F. concolor couguar , and the Florida panther, F. concolor

-8­

\

\

\ I
\

I

\ I
I. F c . concotor
2. F c. couguar
:I. F c. mtssoulensis
4 . F. c. mppolesies

5. F c. oregonensis
6 . F. c. vancouvere~ 5
7. F.c. olympus
8. F . c. caltfor ruca
9. F . c. kaib abensi s

10. F c brown i

I I. 1-. c . tmprocera
12. F c. azteca
I:1 T. c. stanlevan a
14. T. c. coryr
15. T. c. mavensis
16 .
c. costancenst»
17 T. c. bangsi
18. F. c. soderstromt
19 . F c. captncom enst s
20 F c . anthonvi

r

2 1. F c tn carum

cz.

F. c. borbensts

23. F c. osgoodi
24 . F c. acrocodia

25 . F c. gre eni
2(, 1-. c. puma
27 . F c. cabrerae
28 . F. c . nearsoni
29. F. c. patagom ca
) 0 F c . araucanus

FIGURE 2: DISTRIB UTION OF MOUNTAIN LIO N SUBSPECIES
IN AMERICA (Goldman 1946 )

-9­

Florida panthers remaining. The Eastern cougar was thought to be extinct, but
recent cougar sightings in northeastern Canada and the United States have
scientists hopeful there is still a small, remnant popu lation which may have
survived .

REPRODUCTION

Cougars can reproduce on a year-rou nd basis (Sweanor 1992; Ross et a

1992), although most births are reported to occur in the warmer months of Apr

September in their northern range (Robinette et at 1961; Eaton and Verland 19

Ashman et al. 1983; Lindzey 1987) . Both sexes are sexually mature at 24 mon

with females sometimes becoming sexually mature at 20 months (Lindzey 1987

The time of first breeding probably depends on when a female is able to establi
her territory (Hornocker 1970; Seidensticker et at 1973). Logan et al. (1986)

found that females entered the breeding population at age 3-4 years in their hun
study popul ation in Wyoming.

Females in stable populations rarely breed with more than one male dur

estrus (Hemker et al. 1984) . Estrus is app roximately eight days and the estrus

cycle of female cougars is 23 days (Hansen 1992) with a gestation period of91

4 days . Litter size varies from one to six, but the average is 2.2-2.7 (Anderson
1983; Ross et at 1992) . Lindzey (1987) reported that first litt ers may contain
one kitten, which enables a new mother to develop her parenting skills.

Newborn cougars, called kittens or cubs, are born weighing slightly mo

- 10­

canals are closed, their coats are covered with blackish brown spots and their ta
are dark ringed. This color patte rn provides excellent protective camouflage.

Kittens begin nursing within minutes afte r birth and gain weight rapidly. At tw

weeks of age the kitt ens' eyes and ears are open and they are able to walk. Wi
20 to 30 days the kittens may weigh over two pounds (Hanse n 1992) .

While suckling her young, the mother must occasionally leave the den t
hunt. This is the time of her most restricted movements, but she must hunt to
sustain herself and replenish her milk. The kittens are weaned at two to three
months but are dependent on their mother for food until they disperse.
The interbirth interval, which ends when the last litter disperses, can be

year (Robinette et al. 1961) but more generally is 18-24 months (Lindzey 1987

DISPERSAL
The departure of young cats from their mother's home range is called

dispersal. Dispersal is normally initiated by the mother leaving the cubs 0-3
kilometers (km) from an edge of her home range while she moves to or ju st

beyond the opposite edge of her home range, remaining there for 2-3 weeks (B

1995). The age of dispersal in Beier's (1995) study ranged from 13.5 months t
22 months, with an average of 18 months. Dispersal typically occurred in the

spring months of April, May and June in Alberta, Canada (Jalkotzy et al. 1992
but year around in California (Beier 1995).

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Dispersal is a time when the young cougars are especially vulnerable. T

expose themselves to the danger s of finding and taking prey, and no longer hav

available the alternative food source provided by their mother. Since their hunt

slcills are limited, they risk injury while catching prey, and are vulnerab le as wel

injury or death from cougars and other predators, including humans. Since they
are looking to establish their own territory, these young cougars often end up,

even for a short period of time, in marginal habitats, including urban , rural area
greenbelts and parks. They often face lethal remov al by cont rol agencies or
landowners, even if the cougar has not injured a person. However, due to the
close proximity to people and/or children, the potential for hann is still there.

Even in a non-hunted population, mortality is high for dispersing couga
Beier (1995 ) conducted a study on the dispersal of nine juvenile cou gars in

fragmented habitat in the Santa Ana Mountains in Southern California. Cougar

dispersal is important for recruitment into adjacent local populations. As Table

sets out, only two of the nine cougars survived to recruit into other population

-12­

for Nine Juvenile Cougars, 1990-92:
Sex
F

Dispersal age
(Month)
20

Dispersal
Mo nth
September

Fate
Died (natural cause)

Dispersal
Dista nce (k
48

M

18

January

Died (disease)

19

M

19

August

Died (vehicle)

69

M

19. 5

August

Recruited

75

M

19.5

April

Recruited

56

M

18.5

December

Died (vehicle)

77

M

22

March

Died (vehicle)

66

M

17

January

Died (shot)

69

M

13.5

September

Died (natural cause)

75

(Beier 1995 ).

In an Alberta, Canada study, the fates of eight of 11 cougars that disper
from their maternal home ranges were recorded . Five of the eight were shot
within 25 km and 150 km from their maternal home ranges: tw o, a male and

female, were shot as subadults, while three were shot at 2 years 8 mont hs, 3 ye
five months, and 5 years 6 months of age, respectively. Of the remaining three
cougars, two females died of natural causes as subadults, and only one cougar
subsequently established a home range (Jalkotzy et al. 1992).

-13­

HOME RANGE
While cougars are not territorial in the classic sense, i.e., they do not
defend an exclusive area against all cougars, they are attached to their homes.
Within cougar habitat, adult cougars space themselves out and confine their
movement to individual fixed areas. These areas are known as home ranges.

These areas include hunting areas, water sources, resting areas, lookout positio
and denning sites where young can be safely reared . Cougars without a home
range generally do not breed .

Male cougars will usually overlap several females' home ranges. In mo
cases, male home ranges will not overlap , but female home ranges do . It is not

unusual for female cubs to establish home ranges contiguous with their materna
home ranges (Jalkotzy et al. 1992)
There is a feline social hierarchy consisting of three classes of animals:
resident adult males and females, transient males and females, and dependent

offspring of resident females (Hansen 1992). Resident adults maintain establish

home ranges and do most of the breeding in a pop ulation. Transients constantl

move throu gh homes ranges of residents in search of a vacant home range of th
own . Tran sients are approximately 18 to 24 months old and have left their

mother's home range , but have not established their own home range. They ar

-14­

also known as emigrants and dispersers.
The size of an animal's home range is a function of the abundan ce,
distribution and vulnerability of the prey, to pography, climate and a variety of
ot her factors , including body size. Male cougars are 1.5-2 times larger than
females and have larger home ranges .
Sex differences in home range size, however, may be related to factors
other than body size. The size ofa female 's home range must be large enough

provide for the nutritional needs of herself and her offspring. While she is likel
have metab olic requirements similar to the large males due to the cost of

repro duction and parental care, the female will also have a reduced foraging ra

because she must frequentl y return to her cubs. These constraints may explain

smaller, but more intensive use of a home range (Hopkins et al. 1986). Table 4
summarizes home ranges found in studies conducted in seven states and two

Canadian prov inces and shows the difference in the size of home ranges betwee
male and female cougars.

-15­

Me an home ranges (square kilometers) of both male and female cougars from
seven states and two pro vinces of Canada from radio telemetry and markrecapture studies.
Year
Season
Male
Female
State/
Providence
1981-89
204
97
Alberta'
Winter
314
87
Summer
334
Annual
140

Arizona'

1972-73

Winter

British Columbia]

California"

1978-82

Ida hoS

1970-71

All Year

189

67

151

55

82
88
105
355
301

71
62
70

453

268
8
32
20

Winter

Nevada"

New Mexico"
Utah 8

1972-73
1975-78
1978-79

All year
All year
Varied by
cougar

2 11
480

142

222

79

1975-77
1976-78

All year

182

1979-81

No nwinter
Winter

1981-83

All Year

122
573
503

54
67
91
57
Sources: 'Spreadbury, 1988; 2Shaw 1973; ]Ross et al. 1992; "Sitton et al. 1976
sSeidensticker et al. 1973, Homocker 1969; "Ashman 1976, 1983; 7Bavin 1976
8Hemker 1984; "Logan, 1985, 1986.
Wyo ming"

-16­

370
269

232-556 (
100-42 1 (
190

D ENSITY

Cougar social structure ensures that they seldom overpopulate an area.

range size and the degree of overlap between ho me ranges influences the densi

cougar population. In unhunted or very lightly hunted popu lations, social inter

(Homocker 1969; Seidensticker et al. 1973) or prey densities (Hemker et al. 1

may be more impo rtant in controlling populations . As Table 5 sets out, differe

studies in different areas have varying cougar densities. Beier (1993) stated re
cougar density estimates:

Because many study sites were selected because of expected hi

cougar density, some reported densities are atypically high. Als
all studies report how many of these animals were nonbreeding

transients. Cougar carrying capacity must be estimated by numb

breeding adult males and females, excluding the pool of nonbree
male and female transients that characterize most populatio ns
(Seidensticker et al. 1973) . Categorizing all
individuals over 1 year of age as adult breeders would lead to
overly optimistic predictions.

Therefore, it is important to look at specific cougar populat ions, habitat, and p

availability in order to adequately assess whether cougars are "overpopulated."

also important to remember that cougars are recoveri ng from depressed numbe

-17­

elimination of bounties and the change in game status.

TABLE 5: Cougar Population Densities Reported in Intensive Studies in
North Americ a:
Location

Estimated densitya
(Cougar/1 00 km2)
Annual

Albert a

2.7-4 .7
1.5 - 5.9

Utah

0.3 - 0.5
0.37

Idaho
California

Winter

Jalkotzy et at. (1992)
Management Plan for Couga
In Alberta
0.6

Hemker et al. (1984)
Lindzey et al. (1994)

2.1 -7.4
2.9

Seidensticker et at. (19 73)
Homocker (1970)
Sitton et al. ( 1976, 1977)

3.5 - 4.4
1.2-2.3b
1.7-3 .3

Colorado

Source

Currier et at. (19 77)

Arizona

3.2 - 3.5

Shaw ( 1973, 1979)

Nevada

1.4 - 1.6

Ashman (19 76); Ashman et
(19 83)
Unknown

Washington
Oregon

3.0

Oregon Dept. Of Fish and
Wildlife (1993)

British Columbia

3.5- 3.7

Spreadbury (1988)

aCalculated by dividing the size of the study area by the total number of couga
(including dependent kittens and juveniles) present.

bAdults only.


