j

MOUNTAIN LION COMPENDIUMAND

AN EVALUATION OF MOUNTAIN LION MANAGEMENT IN WYOMING

Prepared by

GREGORY D. JOHNSONM. DALE STRICKLAND

WESTERN ECOSYSTEMS TECHNOLOGY, INC.1406 SOUTH GREELEY HIGHWAY

CHEYENNE, WY 82007

- FEBRUARY 24, 1992

AN EVALUATION OF MOUNTAIN LION MORTALITY QUOTAS

Mountain lion management is receiving considerable attention in the western United

States, primarily as a result of the California lion situation, where the courts determined the

state did not have adequate information to manage lions and closed the season. The

Wyoming Game and Fish Department's mountain lion management program is based on a

literature review completed over ten years ago (Strickland 1980). As a result of this review,

mortality quotas used to control harvest were set for each management unit in the state.

However, quotas have changed over the years without a concomitant analysis of the new

literature and analysis of harvest and wildlife observation system data. The objective of this

study was to review the current literature and data available on mountain lions in Wyoming

and use this review to evaluate Wyoming's current management strategy, and to make

suggestions regarding any necessary changes in the Department's management program.

METHODS

A review of the mountain lion literature was conducted to obtain all available relevant

information on mountain lion research and management (Appendix I). . An annotated

bibliography of relevant literature on mountain lion research and management developed

during the literature review is presented in Appendix II. Relevant literature was obtained

by searching with the computerized reference services maintained by the University of

Wyoming, reviev.nng selected recent journals, and by contacting wildlife management

agencies and scientists involved in mountain lion research and/or management. Mountain

M%~~__~""""",,,_~,. __.._. . ._. .. . ,,"

-2-

lion managers/researchers were also contacted for their personal opinion on lion

management.

The original approach used by the Wyoming Game and Fish to estimate lion

abundance and distribution was evaluated in light of contemporary literature and views of

current lion researchers. Mountain lion harvest statistics and observations from the

Department's Wildlife Observation System were compiled and analyzed !o estimate relative

abundance and distribution of mountain lions within each management unit. A revised index

of lion abundance and a recommended mortality quota was produced based on the literature

review and analysis of the available data. The index was based on published lion densities

in high, medium, and low density habitat. The original habitat classification system based

on Kuchler potential vegetation types used by Strickland (1980) was used as the basis for

identifying lion density strata.

Density estimates of Wyoming mountain lion populations include t~e study conducted

by Logan (1983) in the Bighorn Mountains, where density was estimated at 3.4-4.5/100 km2,

and the ongoing study in North Yellowstone (Murphy and Felzien 1991) where density is

estimated at 3.6Ikm2. These densities were selected to represent high density habitat.

Because these are winter densities, when lions are restricted to ungulate winter range, they

are higher than yearlong densities, although the extent to which they are higher cannot be

determined. Therefore, the density selected to represent yearlong high density habitat was

SUbjectively set at 2.5;100 km2. No estimates of density in medium or low density lion. .habitats are availabJefrom Wyoming. Based on mountain lion studies conducted in Utah

-3-

in areas considered low and high density, Utah stratified the state into several lion density

strata (Bates 1989). Medium and low lion densities in Utah were selected to represent

medium and low density in Wyoming. The medium density was set at 1.2/100 km2 and the

low density was set at 0.4/100 km2. Density and distribution data were used to estimate lion

abundance by management unit. Mortality quotas were estimated for each management unit

based on a conservative level of 10%.

The index for lion abundance was calculated using the following formula:

MAI-L (Ah*Dh ) + (Am*Dm) + (A1*D1)

where:

MAl = Total Management Area Index (estimated number ofadult lions in each management area)A = area of the density stratumD = density of lionsh = high density stratumm = medium density stratumI = low density stratum

Mortality quotas were calculated using the following formula:

MAMQ-MAI*Hr

where: MAMQ = Management Area Mortality QuotaH, = selected harvest rate

-4-

RESULTS

The proportion of each mountain lion management unit composed of high, medium

and low density mountain lion habitat, estimated population size, and recommended

mortality quota is presented in Table 1. The estimated statewide lion population estimate

was 1696.

RECOMMENDATIONS

Harvest

Regulating harvest through use of a mortality quota system appears to be a

reasonable method of controlling harvest and regulating distribution of the harvest. Lion

harvest in the state should continue to be regulaied through use of the mortality quota

system. The quota should initially be set based on the estimated population size for each

area. We determined quotas for all areas using a conservative 10% level. However, quotas

can vary with each popUlation, depending on population objectives. The population

estimates are very subjective, and are sensitive to density estimates selected and

stratification. Therefore, population estimates should be verified and harvest should remain

conservative until verifications are complete. We elected to stratify the state into lion

density categories using Kuchler vegetation types. The mountain lion abundance map

developed by the Game and Fish using wildlife observation system data appears to have

several inconsistencies, the most notable being the classification of west-central Wyoming as

low density lion habitat, when in fact a relatively large number of observations have occurred

in this area. To compare the two stratifications, we will estimate lion abundance in the state

-5-

based on the Wyoming Game and Fish abundance as soon as areas are determined.

If the objective is to maintain lion numbers, than the quota should be set at no more

than 10% of the estimated population size. Where lionl1ivestock or lionlhuman conflicts

mandate that the lion population be reduced, quotas of 25-30% or even higher may be

required. Several studies have shown that mortality levels even this high may not initially

reduce populations if large unhunted or lightly hunted lion populations occur in adjacent

areas. Therefore, lion management should be at the population level but recognize the

importance of coordinating management strategies among adjacent populations. It may be

necessary to establish a management strategy regionally to meet individual population

objectives.

Where concern for a population exists, implementing a female mortality subquota

should be considered as a viable means of protecting the population while still allowing some

recreational harvest. When female subquotas are adopted, we recommend setting the

quotas at 3Q% of the total quota. The season should be terminated if the female quota is~A::::::-~::~-';:."'"

reached, even if the total quota has not been met.

In general, harvesting of females should be discouraged to maintain stable

populations and reduce the number of kittens orphaned. Treed mountain lions can generally

be sexed by experienced observers. In many cases, sex may also be determined based on

size of tracks. A brochure describing methods to determine sex of treed lions as well as

tracks should be developed and distributed to all mountain lion outfitters, guides and license

holders. The brochure should provide information on the percentage of females which may

-6-

have kittens during the hunting season and the percentage of the time kittens are not with

their mother. This information should help discourage harvest of female lions.

Monitoring Populations

Effects of harvest on the population should be carefully monitored using age data of

harvested lions, and adjustments to the quota should be made as required. Lightly exploited

lion populations will generally contain more resident adults and should have older age

composition than heavily exploited lion populations. Heavily exploited populations will

contain more transient, younger lions. To obtain age data, mandatory check of harvested

lions should be continued. Explicit aging techniques are not available, but lions can be

placed into broad age categories. Criteria for aging lions are described in detail in Ashman

et a1. (1983). All personnel responsible for examining lion carcasses should be trained in

aging techniques to ensure age data obtained from harvested lions are standardized.

Potentially valuable trend information may also be obtained through

interviewing/surveying 100% of lion license holders and lion guides/outfitters: Information

to be obtained should include number of lions treed, estimated sex and age of lions treed,

number of days spent hunting, location of lions and lion sign observed, method of hunting,

and hunting conditions. Because guides and outfitters spend considerable amounts of time

in pursuit of cougars each year, they should be considered a valuable source of information

on perceived lion population trends in their respective areas. To increase accuracy of data

co]lected from guides and hunters, a survey form could be provided prior to the hunting

-7-

season so that the information could be collected during the hunt, rather than retrieved from

memory at a later date.

A method should be developed to periodically census mountain lions in areas of

special interest or concern. The method described by Van Sickle and Lindzey (1991) using

probability sampling of tracks in snow located with helicopters appears to have the most

utility for Wyoming. Some measure of trend should be considered statewide every 3 to 5

years. Roadside track counts or track counts using snowmobiles could be considered as an

indicator of trend. Ideally, the methodes) chosen should be tested on a population of known

size in Wyoming. This would require conducting an intensive mark-recapture study with

telemetry in conjunction with the census/trend technique(s) chosen. A mail questionnaire

similar to the one conducted by Berg (1981) should be considered to update information on'

status, distribution, and numbers of mountain lions in the state.

Controlling Depredation

Most studies have indicated that largescale reductions of lion populations do not

significantly reduce depredation problems. Although damage complaints and subsequent

harvest of lions in the south end of the Big Horn Mountains were reduced following heavy

harvest of lions in 1987 and 1988, other factors may have also been partly responsible for

the reduced damage and harvest. Data on number of sheep, distribution of sheep, deer

populations, and hunting conditions should be examined to determine what role these factors

may have played. Even if significant reductions of lions are found to reduce damage in the

-8-

Bighorns, largescale reductions of lion populations may become politically infeasible in the

not-to-distant future. Therefore, lion depredation problems should probably be handled by

targeting the offending individual.

-9-

LITERATURE CITED

Ashman, D., G. C. Christensen, M. C. Hess, G. K. Tsukamoto, and M. S. Wichersham.1983. The mountain lion in Nevada. Nev. Dep. Wildl., Reno. 75pp.

Bates, B. 1989. Status of cougar in Utah, 1988. Pp. 32-34 in R.H. Smith, ed. Trans. 3rdMountain lion workshop, Arizona Chapter, The Wildlife Society, and Arizona Gameand Fish Dept.

Berg, R. L. 1981. A mail survey for information on the distribution of mountain lions in theState of Wyoming. M.S. Thesis. Univ. Wyo., Laramie. 94pp.

Logan, K. A. 1983. Mountain lion population and habitat characteristics in the Big HornMountains of Wyoming. M.s. Thesis, Univ. Wyo., Laramie. 101pp.

Murphy, K. M., and G. S. Felzien. 1991. The ecology of the mountain lion (Felis cancolormissouJensis) in the northern Yellowstone Ecosystem. Cumulative Progress Rep!.No.4. Wildlife Research Institute, Moscow, Idaho. 17pp.

Strickland, D. 1980. Mountain lion populations. Wyoming Game Fish Dep. Unpubl. Rep.,Cheyenne. 5pp.

Van Sickle, W. D., and F. G. Lindzey. 1991. Evaluation of a cougar population estimatorbased on probability sampling. J. Wildl. Manage. 55:738-743.

Table 1- Proportion of high, medium, and low density mountain lion habitat, estimatedpopulation size, recommended mortality quota, and current mortality quota by management unit.

Estimated Recommended CurrentHanagement % of unit in each density strata population Mortality MortalityUnit High Medium Low Size Quota Quota

1 0 0 100 20 2 12 0 0 100 18 2 03 0 0 100 38 4 04 0 5 95 43 4 15 0 49 51 39 4 46 0 35 65 97 10 167 24 46 30 105 10 68 5 28 67 54 5 89 0 44 56 51 5 210 47 0 53 17 2 211 12 28 60 91 9 512 0 0 100 13 1 213 14 43 43 62 6 214 15 45 40 50 5 215 a 15 8 76 159 16 2816 0 7 93 47 5 217 35 41 24 388 39 218 26 55 19 46 5 419 17 59 25 157 16 620 8 28 64 47 5 621 78 22 0 47 5 822 22 22 56 107 11 15

Total 1696 171 122

a includes current units 15 and ~3

APPENDIX 1. MOUNTAIN LION RESEARCH AND MANAGEMENTA LITERATURE REVIEW

HABITAT USE AND PREFERENCE

An area's suitability as mountain lion habitat depends primarily on a combination of

three factors: vegetation type and structure, topography, and abundance and vulnerability

of prey (Seidensticker et a1. 1973). Several mountain lion researchers have qualitatively

described habitats in their respective study areas. Only two studies, one in Wyoming (Logan

1983) and one in Utah (Laing 1988) have quantitatively exami~ed lion habitat use. The

following section summarizes descriptions of mountain lion habitat and habitat use from

Wyoming and surrounding states. Because of sample size considerations, most mountain lion

studies have been conducted in areas considered to have relatively high mountain lion

densities. Habitats in these study areas should therefore be considered at least above

average to good mountain lion habitat in the respective states.

Ver1etation

Mountain lion habitat in the Canon City, Colorado area studied by Currier et a1.

(1977) consisted of pine/Douglas fir in the upper elevations and pinyon-juniper woodlands

in the lower elevations. The two habitat types in this area that most mountain lions were

captured in were pinyon-juniper-Douglas Fir and pinyon-juniper-ponderosa pine (Currier

]976). Principal vegetation in the western Montana area studied by Murphy (1983) was the

Douglas fir climax series. The ponderosa pine/bluebunch wheatgrass type dominated the

most xeric south exposures at lower elevations, while scree habitat types were common on

steep slopes of southerly and westerly orientation. Douglas fir/ninebark, Douglas

-2-

fir/bluebunch wheatgrass and Douglas fir/beargrass habitat types covered most of the

remaining winter range slopes. Vegetation in the Idaho Primitive Area was composed

primarily of ponderosa pine and Douglas fir, with Engleman spruce and subalpine fir

dominating the higher elevations (Hornocker 1970).

Mountain lion habitat in the Boulder-Escalante area of south-central Utah, where the

mountain lion population has been studied extensively for several years, consists of desert

grass and shrub communities, sparse pinyon pine, and juniper at the lower elevations, dense

pinyon-juniper with a sagebrush understory, ponderosa pine and oakbrush at mid elevations,

and subalpine meadows, spruce, aspen, and white fir at the higher elevations (Van Sickle

1990). Laing (1988) described mountain lion habitat use in the Boulder-Escalante area

based on 3580 locations of 52 radiomarked mountain lions. Mountain lions were found to

select pinyon-juniper woodlands with lava boulders, ponderosa pine/oakbrush, mixed

aspen/spruce-fir, and spruce-fir habitats. These areas offer advantages to cougars by

providing cover for resting, stalking prey, and feeding on kills (Lindzey et a1. 1989). Cougars

avoided sagebrush bottomlands, agricultural and pasture lands, pinyon-juniper/ponderosa

pine/sagebrush mixtures, slickrock sandstone canyons, and open meadows in this area,

presumably due to lack of sufficient cover. Discriminant function analysis indicated that

overstory cover, overstory height, and slope were the most important structural

characteristics in highly used habitats, ",ith mountain lion use increasing with each of these

parameters.

-3-

In the Bighorn Mountains of Wyoming, mountain lion habitat use was determined

based on 191 track locations, 118 radiolocations, 52 cache sites and 84 scratch sites (Logan

1983, Logan and Irwin 1985). Use and availability of habitats were found to be significantly

different. Mixed conifer and curlleaf mountain mahogany were preferred; Douglas fir,

juniper/sagebrush, lodgepole pine, and riparian zones were used in proportion to their

availability; and sagebrush-grass was avoided. Also in Wyoming, a survey of Yellowstone

Natiomil Park for presence of mountain lions during winter revealed 13 sets of tracks as well

as several kills, scats, and scrapes, all of which occurred in Douglas fir forest associations

(Koehler and Hornocker 1986).

Berg et aJ. (1983) obtained 2,574 observations of mountain lions or lion sign in

Wyoming from a mail questionnaire distributed to trappers and mountain lion hunters,

outfitters, biologists and foresters, and ranchers and farmers, and plotted the locations on

maps of Kuchler potential vegetation types. The five habitat types with the highest

mountain lion use, based on density of sightings for each type, were Douglas fir forest,

Eastern ponderosa forest, pine-Douglas fir forest, black pine forest, and wheatgrass­

needlegrass shrubsteppe. Twenty-three percent of all observations occurred in the Douglas

fir forest type. The Wyoming Game and Fish Department's wildlife observation system data

base has 614 observations of mountain lion sightings, sign or harvest where habitat was

recorded. Most observations (14.5%) occurred in sagebrush-grassland. Other habitat types

where mountain lion observations commonly occurred were ponderosa pine (10.4%),

lodgepole pine (8.8%), ponderosa pine/Douglas fir (3.7%), riparian shrub-shrub steppe

-4-

(3.7%), limber pine (3.4%), pine-juniper (3.3%), mountain mahogany (2.9%), and juniper

(2.8%).

Topographv

In Colorado, Currier et al. (1977) described the study area near Canon City as being

mountainous and the terrain rough. This appears to be typical topography of mountain lion

habitat. The Idaho Primitive Area is characterized by high peaks and ridges dissected by

deep, narrow valleys, and has the greatest relief of any area in Idaho (Hornocker 1970).

Terrain in the area studied in Montana by Murphy (1983) was steep, with most slopes being

>50%. In the Bighorn Mountains of Wyoming, mountain lions preferred steep and rugged

topography (50-100% slope). \10derate (20-40%) slopes were used as available, and gentle

« 20%) slopes were avoided (Logan 1983). The author concluded that, in the Bighorn

Mountains, canyonlands contained the most important mountain lion habitat features. Of

124 mountain lion kills found in the Bighorn Mountains of Wyoming where topographic

information was recorded, 81% were in rimrock or canyon type topography (Norelius and

Bruscino 1988). Observations in Yellowstone National Park were also associated with

canyons and rugged terrain (Koehler and Hornocker 1986). Although topography is

considered important, presence of timber is thought to be more important than topographic

cover in Oregon (Harcombe 1976).