-18­

DIET AND ENERGETICS

The known prey of cougars thro ughout their range includes insects, bird
mice, up to porcupine, capybara, pronghorn antelope, elk, deer, bighorn sheep

moose . Large ungu lates, particularly deer, are the cougars ' principal prey in N

America. Studies on food habitats for cougars in sout hern Chile found that ung
made up 68% of the average diet (Iriarte et al. 1991). Mu le deer, Odocoileus

hemionus, are the primary prey in Oreg on, Alberta and Utah, while in Florida w
tailed deer, 0. virginianus, and wild hogs, Sus serof a, are the preferred prey .

Hornocker (1970) found that cougars in the Idaho Primitive Area fed primarily

mule deer and elk, Cervus canadensis. In Nevada, the adaptable cougar occasi

augments its diet with wild horses , Equu s cabal/us, and desert bighorn sheep, O

canadensis (Hansen 1992). Moose, A Ices alees, also are taken by cougar in B
Columbia and Alberta.

Cougars are opportunistic predators and also feed on a variety of smalle

prey, especially in times of seasonal abundance. Columbian ground squirrels, C

columbianus, are frequently the cougars' main course during the warmer summ

months in Idaho, while peak years of snowshoe hare, Lep us americanus, abund

in British Columbia can comprise over one-quarter of the cougars' diet (Seiden
et aI. 1973; Spalding et al. 1971).

- 19­

In the southern parts of cougar range, and particularly in the tropics, sm

medium-sized prey appear to be more important. Studies by Iriarte et al. ( 199 1

suggested cougars' smaller body size in the tropics , and their low rate of predat

larger prey, such as tapirs, Tapirus sp., may be linked to interspecific co mpetiti
with the jaguar, Panthera onca .

In Washington, the major prey species include mule deer, white-tailed d
and elk east of the Cascades, and Washington hare, Lep us washingtonii, black-

deer and elk west of the Cascades (Brittell and Pierce in Washington Departme
Fish and Wildlife (WDFW) 1997). Other important food sources include the

snowshoe hare, 1. americanus, black-tailed hare, 1. califam icus, white-tailed h

tawnsendii, and porcupine, Erethizon dorsatum . Table 6 shows the wide variet
food sources which cougars utilize.

-20­

TABLE 6: Prey Items Reported Taken by Cougars:
Large
Wild Animals

Bear, Ursus sp.
Mule deer Odoco ileus hemionus
Bobcat, Lynx rufus
White-tailed deer, 0. virginianus
Cougar, F. concolor
Elk, Cer vus canadensis
Bighorn sheep, Ovis canadensis
Coyote, Canis latrans
Moose, Alces alces
Pampas deer, Ozotoceros sp.
Pro nghorn, Antilocapra americana
Caribou, Rangifer caribou
Mountain goat, Oreamnos americanu s
Huemul, Hippocamelus sp
Wild boar, Sus scrofa
Peccary, Pecari angulatus
Porcupine, E. dorsatum
Beaver, Castor canadensi
Badger, Taxidea taxus
Armadillo, Dasypus
no vemcinctus
Guanaco, Lama guanicoe
Small
Wild Animals

Jackrabbits and Hares, Lepus sp

Meadow vole, Microtus
Turkey, M eleagris sp.

pennsylvanicus
White-footed vole, Ph enacomys albipes

Raccoon, Procy on lotor
Other rabbits, Sylvilagus sp.

Pika, Ochotona princeps
Marmot, Marmota sp.

Fox, Vulpes sp.
Skunk, Mephitis sp.

Coati, Nasua narica
Ground squirrels, Citel/us, Otospermophilus sp.
Squirrels, Tamiasciurus, Eutamias, Tamias, Sci urus sp.
Flying squirrel, Glaucomys sp.
Opossum, Didelphis mars
Pocket gopher, Thomomy s sp.
Agouti, Dasyprocta sp.

Mountain beaver, Ap lodontia rufa
Woodrat, Neotoma sp.

Cotton rat, Sigmodon sp.

Ruffed grouse, Bonasa umbel/us

Domestic

Animals

Sheep, Ovis sp.
Pig, Sus sp
Horse, Equus caba l/us
Peafowl, Francolinus sp.
Burro, Equus asinus

Other


Items

Dog, Canis f am iliaris

Cattle, Bos sp.

Goat, Capra sp.

Rhea, Rhe a sp.

Cat, Felis catus


Grass Berrie
Fish
Insect
Domestic Fow

(Hansen 1992)
When the abundance of their primary prey declines, cougars have been
-21­

ational Park in western Texas , cougars were forced to switch to peccaries and
lagomorphs, the next largest prey (Hansen 1992).

The diet of a cougar also varies according to its sex and age. Solitary c

seem to consume more small prey than do females with young, while transients

on smaller prey until they have developed their skills to stalk and kill larger prey
The frequency of prey kills depends upon several factor s:








Sex and reproductive status
Size of the dependent young
Social status
Abundan ce of alternate prey
Rate of spoilage of kill
Time of year

Many of the studies on food habits of cougars have concentrated on dee
elk since those species are managed for human use and cougars are seen as

competitors for the resource. A stud y in southern Utah (Ackerman et al. 1984
estimated the following kill rate s of deer for resident females:
Solitary resident female

1 deer per 16 days

Resident female with three
3-month old kitten s

1 deer per 9 days

Resident female with three
15-month-o ld kittens

1 deer per 3 days

These kill rates are similar to other studies conducted to determine daily
con sumption rates and annual prey kills of cougars (Table 7).

-22­

Source
and Area

Estimated
Consumption

(Kg/day)

Computed*
prey biomass
killed
(kg/yr)

Computed**
animals
killed
(#/yr)

Estimated
kill rate
(1 animal!
# of days)

Resident female: 1
deer/ 16 days
Female w/three,
3-month old cubs :
1 deer/9 days

Ackerman et al.
(1984 ) southern
Utah

Female w/three,
IS-month old cubs
1 deer/3 days
You ng & Goldman
(1946) throughout
West

Single adult: 1 dee
7 days
Female w/cubs:
1 deer/3+ days

Conno lly (1949)
central Utah

Single adult: 1
deer/9 .7 days &
1 porcupine/7.2
days

Robinette et al.
( 1959) Utah
and Nevada

2.3-5.5

Hornocker (1970)
central Idaho

1.8-2.7

860- 1300

14-20 deer
(@ 64 kg)
or
5-7 elk
(@ 175 kg)

Single adult: 1 dee
0-14 days

Shaw (1977)
central Arizona

1,8-2 .7

860-1300

17-25 deer
(@ 51 kg)

Single female:
1 deerlI 0.4 days
Female w/cubs:
1 deer/6 .8 days

Single adult: I dee
4-1 0 days

*Daily consumption x 1.3 to include carcass wastage.
**Prey biomass killed divided average prey weight.

-23­

examined in six studies were empty . Cougars exhibit a gorge-and-fast type of f

behavior and will shift prey selection in response to prey availability. Therefore
cougars may go without eating for days at a time.

There are other variables which dictate the number of prey taken. Spo

of carcasses in summer may lead to an increase in the number of animals killed.

the other hand, scavenging of winter- killed deer and elk (Ackerman et al. 1984)

reduce the predicted impact of cougars on ungulate populations, as could increa

use of unusu ally abundant small prey such as snowshoe hares (Spalding et aI . 19

Changes in structure of the cougar population, e.g. caused by intensive
harvest, wou ld alter the population's energetic needs, and thus alter the cougar

population's impact on prey species. Smuts (1978) showed that harvesting of A

lions caused an increase in the number ofjuveniles, presumably with an attendan

increase in energetic demand by the population (Ackerman et al. 1986). Shaw (

suggested when stable social systems are disrupted with the removal of resident

cougars out of a population, the removal can cause an increase in the populatio
through immigration and establishment of territ ories by tran sient cougars.

Female cougars are clearly in an unusual ecological role. Cubs are dep

on the female for longer periods than juveniles of most vertebrate species . With
exception of ot her large felids, juveniles of most species are able

-24­

to obtain a portion of their food within a few months (Ackerman et al. 1986).

Although it is rare , cougar cubs may be capable of surviving on their own after 6

months ; however, they ordina rily rely heavily on the female for their food until 1
months of age (Ackerman et al. 1986). Females may hunt as far as 16 km from
kittens and be gone for several days (Hemker et al. 1984).

Needs of a single adult could be met by 1-2 black-tailed jackrabbits, but

would tak e 7-8 jackrabbits a day to feed a family group with three, 15-month old

cubs. Ackerman's (1986) study suggests that, while individual cougars can prob
live and supp ort themselves in many areas , an adult female with cubs may be so

dependent on large prey that viable, breeding cougar popu lations cannot exist in
absence of deer-sized ungulates.

Cougars also cache excess food . This enables them to return to the foo
source for several days, thus expending less energy . In a study conducted by
Thompson et al. (1994) in Montana, a cougar was observed feeding on an adult

female elk for 27 days. The study estimated the elk to weigh 236 kg. Assuming

unconsumed remains of the elk constituted 30% of its live weight (Hornocker 1
the cougar consumed approximately 165 kg in 27 days, or an average of 6.1 kg
day. It is unusual for a cougar to consume a kill so slowly, but Thompson et al.

(I 994) theo rized the long period may have been due to low densities of potentia

competitors for the meat.

-25­

MANAGEMENT: BOUNTIES AND HUNTING:

It has taken a long time for the cougar to be viewed as more than verm
When the Europ eans arrived in the New World, they brough t an att itud e of
intolerance for animals which competed with them for land and game.

The first bounty on cougars was established by Jesuit priests in Califor

the 1500s when they offered natives a bull for each cougar killed (Busch 1996)

1694, Connecticut offered a bounty of twenty shillings for each dead "catamou
South Caro lina's "Act for Destroying Beasts of Prey," enacte d in 1695, forced

American hunt ers to submit a predator's hide annually or be whipped in punish

Massachusetts enacted a bounty payment offorty shillings per cougar in 1742 (
1996) .

A new perspective on predators began in the 1960s and a limited degre

protection for the cougar began as state and provincial legislatures and wildlife

managers in most of the West, shifted cougars' official status from injurious pre

to game animal. Alberta, Canada, removed the bounty on cougars in 1964. In

United States, Nevada reclassified the cougar as a game animal in 1965, Wash

in 1966, Utah in 1967, California in 1969 and Arizona was the last state to hav

cougar bounty and did not remove it until 1970. Table 8 sets out the legal stat
hunting and trapping status of cougars in the United States and Canada.

-26­

TABLE 8: Legal Statu s of the Cougar in the United States and Canada:

State/
Province

Hunting
Season
Yes
No

Legal status

X

Trappin
Seaso
Yes
N

X

Alabama

Game

Alberta

Game

X

X

British Columbia

Game

X

X

California

Specially Protected
Mammal


Colorado

Game

Florida

Endangered

Idaho

Game

Lo uisiana

Protected

X

X

Manitoba

Protected

X

X

Mississippi

Endangered

X

X

Montana

Game

Nebraska

Unprotected

Ne vada

Game

X

X

New Mexico

Game

X

X

N . Carolina

Endangered

N . Dakota

Unprotected

X

X

X
X

X

X

X

X

X
X

X
No closed season

-27­

X

X

X

TABLE 8 (continued): Legal Status of the Cougar in the United States an
Canada:

Hunting
Season
Yes
No

Trappi
Seaso
Yes
N

Statel
Province

Legal status

Oklahoma

Protected

Oregon

Game

Saskatchewan

Protected

X

X

S. Dakota

Thr eatened

X

X

Texas

Unprotected


Utah

Game

Virginia

Endangered

Washington

Game

X

X

Wyoming

Game

X

X

X
X

X

X

X

X
X

X

(Green 1991)

In addition to hunting, cougars are killed under the United States' Fede
Animal Damage Control Program (ADe) which was established in 1915. The

program was given sweeping powers for "the destruction of mountain lions, wo
coyotes, ... and oth er animals injurious to agriculture, horticulture, forestry,
husbandry, game or domestic animals, or that carrie d disease". Between 1937

-28­

1977, U. S. federal authorities officially killed over ten thousand cougars, altho
some claim the actual toll was much higher (Busch 1996). In addit ion to the

removal, over two hundred thousand cougars were killed by bounty hunters be
1900 and 1970 (Busch 1996).
Little was known about predator management prior to the 1960s, and
result of large-scale removal of predators was often catastrophic. In the early

Arizona officials decided to kill off the cougars, coyotes, and bobcats from the
Kaibab Plateau in order to "improve the hunting for humans" (Busch 1996).
Without the limiting effect of natural predation, the deer population exploded

3,000 in 1910 to over 100,000 by 1924. The area was soon overgrazed and d
by the thou sands. The cougar did not return for almost fifty years.
COUGAR H ABIT ATIN W ASH INGTON STAT E

Cougars occur throughout Washington State except in areas of the Co

Basin which are devoid of either shrub steppe or forests . The cougar is associ

with coniferous forests with a mixture of vegetative types and seral stages pro

both abundant cover and abundant prey. In Washington, the greatest concentr
of cougars are located in the Cascade Mountains, Blue Mou ntains, Okanogan

Highlands, the northeastern quarter of the state and the Olympic Mountains (W
1997a).
Population and harvest data for game species in Washington State are
analyzed by different geographical areas. WDFW recently developed new

-29­

cougar management units (CMU s). CMU s are based on game management un

ecoregions which were identified using a program developed by the University

Washington (WDFW 1997a). This program determines the location of gaps i

biodiversity of wildlife species and their geographical range and is known as G

Nine CMUs were designated by WDFW and Table 10 out lines the units and th
boundaries :

TABLE 9: Geographic Description of Cougar Ma nagement Units (CMUs
UNIT

Coastal

DESCRlPTION

The western boundary is the Pacific Ocean , the northern
boundary is the Strait of Juan de Fuca, the eastern bounda
the Hood Canal and Capitol State Forest, and the souther
boundary is the Columbia River. All or portions of the Ch
Elwha S'Klallam (Port Angeles), Hoh, Jamestown S'Klall
Makah, Ozette, Quileute, Quinault, Shoalwater, and Skok
Indian reservations lie within this CMU . The Olympic Na
Park and the Willapa Wildlife Reserve are also within this

The western boundary is the Hood Canal, the northern
boundary is the Canad ian border, the easte rn boundary is
western foothills of the Cascade Range, and the southern
boundary is approximately the Cow litz River . All or porti
the Lower Elwah, Lummi, Muckleshoot, Nisqually, Nook
Port Gamble, Puyallup, Skokomish, Squaxin Island,
Stillaguamish, Sauk-Suiattle, Suquamish, Swinomish, Tul
and Upper Skagit Indian reservations lie within this CMU
North
Cascades

The western boun dary is Highway 9 and Highway 203 ; th
northern boundary is the Canad ian border; the eastern bou
is the Mt. Baker National Forest, and the Skagit, Snohom
and King county lines to Snoqualimie Pass; and the southe
boundary is approximately the 1-90 corridor. The North
Cascades National Park lies within this CMU .
-30­

TABLE 9 (continued): Geographic Description of Cougar Management
Units (eMUs) :
UNIT

DESCRIPTIO N


South
Cascades

The western boun dary is Interstate 5, Highway 508, Highwa
7 and the Burlington-Nort hern railroad lines; the nort hern
boundary is the 1-90 corri dor; the eastern boundary is the
Skamania, Lewis, Pierce, and King county lines; and the
southern boundary is the Columbia River at the Oregon bor
The Mount Rainier National Park and part of the Yakama In
Reservation lie within this CMU .

East
Cascades
No rth

The western boundary is the Mt. Baker National Forest, and
the Skagit, Snohomish, King, and Pierce county lines to the
Mt. Rainier National park; the southern boundary is the
Umtaneum Creek , south fork Manast ash Creek, and
American River; the eastern boundary is the Met how River
the Columbia River, Colockum Pass, and North Branch
Canal; and the northern boundary is the Canadian border.

East
Cascades
South

The western boundary is the Skamania, Lewis, and Pierce
county lines; the southern boundary is the Columbia River a
the Oregon border; the eastern boundary is the Yakima Rive
and the Yakama Indian Reservation; and the northern bound
is the Umtanum Creek, South Fork Manastash Creek, and t
American River. Most of the Yakama Indian Reservation li
within this CMU .

Columbia
Basin

The western boundary is the Methow River to the Colu mbia
River ; the sout hern boundary is Oregon bord er; the eastern
boundary follows the Columbia River north to Trinidad, alo
the Douglas county line, up the Columbia River and then th
Okanogan River ; and the northern boundary is the Canadian
border.

North­
eastern

The western boundary is the Okanogan River; the northern
boundary is the Canadian border; the eastern boundary is th
Idaho border; and the southern boundary is Highway 174,
Highway 2, and Highway 231 east to the Spokane County
line. Allor portions of the Colville, Kalispell, and Spokane
Indian reservations lie within this CMU .

-3 1­

TABLE 9 (continued): Geographic Description of Cougar Management
Units (CMUs) :
UNIT

Blue
Mountains

DESCRIPTION


This unit encompasses the entire Blue Mountain range in
Washington from the Snake River on the north, to the Idah
border on the east, the Oregon border on the south, and th
Columbia River to Burbank on the west.

(WDFW 1997a)

The GAP analysis provided WDFW with a habitat inventory which estim

current available cougar habitat in each CMU in the state (Table 10). All forest

land was considered cougar habitat , including deciduous forest, conife rous fore
deciduous/coniferous mixed forest. Forested lands which were devoid of trees
fire, timber harvest, or some other event , were also considered cougar habitat.
Typically these non-forested sites still had an understory of shrubs, grasses and

which pro vide foraging and trave l cover, and serve as year-round habitat once t
trees grow.

-32­

in Washington, 1997 (WDFW 1997a):
Current
Habitata

CMU

Additional
Habitatb

Total
Habitatc

Total as a perc
of statewide

E . Cascades North 1,648,221

11 ,654

1,659,875

19

Coastal

1,433,780

33, 126

1,466,906

17

Northeastern

1,499, 6 19

16,850

1,5 16,469

17

S. Cascades

1,143,562

66,3 86

1,209,948

14

Puget Sound

991,467

60,943

1,052,4 10

12

N. Cascades

900,694

26,770

927,464

10

E. Cascades South

568,011

15,451

583,462

7

Columbia Basin

274,306

133

274,439

3

Blue Mountains

158,371

324

158,69 5

2

8,618,032

231,637

8,849,668

101

TOTALS

a Currently forested refers to land currently covered with deciduous, coniferous, or

deciduous/coniferous mixed forest in any age or size class and any canopy coverage.

b Additional habitat refers to land normally covered with deciduous, coniferous, or

deciduous/coniferous mixed forest that has recentl y had trees removed thro ugh fire, timber h
or some other event. Th ese areas are managed as forest land; tree s are not permanently rem
c Percentages may not add up to 100% because of roundi ng


COUGAR HUNTING IN W ASHINGTON

Washington reclassified the cougar from predator to game animal in 19
which required hunters to possess a hunting license to take cougar. The coug

season was year-round in western Washington and 244 days in eastern Washin

One cougar was allowed to be taken in most count ies, with no limit in four cou

(Clallam, Jefferson , Grays Harbor and Mason). In 1973, the harvest limit was

-33­

one cougar per hunter per year for the state. In 1987, the Washington State

Legislature passed a bill requiring hunters to have a couga r tag in their possess

before hunting for or killing a cougar. Only those hunters who applied and we

drawn for a permit were authorized to hunt that season (WD FW 1997a). Tabl

summarizes the cougar permits in Washington from 1987- I 996 . Permits availa

hunt cougar gradually increased from 170 in 1987 until 1994 whe n it jumped to

and 892 in 1995. Even though the number of permits were increased five-fold ,

number of cougars removed during the ten year period did not jump significant
even with the use of hounds. Hunting seasons are set to maximize hunting

opportunities without negatively affecting the cougar popul ation (WD FW 1997
However, in order to set the number of " harvestable" cougars, it is essential to

what the cougar population is. There is no indication that the cougar populatio
Washington significantly increased to justify a five-fold increase in the number
cougar permits available in 1994, 1995 and 1996.

-34­

TA BLE 11: Cougar Permits Issued in Washington From 1987 to 1997:

Year

Permits
Available

Permit
Applications

1987

]70

803

]70

60

]988

2 10

1,082

2]0

89

1989

210

1,588

210

85

]990

22 5

1,790

225

107

199]

23 5

2,] 88

235

135

]992

262

2,773

262

] 56

1993

307

3,230

307

121

1994

730

3,855

365

177

]995

892

5,073

446

283

1996

506

3,970

506

112

]997

Permits
Issued

Total permit
Harvest

132

(WD FW 1997a)

In 1970, WDFW began a mandatory reporting of cougar kills and in 19
required the inspection and sealing of cougar pelts. This mandatory reporting

inspection allowed WDFW to monitor the number, sex and locat ion of cougar

From 1987 to ] 996, all permit hunters were required to complete a Cougar Hu

Report . Beginning in the 1998-99 general season, both successful and unsucce

cougar hunters will be required to submit a game harvest report card to the W
(WDFW 1997a) .