-5-

Deer Abundance

Mountain lion habitat in the West is essentially that of mule deer (Dixon 1982), and

the most consistent habitat feature of mountain lion habitats studied in the West is the

presence of mule deer (Russell 1978). In Arizona, a total of 144 track count routes

representing 11 habitat complexes were surveyed by Shaw et a1. (1988). Regression of

mountain lion tracks per route and mule deer density was significant (r=0.765, P<0.027).

Because the number of mountain lion tracks went to 0 at a mule deer density of 2.3/mi2, the

authors suggested that this is the minimum deer density capable of supporting mountain

lions. Hemker (1982) found that density of resident mountain lions in Utah was also limited

by abundance of mule deer.

Other factors also influence mountain lion abundance and should be considered when

evaluating an area's suitability as mountain lion habitat. In Utah, mountain lions avoid

logged areas, establish territories in areas "ith less roads than average, and avoid human

residences and other areas of high human activity (Van Dyke et a1. 1986).

In summary, mountain lion habitat in Wyoming is probably concurrent 'With that of

mule deer. Within the range of mule deer, mountain lion densities 'Will be highest in areas

dominated by ponderosa pine, Douglas Fir, pinyon pine, juniper, mountain mahogany and

other areas of open woodland. The more rugged the topography "ithin these habitats, the

higher the mountain lion density will be. Areas lacking timber (e.g. sagebrush communities,

grasslands, agricultural lands), areas with low topographic diversity, areas with low mule deer

-6-

densities, and areas where the number of human dwellings or other human activity is high

will have relatively low mountain lion densities.

MOUNTAIN LION POPULATION CHARACTERISTICS

Age and Sex Structure

Mountain lion management decisions are ideally based on a knowledge of sex and age

composition of populations as well as population size and trend (Barnhurst 1986).

According to Anderson (1983) sex ratios of adult mountain lions in the literature he

reviewed never differed significantly from 50:50. The age structure of mountain lion

populations has been determi l1ed for some lion populations, however, most age data should

not be considered highly accurate, as there is no reliable method currently available for aging

wild mountain lions (Phelps 1989).

In Wyoming, a mountain lion population in the Bighorn Mountains contained up to

50% juveniles « 1 yr) (Logan 1983). The proportion of the population in each of 6 age

classes during the 2 years of study was as follows:

Age0-12 months12-23 months2 years3-4 years5-6 years7-9 years

1981-820.120.000.080.310.190.00

1982-830.500.090.090.170.120.03

-7-

The male:female sex ratio in the Bighorns was 39:61 for 28 juveniles and 45:55 for 22 adults,

neither of which differed significantly from 50:50. There were 2 to 3 times more resident

females than resident males in this area. Also in Wyoming, the sex and age composition of

34 lions in North Yellowstone was: 10 adult females, 6 adult males, 2 adults of unknovm sex,

and 16 kittens (Murphy and Felzien 1991).

Ross and Jalkotzy (1988) determined the following age structure for an intensively

studied lion population in Alberta:

Resident IndependentAdults Subadults Dependent Juveniles

Year male female male female unk male female unk

1984 4-6 8 0 3 0 0-1 6 01985 4 9-10 2-3 7 0 2 4 21986 4 10 2 4 0 2 3 5-71988 5 12 0 2 0 9 4 3-5

In New Mexico, mountain lions less than 1.5 years of age comprised 30.2-51.6% of the

population, depending on year, and adults (> 2 years) comprised 48.4-69.8% (Logan et a1.

1990). The average age of adult female mountain lions in the New Mexico population,

which was not hunted, was 3.6-4.7 years, whereas average age of adult males was 5.1-5.9

years. In the Idaho Primitive Area, the mountain lion population was composed of 50%

adult females, 20% adult males, and 30% young of the year (Hornocker 197]). In Alberta,

over a four year period, resident adults comprised 43-61 % of the population and juveniles

comprised 22-49% (Jalkotzy ]989). Over a 9-year period in Utah, resident females

comprised an average of 32.9% of the population, resident males comprised 8.8%, juveniles

-8-

comprised 42.4%, and transients comprised the remaining 15.9% (Calculated from Lindzey

et al. 1989). In New Mexico, of 33 lions captured for study, 45.5% were adult female,

15.2% were adult male, and 39.4% were less than one year old (Donaldson 1975). Of 57

cougars captured in southwest Colorado, 15.8% were adult (>24 months) males, 22.8% were

adult females, 29.8% were juvenile «24 months) males, and 31.6% were juvenile females

(calculated from Anderson 1988). The captured population was not completely

representative of the actual population, as only tracks of puma estimated to weigh> 27.4

kg were followed. The population composition of lions calculated using results from four

intensive cougar studies conducted in North America was 49% resident female, 22% resident

male, 8% transient female, 8% transient male, 7% female kitten (13-18 months), and 7%

male kitten (Lindzey et al. 1989).

Fecundity.

The age of first reproduction was determined to be 29·48 months in Alberta (Alberta

Forestry, Lands and Wildlife 1991), 3·4 years in the Big Horn Mountains of Wyoming

(Logan 1983, Logan et al. 1986), 23-36 months in North Yellowstone (Murphy and Felzien

1991),20-34 months in Utah (Lindzey et al. 1989), and 32 months in Nevada (Ashman et

al. 1983). Eighty-two percent of females were greater than 2 years old when they produced

their first litter in New Mexico (Logan et al. 1990). In the Bighorn Mountains of Wyoming,

55% of the adult females had kittens and 44% of them gave birth the first study year. The

second year, 86% of the females gave birth to kittens (Logan 1983). In New Mexico, the

-9-

proportion of females producing litters each year ranged from 27-47% (Logan et al. 1990).

Mean interbirth intervals for successful females were determined to be 19 months in Utah

(Lindzey et al. 1989), 16.7 months in New Mexico (Logan et al. 1990) 17.4 months in Nevada

(Ashman et al. 1983), and approximately once every 2 years in Montana (Murphy 1983).

The average number of kittens per family group was 2.7 in the Bighorn Mountains of

Wyoming (Logan 1983) as well as in North Yellowstone (Murphy and Felzien 1991). In

New Mexico, eight female mountain lions produced 10 litters over a 39 month period, for

a total of 29 kittens (Logan et al. 1989). In a Montana lion population, the mean litter size

was 2.6 (Murphy 1983). Average litter size in Utah from a sample size of 20 was 2.4

(Lindzey et al. 1989). In the Idaho Primitive Area, mean litter size ~as 2.5 (Hornocker

1971). Mean litter size of a mountain lion population studied in British Columbia during a

4-year period was 3.14 (Spreadbury 1989). Ashman et al. (1983) present some of the only

litter size data by age:

Prenatal 3.084 months 2.595-11 months 2.3112 months 2.23

The only study where annual recruitment was determined was conducted in Alberta.

Recruitment to independence was 1.0-1.3 kittens/female/year (Alberta Forestry, Lands and

Wildlife 1991).

-10-

Survival

Several studies have examined mountain lion survival. Lindzey et al. (1988) measured

survival rates of an unhunted mountain lion population in Utah from 1980-86 using

radiomarked individuals. Each mountain lion used in survival calculations was monitored

for an average length of 821 days. Mean annual survival of adult females estimated with the

program Micromort (Heisy and Fuller 1985) was 71%, whereas a product-limit analysis

estimated 73% survival. For adult females and males combined, mean survival was 72 and

73%, respectively for the 2 estimators. In southwestern Colorado, 20 male and 21 female

mountain lions ranging in age from 2-84 months were monitored from 1981-1988 (Anderson

et al. .1989). Fifty percent had died by the age of 65.9 months. Annual survival was

estimated at 88.1%. Although the study area was closed to hunting, 11 marked mountain

lions were killed during the study, three in the study area and the rest adjacent to the study

area. In an unhunted mountain lion population in New Mexico, a total of 33 mountain lions

deaths or disappearances were documented during a 5-year study. The most prevalent cause

of mortality for adults and subadults was intraspecific strife, whereas for juveniles the most

important mortality factor was cannibalism. Mortality of adult females was 3 times that of

adult males. (Logan et al. 1990).

Hemker et al. (1986) calculated cub survival of ten mountain lion family groups in

an unhunted population in Utah. Survival from 3-10 months of age (n=lO) was 72% and

from 10 months to age at dispersal (16-19 months) was 92% (n=13). Overall survival from

3 months to dispersal was 67%. From data collected by Robinette in Utah, Hemker et al.

-11-

(1986) calculated a survival rate of 82% from birth to age 10 months. Of 66 mountain lion

progeny monitored in an unhunted population in New Mexico, 22 died during the study, for

a survival rate of 66% (Legan et aJ. 1990).

Density

Density has been determined for several mountain lion populations in the western

United States and Canada (Table 1). Techniques used to estimate density include track

counts from ground and air, mark-recapture, intensive hunting, and telemetry. Densities

estimated using all techniques range from < 1/100 km2 in areas of California, Idaho, Texas,

and Utah to 7.1/100 km2 in Montana and 9.2/100 km2 in California (Table 1). Most

mountain lion density estimates are probably underestimates (F.G. Lindzey, Univ. Wyo.,

Pers. comm.). According to Kenny Legan (Wildlife Research Institute, Moscow, ID, pers.

comm.), the only reliable density estimates are those calculated using data gathered during

intensive mark-recapture studies in combination with telemetry: According to Legan, these

include the study conducted in the Bighorn Mountains of Wyoming, where. density was

estimated at 3.4-4.5/100 km2 (Legan 1983), the work by Hornocker (1970) in the Idaho

Primitive Area, where density was 2.9/100 km2, a study in Alberta, where density was

estimated at 2.7-4.1/100 km2 (Pall et al. 1988), and studies currently being conducted in New

Mexico (Legan et aJ. 1990), where density is estimated at 1.7-3.3/100 km 2 and Yellowstone

National Park, where density is estimated at 3.6/100 kn/ (Murphy and Felzien 1991). The

main problem with other density estimation techniques is that they have never been

-12-

validated on a known density population. Until this is done, their usefulness is e:X1remely

limited for anything other than an index to population abundance (K. Logan, Pers. comm.).

ESTIMATING POPULATION SIZE

Due to their relatively low density, secretive nature, and solitary lifestyle, mountain

lions are not easily censused. Only 20% of 15 mountain lion "experts" surveyed by the

Mountain Lion Preservation Foundation (1987) believed that it is possible to develop an

accurate mountain lion population estimate. Forty-five percent thought it possible in

localized areas only. Fifteen percent responded that it is not possible at all to accurately

determine a mountain lion population size. When this same group was asked if they thought

statewide mountain lion population estimates were of any value, 47% said only when

determining trends, 20% said yes, and 26% responded no. The experts thought that local

population estimates are valuable, but stressed that these should not be compared to other

areas or e:X1rapolated to an entire state. Most management agencies rely on indirect

measures of abundance and trends including harvest, livestock damage complaints, number

of mountain lions killed for damage, and numbers killed by vehicles on roads. Harley Shaw

(General Wildlife Services, Chino Valley, Arizona, pers. comm,) believes that management

agencies will eventually be forced to conduct some kind of field survey to estimate mountain

lion abundance to justify and defend their management programs to the public. This has

already happened in California.

-13-

Techniques currently employed by mountain lion researchers and some management

agencies to estimate mountain lion abundance include roadside track counts, aerial transect

surveys for tracks in snow, mark-recapture techniques using toe-clipped or radiomarked

mountain lions, and attempting to mark all mountain lions in a given area. FoUowing his

review of literature on mountain lions, Anderson (1983) concluded that there is no best

method of estimating mountain lion numbers, and that at least two census techniques should

be used to estimate mountain lion populations. Of the mountain lion experts surveyed by

the Mountain Lion Preservation Foundation (1987), 53% thought capture-recapture \vith

radiotelemetry was the most accurate population estimator, 33% suggested using a

combination of radiotelemetry and track counts, and 7% suggested use of long-term harvest

data. Barnhurst (1986) attempted to correct harvest age and sex data with a vulnerability

index based on frequency of road crossing. The results were not accurate, however, as it

could not account for hunter selectivity. The foJ]o",~ng section describes techniques currently

used by researchers to estimate mountain lion abundance and trend.

Track Counts

Several attempts have been made to estimate mountain lion abundance by conducting

roadside track surveys. Use of mountain lion track counts is the only method currently

available that doesn't require expensive mark-recapture or telemetry for direct estimates of

mountain lion abundance and trend (Shaw 1990). Van Sickle (1990) examined the use of

roadside track surveys .to estimate mountain lion abundance in south-central Utah. There

-14-

was a positive relationship between mountain lion density and the number of track sets

found per total km of road searched, and track counts were found to be reasonable

estimators of the relative number of mountain lion home ranges crossed. Track counts were

not considered by the author to be a valid technique for detecting changes in populations,

however.

Mountain lion abundance has also been estimated in Colorado using track counts

(Currier et al. 1977). In three years of searching, the mean length of road searched per set

of tracks found ranged from 9 to 12.7 km. Population density based on results of the track

counts ranged from one mountain lion/30-56 km2 to one lion/30-60 km2. A population

estimate using marked mountain lions and the Lincoln-Peterson estimator indicated a density

of 1 lion/13-55 km2 and a population estimate of 95 mountain lions in the study area.

Van Dyke et al. (1986) examined the use of track counts as indices of mountain lion

presence in Utah. Under "good dirt" tracking conditions, they noted a direct relationship

between track finding frequency and mountain lion density, and concluded that track counts

can be used to estimate mountain lion relative abundance. Resident females required the

least effort to detect. Searching 360 km of road for every 500 km2 was considered the

maximum effort required to find the track of any mountain lion remaining in the area when

searches are conducted, however, under ideal tracking conditions, < 90 km would need to

be searched.

Based on his study of track counts to estimate mountain lion abundance in Arizona,

Shaw (1980) concluded that to get the required minimum sample size of 40 (based on

-15-

Burnham et a1. 1980), 1200 miles of road would have to be searched to track densities. The

highest frequency of track locations was 0.08 tracks/mile of road driven. Based on this

success, this approach was considered unrealistic, as 60 man-days would be required to

obtain the required sample size. In another Arizona study, Shaw et a1. (1988) ran 144 track

counts representing 11 habitat types. Walking routes of 10 kID were considered feasible

sampling units and were found to be superior to vehicle or horseback searches. The authors

conceded that this technique woufd be infeasible on an annual basis due to time constraints,

but recommended periodic surveys every 3-5 years for continued monitoring of mountain

lion populations. Sitton (1977) also recommended use of track counts every 5 years to

monitor trends.

Smallwood and Fitzhugh (1987) described a mountain lion track survey used to index

mountain lion populations in the state of California in 1985 and 1986. The entire state was

divided into 31 mi2 survey blocks. Each block had three 7-mile routes separated by 10 miles.

Survey blocks were randomly se](~cted for sampling, and routes were surveyed by volunteer

biOlogists. The results were calibrated by one experienced tracker. All routes were

conducted in one day by 2 searchers from a vehicle going 3-5 mph. Volunteers found tracks

in 21 of 48 blocks surveyed, whereas the calibrator found lion tracks in 21 of 41 blocks

surveyed. Based on survey results, a sample size of 44 blocks was considered sufficient to

detect significant differences benveen years. Fitzhugh and Smallwood (1989) described

another technique for monitoring mountain lions on a statewide basis. They suggested

stratifying the state into "bioregions" based on climate, vegetation, and topography. In each

-16-

stratum, a "representative" area of several hundred square miles should be selected and

censused using track counts. According to the authors, a statewide (California) mountain

lion track count designed to provide only an index to the statewide population could be done

for approximately $1,000 plus 82 man-days of expert tracking labor. Based on their work

in California, Kutilek et a1. (1980) recommended that track surveys be repeated on the same

transects over several days to increase the chance of detecting lions. Transects of 40 miles

were considered to be of sufficient length.

Track counts have also been used to estimate lion abundance in Oregon, where it

took 2.7 days of searching per track found (Harcombe 1976). A total of 15 mountain lions

were located during 41 days of tracking. Based on this result, the author believed track

counts could be used to look at yearly differences, and recommended that lion track counts

be conducted in conjunction with other activities (e.g. big game classifications).

Based on a workshop held in Arizona, preliminary recommendations were made for

conducting track counts in that state (Shaw 1990). Recommendations included using

experienced people to search standard 5-mile long routes. The track counts could be

conducted from foot, horseback or vehicle, however, foot routes were preferred. Additional

recommendations included maintaining a minimum of 10 miles between routes, stratifying

on vegetation rather than management unit, and surveying routes at the same time each

year. The group attending the workshop believed conducting the survey on alternate years

would probably be sufficient to monitor trends.

-17-

Smith et al. (1989) described a technique to monitor mountain lion populations in

roadless areas by recording occurrence of tracks, scats, scrapes and prey remains during

biannual surveys conducted on a watershed basis. The authors are currently evaluating the

value of this method for monitoring large-scale changes in populations.