-35­

co mposition and median age, and nuisance cougar complaints to assess cougar

populations (WDFW ] 997a). However, manage ment of cougars based only on

submission of harvest reports by hunters and nuisance complaints does not give

accurate assessment of cougar numbers statewide. Population estimates are on

good as the data which is collected and analyzed. Between ]987 and] 996, ret

rates of harv est information from cougar hunters range d from a high of 98% in

to a low of 57% in 1996. Additionally, the number of cougars lethally removed

WDFW personnel and/or landowners due to depredation or public safety, is no

consistently documented and submitted for inclusion for state wide mortality fig

Tribal members, who can still use hounds to hunt cougar, are not required to h

cougar pelts sealed, and submission of accurate harvest level data by tribal mem
regarding cougar is minimal. Cougar mortality from natural causes, disease,
intraspecific conflicts, kitten mortality, road kills and poaching also need to be
included to more accurately assess cougar populations.

In 1994, the hunting season for eastern Washington and western Washi
was January 1- 31, in all 1993 permit areas; October 15 to November 22 with

hound s not allowed; and November 23 to December 31. Hunters were require

obtain a cougar tag prior to the cougar season . The cougar season did not ove
the general deer and elk season .
When the ban on hound hunting went into effect, WDFW significantly

changed and lengthened the hunting season for cougar. In 1997, the cougar pe

-36­

season was eliminated and the general season extended from August I to Marc

except for closure in three game management units. The general season now o
the deer and elk seasons, increasing the opportunity of encountering a cougar.

witho ut the use of hounds , 132 cougars were killed during the 1997 hunting se
(Table 12).

TABLE 12: Cougar Harvest Statistics for 1997 Hunting Season:

CMU

FEMALE
Hunting

MALE
Hunting

T I

T ~

TOTAL

Coastal

5

6

11

Puget Sou nd

7

3

10

North Cascades

7

4

II

South Cascades

8

4

12

14

10

24

Colu mbia Basin

2

4

6

East Cascades S.

3

0

3

22

20

42

Blue Mou ntains

9

4

13

Total
(WD FW 1997b)

77

55

132

East Cascades N .

Northeastern

-37­

ADDITIONAL MORTALITY

Survival rates are commonly used in managem ent pro grams, but are diff

to estimate for long-lived, secretive mammals occurring at low densities (Lindz

al. 1988). In addition to sport hunting, trapping and predator control, injuries a

death occur within populations from intraspecific killings, injuries suffered durin
attempts to capture prey, starvation and road mort ality.

Problem Cougars: Washington State does not normally utilize the U. S

Department of Agriculture's Animal Damage Control (ADC) program for letha

removal of cougars. Removal of cougars for public safety or depredation is do

the WDFW and/or the reporting party . Under Title 77 RCW, Chapter 36, the o

the owner's immediate family member, the owner's documented employee, or a

tenant of real property may trap or kill on that pro perty any cougar damaging p

pro perty. In 1995, \VDFW and private landowners lethally removed 10 couga

1996, 43 cougars were lethally removed ; and in 1997, 47 were lethally remove

Altho ugh the number of cougars lethally remove d appears to have drastically ju

between 1995 and 1997, the figures are misleading. What has changed is the n

of cougars relocated versus cougars lethally removed . In 1995, 14 cougars we

relocated, in 1996 eleven were relocated and only 2 were relocated in 1997. T

increase in lethal removal numbers shows a change in the method for dealing w
problem cougars away from relocation and toward lethal removal.

Illegal Harvest: Poaching also removes cougars from the population, b

-38­

actual number of cougars poached is unknown. Estimates can be made based o

game violations; however, accurate numbers of cougars poached cannot be acc
assesse d (WDFW 1997a). Not know ing the number of cougars illegally taken

popu lation can skew population figures toward higher cougar numbers. Differ

pop ulations may have different poaching pressures, e.g. roaded areas afford po
with an easier opportunity to find cougars (Neal et al. 1987).

Nativ e American Harvest: Tribal harvest of cougars is not well-docum
The state ban on hound hunting does not affect tribal members. WDFW is

continually working with tribes to obt ain accurate harvest information but some

tribes' data is incomplete and/or are not even submitted to WDFW. Non-triba

hunters can still hunt cougar on reservations with a tribal member using his/her

hounds as a guide (Pozzanghera 1998). Cougar populations cannot be proper
managed without accurate harvest data from tribal hunters .

M otor Vehicle Mortality: Deaths from motor vehicles are increasing
nationwide as more roads are built in and around cougar habitat.

From 1979

1991, almost 50% of documented mortality of Florida panthers was due to co

with cars . In California, 22 cougars were killed by collisions between 1971 and

Cougars seemingly do not avoid roaded areas . He mker et al. (19 84), and Barn

et al. (1989) suggested that the vulnerability of cougars to hunters increases wi

increasing road density. As Washington's population increases and road densi
increases, mort ality of cougars due to collisions will also increase .

Natural Causes: Intraspecific mortality, cougars killing other cougars,

-39­

also have a significant impact on a population. Cougars killing cougars have be
documented in studies conducted by Sitton et al. ( 1976), Robinette et al. (1961
Ho rnocker ( 1970). In a study conducted in Sheep River, Alberta, intrasp ecific

killings acco unted for 14% of the cougar mortality from 1981- 1988 (Jalkotzy e

1992) . In addition, cougar mortality can be caused by disease, accidental death

unknown natural causes (non-human relate d mortality) . In Washington, the ma
of all natural deaths of radio-collared cougars were due to intraspecific killings
injuries sustained while capturing prey (WDF W 1997a).

Cub survival: Although it is illegal to kill a female with cubs in Washing

State, the reproductive status of a female may not be obvious. Females may hu
far as 16 km (Hemker et al. 1984) from their cubs and may be gone for several

Cougar kittens seldom survive if orphaned before six months of age and are stil

dependent on their mother until the age of dispersal, which is approximately 1.5

years of age. Regulations that prohibit killing female cougars with kittens have

limited effectiveness: hunters cannot easily recognize these females since young

kittens (up to 6 months of age) rarely accompany their mother. Juvenile couga

accompany their mother to kills more often than cubs, however, they are also ra

seen but are still dependent. In a study on detecting female cougars with kitten
conducted in the Boulder, Escalante, and Canaan Mountains in southern Utah,

Barnh urst et al. (1989) found kittens age 0-6 months were found with their mo
only 19% of the time and juveniles, 7-12 months of age, were fou nd with their

only 43% of the time. With the new expanded cougar season in Washington St

-40­

August through March , it will be very difficult to determine the mortality rate fo

cou gar kittens . Kittens born in the spring and summer will still be dependent up

their mother for survival during most of the season. If they are orphaned prior

dispersal age, survival of the cubs will be greatly reduced . In a study conducted

female reproductive biology in New Mexico, Sweanor et al. ( 1997) stated: "Sp

hunting may adversely affect a cougar population by killing the most productive
females and/or orphaning cubs ."

UNDERSTANDING MYTH AND LEGEND

Rat her than concentrating on myth, legend and unsupported data, the pu

needs to be educated about cougars, their importance in an ecosystem, and how

live with these large carnivores. For the most part , human att itudes have chang

regarding predators; mass persecution has ended, and real efforts to manage an

preserve these splendid animals have been initiated . The grizzly bear and wolf a

protected in the 48 contiguous states and reintrodu ction of wolves to their form

range has begun. The change in attitude tow ard wo lves and grizzly bears has b
the result of increased research , and intensive educational programs directed at

understanding these animals while dispelling myths. This same educat ion needs
conducted for cougars, as well as other predators.

COUGAR ATTAC KS ON HUMANS IN THE UNITED STATES

One of the biggest myths perpetrated on the cougar is that of indiscrimi

killer of livestock and humans . Paul Beier, a scientist dealing with cougars, did

depth examination of historical and present records and documented all att acks

-4 1­

10 l -year period , from January 1, 1890 through December 3 I, 1990 (Beier 199

The work was updated in 1992 (Beier 1992) . Beier searched both scientific an

popular literature, including hunters' magazines and newspapers, for reports of

unprovoked att acks by wild cougars. An "att ack" was defined as an incident in

the cougar bit, clawed , or knocked do wn a human. Mauling s by captive couga
case s in which a person deliberatel y approached or harassed a wild cougar wer
included .
The following table is a summary of bot h fatal and non-fatal attacks on
humans from 1890 through 1997.

-42­

TABLE 13: Cougar and Human Interactions Both Fatal and Non-Fatal
from 1890 to 1997:
Fatal Attacks
Victi m

Cougar

Month
Jun e

Year
1890

State Sex Age
7
CA M

July

1909

CA

MIF

10/A dul

Dec.

1924

WA

M

13

June

1949

VI

M

7

January

197 1

BC

M

12

M

January

1974

1\TM

M

8

July

1976

VI

F

May

1988

VI

September

1989

January

Sex
FC

Age

Shot
Y

Condition
6

Y
M

3

Y
Y

3

12

Y

5

F

3

Y

4

7

M

2

Y

M

9

M

4

Y

MT

M

5

F

1.5

Y

1991

CO

M

18

M

2-3

Y

April

1994

CA*

F

Adult

FC

2-3

Y

Dec.

1994

CA*

F

Adult

M

Adult

Y

July

1997

CO*

M

10

2

\TJ=Vancouver Island, BC; Al.B<Alberta, Cana da; FC=fem aIc with cubs; Condition: l =good
and normal body mass; 2 = mass (measure) was normal for age; 3 = mass (estima ted) was no
for age; 4 = mass (measure) was below normal for age; 5 = mass (estimated) was below norm
age; 6 = rabid.; 7 = cataracts.
* = national park, state park, remote areas or undeveloped recreational area; ** = curre nt
information from WDFW DElS 1997--no other information on cougar listed.

-43­

Non-Fatal from 1890 to 1997:
Non-Fatal Attacks
Month

Year

Victim
State Sex Age

Sept.

1916

BC

M

8


F

11


Sex


Cougar
Age
Shot

2.5

Y

May

1934

VI


M

Adult


y

May

1935

VI


M

Adult


Y

January

1951

VI


M

63


F


July

1951

BC

M

29


FC


March

1953

VI


M

43


April

1953

TX* M

June

1953

VI


F

24


March

1962

ALB M

6


June

1963

BC

M

6


March

1965

BC

M

October

1966 BC

Sept.
June

1.5

2

Adult


Condition

7


Y

4


y


5


Y

2


Y

Y

5


1.5

Y

5


M


Adult

y


5


15


F


Adult

Y

5


M

Adult


M


Adult Y

5


1969 VI


M

13


1970 CO

M

2


F

Y
M


Adult Y

2


VI=Vancouvcr Island, BC; ALB=Alberta, Canada; FC=fcmaIc with cubs; Condition: I=good
and normal body mass; 2 = mass (measure) was normal for age; 3 = mass (estimated) was no
for age; 4 = mass (measure) was below normal for age; 5 = mass (estimated) was below norm
age; 6 = rabid ; 7 = cataracts.
* = national park, state park, remote areas or undeveloped recreational area ; ** = current
information from WDFW DEIS 1997--no other information on cougar listed.