Mark - Recapture

This technique is considered the most accurate for determining population size. Van

Sickle (1990) toe clipped several individuals in Utah and used a mark-recapture study design

based on the number of marked and unmarked tracks located using road surveys. This

method esti~ated a population size of 22.5 (SE 6.6) for a known population of 23. Based

on the effort involved in marking mountain lions, however, the author concluded that this

technique was beyond the scope of most managers. Several investigators have estimated

mountain lion abundance and several other parameters using radiomarked individuals

(Currier et al. 1977, Shaw 1980, Logan 1983, Murphy 1983, Hemker et al. 1984, Laing 1988,

Van Sickle 1990). This method is generally considered too expensive and labor~intensive to

be within the scope of most mountain lion management plans. Utah, however, is proposing

mark-recapture surveys on 3 units a year to assist with determining lion trends (Bates 1989).

Aerial Track Survevs

A technique that appears to have the most utility for estimating mountain lion

numbers where snow cover commonly occurs involves line-intercept probability sampling

-18-

using aerially located mountain lion tracks. This method was tested in Utah and was

considered to be a viable means of estimating cougar numbers (Van Sickle 1990, Van Sickle

and Lindzey 1991). In brief, the technique involves searching transects with a helicopter

after sufficient (8-12 em) snowfall. Recommendations included selecting transects using a

replicated systematic sample design with a minimum distance between transects of 4 km.

Transects should be oriented perpendicular to the majority of water drainages. Surveys

should be conducted within 2 days of a snowfall event (one day if it is windy). The

technique performs best in areas of high mountain lion density because variance is reduced.

The primary drawback to this technique is the logistical problem of having helicopters

immediately available when the right snow conditions occur for conducting the surVey (EG.

Lindzey, pers. comm.).

Mail Questionnaires

A mail questionnaire sent to trappers, mountain lion hunters, outfitters, biologists,

foresters, and ranchers and farmers was used to determine distribution and relative

abundance of mountain lions in the state of Wyoming (Berg 1981, Berg et a1. 1983). A total

of 2,574 locations of mountain lion sign and sightings were reported by 999 individuals. Van

Dyke and Brocke (1987) attempted to determine reliability of mountain lion sightings by

campers, deer hunters, and mountain lion hunters. They found that most campers and

hunters could not identify a mountain lion track shown to them or describe diagnostic

features of a mountain lion track. Based on this information, the authors concluded that

~1II1!I!!......... , ----

-19-

solicitation of mountain lion sightings from inexperienced observers by management agencies

is an inefficient and unreliable method of determining mountain lion status.

Modelling

To successfully model a population, data on population size, age structure, sex ratio,

birth rate, death rate and age at first reproduction are required (Trulio 1989). A few

attempts have been made to model mountain lions, the most notable being that of Sherriff

(1978). Sherriffs model, called LIONPOP, is a modification of ONEPOP. California has

also attempted to model lion populations (see Barrett 1986). According to Hopkins (1984),

the usefulness of mountain lion models is currently limited by th~ Jack of data required for

a model to perform.

MOUNTAIN LION HARVEST

Effects of Harvest

In Montana, Murphy (1983) reported an annual average harvest of 22% of the adult

females and 48% of the adult males available. According to the author, at these rates, all

adult females would have been removed in 4.5 years and all males in 12.1 years in the

absence of recruitment. Due to recruitment bv transient lions, however, sex ratios and

numbers of mountain lions in the study area remained stable during the 3-year study despite

an average annual harvest of approximately 32% of the adult population. In the Canon City,

Colorado mountain lion population studied by Currier et aI. (1977), 6 male and 3 female

-20-

mountain lions were harvested the second year of the study, and 12 males and 6 females

were harvested the third year. The mountain lion population was estimated during the third

year only. Based on their population estimate of 95 mountain lions, then 19% of the

population was harvested the third year. The authors determined that at least 187 mountain

lions were harvested in the study area during the 10 years prior to the study, but believed

that this harvest did not jeopardize the population. Assuming the mountain lion population

was stable for the 10 years preceding the study, then an average of 19.7% of the population

was harvested annually during this time period.

In Arizona, Shaw (1980) estimated that no more than 40 adult or transient mountain

lions were present on his study area at the start of a 3-year study. By the end of the second

year, this was reduced to 15, and at the end of the third year, 16-18 mountain lions were

present. A total of 28 mountain lions were known to have died during this period, 21 of

which were harvested and two of which were killed by research activities. The remaining

mortality was considered natural. The author concluded that this level of Iiarvest, combined

with natural loss, reduced the mountain lion population over the 3-year period. Assuming

the author's population estimates were accurate, then mortality due to hunting was

approximately 35% the first year of the study, 27% the second year, and 17-19% the third

year.

Total mortality of a mountain lion population in the Bighorn Mountains of Wyoming

was 27% the first year of a 2-year study (Logan 1983). The natural mortality rate was 12%

with the rest due to hunting. The proportion of adults and juveniles dying was 31 % and

-22-

harvest above this level would reduce population levels. In Utah, Robinette et al. (1977)

calculated that annual recruitment and mortality in their study area was 32%. Recorded

deaths during a several-year study in Alberta ranged from 3-14% of the population. Density

increased or remained stable during this time period (Ross and Jalkotzy 1989). Harvey and

Stanley Associates (1988) documented annual adult mortality ranging from 33-50%, and

annual mortality of juveniles/subadults ranging from 58-83% in the vicinity of Carlsbad.

Caverns and Guadalupe National Parks. Overall impact to this population was diffused due

to immigration from less exploited populations. In Utah, an estimated 30% of a cougar

population was experimentally removed to simulate normal harvest with regards to sex and

age classes. The cougar population had recovered numerically by the following year

(Lindzey et al. 1989). A study currently being conducted in New Mexico is evaluating

population response to a 70% removal carried out by researchers in 1991, but it is too early

to evaluate effects of the removal (K. Logan, Wildlife Research Institute, Moscow, Idaho,

pers. camm.).

Mountain lion densities may increase temporarily as a result of killing residents due

to influx of transients (Russell 1978). Unexploited mountain lion populations have been

found to contain up to 33% transients, and the ability of this species to disperse is so great

that differences in density caused by concentrated harvest are soon compensated for

(Murphy 1983). In Idaho, 26 of 30 mountain lions were removed from a 1600 mi2 areas

followed by removal of 39 more over the next six years. Reoccupancy was so rapid that no

annual changes in densities were detected (See Anderson 1987). An estimated 50%

-23-

reduction of cougar numbers in Utah had no effect on the proportion of deer carcasses

attributed to lion predation located in the study area (Robinette et a1. 1977). Similarly,

efforts to control depredation by eliminating large numbers of mountain lions have failed to

affect depredation in New Mexico (Evans 1983). Based on this, Evans suggested that

mountain lions should be managed to maintain their social structure, as excessive losses

result in higher reproductive rates and a rapid influx of transient lions. California estimated

annual mountain lion growth at only 8% with no hunting (Schulenberg 1984).

Because all mountain lion populations are dependant on adjacent populations for

recruitment of transients, Hornocker (1971) suggests that management should be uniform

in all areas, and adjacent populations should be harves~ed at similar levels, depending on

relative densities. Recruitment of transients is dependent on proximity of other lion

populations. Based on extensive surveys in the Henry Mountains of Utah, which showed a

much smaller than expected mountain lion population, it was concluded that isolated ranges

surrounded by large expanses of unsuitable mountain lion habitat are very slow to recover

from harvest (Ackerman et a1. 1981).

Because harvest may result in an mcrease of transients, managers may evaluate

harvest pressure by looking at age classes of harvested lions. In wildlife management units

in Alberta which sustained the highest lion harvest, subadult «2 yrs) males and females

made up a higher percentage of the population than in units which received less harvest

(Alberta Forestry, Parks and Wildlife 199]). According to K. Murphy (pers. comm.), effects

of harvest can be monitored by determining the percentage of mature lions in the harvest.

i

f~mailllilll_Mi\IIl!W.~illI&illI.!!il£!!l;;;;IIl:mi_i ;;;li!i!!iliiTl\lJl; jj!!l,IIl!; ~""_..3!fj__~JJ.;Y,1'''';w. 1 lt~.iiJ!i,\Ji;jfjjM;["JSMWilWli

-24-

A large percentage of older lions, especially males, generally indicates light harvest, whereas

an abundance of young lions in the harvest indicates heavy harvest. Average age of alJ lions

harvested in Wyoming over the last 5 years ranged from 3.3 years for area 7 (n = 17) to 4.6

years in area 9 (n = 7) (Calculated from Wyoming Game and Fish harvest records). Of 14

areas with at least 5 harvested lions where age was reported, mean age was 4.0 or greater

in 10 of them. These ages generally fall within or slightly below the mean age of lions in an

unhunted population in New Mexico, where average age of adult females was 3.6-4.7 years

and average age of adult males was 5.1-5.9 years (Logan et al. 1990), indicating that harvest

is apparently not excessive in any of these areas.

ReR:ulating Harvest

Female mountain lions appear more susceptible to hunting than males. Females with

kittens leave more tracks in a smaller area, making it easier for hunters to locate them

(Logan 1983). In Montana, 72% of tracks encountered by hunters and researchers in the

study area were made by females, yet females comprised only 62% of the population

(Murphy 1983). Based on this, the author believed that females may be more susceptible

to hunting because it appeared they crossed roads more frequently than males.

The resident female segment is essential to maintenance of mountain lion

populations, whereas resident males are expendable if adjacent populations produce

transients (Hornocker 1971). Based on this, Hornocker recommended that the resident

female segment be left intact, and that hunters be trained to distinguish males from females.

-25-

According to Lindzey et al. (1989), because resident female offspring replace deceased

resident females more often than transients, management to maintain female residents in

the population should enhance replacement of harvested cougars. It is the consensus of

most researchers that harvest of male mountain lions is compensatory. Therefore,

concentrating harvest on the male segment of the population would maximize the number

of mountain lions which could be harvested yet reduce potential impacts to the population.

Recent harvest in Wyoming has averaged about 50% male and 50% female. Of 302

mountain lions legally harvested state\\1de from 1987-1991,46% were females and 54% were

males (calculated from Wyoming Game and Fish harvest statistics). Variations of this may

occur in localized areas; of 39 mountain lions killed in the Bighorn Mountains of Wyoming,

only 20% were adult males, and a full 49% of all mountain lions killed were adult females

(Logan 1983).

Another reason commonly cited to protect females is that many adult females killed

by hunters have dependent kittens. Many mountain lion studies in the Rocky Mountain

region have documented a fall birth peak. In Utah, 10 of 16 litters documented were born

from October-December, and most kittens produced in the area were very young during the

hunting season (Roberson 1984). Also in Utah, 16 of 17 mountain lion litters were born

from late August through November (Barnhurst and Lindzey 1984). In Montana, mountain

lions produced litters all seasons of the year except Spring (Murphy 1983). In Yellowstone

National Park, three of six litters were born in June and three in July (Murphy and Felzien

~1&&: ,iii I! __ ,aCZllJCllll"

-26-

1991). According to K. Murphy (pers. comm.) 8 of 9 adult females have dependent kittens

during the hunting season in this area.

Females with kittens in the Bighorn Mountains of Wyoming were found to be away

from their kittens 50% of the time, and oftentimes displayed little evidence visible to hunters

(e.g. swollen teats) that they had kittens (Logan 1983). Barnhurst (1986) reported that

kittens 0-6 months of age are the most vulnerable to hunting, and that this age class is with

their mother only 19% of tbe time. Kittens 7-12 months old are \\1th their mother 43% of

the time. In their study area in Utah, they estimated that 50% of all resident females should

have 0-6 month old cubs during the winter hunting season. Based on track data, kitten

tracks were found with their mother only 25% of the time (Barnhurst and Lindzey 1989).

Therefore, 75% of females with kittens would not be recognized by hunters searching for

tracks. The authors suggested that it is possible to regulate loss of females with kittens by

adjusting the season so that kittens are older and \\1th their mothers more often. In addition

to orphaning kittens as a result of hunting, mauling of kittens by dogs chasing mountain lions

is considered a severe problem. According to Hemker et a1. (1986), kitten mortality from

dog mauling and orphaning during hunting seasons is probably as significant as adult harvest.

One way of limiting female harvest is to restrict harvest to males. According to

Poelker (1976), however, mountain lion hunters generally cannot determine sex, and it isn't

feasible or enforceable to limit harvest to males. Limiting harvest to males also may not

hold up in court because of the difficulty in distinguishing the sexes (Coles 1984). Other

researchers disagree, however. Tully (1976) feels that a "good hunter" has a 95% probability

j

~I~_~~'\!fl~_~m~'lit~

-27-

of distinguishing and harvesting a male mountain lion. According to K. Logan (pers.

comm.), once a mountain lion is treed, it is usually possible to observe it for several hours,

if necessary, and, with experience, it is possible to correctly sex 98% of treed mountain lions.

Colorado has had some male only seasons in some management areas, but do not know the

e~1ent to which female mountain lions are inadvertently shot and left or snuck out of the

area (Tully 1976, AE. Anderson, Colorado Division of Wildlife, pers. comm). Another way

to reduce female harvest is with a female subquota system such as that used in Alberta. The

female quota is set at half the full quota, and the season is closed as soon as either the full

quota or the female quota is reached (Alberta Forestry, Parks and Wildlife 1991). Female

harvest may also possibly be regulated by adjusting season length. In portions of Alberta,

a reduction in season length resulted in an increase in the number of female mountain lions

harvested (Jalkotzy and Ross 1989). Presumably, with a shorter season, hunters could not

afford to be as selective with regards to sex of mountain lion harvested.

The most serious management problem confronting biologists is avoiding overkill in

mountain lion habitats that are easily accessed by roads (Murphy 1984). When snow

conditions are excellent for tracking and persistent, it is possible to harvest more than 50%

of the resident mountain lion population in areas easily accessed by hunters (Murphy 1983).

In Idaho, mountain lion harvest is directly tied to length of time snow covers deer and elk

winter ranges (Power 1985), and in Alberta, variability in annual harvest is tied directly to

snow conditions (Pall et a1. 1988). The two most important factors affecting vulnerability

of mountain lion populations are road access and seasonal tracking conditions (Barnhurst

I

~i'.iila.";i;;;;;;JJ1i'=;;;iiiiUL_~~

-28-

1986). Therefore, the author concluded that it is possible to regulate harvest by varying

season dates and controlling road access.

MANAGEMENT STRATEGIES IN OTHER STATES/PROVINCES

Alberta

Alberta manages harvest of their cougar population using a mortality quota system

for management areas. The quota is set at 10% for hunting mortality, and a maximum of

15% man-caused mortality (Alberta Forestry, Lands and Wildlife 1991). There is a female

subquota, which is set at half the total quota. The hunting season is 3 months long. Cougar

population estimates are determined based on habitat and density estimates from an

intensive study carried out in southern Alberta.

Arizona

Arizona has a year-round season, with a limit of one lion per hunter per year.

Ranchers are allowed to shoot any lions suspected of causing damage. There are currently

no mortality quotas, although the state is considering implementing a quota system Harley

Shaw (peTS. comm.).

British Columbia

According to Fred Harper. of the British Columbia Fish and Wildlife (pers. comm.)

British Columbia has no management strategy and no quotas. The season in most areas is

l~l"""'m·m;mr~~i~_:llIl!!!iII?lli?ilill~";illlfillfF_ll\.IIIL"'D.1llI'lti

-29-

early winter - March, (i.e. the period of snow cover). It is legal to kill females, but killing

of females with kittens is discouraged. Presently, it isn't even required to bring the carcass

out, but they are trying to change this. The limit is 2 per year.

California

There is no legal hunting of mountain lions allowed in California. Killing lions

believed responsible for livestock loss is allowed. California conducts road transects every

2-3 years to monitor trends (Schulenberg 1984).

Colorado

Mountain lions in Colorado are managed using a quota system.

The quota is not based on a population estimate and percent harvest. Rather, quotas are

set by wildlife managers who make a "best guess" based on whether the population has

increased or decreased from the previous year. This determination is made based on hunter

success data and interviews with professional lion guides (Allen Anderson, pers. comm.).

Idaho uses a quota system in some areas of the state, whereas in other areas there

is no effort to control hal"est. Quotas are based on past hal"est; there are no attempts to

estimate a population size. Areas where harvest is not controlled are generally those where

lion conflicts are occurring (Charles Harris, Idaho Fish and Game, pers. comm.).

-30·

Montana

Montana manages lions using a quota and female subquota system. The female

subquota is set at 30-40% of the total quota, depending on management objectives for the

unit. There is no attempt made to estimate a population size; quotas are based on trends

and perceived population levels (S. Riley, Montana Fish, Wildlife and Parks, pers. comm.).

Nevada

From Ashman et al. (1983) and Tsukamoto (1984), mountain lion populations are

monitored using harvest data, track counts, and home ranges. Estimated annual recruitment

is 30%. Therefore, harvest objectives are determined by estimating the mountain lion

population in a mountain range and setting a 25·30% harvest rate. Population estimates are

based on analysis of habitat based upon the mountain lion study conducted in the state by

Ashman et al. (1983). Harvest data and deer densities are used to "form opinions" as to

the quality of the area as mountain lion habitat. Population estimates generally include only

residents. Areas are classified as high and moderate-low density areas. The density estimate

used for high density areas is 1 mountain lion/25 mi2, whereas the density assigned

moderate-low density areas is 1 mountain lion/40 mi2. Major management problems are

population estimation refinement, mountain lion depradation analysis and research, and

population monitoring (Tsukamoto 1984). The harvest is now controlled by hunt unit and

type of harvest for each management area (Stiver 1989). Nevada feels they can set harvest

quotas up to 35% because most mountain lion har/est occurs in localized areas, therefore

~\~~Jtm1iK"3Vi~~~~ II" refLB~E~.~

-31-

they are quickly repopulated by transients from areas hunted less (Coles 1984). Mountain

lions killed for depradation are included in the harvest quotas, and some areas are closed

before the hunting season even starts because quotas have been filled taking depredating

mountain lions (Coles 1984).