-44­

TABLE 13 (continued): Cougar and Human Interactions Both Fatal and
Non-Fatal from 1890 to 1997:
Non-Fat al Attacks
Vict im
Sex Age

50

Sex
M

M

29

F

M

Adult

F

Co ugar
Age
Shot
Adult Y

Mont h
June

Year

1970

State
BC* F

De cember

1970

BC

July

1971

NY

June

1972

VI*

June

1972

VI*

M

25

N

July

1972

VI

M

8

N

June

1975

BC

M

8

M

2

Y

December

1976

CO

M

14

F

1.5

Y

June

1977

WA

F

28

M

1.7

Y

F

4

2
Adult

Condition

2

Y

3

Y

3

N

November

1978

TX*

February

1979

BC

F

9

F

August

1979

VI

F

4

May

1981

ALB M

1981

VI

M

4

N

Child

5

Y

4

F

Y

5

Child

FC

N

Adult

F

1.5

Y

V1=Vancouver Island, BC; ALB=AIberta, Canada; FC=female with cubs; Condition : l =good
and normal body mass; 2 = mass (measure) was normal for age; 3 = mass (estimated) was no
for age: 4 = mass (measure) was below normal for age; 5 = mass (estimated) was below norm
age: 6 = rabid ; 7 = cataracts.
* = national park, state park, remote areas or undeveloped recreatio nal area; ** = current
information from WDFW DEIS I997--no other information on cougar listed.

-45­

TABLE 13 (continued): Cougar and Human Interactions Both Fatal and
Non-Fatal from 1890 to 1997:
Nonfata l Attacks
Month
Augu st

April

1982

Victim
State Sex Age
ALB ~ M
9

1983

VI

M

Adult

Y

1983

VI

M

10

Y

M

11

M

16

Year

Sex
M

Cougar
Age
Shot
0.5
Y

VI

April

1984

TX* F

Adult

August

1984

TX* M

9

May

1985

VI*

M

12

August

1985

VI

F

10

M

1.5

Y

2

March

1986

CA*

F

5

M

2

Y

1

October

1986

CA*

M

6

April

1987

TX * F

31

M

1.2

Y

4

May

1988

AZ

F

6

F

1.5

Y

4

January

1989

VI

M

28

F

2

Y

4

April

1989

AZ

M

5

M

Y

1

N

1.8

Y

2

1983

M

3

Condition

4

N

N

N

V1=Vancouver Island, BC; ALB=Albena, Canada; FC=female with cubs; Condition: l =good
and norma l body mass; 2 = mass (measure) was normal for age; J = mass (estimated) was no
for age; 4 = mass (measure) was below normal for age; 5 = mass (estima ted) was below norm
age; 6 = rabid ; 7 = cataracts.
* = national park, state park, remote areas or undeveloped recreation al area; ** = current
information from WDFW DEIS 1997--no other information on cougar listed.

-46­

TABLE 13 (continued: Cougar and Human Interactions Both Fatal and
Non-Fatal from 1890 to 1997:

Month
June

Year
1990

Non fatal Attacks
Victim
Cougar
Age
State Sex Age
Sex
Shot
28
CO
F
FC
N

July

1990

MT*

M

9

July

1991

BC

M

2

F

1.5

M

12

1992

CA*

1992

WA** F

1992

WA**

Sept.

1993

CA*

F

August

1994

CA*

M

August

1994

WA* M

March

1995

CA*

1996

WA** M

March

M

1.3

N

Condition

4

N

M

Adult

10

F

1-2

50's

F

2

5

F

1.5

28

F

Adult

N

5
Child

M

Y

Adult

VI=Vancouver Island, BC; Al.H<Alberta, Canada; FC =femal e with cubs; Condition: l =good
and normal body mass ; 2 = mass (measure) was normal for age; 3 = mass (estimated) was no
for age; 4 = mass (measure) was below normal for age ; 5 = mass (estimated) was below norm
age; 6 = rabid; 7 = cataracts.
* = national park., state park., remote areas or undeveloped recreational area ; ** = current
information from WDFW DEIS 1997--no other informati on on cougar listed.

(Beier 1991, 1992; Torres et al. 1996; Busch 1996; WDFW 1997a)

-47­

fatal attacks and 57 non-fatal attacks over the past 107 years (Table 13). The

of deaths att ributed to cougars over the past centu ry is far less than the annual

of people killed by lightning strikes, rattlesnake bites, or bee-sti ngs (Beier 199
Attacks are especially rare when one considers that cougars forego thousands

opportunities to attack humans. In most cases, cougars were merely seen, or a
att ack was prevented by the action of the human.
Table 13 also shows that 74.6% of cougar attacks occurred during the
and summer months by young, inexperienced juvenile cats . This time period

coincides with the time when many young cougars leave their mother' s home r
and set out on their own . Beier (1991) in studying cougar attacks on humans
"The data suggest that yearlings and underweight cougars were most likely to

humans." Juvenile cougars (0-24 months of age) may have difficulty captu ring

prey. The low body mass of most juvenile attackers may be an important facto
attacks on humans (Beier 1991).

Over 63% of cougar attacks are on children . The smaller size of childr

similar to that of the cougar's natural prey . Cats are stimulated to attack by sm

objects moving rapidly across or away from their line of travel (Fitzhugh and F

1997). Children have quicker, more erratic movements compared with adults

make them vulnerable . The vulnerability of children to attacks can be minimiz

keeping them within sight of an adult who may not prevent, but can repulse an
(Beier 199 1).

-48­

Many attacks have occurred in Canada. Fifty-one percent of the attack

children have occ urred in British Columbia, including Vancouver Island, and A

Canada, where cougars are heavily hunted using hounds. Knut Atkinso n ( 1996
carnivore biologist with the British Columbia Wildlife Branch stated:
Many of our attacks, and all of the fatalities, have taken place
on the west coast of the island. This is the area where our
deer populations are the lowest, due to a combination of
poorer habitat, logging, and wolf predation, and where
people and their houses are right against the bush. There is
no buffer zone. . . Occasionally a child is in the wrong place
at the wrong time and we have an attack (83).

Like Vancouver Island, human population growth, destruction of habita
fragmentation, and increased recreational access to isolated areas are the main

reasons for more cougar encounters . Comp laints from the public to WDFW al

have steadily increased as the human population has increased, and they have n

been limited to complaints regardi ng cougars. Complaint s regarding wildlife in

general , have increased, especially for deer and elk damage, opossums, raccoon

bear. In 1995 WDFW received 247 cougar complaints, 495 in 1996 and 563 i
1997. Table 14 shows the number of cougar complaints received in 1996 by t

WDFW (final data on 1997 complaints were not available at the time this thesis

completed). The majority of complaints in 1996 (n=313) were sightings or a ch
encounter with a cougar (WDFW 1997a).

-49­

and the Activity Cou gar Were Engaging in for 1996:

Confi rmed

County

complaints

Encounter
Type(a)

Complaint
Type (b)

SIC

I

A

H LIP

Service
Provided (c)
0

T

L

0

Asotin

6

2

4

0

4

1

1

0

0

3

Benton

4

0

4

0

0

0

4

0

0

0

Chelan

25

18

7

0

19

5

1

0

3

1

Clallam

14

9

5

0

9

3

2

0

1

1

Clark

8

7

1

0

5

2

1

0

0

0

Columbia

4

1

3

0

1

3

0

0

1

2

Cowlitz

9

9

0

0

9

0

0

0

0

3

Douglas

4

4

0

0

3

0

1

0

0

0

21

13

8

0

15

3

3

1

4

2

Garfield

2

1

1

0

1

1

0

0

0

0

Grant

1

1

0

0

1

0

0

0

0

0

Ferry

(a) Encounter type involves three categories: a sighting or chance encounter wi
cougar (SIC), an incident involving direct confrontation between a human and a
cougar (1), and a cougar attacking a human (A) .
(b) Complaint type involves three categories: a human encounter with a cougar (
livesto ck or pet depredation (LIP); or other including cougar entering a croplorc
a nuisance cougar, or an injured cougar (0).
(c) Three categories of service were provided by the WDFW for dealing with
problem cougar in 1996: the offending cougar was trapped and relocated or an
attempt was made to relocate (T) ; the cougar was lethally removed or an attemp
remo val was made either by a Department official or through the issuance of a
removal permit (L); or other, including using repellents , noise makers, hazing,
fencing, referring to a hunter, issuing a preference permit to a landowner, placing
cameras for observation, or the cougar was found sick or dead.

-50­

Enforcement Program and the Activity Cougar
Were Engaging in for 1996:

County

Confirmed
co mplaints

Complaint
Type (b)

Encounter
Type{a)

Service
Provided (c)

SIC

I

A

H LIP

0

T

L

0

Grays
Harbor

8

6

2

0

5

2

1

1

0

0

Island

7

7

0

0

3

. '\

v

4

2

0

0

Jefferson

17

13

4

0

11

4

2

1

0

0

King

56

33

23

0

3 1 21

4

12

3

1

Kitsap

2

2

0

0

1

0

1

0

0

0

Kittitas

12

7

5

0

7

3

0

2 2

Klickitat

2

0

2

0

0

2

0

I

0 0

12

6

6

0

10

2

0

1

I

Lincoln

1

1

0

0

1

0

0

0

0 0

Mas on

5

5

0

0

4

0

1

0

0 0

26

9

17

0

7 18

1

0

13 3

Lewis

Okanogan

2

0

(a) Encounter type involves three categories: a sighting or chance encounter w
cougar (SIC), an incident involving direct confrontation between a human and
cougar (I), and a cougar attacking a human (A) .
(b) Complaint type involves three categories: a human encounter with a cougar
livestock or pet depredation (LIP); or other including cougar entering a croplor
a nuisance cougar, or an injured cougar (0).
(c) Three categories of service were provided by the WDFW for dealing with
problem cougar in 1996: the offending cougar was trapped and relocated or an
attempt was made to relocate (T) ; the cougar was lethally removed or an att em
removal was made either by a Department official or through the issuance of a
removal permit (L) ; or other, including using repelle nts, noise makers, hazing,
fencing, referring to a hunter, issuing a preference penni t to a landowner, placi
cameras for observation, or the cougar was found sick or dea d.