New Mexico

New Mexico has a four month long season. There are no restrictions on number of

lions taken in the state. The limit is one lion per hunter, and approximately 150 lions are

harvested every year (W. Evans, pers. comm.).

Oregon

Oregon controls their lion harvest through use of a permit system. The number of

permits are determined based on expected harvest as determined from previous years. The

age and sex structure of harvested lions is used to determine effects of harvest, and the

number of permits is adusted accordingly. The Oregon Department of Wildlife has

developed an in-house model for mountain lions that is in the process of being published

(W. Van Dyke, Oregon Department of Fish and Wildlife, pers. comm.).

Mountain lions in Texas are classified as a nongame animal. There are no license

requirements, limits or other restrictions placed on their take (Russ] 989, J. Rutledge, Texas

Parks and Wildlife Department, pers. comm.).

-32-

Utah

In Utah, the average number of mountain lions killed per square mile is used to

group management areas into similar densities. Mountain lion density estimates and habitat

similarities from studies in Idaho and Nevada were used to refine estimates. They monitor

population trends through harvest statistics. Utah will use mark-recapture surveys to

estimate mountain lion density in the future. Starting in 1990, 3 units will be sampled per

year, and results will be compared to harvest statistics to determine trends state\\~de (Bates

1989). Harvest is controlled through use of a permit system. The number of permits issued

is based primarily on historical harvest (Randy Radant, Utah Dh~sion of Wildlife Resources,

pers. comm.).

Washington

Washington uses a permit system similar to the ones used for ungulates. Indices they

use to monitor population include sex and age structure of harvest, number of depredation

complaints and other human contact, and hunter effort. Success rate is relatively stable, and

the number of permits issued is based on expected success rate (1. Reick, Washington

Department of Wildlife, pers. comm.)

-33-

LITERATURE CITED

Ackerman, B. B., T. P. Hemker, F. G. Lindzey, and A. J. Button. 1981. Cougarnumbers in the Henry Mountains, Utah. EncycJia 58:57-62.

Alberta Forestry, Lands and Wildlife. 1991. Management plan for cougar in Alberta.Alberta Forestry, Lands and Wildlife Wildlife Management Planning SeriesNo.5. 91pp.

Anderson, A. E., D. C. Bowden, and D. M. Kattner. 1989. Survival in an unhuntedmountain lion (Felis collc%r hippo/estes) population in southwesternColorado. P. 57 in R.H. Smith, ed. Trans. 3rd Mountain Lion Trans, ArizonaChapter, The Wildlife Society and Arizona Game and Fish Dept.

Anderson, A. E. 1988. Mountain lion population dynamics. Fed Aid Job Prog. Rep.W-53-R-2, WP GA-J1. Pp. 193-220 in Wildl. Res. Rep. July, 1988, Part 2, Colo. Div.Wild!., Fort Collins.

Anderson, A. E. 1987. Mountain Lion investigations. Colo. Div. Wildl. Wildl. Res.- Rep. Proj. No. 01-03-048 (FW 26 P) Work Plan No.6. Job No. 1.

Anderson, A. E. 1983. A critical review of literature on puma (Felis collc%r). Colo.Div. Wild!. Spec. Rep. No. 54. 91pp.

Ashman, D., G. C. Christensen, M. C. Hess, G. K. Tsukamoto, and M. S.Wichersham. 1983. The mountain lion in Nevada. Nev. Dep. Wildl., Reno.75pp.

Ashman, D. 1976. Mountain lion investigations. Job Perform. Rep., P-R Proj. W-48­7, Study S&I, Job 5 and Study R-Y, Job 1. Nev. Fish Game Dep_ 19pp.

Barnhurst, D. and F. G. Lindzey. 1989. Detecting female mountain lions with kittens.Northwest Sci. 63:35-37.

Barnhurst, D. 1986. Yulnerability of cougars to hunting. M.S. Thesis, Utah. St. Univ.,Logan. 66pp.

Barrett, R. H. 1986. Population models for black bear and mountain lion inCalifornia. Final Report, Proj. C-1421. Calif. Dept. Fish Game, Sacramento.43pp + appendices.

-34-

Bates, B. 1989. Status of cougar in Utah, 1988. Pp. 32-34 in R.H. Smith, ed. Trans.3rd Mountain lion workshop, Arizona Game and Fish Dept.

Berg, R. L., L. L. McDonald, and M. D. Strickland. 1983. Distribution of mountainlions in Wyoming as determined by mail questionnaire. Wildl. Soc. Bull.11:265-268.

Berg, R L. 1981. A mail survey for information on the distribution of mountain lionsin the State of Wyoming. M.S. Thesis. Univ. Wyo., Laramie. 94pp.

Coles, F. 1984. Harvest strategies. Pp. 230-266 in J. Roberson and F. Lindzey,(eds.). Proc. 2nd Mtn. Lion Workshop. Utah Div. WildJ. Re_s., Salt Lake City.

Currier, M. J. P., S. L. Sheriff, and K. R. Russell. 1977. Mountain lion populationand harvest near Canon City, Colorado. Colo. Div. WildJ. Spec. Rep. No. 42.

. 1')V1+ _pp.

Currier, M. J. P. 1976. Characteristics of the mountain lion population near CanonCity, Colorado. M.S. Thesis. Colo. St. Univ. 51pp.

Dewar, P. 1976. Comments. P. 65 in G. C. Christensen and R. J. Fischer, co­chairmen. Trans. Mountain Lion Workshop. U.S. Fish WildJ. Serv., Portland,Oregon and Nev. Fish Game Dep., Reno. 213pp.

Dixon, K. R 1982. Mountain lion: Felis concolor. Pp. 711-727 in Wild Mammals ofNorth America: Biology, Management, and Economics. J .. A. Chapman andG. A. Feldhamer, eds.

Donaldson, B. 1975. Mountain lion research. Perform. Rep., P-R Proj. W-93-R,17,Work Plan 15, Job 1. New Mex. Game Fish Dep. 18pp.

Evans, W. 1983. The cougar in New Mexico: biology, status, depredation oflivestock, and management recommendations. New Mex. Dept. Game Fish,Santa Fe. 40pp.

FitZhugh, E. L. and K. S. Smallwood. 1989. Techniques for monitoring mountain lionpopulation levels. Pp. 69-71 in RH. Smith, ed. Trans. 3rd Mountain LionWorkshop, Arizona Chapter, The Wildlife Society and Arizona Game andFish Dept.

j -l. . .".@"" ~__'''_ . '1!i-''''--'-'''@.1tl'ij@._'''-'-~_ffilV.IM'J£W:\'~~m"~~~~,jlj({14\iio'i!'./!~~liM;llll;l.'ii:ijli1'" ili ··lfliW«"~;;.~f,tt,k;&,x~all:~ii"!~~~iRi,

-35-

Harcombe, D. W. 1976. Oregon cougar study. Oreg. Dept. Fish Wild!. Portland,Oreg. 62pp.

Harvey and Stanley Associates, Inc. 1988. Mountain lions (Feliscollcolor) in the vicinity ofCarlsbad Caverns National Park, New Mexico and Guadalupe Mountains NationalPark, Texas: An Ecological Study. Final Rep. Submitted to National Park Service,Santa Fe, New Mex. 137pp.

Hebert, D. 1989. The status and management of cougar in British Columbia. Pp. 11­14 in R. H. Smith, ed. Trans. 3rd Mountain lion workshop, Arizona Chapter,The Wildlife Society and Arizona Game and Fish Dept.

Hebert, D. 1976. Comments. P. 99 in G. C. Christensen and R. J. Fischer, co­chairmen. Trans. Mountain Lion Workshop. U.S. Fish Wild!. Serv., Portland,Oregon and Nev. Fish Game Dep., Reno. 213pp.

Heisy, D. M., and T. K. Fuller. 1985. Evaluation of survival and cause-specific mortalityrates using telemetry data. J. Wild!. Manage. 49:668-674.

Hemker, T. P., F. G. Lindzey, B. B. Ackerman, and A. J. Button. 1986. Survival ofcougar cubs in a non-hunted population. Pp. 327-332 in Cats of the World:Biology, Conservation, and Management. S. D. Miller and D. D. Everett, eds.

Hemker, T. P., F. G. Lindzey, and B. B. Ackerman. 1984. Population characteristicsand movement patterns of cougars in southern Utah. J. Wild!. Manage.48: 1275-1284.

Hemker, T. P. 1982. Population characteristics and movement patterns of cougarsin southern Utah. M.S. Thesis, Utah S1. Univ., Logan. 66pp.

Hopkins, R. A. 1984. The role of the puma in a nonmigratory large mammal communityin the Diablo Range of California. Pp. 134-137 in J. Roberson and F. Lindzey, (eds.).Proc. 2nd Mtn. Lion Workshop. Utah Div. Wild!. Res., Salt Lake City.

Hornocker, M. G. 1971. Suggestions for the management of mountain lions astrophy species in the Intermountain region. Annu. Proc. Western Assoc. StateGame Fish Commissioners 51:399-402.

Logan, K. A., L. L. Sweanor, J. F. Smith, B. R. Spreadbury and M. G. Hornocker.1989. Ecology of an unexploited cougar population in a desert environment.P. 74 in R. H. Smith, ed. Trans. 3rd Mountain Lion Workshop, ArizonaChapter, The Wildlife Society and Arizona Game and Fish Dept.

-36-

Hornocker, M. G. 1970. An analysis of mountain lion predation upon mule deer andelk in the Idaho primitive area. Wildl. Monogr. 21:1-39.

Jalkotzy, M. G. 1989. Population ecology of cougars in southwestern Alberta. P. 47in R. H. Smith, ed. Trans. 3rd Mountain lion workshop, Arizona Chapter, TheWildlife Society and Arizona Game and Fish Dept.

Jalkotzy, M. G., and P. 1. Ross. 1989. Cougar hunting regulations and harvest inAlberta between 1973 and 1987. P. 6 in R. H. Smith, ed. Trans. 3rd MountainLion Workshop, Arizona Chapter, The Wildlife Society and Arizona Gameand Fish Dept.

Koehler, G. M. and M. G. Hornocker. 1986. A preliminary survey of mountain lionsin Yellowstone National Park. Unpubl. Rep. Wildlife Research Institute,Moscow, Idaho. 12pp.

Koford, C. B. 1978. The welfare of the puma in California. Carnivore 1:92-96.

Kutilek, M. J., R. A. Hopkins, and T. E. Smith. 1980. Second annual progress report on theecology of mountain lions (Felis collc%r) in the Diablo Range of California. Dept.BioI. Sci., San Jose St. Univ., 21pp.

Laing, S. P. 1988. Cougar habitat selection and spatial use patterns in southernUtah. M.S. Thesis. Univ. Wyo., Laramie. 68pp.

Lindzey, F. G., B. B. Ackerman, D. Barnhurst, T. Becker, T. P. Hemker, S. P. Laing,C. Mecham, and W. D. Van SickJe. 1989. Boulder-Escalante Cougar ProjectFinal Report. Utah Div. Wildl. Res., Salt Lake City, Utah. 92pp.

Lindzey, F. G., B. B. Ackerman, D. Barnhurst, and T. P. Hemker. 1988. Survivalrates of mountain lions in southern Utah. J. Wildl. Manage. 52:664-667.

Logan, K. A., L. L. Sweanor, J. F. Smith, B. R. Spreadbury, and M. G. Hornocker.1990. Ecology of an unexploited mountain lion population in a desertenvironment. Wildlife Research Institute Unpubl. Rep. to New Mexico Dept.Game Fish, Santa Fe. BOpp.

II~-mfi£"W®J%".'i'" j ,: ,,:,', , 'W);, _,~,"!;!~'l~;, ',:.' r" "'J.!'>W,''''::., "l,.'" ""'%'1:, 'JO((d,.,i!!iJ,.I!Si1_liiIillll '" ~iLi!i!i£l'lT.k,,,ll'I,._,,,i!fll!lllt~R,liiIi!liiIi!i!i"Ill'JJJim~~.li\.

-37-

Logan, K. A., L. L. Irwin, and R. Skinner. 1986. Characteristics of a huntedmountain lion population in Wyoming. J. Wildl. Manage. 50:648-654.

Logan, K. A. and L. R. Irwin. 1985. Mountain lion habitats in the Big HornMountains, Wyoming. Wildl. Soc. Bull. 13:257-262.

Logan, K. A. 1983. Mountain lion population and habitat characteristics in the BigHorn Mountains of Wyoming. M.S. Thesis, Univ. Wyo., Laramie. 101pp.

Mansfield, T. M. 1986. Mountain lion management in California. Trans. N. Am.Wildl. Nat. Resour. Conf. 51:178-182.

McBride, R. T. 1976. The status and ecology of the mountain lion Felis concolorstanleyGnG of the Texas-Mexico border. M.S. Thesis, SuI Ross St. Univ.,Alpine, Texas.

Mountain Lion Preservation Foundation. 1987. Summary of information fromquestionnaires mailed to mountain lion experts. Mountain Lion PreservationFoundation, Sacramento, California. 11pp.

Murphy, K. M., and G. S. Felzien. 1991. The ecology of the mountain lion (Felis concolormissoulensis) in the northern Yellowstone Ecosystem. Cumulative Progress Rept. No.4. Wildlife Research Institute, Moscow, Idaho. 17pp.

Murphy, K. M. 1984. Montana. Pp. 39-43 in J. Roberson and F. Lindzey, (eds.).Proc. 2nd Mtn. Lion Workshop. Utah Div. Wildl. Res., Salt Lake City.

Murphy, K. M. 1983. Relationships between a mountain lion population and huntingpressure in western Montana. Montana Dept. Fish, Wildl. Parks Proj. W-120·R-13 and 14. 48pp.

Neal, D. L. 1989. Mountain lion density and movement in the central Sierra Nevada(California). P. 72 in R. H. Smith, ed. Trans. 3rd Mountain lion workshop,Arizona Chapter, The Wildlife Society and Arizona Game and Fish Dept.

Norelius, S. and M. Bruscino. 1988. Domestic sheep predation from mountain lionsin the southern Bighorns: A field study and literature review. Wyoming GameFish Dep., Cheyenne. 42pp.

Pall, 0., M. Jalkotzy, and J. Ross. 1988. The cougar in Alberta. Fish Wildl. Div.Alberta For. Lands Wildl. 145pp.

-38-

Parsons, J. W. 1976. Comments. P. 30 in G.c. Christensen and RJ. Fischer, co­chairmen. Trans. Mountain Lion Workshop. U.S. Fish Wildl. Serv., Portland,Oregon and Ney. Fish Game Dep., Reno. 213pp.

Phelps, J. S. 1989. Status of mountain lions in Arizona. Pp. 7-9 in R H. Smith, ed.Trans. 3rd Mountain Lion Workshop, Arizona Chapter, The Wildlife Societyand Arizona Game and Fish Dept.

Poelker, R J. 1976. The status and management of the puma in Washington.World's cats. 3:20-27.

Power, G. D. 1985. Mountain lion management plan 1986-1990. Idaho Dept. FishGame. 16pp.

Power, G. D. 1976. Comments. Pp. 50, 55, 89-90 in G. C. Christensen and R J.Fischer, co-chairmen. Trans. Mountain Lion Workshop. U.s. Fish Wildl. Serv.,Portland, Oregon and Nev. Fish Game Dep., Reno. 213pp.

Rasmussen, D. I. 1941. Biotic communities of Kaibab Plateau, Arizona. Ecol.Monogr. 11:229-275.

Roberson, J. 1984. Utah mountain lion status report. Utah Diy. Wildl. Res. 23pp.

Robinette, W. L., N. V. Hancock, and D. A. Jones. 1977. The Oak Creek mule deerherd in Utah. Utah Diy. Wildl. Res. Publ. 77-15. 148pp.

Ross, I. and M. Jalkotzy. 1989. The Sheep River cougar project phase II. FinalReport 1987-1989. Arc A~sociated Resource Consultants, Ltd. Calgary,Alberta. 56pp.

Ross, I. and M. Jalkotzy. 1988. The Sheep River cougar project phase II. 1987-1988Prog. Rep. Arc Associated Resource Consultants, Ltd. Calgary, Alberta. 21pp.

Russ, W. B. 1989. Status of the mountain lion in Texas. Pp. 30-31in RH. Smith, ed. Trans. 3rd Mountain Lion Workshop, ArizonaChapter, The Wildife Society and Arizona Game and Fish Dept.

Russell, K. R. 1978. Mountain lion. Pp. 207-225 jn J. L. Schmidt and D. L. Gilbert,eds. Big Game of North America: ecology and Management. Stackpole Co.,Harrisburg, Pa. 494pp.

~

-39-

Schulenberg, B. 1984. California. Pp. 23-26 in 1. Roberson and F. Lindzey, (eds.).Proc. 2nd Mtn. Lion Workshop. Utah Div. Wildl. Res., Salt Lake City.

Seidensticker, J. c., IV., M. G. Hornocker, W. V. Wiles and 1. F. Messick. 1973.Mountain lion social organization in the Idaho primitive area. Wildl. Monogr.35.60pp.

Shaw, H. G. 1990. Preliminary recommendations for mountain lion track counts inArizona. Unpubl. Rept. General Wildlife Services, Chino Valley, Arizona.7pp.

Shaw, H. G., N. G. Woolsey, J. R. Wegge and R. L. Day, Jr. 1988. Factors affectingmountain lion densities and cattle depredation in Arizona: A final Repon.Ariz. Game Fish Dep. Proj. W-78-R, Work Plan 2, Job 29. 16pp.

Shaw, H. G. 1980. Ecology of the mountain lion in Arizona. Final Rep., P-R Proj.W-78-R, Work Plan 2, Job 13. Ariz. Game Fish Dep. 14pp.

Shaw, H. G. 1977. Impact of mountain lion on mule deer and cattle in northwesternArizona. Pp. 17-32 in R. L. Phillips and C. Jonkel, eds. Froc. 1975 PredatorSymp. Univ. Mont., Missoula.

Sheriff, S. L. 1978. Computer model for mountain lion populations. M.S. Thesis.Colo. St. Univ., Ft. Collins. 287pp.

Sitton, L. W. 1977. California mountain lion investigations with recommendations formanagement. Final Rep. to the State Legislature, P-R Proj. W-51-R,Sacramento: California Fish Game Dep. 35pp.

Sitton, L. W. 1972. Investigations on the status of the California mountain lion. Cal­Neva Wildl. 1972:31-34.

Smallwood, K. S. and E. L. Fitzhugh. 1987. A statewide mountain lion populationindex technique. Wildl. Ext., Univ. Calif., Davis. 26pp.

Smith, T. E., R. R. Duke, M. J. Kutilek, and J. Walters. 1989. A non invasivemethod for monitoring mountain lion population trends in roadless areas. P.73 in R. H. Smith, ed. Trans. 3rd Mountain Lion Workshop, Arizona Chapter,The Wildlife Society and Arizona Game and Fish Dept.

-40-

Smith, T. E., R. R. Duke, and M. J. Kutilek. 1984. The ecology of the mountain lionin the Guadalupe Mountains of Texas and New Mexico. Pp. 174-175 in J.Roberson and F. Lindzey, (eds.). Proc. 2nd Mtn. Lion Workshop. Utah Div.Wildl. Res., Salt Lake City.

Spreadbury, B. 1989. Cougar ecology in southeastern British Columbia. P. 48 in R.H. Smith, ed. Trans. 3rd Mountain Lion Workshop, Arizona Chapter, TheWildlife Society and Arizona Game and Fish Dept.

Stiver, S. J. 1989. Status of mountain lions in Nevada. Pp. 26-29 in R. H. Smith, ed.Trans. 3rd Mountain lion workshop, Arizona Chapter, The Wildlife Societyand Arizona Game and Fish Dept.

Trulio, L. A. 1989. What mountain lion scientists say about their research: results ofa survey on mountain lion research methods. Mountain Lion PreservationFoundation. llpp.

Tsukamoto, G. 1984. Nevada. Pp. 44-48 in 1. Roberson and F. Lindzey, (eds.). Proc.2nd Mtn. Lion Workshop. Utah Div. Wildl. Res., Salt Lake City.

Tully, R. J. 1976. Comments. P. 120 in G. C. Christensen and R. J. Fischer, co­chairmen. Trans. Mountain Lion Workshop. U.S. Fish Wildl. Ser., Portland,Oreg. and Nev Fish Game Dep., Reno. 213 pp.

Van Dyke, F. G. and R. H. Brocke. 1987. Searching technique for mountain lionsign at specific locations. Wildl. Soc. Bull. 15:256-259.

Van Dyke, F. G., H. B. Brocke, and H. G. Shaw. 1986. Use of road track counts asindices of mountain lion presence. 1. Wildl. Manage. 50:102-109.

Van Sickle, W. 1990. Methods for estimating cougar numbers in southern Utah.M.S. Thesis. Univ. Wyo., Laramie. 73pp.

Van Sickle, W. D., and F. G. Lindzey. 1991. Evaluation of a cougar populationestimator based on probability sampling. 1. Wildl. Manage. 55:738-743.

;;~~~~j'E~~am;;m~!W~~1iiWmr~"wrn*i:lrM'M!111Mw.L"Wl!I~~~",m:~tWI';;&,\l\'./i!lW«llif,:r!ril M.*",,$~~#j"lilm,\>mf"

Table 1. Mountain lion Densities in the Western United States and Canada

Location

Alberta

Arizona

B.C.

California

Colorado

Idaho

Montana

Nevada

New Mexico

Texas

Texas/New Mex.

Utah

Wyoming

Density (per 100 km')

2.7-5.9

2.2"5.01.73.6

4.0-4.4

1.9-3.21.5

3.5-3.7

9.23.5-4.4

3.83.41.81.40.34.4

1.9-2.33.94.1

4.91.8-3.3

1.2

2.90.9

7.1

1.4-1.61. 4-1. 7

1.41.7-3.3

0.70.5

5.2-6.4

2.3b

2.2-2.90.451.19

0.3-0.5

3.4-4.53.6

Source

Alberta Forestry,Lands, Wildlife (1991)

Shaw 1979, 1981Rasmussen 1941Rasmussen 1941Shaw 1977Phelps 1989

Hebert 1976Dewar 1976Spreadbury 1989

Sitton 1972Sitton 1972Sitton 1972Sitton 1972Sitton 1972Koford 1978Koford 1978Sitton 1977Mansfield 1986Mansfield 1986Neal 1989

Currier et ale 1977Currier et a1. 1977Currier et ale 1977

Hornocker 1970Power 1976

Murphy 1983

Ashman 1976, 1981Ashman 1976

Donaldson 1975Logan et a1. 1990

McBride 1976McBride 1976Parsons 1976

Smith et al. 1984

Robinette et a1. 1977Hemker 1982Laing 1988Hemker et a1. 1984

Logan 1983Murphy and Fe1zien 1991

: Resident adults onlyMaximum density of adults only

APPENDIX II. MOUNTAIN LION RESEARCH AND MANAGEMENTANNOTATED BIBLIOGRAPHY

Ackerman, B. B., F. G. Lindzey, and T. P. Hemker. 1986. Predictive energetics model forcougars. Pp. 333-352 in Cats of the World: Biology, Conservation, and Management.S. D. Miller and D. D. Everett, eds.

Describes model to predict required energy intake and number of prey animals consumedby free-ranging cougar in Utah.

Ackerman, B. B. 1982. Cougar predation and ecological energetics in south-central Utah.M.S. Thesis, Utah St. Univ., Logan. 95 pp.

Presents data on cougar consumption rates, activity patterns, estimated metabolic costs andprey requirements.

Ackerman, B. B., T. P. Hemker, F. G. Lindzey, and A. J. Button. 1981. Cougar numbers inthe Henry Mountains, Utah. Encyclia 58:57-62.

Describes results of eX1ensive lion surveys in an isolated mountain range in Utah. Based onlower than expected population, the authors concluded isolated ranges may be slow torecover from harvest.

Anderson, A. E. 1988. Mountain lion population dynamics. Fed Aid Job Prog. Rep. W-53­R-2, WP GA-J1. Pp. 193-220 in Wild], Res. Rep. July, 1988, Part 2, Colo. Div. Wild]',Fort Collins.

Presents results of telemetry study on lions in Colorado to assess effects of sport hunting.Data on mortality, age and sex composition, survival, home range, and prey are presented.

Anderson, A. E., and R. J. Tully. 1989. Status of the mountain lion in Colorado. Pp. 19-23in R. H. Smith, ed. Trans. 3rd Mountain lion workshop. Arizona Chapter, TheWildlife Society and Arizona Game and Fish Dept.

Discusses current status of, management programs, and research being conducted onColorado mountain lions.

-2-

Anderson, A. E, D. C. Bowden, and D. M. Kattner. 1989. Survival in an unhuntedmountain lion (Felis concolor hippolesles) population in southwestern Colorado. P. 57in R. H. Smith, ed. Trans. 3rd Mountain Lion Trans, Arizona Chapter, The WildlifeSociety, and Arizona Game and Fish Dept.

Provides survival estimates based on monitoring of 20 male and 21 females. Annual survivalestimated at 88.1 %.

Anderson, A. E, D. C. Bowden, and D. M. Kattner. 1989. Dynamics of home range size ofunhunted mountain lions (Felis concolor hippolestes) in southwestern' Colorado. P.57 in R. H. Smith, ed. Trans. 3rd Mountain Lion Trans, Arizona Game and FishDept.

Discusses temporal changes in home range size.

Anderson, A. E. 1987. Mountain Lion investigations. Colo. Div. Wildl. Wildl. Res. Rep.Proj. No. 01·03-048 (FW 26 P) Work Plan No.6. Job No. 1.

Presents results of telemetry study on lions in Colorado to assess effects of sport hunting.Data on mortality, home range, and prey are presented.

Anderson, A. E 1986. Mountain Lion investigations. Colo. Div. Wildl. Wifdl. Res. Rep.Proj. No. 01·03·048 (FW 26 P) Work Plan No.6. Job No. 1. 32pp.

Presents results of telemetry study on lions in Colorado to assess effects of sport hunting.Data on mortality, home range, activity and movement, and prey are presented.

Anderson, A. E. 1984. Mountain lion investigations - mountain lion population dynamics.Colo. Div. WildL Res. Rep. July, Part 2: 221-268.

Presents results of telemetry study on lions in Colorado to assess effects of sport hunting.Data on mortality, activity and movement, grov.1h and prey are presented.

-3-

Anderson, A. E., C. R. Anderson, and D. M. Kattner. 1984. Mountain lion populationdynamics. Pp. 147-148 in J. Roberson and F. Lindzey, (eds.). Proc. 2nd Mtn. LionWorkshop. Utah Div. WildJ. Res., Salt Lake City.

Presents preliminary information from telemetry study in southwest Colorado to monitoreffects of harvest. Gives fates of cougars marked to date.

Anderson, A. E. 1984. Colorado. Pp. 27-28 in 1. Roberson and F. Lindzey, (eds.). Proc.2nd Mtn. Lion Workshop. Utah Div. WildJ. Res., Salt Lake City.

Provides list of Colorado mountain lion literature.

Anderson, A. E. 1983. A critical review of literature on puma (Felis cOllcolor). Colo. Div.WildJ. Spec. Rep. No. 54. 91pp.

Comprehensive review of mountain lion literature covering all aspects of mountain lionecology, biology, and management.

Anderson, A. E. 1982. Mountain lion investigations - mountain lion population dynamics.Colo. Div. WildJ. Res. Rep. July, Part 2: 143-159.

Presents preliminary information of cougar telemetry study in Colorado.

Ashman, D., G. C. Christensen, M. C. Hess, G. K. Tsukamoto, and M. S. Wichersham.1983. The mountain lion in Nevada. Nev. Dep. WildJ., Reno. 75pp.

Describes population dynamics, effects of harvest, population monitoring techniques, andmanagement of mountain lions in Nevada;gives criteria for aging lions.

Ashman, D. 1979. Mountain lion investigations. lob Perform. Rep., P-R Proj. W-48-1O,Study S&I, lob 5 and Study R-Y, lob 1. Nev. Fish Game Dep. lOpp.

Presents data on mortality rates, sources of mortality, and depredation from radiotelemetrystUdy. Describes track counts and scratch site monitoring and presents harvest data.

-4-

Ashman, D. 1978. Mountain lion investigations. Job Perform. Rep., P-R Proj. W-48-9,Study S&I, Job 5 and Study R-V, Job 1. Nev. Fish Game Dep. 10pp.

Presents data on mortality and sources of mortality from radiotelemetry study. Discussesresults of track surveys from ground and helicopter and presents harvest data.

Ashman, D. 1977. Mountain lion investigations. Job Perform. Rep., P-R Proj. W-48-8,Study S&I, Job 5 and Study R-V, Job 1. Nev. Fish Game Dep. llpp.

Presents data on mortality, sources of mortality, productivity, and population structure fromradiotelemetry study. Discusses results of track and other sign surveys from ground and air.

Ashman, D. 1976. Mountain lion investigations. Job Perform. Rep., P-R Proj. W-48-7,Study S&I, Job 5 and Study R-V, Job 1. Nev. Fish Game Dep. 19pp.

Presents data on mortality and sources of mortality from radiotelemetry study. Discussesresults of track and other sign surveys.

Ashman, D. 1975. Mountain lion investigations. Job Perform. Rep., P-R Proj. W-48-6,Study S&I, Job 5. Nev. Fish. Game Dep. 18 pp.

Presents data on mortality and sources of mortality from radiotelemetry study. Discussesresults of track and other sign surveys.

Barnhurst, D., and F. G. Lindzey. 1989. Detecting female mountain lions with kittens.Northwest Sci. 63:35-37.

Presents data on time kittens spend with mother and implications for management byadjusting season timing.

Barnhurst, D. 1986. Vulnerability of cougars to hunting. M.S. Thesis, Utah. St. Univ.,Logan. 66pp.

Describes vulnerability of kittens to orphaning by age class based on time spent with mother.Discusses effects of area accessibility and snow conditions on cougar harvest, and providesrecommendations for management.

-5-

Barnhurst, D., and F. Lindzey. 1984. Utah. Pp. 185-188 in J. Roberson and F. Lindzey,(eds.). Proc. 2nd Mtn. Lion Workshop. Utah Div. Wild!. Res., Salt Lake City.

Summarizes results of Boulder-Escalante study of cougar ecology, behavior, populationdynamics, movement patterns, food habits, and response to harvest.

Barrett, R. H. 1986. Population models for black bear and mountain lion in California.Dept. For. Resour. Manage., Univ. Calif., Berkeley. 43pp + appendices.

Describes mountain lion model and assumptions of model to determine population dynamicsand effects of harvest. Model indicates that 25% harvest is maximum sustained yield.

Barrett, R. H. 1987. Population model for mountain lion in California. Dept. For. Resour.Manage., Univ. Calif., Berkeley. lOp + appendix.

Describes mountain lion model and assumptions of model to determine population dynamicsand effects of harvest.

Bates, B. 1989. Status of cougar in Utah, 1988. Pp. 32-34 in R.H. Smith, ed. Trans. 3rdMountain lion workshop, Arizona Chapter, The Wildlife Society, and Arizona Gameand Fish Dept.

Describes status, management, economic values and research being conducted on mountainlions in Utah.

Bavin, R. 1978. Mountain lion research. Perform. Rep., P-R Proj. W-124-R-l, Job 1. NewMex. Game Fish Dep. 5 pp.

Presents data on home range, daily activities and movement patterns, dispersal, and speciescomposition of lion kills in New Mexico.

Bavin, R. 1976. Mountain lion research. Perform. Rep., P-R Proj. W-93-18, Work Plan 15,Job 1. New Mex. Game Fish Dep. 3pp.

Presents data on home range, daily activities and movement patterns, dispersal, and speciescomposition of lion kills in New Mexico.

-6-

Belden, R. C 1988. The Florida panther. Audubon Wild!. Rep. 515-532.

Provides description of subspecies, natural history, significance, historical perspective, currenttrends, management, prognosis, and recommendations.

Belden, R. C, W. B. Frankenberger, R. T. McBride, and S. T. Schwikert. 1988. Pantherhabitat use in southern Florida. J. Wild. Manage. 52: 660-663.

Provides horne range, habitat preference and projects population size based on 1,630locations of 6 panthers.

Belden, R. C 1986. Florida panther recovery plan implementation - a 1983 progress report.Pp. 159-172 in Cats of the World: Biology, Conservation, and Management. S. D.Miller and D. D. Everett, eds.

Presents goals of recovery plan and results of research conducted to date includingdistribution, habitat use, food habits, and characteristics. Presents recommendations formanagement and protection and future research.

Belden, R. C, J. C Robski, and D. K. Jansen. 1984. Florida mountain lion research. Pp.149-166 in J. Roberson and F. Lindzey, (eds.). Proc. 2nd Mtn. Lion Workshop. UtahDiv. Wildl. Res., Salt Lake City.

Description of studies being conducted as part of the Florida Panther Recovery PlanImplementation Project.

Berg, R. L., L. L. McDonald, and M. D. Strickland. 1983. Distribution of mountain lionsin Wyoming as determined by mail questionnaire. WildJ. Soc. Bull. 11:265-268.

Describes lion distribution, relative densities, and habitat based on survey of trappers, lionhunters, outfitters, biologists, foresters, and ranchers.

-7-

Berg, R. L. 1981. A mail survey for information on the distribution of mountain lions in theState of Wyoming. M.S. Thesis. Univ. Wyo., Laramie. 94pp.

Describes lion distribution, relative densities, and habitat based on survey of trappers, lionhunters, outfitters, biologists, foresters, and ranchers.

Boddicker, M. L. 1983. Mountain lions. Pp. C65-C67 in Prevention and Control of WildlifeDamage. R.M. Timm, ed.

Presents brief description of lion biology and recommends control measures.