-51­

TABLE 14 (continued): Cougar Complaints Reported by WDFW
Enforcement Program and the Activity Cougar
Were Engaging in for 1996:

County

Continn ed
complaints

Complaint
Type (b)

Encounter
Typera)

Service
Provi ded (c)

0

T

L

0

0

2

1

1

0

16

12

3

2

4

0

0

16

14

8

5

7

0

8

0

10

6

0

0

0

1

31

11

0

21

11 10

4

2

0

33

29

4

0

29

4

0

2

0

0

Stevens

32

22

10

0

16

9

7

1

8

0

Thurston

17

9

8

0

11

5

1

2

0

0

Wahkiakum

3

2

1

0

2

1

0

0

0

0

Walla Walla

4

2

2

0

3

0

1

0

2

1

H LIP

SIC

I

A

3

3

0

0

1

Pend Oreille

31

13

18

0

Pierce

38

23

15

Skagit

16

8

Snohomish

42

Spokane

Pacific

(a) Encounter type involves three categories: a sighting or chance encounter w
cougar (SIC), an incident involving direct confrontation between a human and
cougar (1), and a cougar attacking a human (A).
(b) Complaint type involves three categories: a human encounter with a cougar
livestock or pet depredation (LIP); or other including cougar entering a croplo
a nuisance cougar, or an injured cougar (0).
(c) Three categories of service were provided by the WDFW for dealing with
problem cougar in 1996: the offending cougar was trapped and relocated or an
attempt was made to relocate (T) ; the cougar was lethally removed or an attem
removal was made either by a Department official or through the issuance of a
removal permit (L); or other, including using repellents , noise makers, hazing,
fencing, referring to a hunter , issuing a preference permit to a landowner, placi
cameras for observation, or the cougar was found sick or dead.

-52­

TABLE 14 (conti nued): Cougar Complaints Reported by WDFW
Enforcement Prozram and the Activity Couzar
Were Engaging ~n for 1996:
Confirmed
complaints

Co unty

Encounter
TypeCa)

Complaint
Type (b)

SIC

1

A

Service
Provided (c)

H LIP

0

T

L

0

Whatcom

19

11

8

0

11

-4

4

2

1

0

Whitman

2

2

0

0

2

0

0

0

0

0

Yakima

,)

1

2

0

1

1

1

0

0

0

Oly Nat'l
Park

1

0

0

1

0

0

1

0

0

1

Unknown

1

1

0

0

0

0

0

0

0

0

,.,

TOT AL
495
313 18 1 1
290 100 68
38 54 20

(a) Encou nter type involves three categories: a sighting or chance encounter w
co ugar (SIC), an incident involving direct confrontation between a human and
cougar (I), and a cougar attacking a human (A).
(b) Compl aint type involves three categories: a human encounter with a couga
livestock or pet depredation (LIP); or other including cougar entering a croplo
a nuisance cougar, or an injured cougar (0).
(c) Three categories of service were provided by the WDFW for dealing with
problem cougar in 1996: the offending cougar was trapped and relocated or an
attempt was made to relocate (T); the cougar was lethally removed or an att em
removal was made either by a Department official or through the issuance of a
removal permit (L); or other, including using repellents , noise makers, hazing,
fencing, referring to a hunter, issuing a preference permit to a landowner, plac
cameras for observation, or the cougar was found sick or dead.
(WDFW 1997a)

Over 62% of the complaints received in 1996 were "sightings or chanc

encounter" with a cougar. Reports of cougar sightings have been heavily criti

as indices of cougar population trends (Beier et al. 1996; Van Dyke et al. 1986

-53­

Van Dyke et al. (1987) wrote:
Indiscreet solicitation of lion sightings by management
agencies is an inefficient, inappropriate, and unreliable method
of determining lion status . . . sightings alone should never
be used for describing cougar distribution and abundance . .
[sightings] should never serve as a basis for describing the
distribution or abundance of mountain lions.
Coug ar reports by the public are notoriously unreliable. California

investigations have found that bobcats, large house cats, coyotes, deer, raccoon

domestic dogs, especially yellow Labrador retrievers, are often reported as cou
sightings (PDI 1997) . In addition , field studies have shown that experienced

campers and hunters cannot reliably identify cougar tracks. Only 19% of west

deer hunters and 20% of campers, with an average of 21 years of experience, c

distinguish a drawing of a cougar track (Appendices 3 and 4) from among othe

of tracks (PDI 1997). Finally, auditory reports ascribed to cougars have been

to be noises made by bobcats, domestic cats, and several species of owl (Yo un
Goldman 1946).

HUMAN POPU LATION TRENDS

Starting in the early 1990s, there has been a surge in rural population gr
in Washington due to people's desire to have a simpler lifestyle and less stress

urban crime and congestion. This urban flight fueled a demand for more house

built in previously undeveloped areas. Due to the continued conversion of wild

-54­

species are no longer available.

Table 15 identifies the ten Washington cou nties with the highest number

cougar comp laints. A review of the growt h in population and where that growt

occu rring in these ten counties shows each has experienced rapid population gro
over the past seven years .

TABLE 15: Counties with Highest Number of Couga r Complai nts and
Populati on Change:
COUNTY

CONFIRMED
COMPLAINTS

POPULATION NUMBER % POPULATIO
INCREASE INCREASE IN
% CHANGE
1990-1997
UNlNCORPOR
AREAS
25.63
2,285
87.6

Pend Oreille

31

Whatcom

19

22.24

28,420

46 .5

Stevens

32

20.85

6,452

82.5

Chelan

25

19.04

9,9 50

52.1

Snohomish

42

18.38

85,572

18.7

Ferry

21

15.97

1,005

90.0

Okanogan

26

15.14

5,050

71.6

Pierce

38

15.03

88,097

**

Spokane

33

13.44

48,567

69.2

King

56

9.21

138,895

**

**Population increase was only in incorporated areas because of annexations an
incorporations of cities and towns. (Office of Financial Management-- 1997
Population Trends)

-55­

Total population numbers in the counties in eastern Washington, except Spoka

County, first appear to be relatively small. However, the percent of population
change shows these cou nties are also experiencing rapid growt h. What is also

significant, is the population growth in the eastern Washington cou nties of Pen

Oreille, Stevens, Okanogan, Chelan, and Ferry was in unincorp orated areas (Ta
15).

In the western Washington cou nties, Whatcom County had 46.5% of t
growth in unincorporated areas . Figures from Pierce and King Counties show
reduction in population in the unincorporated areas and Snohomish County's
unincorporated growth appears low at 18.7% . However, from 1990 to 1997,
those counties increased the incorporated areas (cities and towns) through an

increase in annexations and incorporations. Their population increases are still
significant.

Counties with good opportunities for recreat ion and retirement living a
the fastest growing counties (OFM 1997). With an increase in Washington's
population, also comes an increase in recreational use of state and federal

wildlands where development is limited. With increased access to historic coug

habitat, there is going to be an increase in cougar-human interactions. All of th
top ten counties with the highest cougar encounters have large tracts of

undeveloped land designated as state or federal forests, wilderness areas and pa

used for recreation. The statistics on cougar encounters do not designate whet
the cougar encounter was in a residential area or a wilderness area . This

-56­

information is important to obtai n in order to properly evaluate the number of
cougars which are entering residential areas and creating a public safety issue.

EXPLORING CALIFORNIA'S COUGAR HISTORY

When looking at the impacts associated with the ban on hound hunting an

cou gar-human interactions, a review of California's cou gar management is useful
since they have had a complete hunting ban on cougars for over 25 years.

In California, cougars were bountied from 1907 to 1963 and state records
indicated that 12,46 1 were killed during that 57-year peri od (Torres et al. 1996).
During 1963- 1968, cougars were managed as a non-game and non-protected
mammal; take was not regulated and no state records were kept. In 1972,

Californians passed a moratorium on cougar hunting. In 1986, co ugars were aga
classified as game mammals . The California Department of Fish and Game
recommended regulated cougar hunts , but were challenged in court . In 1990 ,
another ballot initiative, Proposition 117, designated the cougar as a "specially
protected mammal " . Cougar hunting has not occurred in California since 1972 .

California has continued to experience increased human population, habit

destruction, and fragmentation, as well as an increase in cou gar populations . As

Beier stated: "Five years after shooting every cougar seen, of course there' s been

an increase in numbers" (qtd. in Lyons 1996). Over the past 24 years, up to 199
the human population in California has increased to 32 million people and record
indicate annual housing development reached a high of 163,000 new, single-unit
homes in 1989 (Torres et al. 1996). Population centers have expanded, and

-57­

peop le now live in many previously undeveloped areas . This has caused an
increase in encroachment into wildlife habitat. Torres' study concluded that
observed conflicts between cougars and humans vary regionally for different

reasons. Regional differences include diversity of cougar habitat, prey availabil
and human impacts . Management go als fo r cougars in California include: ( 1)
maintaining viable populations of cougars; (2) minimizing conflicts related to
public safety, property damage, and other wildlife; (3) protecting important
habitats; (4) recognizing their ecological role and value; (5) monitoring

popu lations and conducting research; and (6) improving public awareness (Tor
et al. 1996).
California still removes cougars under their cougar depre dation policy.
Between 1990 and 1996, they removed on average 88 cougars per year for
livestock depredation or public safety. Even with a total hunting ban on couga

the California cougar population, estimated between 4,000 to 6,000, is conside
stable and there is no detectable change in population over the last five years
(Torres 1997).

WASHINGTON'S NEW COUGA R SEASON

Shortly after Initiative 655 (page 1) was passed, media reports on coug

sightings or encounters stressed the fact that the increase was due to the ban on
the use of hounds to hunt cougars. Media reports placed the blame for the

increase in cougar encounters squarely on the shoulders of the public for bannin

the use of hounds . The public was told that without the use of hounds, huntin

-58­

cougar wou ld be almost impossible, causing overpopulation of cougars. In an

article in the Yakima Herald Republic, on Oregon's debate on whether to reins
hound hunting, it stat ed:

, , , unless hunters are given more of a sporting chance, the state

[Oregon] is headed for a deadly confrontation like the one in wh
a California jogger was killed by a cougar in 1994 (Hall 1997).
On Augu st 22, 1998, The News Tribune in Tacoma, Washington, gave

quote from the Thurston County Sheriff, Gary Edwards: "If you spot a cougar

or near your property, shoot it. I'm not going to wait until they eat some kid to
shoot them ." Sheriff Edwards went on to state he believed the passage of

Initiative 655, passed in 1996, had an effect on the cougar population becaus e o

the lack of hunting pressure. Even though the indiscriminate kiIIing of cougars

illegal, the sheriff still advocated this "shoot on sight" policy (Hucks 1998), Th

type of media hype fuels the public's fear and puts forth inaccurate information
is clear from a review of California 's management history, cougar biology and

research on hunted and non-hunted cougar popul ations, that the informatio n be
disseminated is inaccurate.