Bowns, J. E. 1984. Predation-Depredation. Pp. 204-215 in J. Roberson and F. Lindzey,(eds.). Proc. 2nd Mtn. Lion Workshop. Utah Div. Wild!. Res., Salt Lake City.

Discussion of cougar livestock predation and potential solutions.

BrittreJl, J. D. 1984. Washington. Pp. 80-87 in J. Roberson and F. Lindzey, (eds.). Proc.2nd Mtn. Lion Workshop. Utah Div. WildJ. Res., Salt Lake City.

Provides information on mountain lion distribution, harvest information, and managementprograms in Washington.

Brown, D. E. 1984. A lion for all seasons. Pp. 12-22 in J. Roberson and F. Lindzey, (eds.).Proc. 2nd Mtn. Lion Workshop. Utah Div. Wildl. Res., Salt Lake City.

Reports that the age class of harvested mountain lions can be used to examine popUlationdynamics. A dominance of younger age animals indicates a heavily exploited population.Large numbers of prime age animals and few yearlings and kittens suggest resident lions arebeing harvested.

Brown, D. E. 1984. Arizona. Pp. 9-12 in J. Roberson and F. Lindzey, (eds.). Proc. 2ndMtn. Lion Workshop. Utah Div. Wild!. Res., Salt Lake City.

Summarizes mountain lion harvest statistics in Arizona.

!Il!l1l1ll••,~I.•••••••••••••~m~~~~~;0t "', " m.~~.;l!tl v "",,""(0WJ~~fMif~~"" ~&illlMi'~lt"

-8-

Bruscino, M. 1989. Evaluating mountain lion depredation of domestic sheep. U.s. For. ServoGen. Tech. Rep. RM-171. Pp 33-33.

Describes condition and locations of 71 domestic sheep killed by mountain lions in theBighorn Mountains of Wyoming.

Burriss, J. S. 1979. The Utah cougar harvest book 1977-1978. Utah Div. Wildl. Res. Publ.No. 79-1. 17pp.

Summarizes cougar management and describes harvest and harvest statistics for 1977-1978season.

Christensen, G. C. and R. J. Fischer, Co-chairmen. 1976. Trans. Mountain Lion Workshop.U.S. Fish Wildl. Ser., Portland, Oreg. and Nev. Fish Game Dep., Reno. 213pp.

Compilation of research and status reports and discussion of cougar research andmanagement in the U.S. and Canada.

Clarke, J. and H. Vriend. 1980. History of cougar management in Alberta. Alberta FishWildl. Div., Lethbridge. Unpub. Rep. 20pp.

Chronicles history of cougar management in Alberta from bounty years to 1980.

Coles, F. 1984. Harvest strategies. Pp. 230-266 in J. Roberson and F. Lindzey, (eds.). Proc.2nd Mtn. Lion Workshop. Utah Div. Wildl. Res., Salt Lake City.

Discussion of lion harvest strategies, including hunting seasons and dates, birth peaks,depradation and hunting, setting quotas, harvest of depredating lions, and male-only hunting.

Currier, M. J. P. 1983. Felis COlleolor Linnaeus, 1771 Mountain Lion. Am. Soc. Mammal.Mammal. Species. No. 200. 7pp.

Describes taxonomy, general characteristics, distribution, reproduction, ecology, genetics andbehavior.

-9-

Currier, M. J. P. 1979. An age estimation technique and some normal blood values formountain lions (Felis concolor). Ph.D. Thesis. Colorado St. Univ., Ft. Collins. 81pp.

Provides aging criteria and blood values from captive and wild cougars. Blood values werenot correlated ",,'ith sex or age.

Currier, M. J. P., S. L. Sheriff, and K. R. Russell. 1977. Mountain lion population andharvest near Canon City, Colorado. Colo. Div. Wildl. Spec. Rep. No. 42. vi+12pp.

Describes mountain lion habitat, population structure, densities, and census results.Summarizes harvest in study area over 10 year period and describes effects of harvest onpopulation. .

Currier, M. J. P. 1976. Characteristics of the mountain lion population near Canon City,Colorado. M.S. Thesis. Colo. St. Univ. 51pp.

Describes mountain lion habitat, population structure, densities, and census results.

Deems, E. F., Jr. and D. Pursley, eds. 1978. North American furbearers: their management,research and harvest status in 1976. Int. Assoc. Fish Wildl. Agencies, Washington,D.C. 177pp.

Summarizes mountain lion management in U.s. and Canada.

Dewar, P., and P. Dewar. 1976. The status and management of the puma in BritishColumbia. World's Cats 3:4·19.

Discusses history, status and management of cougars in British Columbia.

Dixon, K. R. 1982. Mountain lion: Felis concolor. Pp. 711-727 in Wild Mammals of NorthAmerica: Biology, Management, and Economics. l.A. Chapman and G.A.Feldhamer, eds.

Review of mountain lion literature.

-10-

Donaldson, B. 1975. Mountain lion research.,Perform. Rep., P-R Proj. W-93-R-17, WorkPlan 15, Job 1. New Mex. Game Fish Dep. 18pp.

Presents results of study to estimate population numbers and harvest and to study mountainlion activities. Estimates annual recruitment into breeding population of 15% in NewMexico.

Donaldson, B. 1974. Mountain lion research. Perform. Rep., P-R Proj. W-93-16, Work Plan15, Job 1. New Mex. Game Fish Dep. l1pp.

Presents results of study to estimate population numbers and to study mountain lionactivities.

Eaton, R. L., and J. R. van Oosten. 1974. The status and conservation of North America'scats. Symp. on Endangered and Threatened Species of North America. Wild CanidSurvival and Research Center, Washington, D.C. 339pp.

Discusses endangered and threatened subspecies of cougar.

Eaton, R. L. 1973. The status, management and conservation of the cougar in the UnitedStates. World's Cats 1:68-89.

Presents information on status and management of cougar by state.

Evans, W. 1983. The cougar in New Mexico: biology, status, depredation of livestock, andmanagement recommendations. New Mex. Dept. Game Fish, Sante Fe. 40pp.

Reports that cougars should be managed to maintain social structure. Excessive harvest tocontrol depredation results in large influx of transients and higher reproduction rates.Efforts to control depredation by eliminating large numbers of cougars have failed in NewMexico.

-11-

Fair, J. S. 1977. Utah cougar harvest 1976-1977. Utah Dept. Nat. Res. Div. Wild!. Res.Pub!. No. 77-10. 16pp.

Summarizes cougar management and describes harvest and harvest statistics for 1976-1977season.

Fitzgerald, J. W. 1981. The Utah cougar harvest book 1980-1981. Utah Dept. Nat. Res. Div.Wild!. Res. Pub!. No. 81-11. 19pp.

Sumrr:arizes cougar management and describes harvest and harvest statistics for 1980-1981season.

Fitzhugh, E. L. 1989. Managing with potential for lion attacks against humans. Pp. 74-77in R. H. Smith, ed. Trans. 3rd Mountain Lion Workshop, Arizona Chapter, TheWildlife Society, and Arizona Game and Fish Dept.

Discusses ways to minimize probability of attacks and provides suggestions for studyingattacks.

Fitzhugh, E. L., and K. S. Smallwood. 1989. Techniques for monitoring mountain lionpopulation levels. Pp. 69-71 in R. H. Smith, ed. Trans. 3rd Mountain Lion Workshop,Arizona Chapter, The Wildlife Society and Arizona Game and Fish Dept.

Describes a technique for monitoring lions on a statewide basis by conducting track countson randomly selected units. A statewide track count could be conducted for approximately$1,000 plus 82 man-days of tracking labor.

Fitzhugh, E. L. 1984. Lion track counts in California. Pp. 130- 133 in J. Roberson and F.Lindzey, (eds.). Proc. 2nd Mtn. Lion Workshop. Utah Div. Wild!. Res., Salt LakeCity.

Describes use of track counts to detect lions and describes method to identify individual lionsthrough tracing tracks.

-12-

Fjelline, D. P., and T. M. Mansfield. 1989. Method to standardize the procedure formeasuring mountain lion tracks. Pp. 49-51 in R. H. Smith, ed. Trans. 3rd MountainLion Workshop, Arizona Chapter, The Wildlife Society and Arizona Game and FishDept.

Describes technique to establish a standard procecure for mountain lion track surveys.

Greer, K. G. 1980. Mountain lion studies (1979-1980). W-120-R-11, Study L-Ll, Job 3.Mont. Game Fish Dep. 14pp.

Provides information on harvest statistics, age and sex structure of harvest, parasites, andgives management recommendations.

Greer, K. G. 1979. Mountain lion studies (1978-1979). W-120-R-10, Study L-Ll, Job 3.Mont. Game Fish Dep. 14pp.

"Provides information on harvest statistics, age and sex structure of harvest, parasites, andgives management recommendations.

Greer, K. G. 1978. Mountain lion studies (1977-1978). W-120-R-9, Study L-Ll, Job 3.Mont. Game Fish Dep. 14pp.

Provides information on harvest statistics, age and sex structure of harvest, parasites, andgives management recommendations.

Greer, K. G. 1977. Mountain lion studies (1976-1977). W-120-R-8, Study L-Ll, Job 3.Mont. Game Fish Dep. lOpp.

Provides information on harvest statistics, age and sex structure of harvest, parasites, andgives management recommendations.

Greer, K. G. 1976. Mountain lion studies (1975-1976). W-120-R-7, Work Plan 4, Job L­1.2R. Mont. Game Fish Dep. 16pp.

Provides information on harvest statistics, age and sex structure of harvest. parasites, andgives management recommendations.

I.1"liIiii•••il'lIJl!IJI!IlIIUllilil.lltlil.llil!iJl.illi.iIlIIILI~IIMIIlii1Il;li'III£llliliIlI•••I!tilliiliJlill_!ilIII•••5iIl~_

-13-

Greer, K. G. 1975. Mountain lion studies (1974-1975). W-120-R-6, Work Plan 4, Job L­1.2R. Mont. Game Fish Dep. 10pp.

Provides information on harvest statistics, age and sex structure of harvest, and parasites.

Greer, K. G. 1974. Mountain lion studies (1973-1974). W-120-R-5, Work Plan 4, Job L­1.2R. Mont. Game Fish Dep. 13pp.

Provides information on harvest statistics, age and sex structure of harvest, parasites, andgives management recommendations.

Greer, K. G. 1974. Mountain lion studies (1972-1973). W-120-R-4, Work Plan 4, Job L­1.2R. Mont. Game Fish Dep. 12pp.

Provides information on harvest statistics, age and sex structure of harvest, parasites, andgives management recommendations.

Guenzel, R. J. 1987. Estimating mountain lion depredation on domestic Sheep: a review ofthe literature. Wyo. Game Fish Dep. Unpubl. Rep., Cheyenne. 10pp + attachments.

Describes problem of lion predation on sheep, reviews procedures for estimating proportionof missing sheep killed by lions and gives recommendations for estimating actual losses tolions.

Guggisberg, G. A. W. 1975. Wild cats of the world. Taplinger Pub. Co., New York. 328pp.

Presents chapter on biology and ecology of all cats in the world, including cougar.

Hancock, L. 1980. A history of changing attitudes to Felis collc%r. M.S. Thesis, Simon. Fraser Univ., Vancouver, B.C.

Describes results of survey to determine public attitudes toward cougars.

-14-

Harcombe, D. W. 1976. Oregon cougar study. Oreg. Dept. Fish WildJ. Portland, Oreg. 62pp.

Describes mountain lion habitat use and requirements in Oregon, presents results of liontrack counts.

Harris, C. E. 1991. Mountain lion management plan 1991-1995. Idaho Department of Fishand Game, Boise. 43 pp.

Describes proposed five year management plan for mountain lions in Idaho.

Harrison, S. 1989. Predation rates of cougar within the Junction Wildlife Management Area.P. 48 in R. H. Smith, ed. Trans. 3rd Mountain Lion Workshop, Arizona Chapter, TheWildlife Society, and Arizona Game and Fish Dept.

Provides data on predation rates of female cougars with kittens in British Columbia. Averagepredation rate on bighorn sheep and mule deer was O.7-2.1/week.

Harvey and Stanley Associates, Inc. 1983. Mountain lions (Felis collc%r) in the vicinity ofCarlsbad Caverns National Park, New Mexico and Guadalupe Mountains NationalPark, Texas. UnpubJ. Rep. Submitted to National Park Service, Santa Fe, New Mex.34pp.

Presents data from telemetry study on home range, dispersal, movement, activity, mortality,reproduction, density, scrape sites, and food habits.

Harvey and Stanley Associates, Inc. 1988. !'4ountain lions (Fe/is collc%r) in the vicinity ofCarlsbad Caverns National Park, New Mexico and Guadalupe Mountains NationalPark, Texas: An Ecological Study. Final Rep. Submitted to National Park Service,Santa Fe, New Mex. 137pp.

Presents data from telemetry study on home range, dispersal, movement, activity, mortality,reproduction, density, scrape sites, and food habits.

-15-

Hebert, D. 1989. The status and management of cougar in British Columbia 1988. Pp. 11-14in R.H. Smith, ed. Trans. 3rd Mountain lion workshop, Arizona Chapter, The WildlifeSociety, and Arizona Game and Fish Dept.

Describes results of lion research and lion management in British Columbia. Estimate ofannual harvest is 6-10%. Over a 12 year period harvest composed of 56.3% male and 43.5%female.

Hemker, T. P., F. G. Lindzey, B. B. Ackerman, and A. J. Button. 1986. Survival of cougarcubs in a non-hunted population. Pp 327-332 in Cats of the World: Biology,Conservation, and Management. S. D. Miller and D. D. Everett, eds.

Presents cub survival data based on ten cougar family groups monitored in a non-huntedpopulaiton. Survival of cubs to dispersal was estimated at 67%. Reported that cub mortalityfrom maulings by dogs and orphaning during hunting seasons is probably as significant asadult harvest.

Hemker, T. P., F. G. Lindzey, and B. B. Ackerman. 1984. Population characteristics andmovement patterns of cougars in southern Utah. J. Wildl. Manage. 48:1275-1284.

Describes population dynamics, densities, and movements of mountaion lions in Utah.Dispersal of cubs was found to be independant of adult resident density. Density regulatedby social structure but limited by abundance of mule deer.

Hemker, T. P. 1982. Population characteristics and movement patterns of cougars lD

southern Utah. M.S. Thesis, Utah St. Univ., Logan. 66pp.

Describes population dynamics, densities, and movements of mountaion lions in Utah.Dispersal of cubs was found to be independant of adult resident density. Density regulatedbv social structure but limited bv abundance of mule deer.

~ .

Hess, M. 1981. Trophy big game investigations and hunting season recommendations. Nev.Dept. Wild!. 37pp.

Presents information on hunting pressure, harvest trend, population trends, and managementrecommendations for mountain lions in each of the state management regions.

-16-

Hopkins, R. A, M. J. Kutilek, and G. L. Shreve. 1986. Density and horne rangecharacteristics of mountain lions in the Diablo Range of California. Pp. 223-235 inCats of the World: Biology, Conservation, and Management. S. D. Miller and D. D.Everett, eds.

Presents data on lion horne range and density based on 543 locations of 6 lions.

Hopkins, R. A 1984. Current techniques used in the research of pumas. Pp. 216-229 in J.Roberson and F. Lindzey, (eds.). Proc. 2nd Mtn. Lion Workshop. Utah Div. Wildl.Res., Salt Lake City.

Describes status of methods used in cougar research and outlines research needs.

Hopkins, R. A 1984. The role of the puma in a nonmigratory large mammal communityin the Diablo Range of California. Pp. 134-137 in J. Roberson and F. Lindzey, (eds.).Proc. 2nd Mtn. Lion Workshop. Utah Div. Wildl. Res., Salt Lake City.

Presents results of past studies conducted in Diablo Range and describes proposed study todetermine relationship between mountain lions and their prey.

Hopkins, R. A, M. J. Kutilek, and J. Shreve. 1982. The density and horne rangecharacteristics of mountain lions in the Diablo Range of California. Proc. Intl. CatSymp., Kingsville, Texas. (abstract).

Presents data on lion horne range and density.

Hornocker, M. G., K. M. Murphy, and J. W. Tischendorf. 1989. The ecology of themountain lion (Felis concolor missoulensis) in the North Yellowstone Ecosystem. P.56 in R. H. Smith, ed. Trans. 3rd Mountain Lion Workshop, Arizona Chapter, TheWildlife Society and Arizona Game and Fish Dept.

Presents preliminary results of study intitiated in Yellowstone to document populationparameters, spatial relationships, predator-prey relationships, and impacts of sport harvest.

III!

I,I

I

-17-

Hornocker, M. and T. Bailey. 1986. Natural Regulation in three species of felids. Pp.211­220 in Cats of the World: Biology, Conservation, and Management. S. D. Miller andD. D. Everett, eds.

Describes regulatory mechanisms for mountain lions, bobcats and jaguars. Concludes socialbehavior, not prey abundance, limits density.

Hornocker, M. G. and G. K. Koehler. 1984. 1984. Reintroducing orphaned mountain lionkittens into the wild. Pp. 167-169 in J. Roberson and F. Lindzey, (eds.). Proc. 2ndMtn. Lion Workshop'. Utah Div. Wildl. Res., Sal! Lake City.

Describes fate of 3 lion kittens released at 6-7 months of age. Results implied that lionkittens must be at least this age to survive on their own.

Hornocker, M. G. and T. N. Bailey. 1982. Natural regulation in 3 species of Felidae. Proc.IntI. Cat Symp., Kingsville, Texas. (abstract).

Describes regulatory mechanisms for mountain lions, bobcats and jaguars. Concludes social'behavior, not prey abundance, limits density.

Hornocker, M. G. 1976. The possible influence of the mountain lion on mule deerpopulations. Pp. 107-109 in G.W. Workman and J.B Low, eds. Mule deer decline inthe West: A symposi.um. Utah St. Univ., Utah Agric. S:p. Stn., Logan. 134pp.

Discusses role of predation in general. Conclusion is that there is not enough data yet toevaluate effects of mountain lion predation on mule deer populations, althoug it is likely lionpredation is not a factor in the mule deer decline, but habitat loss is.

Hornocker, M. G. 1971. Suggestions for the management of mountain lions as trophyspecies in the Intermountain region. Annu. Proc. Western Assoc. State Game FishCommissioners 51:399-402.

Summarizes results of mountain lion study in Idaho Primitive Area. Recommends leavingfemale portion of lion population intact, uniform management over large areas, andmanaging lions as a trophy species with mandatory reporting of kills.

-18-

Hornocker, M. G. 1970. An analysis of mountain lion predation upon mule deer and elkin the Idaho primitive area. Wild!. Monogr. 21:1-39.

Presents results of study on lion predation in Idaho primitive areas. Conclusion is that lionsocial structure limits their population size. Elk and deer are limited by winter habitat, notlions.

Hornocker, M. G. 1969. Winter territoriality in mountain lions. J. Wild!. Manage. 33:457­464.

Presents data on home range SIze, factors influencing home range SIze, and mutualavoidance behavior in lions.

Howard, W. E. 1989. Why lions need to be hunted. Pp. 66-68 in R. H. Smith, ed. Trans.3rd Mountain Lion workshop, Arizona Chapter, The Wildlife Society and ArizonaGame and Fish Dept.

Discusses reasons why sport harvest of lions IS necessary to maintain healthy breedingpopulations.

Ingram, R. 1984. Oregon. Pp. 53-55 in J. Roberson and F. Lindzey, (eds.). Proc. 2nd Mtn.Lion Workshop. Utah Div. Wilen. Res., Salt Lake City.

Describes mountain lion status, harvest information, and management programs in Oregon.

Jalkotzy, M. G., and P. 1. Ross. 1989. Population ecology of cougars in southwesternAlberta. P. 47 in R. H. Smith, ed. Trans. 3rd Mountain lion workshop, ArizonaChapter, The Wildlife Society and Arizona Game and Fish Dept.

Provides information on litter size, birth peak, age of independence, population structureand density of a mountain lion population in Alberta.

-19-

Jalkotzy, M. G. and P. 1. Ross. 1989. Cougar hunting regulations and harvest in Albertabetween 1973 and 1987. P. 6 in R. H. Smith, ed. Trans. 3rd Mountain LionWorkshop, Arizona Chapter, The Wildlife Society and Arizona Game and Fish Dept.

Describes regulations and effects of harvest of lions in Alberta. Reducing season length insome areas was found to increase female harvest. Apparently, lion hunters are less selectiveduring short seasons.

Johnson, J. F. 1982. Mountain lion research. Job Final Rep. P.R. Proj. W-124-R-4. NewMexico Dept. Game Fish. Santa Fe. 16pp.

Describes home range size and overlap, daily activities and movement patterns, and dispersalof mountain lions in New Mexico.

Johnson, K. A., and W. L. Franklin. 1984. Ecology and management of the Patagonia puma(Felis concolor patagonica) in southern Chile. Pp. 141-146 in J. Roberson and F.Lindzey, (eds.). Proc. 2nd Mtn. Lion Workshop. Utah Div. Wild!. Res., Salt LakeCity.

Describes proposed research to investigate ecology of a protected cougar population inChile.

Jorgensen S. E., and L. D. Mech, eds. 1971. Proc. Symp. on the Native Cats of NorthAmerica: Their Status and Management. U.S. Fish Wild!. SerV., Bur. Sport FishWild!., Twin Cities, Minn. 139pp.

Summarizes status, management and conservation of cougars by state and Canadianprovince.

Koehler, G. M., and M. G. Hornocker. 1986. A preliminary survey of mountain lions inYellowstone National Park. Unpub!. Rep. Wildlife Research Institute, Moscow,Idaho. 12pp.

Describes results of cursory survey of Yellowstone in winter for lion sign. Thirteen sets oftracks, kills, scats, and scrapes found.

-20-

Koford, C. B. 1978. The welfare of the puma in California. Carnivore 1:92·96.

Presents data on location and relative abundacne of cougar populations in California basedon kill data and sightings.

Kohls, R. 1989. Mountain lion law enforcement problems in Arizona. P. 42 in R.H. Smith,ed. Trans. 3rd Mountain lion workshop, Arizona Chapter, The Wildlife Society, andArizona Game and Fish Dept.

Discusses prob!ems involved in enforcing mountain lion regulations and suggests theseproblems be considered when formulating management recommendations.

Kutilek, M. J., T. E. Smith, R. A. Hopkins, and E. W. Clinite. 1980. California Mountain liontrack transect survey, 1980. Harvey and Stanley Assoc., Inc. Alviso, Calif. A reportto Calif. Dept. Fish Game. 30 pp mimeo.

Presents results of lion track surveys conducted In four national forests -in northernCalifornia.

Kutilek, M. J., R. A. Hopkins, and T. E. Smith. 1980. Second annual progress report on theecology of mountain lions (Felis concolor) in the Diablo Range of California. Dept.BioI. Sci:, San Jose St. Univ., 21pp.

Provides information on mountain lion population dynamics and density.

Kutilek, M. J., R. A. Hopkins, and T. E. Smith. 1979. Progress report on the ecology ofmountain lions (Felis concolor) in the Diablo Range of California. Dept. BioI. Sci.,San Jose St. Univ., 21pp.

Provides information on mountain lion population dynamics and density.

Laing, S. P. 1988. Cougar habitat selection and spatial use patterns in southern Utah. M.S.Thesis. Univ. Wyo., Laramie. 68pp.

Describes mountain lion habitat use based on 3580 locations of 52 radiomarked cougars.

-21-

Leatham, J. P. 1983. The Utah cougar harvest book 1982-1983. Utah Dept. Nat. Res. Div.Wild!. Res. Pub!. No. 83-11. 20pp.

Summarizes cougar management and describes harvest and harvest statistics for 1982-1983season.

Lindzey, F. G., W. D. Van Sickle, S. P. Laing, and C. Mecham. In Press. Simulated cougarharvest in southern Utah. J. Wild!. Manage.

J:?iscusses population effects resulting from a 27% simulated harvest of cougars in Utah.

Lindzey, F. G., B. B. Ackerman, D. Barnhurst, T. Becker, T. P. Hemker, S. P. Laing, C.Mecham, and W. D. Van Sickle. 1989. Boulder-Escalante Cougar Project FinalReport. Utah Div. Wild!. Res., Salt Lake City, Utah. 92pp.

Presents information on population characteristics, movements and dispersal patterns,replacement of residents, food habits, energetics, cougar-mule dlOer interactions, habitat use,and cougar harvest in the Boulder-Escalante area of Utah.

Lindzey, F. G., B. B. Ackerman, D. Barnhurst, and T. P. Hemker. 1988. Survival rates ofmountain lions in southern Utah. 1. Wild!. Manage. 52:664-667.

Lion annual survival rate in Utah study area estimated at 71-73%. Hunting mortality is notconsidered compensatory.

Lindzey, F. G. and W. Van Sickle. 1989. Mountain lion population dynamics (Utah). P. 56in R. H. Smith, ed. Trans. 3rd Mountain lion workshop, Arizona Chapter, TheWildlife Society and Arizona Game and Fish Dept.

Discusses effects of 30% removal to stimulate harvest in Utah.

I

-22-

Lindzey, F. 1987. Mountain Lion. Pp. 656-668 in Wild Furbearer Management andConservation in North America. M. Novak, J.A Baker, M.E. Obbard, and B.Malloch, eds.

Review of mountain lion literature including distribution, life history, ecology, food habitsand management.

Logan, K A, L. L. Sweanor, J. F. Smith, B. R. Spreadbury, and M. G. Hornocker. 1990.Ecology of an unexploited mountain lion population in a desert environment.Wildlife Research Institute Unpubl. Rep. to New Mexico Dept. Game Fish, Santa Fe.BOpp.

Describes population characteristics of an unexploited mountain lion population in NewMexico. Results are based on 50 radiomarked lions located over 5,000 times.

Logan, K A, L. L. Sweanor, J. F. Smith, B. R. Spreadbury and M. G. Hornocker. 1989.Ecology of an unexploited cougar population in a desert environment. P. 74 in R.H. Smith, ed. Trans. 3rd Mountain Lion Workshop, Arizona Chapter, The WildlifeSociety and Arizona Game and Fish Dept.

Provides preliminary information on population characteristics of an unexploited mountainlion population in New Mexico.

Logan, K, F. Smith, and L. Sweanor. 1986. Ecology of an unexploited cougar populationin a desert environment. First Annual Report to New Mexico Dep. Game and Fish.Wildl. Res. Instit., Moscow, Idaho. 43pp.

Provides preliminary information on population characteristics of an unexploited lionpopUlation in New Mexico. Results are from caputre and marking of 57 cougar.

Logan, K. A., L. L. Irwin, and R. Skinner. 1986. Characteristics of a hunted mountain lionpopulation in Wyoming. J. Wildl. Manage. 50:648-654.

Presents data on density, home range size, sex and age structure, dispersal, and mortality oflion populatio in Bighorn Mountains of Wyoming.

-23-

Logan, K. A. and L. L. Irwin. 1985. Mountain lion habitats in the Big Horn Mountains,Wyoming. WildJ. Soc. Bull. 13:257-262.

Describes habitat use versus availability based on 191 track locations, 118 radiolocations, 52cache sites and 84 scratch sites.

Logan, K. A., and L. L. Irwin. 1984. Mountain lion population and habitat characteristicsin the Big Horn Mountains, Wyoming. Pp. 189-190 in J. Roberson and F. Lindzey,(eds.). Proc. 2nd Mtn. Lion Workshop. Utah Div. Wild!. Res., Salt Lake City.

Describes population characteristics (density, home range, age and sex structure, dispersal,and mortality) and habitat use of mountain lion population in Wyoming.

Logan, K. A. 1983. Mountain lion population and habitat characteristics in the Big HornMountains of Wyoming. M.s. Thesis, Univ. Wyo., Laramie. 101pp.

Compares habitat use versus availability, describes population dynamics, density, andmortality, provides management recommendations.

MacGregor, W. G. 1972. The changing status of the mountain lion in California. Procs.West. Assoc. State Game Fish Comm. 259-263.

Discusses status, history, and trends; provides a populatiDn estimate for state and presentspreliminary information from radiotelemetry study.

Maehr, D. S., J. W. McCov.TI, E. D. Land, and J. C. Roof. 1989. Panther research updatefor southwest Florida. P. 52 in R. H. Smith, ed. Trans. 3rd Mountain Lion Workshop,Arizona Chapter, The Wildlife Society and Arizona Game and Fish Dept.

Presents results of study to determine panther horne range size and prey (deer) populationcharacteristics.

-24-

Mansfield, T. M., and R A. Weaver. 1989. The status of mountain lions in California. Pp.15-18 in RH. Smith, ed. Trans. 3rd Mountain lion workshop, Arizona Chapter, TheWildlife Society and Arizona Game and Fish Dept.

Discusses status of lions in California and presents research results. The most commmonelement of mountain lion habitat in California is presence of mule deer. Mountain liondensities (adults/lOO mi2) in 4 study areas were 6-9, 5-10, 5-7, and 3-6.

Mansfield, T. M. 1989. Legal challenge to mountain lion hunting in California. P. 42 in R.H.Smith, ed. Trans. 3rd Mountain lion workshop, Arizona Chapter, The Wildlife Societyand Arizona Game and Fish Dept.

Discusses issue of mountain lion hunting in California and California Department of Fishand Game response to legal challenge.

Mansfield, T. M. 1986. Mountain lion management in California. Trans. N. Am. Wildl. Nat.Resour. Conf. 51:178-182.

Presents results of mountain lion research conducted in Calilfornia and describesmanagement. Trends are determined by number of damage complaints, number killed fordamage, and number killed on roads.

McBride, R T. 1976. The status and ecology of the mountain lion Felis concolor sianleyanaof the Texas-Mexico border. M.S. Thesis, SuI Ross St. Univ., Alpine, Texas.

Presents information on mountain lion ecology in Texas from radiotelemetry study.

Morrison, B. 1984. New Mexico. Pp. 49-52 in J. Roberson and F. Lindzey, (eds.). Proc. 2ndMtn. Lion Workshop. Utah Div. WildJ. Res., Salt Lake City.

Review mountain lion status, harvest information, and management and research programsin New Mexico.

-25-

Morse, S. C. 1989. Begging the question: what is mountain lion management. Pp. 3-4 inR.H. Smith, ed. Trans. 3rd Mountain lion workshop, Arizona Chapter, The WildlifeSociety and Arizona Game and Fish Dept.

Philosophical discussion of mountain lion management.

Mountain Lion Preservation Foundation. 1987. Summary of information fromquestionnaires mailed to mountain lion experts. Mountaion Lion PreservationFoundation, Sacramento, California. llpp.

Presents results of a survey of 15 mountain lion experts covering census techniques andstatus of lions in their respective states.

Murphy, K. M., and G. S. Felzien. 1991. The ecology of the mountain lion (Felis concolormissoulensis) in the northern Yellowstone Ecosystem. Cumulative Progress Rept. No.4. Wildlife Research Institute, Moscow, Idaho. 17pp.

Presents data on abundance, rate of increase, sex and age structure, home ranges andmovements, mortality and survival, and predation dynamics from mountain lionradiotelemetry study in north Yellowstone.

Murphy, K. M. 1984. Montana (Status Reports). Pp.39-43 in J. Roberson and F. Lindzey,(eds.). Proc. 2nd Mtn. Lion Workshop. Utah Div. Wildl. Res., Salt Lake City.

Describes status of mountain lions in Montana. The most serious management. problemconfronting biologists is avoiding overkill in lion habitats that are easily accessed by roads.

Murphy, K. M. 1984. Montana (Research Reports). Pp. 172-173 in J. Roberson and F.Lindzey, (eds.). Proc. 2nd Mtn. Lion Workshop. Utah Div. Wildl. Res., Salt LakeCity.

Presents results of study to determine relationships between hunting pressure and mountainlion population characteristics.

r ""'~1l!""_l'Ii!!'"1!lIt."_""__~~"B'~Jfu}k~"imTI&~4.m~it*~~1fi~tm-~t

-26-

Murphy, K. M., G. R. Power, and J. J. Beecham. 1984. Characteristics of exploited lionpopulations and hunting pressure relation-ships in central Idaho and westernMontana. Procs. 1984 Predator Symposium, C. J. Jonkel and C. L. Markum, eds.Univ. Mont. Forestry School, Missoula.

Report describing population dynamics, harvest, effects of harvest, and factors contributingto harvest of a mountain lion population in Montana.

Murphy, K. M. 1983. Relationships between a mountain lion population and huntingpressure in western Montana. Montana Dept. Fish, WildJ. Parks Proj. W-I20-R-l3and 14. 48pp.

Report describing population dynamics, harvest, effects of harvest, and factors contributingto harvest of a mountain lion population in Montana.

Murphy, K. M. 1983. Relationships between a mountain lion population and huntingpressure in western Montana. M.S. Thesis. Univ. Montana. 45pp.

Report describing population dynamics, harvest, effects of harvest, and factors contributingto harvest of a mountain lion population in Montana.

Neal, D. L. 1989. Mountain lion density. and movement in the central Sierra Nevada. P.72in R. H. Smith, ed. Trans. 3rd Mountain lion workshop, Arizona Chapter, TheWildlife Society and Arizona Game and Fish Dept.

Provides information on adult and kitten density and movements of a lion population inCalifornia.

Neal, D. L., G. N. Steger, and R. C. Bertram. 1987. Mountain Lions: preliminary findingson home-range use and density in the central Sierra Nevada. U.S. For. Servo Res.

~ "Note PSW-392. 6pp.

Provides home range and density information for a California lion population.

-27-

Neal, D. L., G. N. Steger and R. C. Bertram. 1986. A preliminary report on mountain lionhome-range use and density in the central Sierra Nevada. Draft Manusc. PacificSouthwest For. Ran. Exper. Stn., Fresno, Calif. 27pp.

Provides home range and density data. Maximum density in the study area was 1/19.9 km2.

Neal, D. L. 1984. The effect of mountain lion predation on the North Kings deer herd. Pp.138-140 in J. Roberson and F. Lindzey, (eds.). Proc. 2nd Mtn. Lion Workshop. UtahDiv. Wildl. Res., Salt Lake City.

Lions were found to keep low population of deer from increasing. Estimated one lion forevery 75 deer in study area.

Nero, R. W., and R. E. Wrigley. 1977. Status and habits of the cougar in Manitoba. Can.Field-Nat. 91:28-40.