With Washington's new liberal hunting season, it is very doubtful the ba

on hound hunting will be a significant factor in cougar-human interactions. Rat
the real problems are increasing population, encroachment into cougar habitat,
habitat loss and fragmentation, and people's intolerance of wildlife. These are

areas which need to be addressed in order to decrease cougar-human interactio

-59­

One common belief conce rning cougar-human interactions is that, in ord

to prevent or eliminate the se interactions, you must hunt the cougars "to keep t
numbers down"

Hunting does not necessarily red uce cougar-human interactio

British Columbia accounts for 48%, of all recorded attacks from 1890 to 1997
(Ta ble 13). The hot spot has been on Vancouver Island where cougars have

accounted for 30% of all recorded attacks by animals despite having been hunte
relentlessly.

Maurice Hornocker, the founder of the Hornocker Wildlife Institute and
for the past three decades, the dean of cougar research states: "Hunting could

solve the problem by annihilating lions completely . We need instead to educate

people on how to live with lions." He points to human population growth as th

real numbers problem: "All you need to do is fly to Los Angeles and see all the

new subdivisions on the finger ridges. Those ridges are wildlife corridors, criti

grounds for juvenile cougars to establish themselves and try new behaviors whe
they leave their mothers. And this is where humans and cougars will most
frequently encounter each other" (qtd . in Lyons 1996). Beier puts it more

succinctly: "You must accept some risk by living in these shared areas . I accep

that risk. I'm irritated that people who move in want to sanitize the West" (qtd
Lyons 1996).

COUGAR BEHAVIOR DURIN G ENCOUNTERS WITH HUMA NS

AND APPR OPRIATE HUMAN RESPONSES


The possibility of being attacked by a cougar is still very rare, and defini

-60­

not a "major public safety conc ern" if numbers are an appropriate measure

(Fitzhugh and Fjelline 1997). Even so, the small chance of being eaten by a la
carnivore causes fear and anxiety in many people .
To help allay these concerns, it is useful to know that cats, including

cougars, are intimidated by anything unusual that is larger, especially taller, tha
them, and by things that approach rapidly . Positions above a cat are positions
dominance, and those below are positions of subordinance.

Some basic principles of cougar behavior and attacks are : (1) most cou

attacks on humans seem to have been predatory in nature; (2) cat s in general a

threatened and intimidated by large, strange, objects app roaching rapidly and f

above; (3) cats are stimulated to attack by smaller object s moving rapidly acro

away from their line of travel (Fitzhugh and Fjelline 1997). Fitzhugh and Fjel

(1997) developed the following summary (Table 16) from the works of
Leyhausen (1979), Beier (1991), Bogue and Ferrari (19 74), Bogue (n.d.), and

From hold (n.d .), and from discussions with experienced cougar biologists. Th

table summarizes cougar behavior and recommends appropriate human respon

-6 1­

TABLE 16: A Summary of Cougar Predatory Behavior and Suggested
Associated Human Responses:
Ap propriate
Response

Cougar activity

Meaning

Human Risk

Cougar far away and
movmg away

Secretive and
avoidance

Insignificant

Keep children where
they can be obs erved

Slight, provided
human response
is appropriate

Avoid rapid moveme
run ning, loud, excited
Stay in groups; keep
children with adults.
Observe cougar. For
agenci es, this may in
future problems if re

Slight for adults
given proper
response. Serious
for unaccompanied
children

Hold small childre n;
older children close t
adult. Do not tu m b
cougar; assume stan d
position on ground, r
or large equ ipment th
are above the cougar
possible. Look for s
rocks or other weapo
pick them up, using a
aggressive posture w
doing so. Watch cou
all times. However,
couga r sits, looks aw
grooms itself, this is n
predatory situation, a
you should imitate th
cougar, but keep it in
peripheral vision. Fo
agencies, consider w
visitor s and limiting h
to gro ups.

Indifference
Co ugar > 100 yards
away various positions
and movements,
att ention directed away
from people.

Cougar > 50 yards
Curiosity
away ; various body
positions; ears up ; may
be changing positions ;
intent attention toward
people; following
behav ior

-62­

TABLE 16 (continued): A Summary of Cougar Predatory Behavior and
Suggested Associated Human Responses:
Cougar activity

Meaning

Human Risk

Appropri ate
Response

Cougar < 50 yards
away; intense staring
at humans; hiding

Assessing
success of
atta ck

Substantial

All of above steps, p
Place older children
adults. If a safer loc
or one above the co
available, go there.
run. Raise hands an
objects such as jack
above head so as to
image of bulk as hig
poss ible. Prepare to
your self.

Intense staring and
hiding coupled with
crouching and/or
creeping towards
humans

Moving to
attack
position

Serious if within
200 yards

Take all the above a
If possible, move slo
place large objects s
trees, boulders betw
yourself and the cou
but do not lose sigh
cougar. Smile! (Sho
teeth). Make menac
sounds; throw thing
cougar is close enou
hit.

Crouching; tail
twitchi ng; intense
staring at humans;
ears erect; body low
to ground; head may
be up

Pre-attack;
awaiting
opportunity

Grave

Do all of the above
whatever weap ons y
If you have lethal w
take careful aim and
them now . Pepper
may be effective if c
is close enough and
downwind. If you h
rocks or other item
can be thrown, do s

-63­

TA BLE 16 (continued): A Summary of Cougar Predatory Behavior and
Sugg ested Associated Human Responses:
Couga r activity

Mea ning

Imminent
Ears turned so the
"fur" side is forw ard;
Tail twitc hing; body
and head low to
ground; rear legs
may be " pumping" or
"treading" gently up
and down.

Human Risk

Extreme attack;
cougar ready to
leap

Appropriate
Response

Prep are to defend yo
close combat. Fight
Make menacing nois
attack may happen w
seconds. If you hav
chance of averting it
acting aggressivel y t
the cougar. If the di
too great to use a sti
rapidly toward the c
until you can put the
its face and eyes. If
a stick, run toward t
cougar with arms hig
making loud noises.
before you are withi
distance of its paws .
movements towards
cougar, especially fr
it, may still deter an
Avoid positions belo
cougar; do not tum
back on it.

(Fitzhugh and Fjelline 1997) .

CONCLUSIONS AND RECOMMENDATIO NS

Humans all over the world have lived in the presence of large, dangerou

since our time began . We, too, must learn to live with cougars. Cougars have

lived closer to humans than many humans realize. Because of their solitary, se
crepu scular and nocturnal nature , they are rarely seen, but are still present.

-64­

Cougars represent a link between wildland values and civilization and

should be considered a treasured natural and national heritage. Living with the

cougar should be considered an honor and a privilege, but one must understand

risks and accept the anxieties that living among large predators generate (Benso
199 1).

RESEARCH AND MANAGEMENT: Cougar hunting will continue

hunters can be successful without the use of hounds . However, it is imperativ
that additional research be conducted on the population status of cougars in
Washington State. Cougar populations need to be evaluated on a regional

population basis, their management should be biologically motivated and direct
to ensure their survival and viability, while minimizing conflicts with humans.

Therefore, WDFW should adopt a regional approach toward wildlife conservat
that consid ers the differing status of each cougar populatio n and the feasibility
potential management activities .

It is also mandatory that accurate and corroborated data be collected on
cougar and human interactions. As we have seen from previous published
literature, public reports on cougar sightings and encounters are notoriously

inaccurate. Inflated numbers of phone calls to an agency about cougar sighting
encounters, without verification , only leads to an atmosphere offear and

uncertainty. Additionally, accurate information needs to be compiled dealing w
the age, sex and physical conditions of the offending cougar. An effort should
made to collar and relocate cougars, if they have not attacked people . This is

-65­

particularly important since many of the cougars that find their way into fringe

areas are newly disper sed juveniles who are exploring new areas and looking fo
suitable habitat to establish their territories. Important data can be gained by

monitoring these relocated cougars. How success ful is relocating thes e cougar

When do they establish their own territ ory. Do they return to fringe and human
development areas. A dead cougar reveals very little data .

This author is concerned about the liberalization of the hunting seaso n f

cougars without adequate data to justify a hunting season from August I throu

March 15. There is a high probability of mortality of cubs that are not yet able

survive on their own when a female is killed. Tho se kittens born in spring, sum

and fall would probably not survive on their own if orphaned within 9 months o

birth. In addition to liberalizing the season , the cost of a cougar tag is now onl
$5, a very good incentive for deer and elk hunters to obt ain a cougar permit.

Rather than lowe ring the fee, the fee should be increased and the money put int
cougar research . When asked about the lengthening of the cougar season and

potential impacts on cougar populations, Steve Pozzanghera, WDFW, indicate
was a compromise with the legislature . "WDFW is being pressured from
legislators to make the cougar season year round" (Pozzanghera 1998). This

wou ld be like going back to the old bounty days and it woul d be very difficult t

monitor and protect cougar populations. Protection and management of wildli

especially cougars, should be based on facts and accurate data, not political wh

Because it is up to WDFW to provide the best scientific data available o

-66­

cougar populations, the agency must have sufficient funding to conduct adequat
research on cougars on a regional basis to ensure that populations are not

adversely affected by the liberalization of the hunting season . It may take severa

years to determine the impact on a population if a significant number of breedin

females and litte rs are lost. Additionally , accurate data on tribal harvest must be
provided to WDFW to adequately evaluate and manage the population A
cooperative management agreement between tribes and the WDFW should be
secured to ensure data is gathered, compiled and submitted in a timely manner.
Funding and manpower must be increased to more comprehensively

research the cougars ' complex biology . These studies need to be for periods of
least five years in duration to gain adequate data. Research needs to address
habitat issues, prey availability, age, sex and condition of depredating cougars,
population dynamics.

As mentioned above, reports by the public on cougar sightings, encounte

lost pets, or tracks are poor methods for evaluating cougar populations (Van Dy

et al. 1987). Additionally, other states indicate that field staff inconsistently and

incompletely fill out report forms used to document problem activities. Trends
proble m activities from the 1980s to 1997, which are based on the old WDFW

Form 57, are not accurate (WDFW 1997a). Although it appears there are more

cougar encounters, without adequate and corroborated data, the speculation tha

the cougar population is increasing and cougar-human interactions are increasing

due to the ban on hound hunting, is erroneous and deceptive. It is a much more

-67­

comp lex and varied issue and the public needs all the information in order to lea
to live with cougars. WDF W needs to accurately record and evaluate the data

ensure that unsubstantiated complaints do not cloud the real issues of habitat lo
human encroac hment and expanding human population. A standard reporting
policy is needed . This should not necessarily include all reported sightings, but
certainly should include field-verified accounts of interactions between humans
cougars. Field case and laboratory necropsy reports should also be included in
each incident report (Aune ]99]).

HABITAT PROTEC TION:

Since degradation of their habitat is one

the biggest threats to the cougar, their ecological needs must be factored into a
proposed developments in formerly wild back country, and rural areas . This is
especially true for residential developments, such as subdivisions and town

expansions, recreational development such as golf courses and tou rist lodges, a

road building intended to provide access for residential, recreational and indust r
activities (Hummel et al. ] 99] ). Attention must be placed on direct impacts on

cougars themselves, as well as the primary prey species of cougars, deer and elk

As Hu mmel et at. ( 1991), states: "When will we finally understand that the stat u

of top predators, such as cougars, indicates how well we are doing in protecting
the entire wildlife system. . . ."
Habitat acquisition, enhancement, restoration and protection are

fundamental to cougar survival, as well as to their prey base. The following is A

Action Plan for Cougar Management and Preservation developed by the Couga

-68­

Foundation and Kevin Hansen (1992) and addresses very important factors. T
tenets would be applicable for cougar management in Washington State:

I. Protect large, contiguous tracts of cougar habitat as wildlife preserv

These must be large enough to support healthy cougar populations.

2. Provi de funding to acquire habitat and to enhance and resto re habita
including special appropriations, federal funds, and bond acts.

3. Prevent construction ofbarriers--roads, canals, reserv oirs, cropland
residential developments--that separate cougar populations from

portions of their habitat. In those areas where cou gar habitat is mo
fragmented, place land acquisition emphasis on habitat corridors.

4. In areas where there is a decreasing prey base, pro mote the increase

ungu late numbers as part of an overall biodiversity protection progr

5. Develop incentives to protect private land from development, includ

acquisitions, and swaps , transfer of development rights, agreements
easements. Establishing conservation easements and cooperative

wildlife protection projects on private wildlands saves tax dollars an

can be innovative state/private cooperative ventures. There are ma
creative programs that provide incentives to land managers for
voluntary habitat protection efforts .

6. Restore degraded habitats , and, where appro priate, create new habi
For example, many eastern states now have large tracts ofland no
longer suitable for farming that may be able to support cougar

-69­

populations.
7. Wildlife corridors, connecting habitat is essential. Linda Sweanor, a
cougar researcher in New Mexico stat es: "It is apparent that
fragmented habitat may only support lions on a long-term basis if
individuals are allowed to successfully immigrate, hence requiring
dispersal from other local populations. To con serve populations of
cougars over the long term, adequate habitats must be maintained in
effective patchwork composed of relatively large blocks of wildland
reserves interconnected by dispersal corri dors." (76).

These recommendations will also help numerous other species of wildli
since habitat destruction and degradation affects all wildlife.

EDUCATION: One of the most important first steps is public educatio

particularly among the younger generation, in order to preserve cougars for futu
generations. It is essential that people understand carnivores and how they are

indispensable component of a healthy environment. Coexistence with cougars i

possible, but it requires changing our attitude toward the wild animals that share
our landscape.

Living in cougar country poses some risk, but it is a manageable one . Pa

of Florida's success in bringing the alligator back from the brink of extinction w
an aggressive public education campaign about the importance of making room
wildlife that some considered dangerous (Hansen 1992).
Education programs do work. Colorado also is experiencing increased
-70­

population and cougar-human interactions, especially in the Front Range where
human population is intruding into historic cougar habitat (Seidel 1998) . The
State of Colorado Division of Wildlife conducted a number of public
information/education programs, developed a brochure entitled "Living with
Wildlife in Lion County" and developed a standardized reporting system .
Colorado saw a 30% reduction in cougar-human encounters after the first year
its education campaign (Homocker 1996). The broc hure points out that "with
better understanding of cougars and their habitat, we can coexist with these

magnificent animals." The brochure gives a brief overview of cougars: physica

appearance, tracks, habitat, hunting and feeding habits, mating and breeding, bir
to maturity and recommendations for living in cougar country. Brochures on

living with cougars have also been developed for Montana, "Living With Monta

Mountain Lions ; The Predator Defense Institute developed "Living with Couga
for Oregon and the Olympic National Park puts out an informational sheet on

traveling in cougar country. WDFW has just completed their first informationa

brochure dealing with cougars which is now available. It is hoped this will be th
start of an aggressive education program in Washington. This brochure should
provided to cities, counties, parks, realtor s, recreationists, natural resource

agencies and any others who may have the privilege of coexisting with our large
feline predator.

It does not have to be an "either them or us" mentality. There will alwa

be cougar and human encounters, and some cougars will need to be removed fro

-71­

areas where they have caused problems. Many of the cougars that wander into

urban areas are young, transient cougars or old cougars who are looking for eas

prey. If there is no food for them to eat, they will normally leave the area. Pets

are easy prey and, even when not in cougar habitat, they should be under contro
all times. The public needs to be aware that pets which roam are also prey for
coyotes, eagles and great homed owls; poiso ning and motor vehicles collisions
also take a toll. Not all pet disappearances are due to cougars.

It is hoped that this thesis will be a catalyst for accepting and embracing

cougars as the majestic feline predator which personifies strength, movement,

grace, stealth, independence, and the wilderness spirit. The cougar is resilient--

it is vulnerable, e.g. the Florida panther, the Eastern panther , Yuma cougar, Co

Rican cougar. The public needs accurate and non-biased information on cougar
in orde r to coexist with these magnificent creatures .

-72­

BffiLIOGRAPHY

Ackerman, Bruce B., Frederick G. Lindzey, and Thomas P. Hemker, 1986.
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Ackerman, Bruce B., F. G. Lindzey, and T. P. Hemker, 1984. "Coug ar Food H
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Anderson, Allen E., 1991. "Frequency of Mountain Lion Sightings by Resident
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---, 1983. " A Critical Review of Literature on Puma (Felis concolor) . Spec . Re
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Aune, Keith E. , 1991. "Increasing Mountain Lion Populations and Human­
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Beier, Paul, and Stanley C. Cunningham, 1996. "Power of Track Surveys to
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Classifi cat io n of t he Felidae
by W. Chnsropner Wozencraft ( 1993)
Family Feli liae G . Fischer. 1817
Subfami ly AClnonych lnae Po co c k . 19 17
Aa non yx Brookes. 1828
jutJatus Scnreoe r. 1776

Chee ta h

Su bfa m ily Felin ae Fi sch er. 1817
Caracal
cara ca! (S:::nreoe r. 1776)

Cam ca l

Cacopuma s evertz ov, 1858

osau (Gra. 1874)

tem mincx . (Vigors and Horsheld . 1827)


Bomean oay cat

As18UC gOlaen cat


Felis

Chinese rnouruari rc esern ca t
Jungle cat
Sana cat
Biack-toote n cat
W ildcat 01 At nca an d Eurasia

oreti Mllne-::'awaras . 1892

cnaus Scnreoe r. 1T77

m argama i.ocne . 1858

mgnoes Burcnell. 1824

suvesms S:::nreoe r. 1775

Herpa llUllJS

Ja guarundi

yagwuo ncI Lace peae. 1809
Leopardus Gray. 1842

pardallS n.m naeus . 1758)

/lgnnus tScnreoer . 1775)

wledi (Scnmz. 1821 )


Ocelot


Orienta . L ittle tiger cat

Margay


Lepcal/urus severtzov. 1858

serve! (Scnreoer. 1776)


Serval

Lynx Kerr 1792


Canaa a rvnx

EurasIan rvnx

lbe nan tynx


csn eaensis Kerr . 1792

lynx (Lmnaeus. 1758)

paramus iTemmlnck. 1824)

tutus (Scnreoer. 1776)


Bobcat

Oncite/is ssvertzcv . 1858
co /ocata (Molina. 1782)
geoffroy' Id'Orblgny and Gervais . 1844 )
gUlgna (MOlina. 1782)

Pampas cat

Geo1fToys cat

Kodkod


Oreailurus Cabrera . 1940

jacoblfUS IC omaha . 1865)


Ande an mountain cat

Ouxxnoous Brar.::::: 1842


Pallas s ca t

manu! (Pa llas. 1776 )


Proruuiutus SevertZov . 1858
oenasiens 's (Kerr. 1792)
ommceos I VICOrs and HorslJe ld . 1827 i

ruounnosi: s II. GeotTroy Saint-Hilair e 1831 I
vivemnus l o en nen . 1833)

-88­

Leooara ca t

Ftat-neaoeo cal

Rusrv-sooneo cat

r:lsnlng cal


Atncan golden cat


aurata (Temmmck . 1827)
Puma uarcme. 1834 .

concotor (Lmnaeus 1771)

Puma. Cougar. or Mountain bon


SUbtam ll y Panthen nae Po coc k 1917

I
I
\

\

Neofelis Gra y. 1867

netnuoss (Griffith , 162 1)

Clouded leopard


Panrnera Oxe n, 1816 ,

leo (Lmnaeus. 1758)
onca (unnaeus 1758 )
oerau: ILlnnae:.: c 1758 )
uqns (t.mnaeu s. 1758)

Uon

Jaguar

t.eoca rr
TIge r

Pardo tetis Severtzo v . 1858

marmorata Ma nln . 1837

MartJled cat


Uncia Gra v 18~

uricte (Scnteo e: 1758 )

Sn ow reooar d


\

wozencratt. w,e, 1993 Ore e r Carnrvora Po , 25 0·346 In D.E Wilson an d O M , Aeeoe r. ecs Mam
of tne world. a ts xonomic ana aeoc rtux»: reference tS econo ecuuorn . Srrutnsoruan Institution Pres
D.C , and London

1 iecotms. wieo». and tettutnncsu In Wozencra~ (1 993 1amended to iecotmus vaet». and temmmc
with the 1965 Intern ational Cooe of Z oological Nornenctature Arucie 31a mandating that patronym
toll ow th e nnes 01 latin gr ammar.

N ote: Brackets round tne nam e of the autncnrv inc ica te mat me genus nas oeen cnanged since fi
pu blica tion by that autnority.


-89­

APPEN DlX 2: COU GA R MANAGEMENT UNITS (CMUs) IN WASHINGTON.
ARE BASED ON ECOREGIONS DEVELOPED BY THE GAP
PRO G RAM AND GROUPINGS OF WASHIN G TO N DEPARTM
OF FISH AND W ILDLIFE GA ME MA AG EM ENT UNITS (GM
(W DFW 1997 a )

APPENDIX 3: COUGAR TRACKS
(Hansen 1992 )

CO U GA R TRACKS
(nee

(0

scale)

­

\\ 'ALKI t' C CAIT

HEEL WI

'R... rr.-Il. ~ )

I ~ ."

ron al ..

.



....'....

C _ ",

~

,... ...... ,...

n uux

75·

i"",

I::

17.

SrraoJJ :..

~ ~H

fRO J'l

-

~

:,TRlnE

T RA C ~

. 1>

L EFT RE

~R

TRAL r.

-91­

,

DOG AND COYOTE (pandelll992; Murie 1974)

Mounuln L..ion

lifr Jar '" rrnuI