Updates status of cougar in Manitoba; describes habitat, prey, distribution and estimatespopulation size of 50 in province.

Norelius, S. and M. Bruscino. 1988. Domestic sheep predation from mountain lions in thesouthern Bighorns: A field study and literature re\~ew. Wyoming Game Fish Dep.,Cheyenne. 42pp.

Presents results of intensive searches for lion-killed sheep. Provides recommendations toreduce losses.

Nowak, R. M. 1976. The cougar in the United States and Canada. U.s. Dept. Inter., FishWildl. Serv., Washington, D.C. and New York Zool. Soc., New York. 190pp.

Discusses history, status, subspecies and distribution of cougar in the United States andCanada.

-28-

Pall, 0., M. Jalkotzy, and 1. Ross. 1988. The cougar in Alberta. Fish Wild!. Div. AlbertaFor. Lands Wild!. 14Spp.

Presents data on population age and sex structure, density, and mortality. The only annualrecruitment rate estimated for a cougar population is presented (1.0-1.2 kittens/female/year).

Pall, O. 1984. Alberta. Pp. 1-8 in J. Roberson and F. Lindzey, (eds.). Proc. 2nd Mtn. LionWorkshop. Utah Div. Wild!. Res., Salt Lake City.

Describes status of mountain lions in Alberta. Alberta harvests what they believe to be 10%of population, which they consider a conservative harvest.

Pall, O. 1984. The population status of cougars near Sheep River, Alberta. Pp. 116-118 inJ. Roberson and F. Lindzey, (eds.). Proc. 2nd Mtn. Lion Workshop. Utah Div. Wild!.Res., Salt Lake City.

Provides preliminary 'results of cougar telemetry study in Alberta.

Pall, O. and M. Jalkotzy. 1983. The population status of cougars near Sheep River, Alberta,1982-1983. Progress Report. Alberta Fish Game Wildl. Div., Calgary. Unpubl. Rep.38pp.

Provides preliminary results of cougar telemetry study in Alberta.

Pence, D. B., R. J. Warren, D. Waid and M. J. Bavin. 1987. Aspects of the ecology ofmountain lions (Felis concolor) in Big Bend National Park. Final rep. to Natl. ParkServ., Santa Fe, New Mexico. 81pp.

Presents results of 2-year study to determine mountain lion movements, home ranges, foodhabits, and population size in Big Bend National Park.

-29-

Phelps, J. S. 1989. Status of mountain lions in Arizona. Pp. 7-9 in RH. Smith, ed. Trans.3rd Mountain Lion Workshop, Arizona Game and Fish Dept.

Discusses results of mountain lion research in Arizona. Reports that no reliable methodexists to age wild mountain lions. Track counts indicated lion densities directly related tomule deer density.

Poelker, R J. 1976. The status and management of the puma in Washington. World's cats.3:20-27.

Discusses history, status and current management of cougar in Washington.

Power, G. D. 1985. Mountain lion management plan 1986-1990. Idaho Dept. Fish Game.16pp.

Discusses mountain lion management proposed for Idaho. The management plan calls formaintaing harvest age ratios within 5% of current levels (35% < 3yrs, 44% 4-8 yrs, 21 % >8 yrs). Because harvest is determined by snow conditions, they recommend managing on 3­year running averages rather than year to year to reduce variability.

Quigley, H. B., G. M. Koehler, and M. G. Hornocker. 1989. Dynamics of a mountain lionpopulation in central Idaho over a 20-year period. P. 54 in R H. Smith, eg. Trans.3rd Mountain lion workshop, Arizona Chapter, The Wildlife Society and ArizonaGame and Fish Dept.

Presents age and sex structure and population growth rates of mountain lions inhabitingcentral Idaho.

Rieck, J. M. 1989. Status of the cougar in Washington. Pp. 35-37 in R H. Smith, ed. Trans.3rd Mountain lion workshop, Arizona Chapter, The Wildlife Society and ArizonaGame and Fish Dept.

Presents status, management programs, research and economic values of cougars mWashington.

Ill!lf••_"o!i"'''~ .~_~!I1\f!l1I_J!i{

~lr~lf&~.w~~~~I~l~~~~fi&i'b~i~~;

-30-

Roberson, J. 1984. Utah mountain lion status report. Utah Div. Wildl. Res. 23pp.

Describes status of mountain lions in Utah. Discusses problem with kitten mortality due toorphaning and mauling by dogs during hunting seasons.

Roberson, J. 1984. Pursuit seasons. Pp. 191-203 in J. Roberson and F. Lindzey, (eds.). Froc.2nd Mtn. Lion Workshop. Utah Div. Wildl. Res., Salt Lake City.

Discussion of advantages and disadvantages of cougar pursuit seasons.

Roberson, J. and F. Lindzey, (eds.). 1984. Proc. 2nd Mtn. Lion Workshop. Utah Div. Wild!.Res., Salt Lake City.

Compilation of status reports, questionnaire responses, and research reports on cougars fromstate and provinical agencies.

Roberson, J. 1984. Utah. Pp. 60-79 in J. Roberson and F. Lindzey, (eds.). Proc. 2nd Mtn.Lion Workshop. Utah Div. Wild!. Res., Salt Lake City.

Provides information on cougar status, depredation, harvest and research in Utah.

Robertson, M. and M. Bell. 1989. Status of the mountain lion in New Mexico. Pp. 24-25 inR. H. Smith, ed. Trans. 3rd Mountain lion workshop, Arizona Chapter, The WildlifeSociety and Arizona Game and Fish Dept.

Discusses population levels, management, research and economic value of mountain lionsin New Mexico.

Robinette, W. L., N. V. Hancock, and D. A. Jones. 1977. The Oak Creek mule deer herdin Utah. Utah Div. Wild!. Res. Pub!. 77-15. 148pp.

Describes unsuccessful attempt to determine effects of cougar predation on deer population.Cougar density estimated at 1/22-29 square miles. Harvest of 50% or more of cougarpopulation did not reduce number of cougar-killed deer carcasses found on study area.Estimated annual recruitment and mortality of 32%.

-31-

Robinette, W. L., J. S. Gashwiler, and O. W. Morris. 1961. Notes on cougar productivityand life history. J. Mammal. 42: 204-217.

Presents data on Utah and Nevada cougar birth peaks, pregnance rate, interbirth intervals,and litter size.

Roop, L. 1971. The Wyoming mountain lion situation. Wyoming Wildl. 35:16-21.

Describes lion presence, distribution, abundance and welfare in Wyoming.

Ross, I. and M. Jalkotzy. 1989. The Sheep River cougar project phase II. Final Report1987-1989. Arc Associated Resource Consultants, Ltd. Calgary, Alberta. 56pp.

Describes population dynamics, including recruitment, mortality, and density of an intensivelymonitored cougar population in Alberta.

Ross, 1. and M. Jalkotzy. 1988. The Sheep River cougar project phase II. 1987-1988 Prog.Rep. Arc Associated Resource Consultants, Ltd. Calgary, Alberta. 21pp.

Describes preliminary results from an intensive study of cougar population dynamics inAlberta.

Russ, W. 1990. Mountain lion status survey. Fed AId Job Perf. Rep. W-125-R-1, Job 69.Texas Parks Wildl. Dep., Austin, Tex. 7pp.

Describes lion distribution in Texas based on documented sightingsand mortalities.

Russ, W. 1989. Mountain lion status survey. Fed AId Job Final Rep. W-103-R-19, Job 69.Texas Parks Wildl. Dep., Austin, Tex. llpp.

Describes lion distribution in Texas based on documented sightingsand mortalities.

-32-

Russ, W. B. 1989. Status of the mountain lion in Texas. Pp. 30-31 in R.H. Smith, ed. Trans.3rd Mountain Lion Workshop, Arizona Chapter, The WiJdife Society and ArizonaGame and Fish Dept.

Discusses current status, management program, research and economic values of mountainlions in Texas.

Russell, K. R. 1978. Mountain lion. Pp. 207-225 in J. L. Schmidt and D. L. Gilbert, eds. BigGame of North America: ecology and Management. Stackpole Co., Harrisburg, Pa.494pp.

Review of mountain lion literature.

Russo, J. P. and 1. N. Carr. 1970. Status of the lion in Arizona and its future. West. Assoc.St. Game Fish Comm. 50:387-403.

Discusses status, subspecies present in state, history of management, and providesrecommendations for future management.

Sandfort, W. W. and R. J. Tully. 1971. The status and management of the mountain lionand bobcat in Colorado. Pp. 72-85 in S. E. Jorgensen and L. D. Mech, eds. Proc.Symp. on the Native Cats of North America: Their status and management. U.S. FishWild!. Serv., Bur. Sport Fish. Wild!., Twin Cities, Minn. 139pp.

Reviews status and management of mountain lions in Colorado.

Schulenberg, B. 1984. California. Pp. 23-26 in J. Roberson and F. Lindzey, (eds.). Proc. 2ndMtn. Lion Workshop. Utah Div. Wildl. Res., Salt Lake City.

Discusses mountain lion status and research in California. They estimate a current annualgrowth of 8% with no hunting. Road transects are conducted every 2-3 years to monitortrends.

"1l~••••iliilll!l!i!i!IlI-I""••!i!i!.IllII!i!i!i!iI.liIM.I£lilll.iliiiIHli••••IiiiiI.IJlJIII~I•••iIlll.•llil.IIIIII••~

-33-

Seidensticker, J. c., IV. 1973. Mountain lion social organization in th9 Idaho primitive area.Ph.D Diss. Univ. Idaho. 161pp.

Discusses territory size and overlap, role of territory in breeding, and home range dynamics.

Seidensticker, J. c., IV., M. G. Hornocker, W. V. Wiles and J. F. Messick. 1973. Mountainlion social organization in the Idaho primitive area. Wild!. Monogr. 35. 60pp.

Describes effects of residency on breeding status,presents data on home range size andinfluences on size; discusses land tenure system for maintaining population size.

Shaw, H. G. 1990. Preliminary recommendations for mountain lion track counts in Arizona.Unpub!. Rept. General Wildlife Services, Chino Valley, Arizona. 7pp.

Describes need to monitor lion populations statewide to defend management practices.Provides recommendations for conducting track counts to monitor lion trends.

Shaw, H. G. 1989. Soul among lions: the cougar as peaceful adversary. Johnson Books,Boulder, Colo. 140pp.

Popular literature describing cougar biology and author's experience with cougars in Arizona.

Shaw, H. G., N. G. Woolsey, J. R. Wegge and R. L. Day, Jr. 1988. Factors affectingmountain lion densities and cattle depredation in Arizona: A final Report. Ariz.Game Fish Dep. Proj. W-78-R, Work Plan 2, Job 29. 16pp.

Describes results of 144 track count routes conducted in Arizona. Lion track densities wererelated to mule deer density and habitat. Recommended periodic track surveys to monitorlion trends and discussed methods of conducting the counts.

Shaw, H. G. 1984. Cattle growers and lions. Pp. 119-129 in J. Roberson and F. Lindzey,(eds.). Proc. 2nd Mtn. Lion Workshop. Utah Div. Wild!. Res., Salt Lake City.

Presents results of rancher-mountain lion problem mail survey distributed to Arizona CattleGrowers.

-34-

Shaw, H. G. 1980. Ecology of the mountain lion in Arizona. Final Rep., P-R Proj. W-78-R,Work Plan 2, Job 13. Ariz. Game Fish Dep. 14pp.

Describes use of track counts to estimate lion abundance and gives recommended samplesizes. Provides data on mortality and effects of harvest on population in North Kaibab.

Shaw, H. G. 1979. A mountain lion field guide. Spec. Rep. No.9. Ariz. Game Fish Dep.27pp.

Field guide to identification of lion sign, lion kills and provides methods to determine lionpopulation characteristics.

Shaw, H. G. 1977. Impact of mountain lion on mule deer and cattle in northwesternArizona. Pp. 17-32 in R. L. Phillips and C. Jonkel, eds. Proc. 1975 Predator Symp.Univ. Mont., Missoula.

Provides information on food habits based on scat analysis and kill locations; estimatesnumber of prey animals killed per cougar.

Sheriff, S. L. 1978. Computer model for mountain lion populations. M.S. Thesis. Colo. St.Univ., Ft. Collins. 287pp.

Describes mountain lion model derived from ungulate model ONEPOP and presents resultsof model validation.

Shorma, G. 1988. Report on mountain lion status, management, research, and economicvalues. Wyo. Game Fish Dep. Unpub!. Rep., Cheyenne.

Reports that Wyoming lion population appears to be increasing. Provides data on harvestand hunter success for 1983-1987.

-35-

;horma, G. 1989. Status of the mountain lion in Wyoming. Pp. 38-39 in R. H. Smith, ed.Trans. 3rd Mountain lion workshop. Arizona Chapter, The Wildlife Society andArizona Game and Fish Dept.

Discusses current status, management programs, research and economic values of Wyomingmountain lions.

::>itton, L. W. 1977. California mountain lion investigations with recommendations formanagement. Final Rep. to the State Legislature, P-R Proj. W-51-R, Sacramento:California Fish Game Dep. 35pp.

Reviews mountain lion research conducted in California. Recommends track surveys every5 years to monitor statewide trends.

Smallwood, K. S., and E. L. Fitzhugh. 1989. Differentiating mountain lion and dog tracks.Pp. 58-63 in R. H. Smith, ed. Trans. 3rd Mountain lion workshop, Arizona Chapter,The Wildlife Society and Arizona Game and Fish Dept.

Presents ways to discriminate dog from lion tracks when conducting lion track surveys.

Smallwood, K. S. and E. L. Fitzhugh. 1987. A statewide mountain lion population indextechnique. Wildl. Ext., Univ. Calif., Davis. 26pp.

Describes a lion track survey used to index lion populations in California in 1985 and 1986.

Smith, R. H., ed. 1989. Trans. 3rd Mountain lion workshop. Arizona Game and Fish Dept.88pp.

Presents state and provincial status reports, legal problems in mountain lion management,research reports, and population monitoring and harvest strategies.

-36-

Smith, T., R. Duke, and M. Kutilek. 1988. Mountain lion population trends monitoring inCarlsbad Caverns and Guadalupe Mountains National Parks. Harvey and StanleyAssociates, Inc. Report to National Park Service. 33pp.

Presents a technique to monitor lion trends in the National Parks using mountain lion sign(tracks, scats, scrapes).

Smith, T. E., R. R. Duke, M. J. Kutilek, and J. Walters. 1989. A non invasive method formonitoring mountain lion population trends in roadless areas. P. 73 in R. H. Smith,ed. Trans. 3rd Mountain Lion Workshop, Arizona Chapter, The Wildlife Society andArizona Game and Fish Dept.

Describes a technique to monitor lion population trends by recording tracks, scats, scrapes,and prey remains during biannual surveys of watersheds.

Smith, T. E., R. R. Duke, M. J. Kutilek, and H. T. Harvey. 1986. Mountain lions in thevicinity of Carlsbad Caverns and Guadalupe Mountains National Parks - Anecological study. Rep. to Natl. Park Ser. 137pp.

Presents data on home range, dispersal and movements, activity, mortality, reproduction,density, and food habits.

Smith, T. E., R. R. Duke, and M. J. Kutilek. 1984. The ecology of the mountain lion in theGuadalupe Mountains of Texas and New Mexico. Pp. 174-175 in J. Roberson andF. Lindzey, (eds.). Proc. 2nd Mtn. Lion Workshop. Utah Div. Wildl. Res., Salt LakeCity. . .

Provides data on density, food habits, and mortality of a mountain lion population in Texasand New Mexico.

Spreadbury, B. 1989. Cougar ecolo!,')' and related management implications and strategiesin southeastern British Columbia. M.s. Thesis, Univ. Calgary, Alberta. l05pp.

Presents results of telemetry study in British Columbia and gives management implicationsand recommendations.

-37-

Spreadbury, B. 1989. Cougar ecology in southeastern British Columbia. P. 48 in R. H.Smith, ed. Trans. 3rd Mountain Lion Workshop, Arizona Chapter, The WildlifeSociety and Arizona Game and Fish Dept.

Presents results of 3-year study conducted in British Columbia, including density, populationstructure, and litter sizes.

Steger, G. N. 1989. Movement and survival of 14 month old orphaned mountain lionkittens. P. 73 in R. H. Smith, ed. Trans. 3rd Mountain Lion Workshop, ArizonaChapter, The Wildlife -Society and Arizona Game and Fish Dept.

Two female kittens survived being orphaned at 14 months of age and successfully establishedresidence.

Stiver, S. 1. 1989. Status of mountain lions in Nevada. Pp. 26-29 in R. H. Smith, ed. Trans.3rd Mountain lion workshop, Arizona Chapter, The Wildlife Society and ArizonaGame and Fish Dept.

Presents status, management programs, research results and economic values of mountainlions in Arizona.

Strickland, D. 1980. Mountain lion populations. Wyoming Game Fish Dep. Unpub!. Rep.,Cheyenne. 5pp.

Presents method to estimate Wyoming lion population based on habitat and harvest andsighting data, gives recommended mortality quotas by management unit.

Suminski, H. R. 1982. Mountain lion predation on domestic livestock in Nevada. Proc.Vertebr. Pest Conf. 10:62-66.

Discusses depredation problem and estimates annual livestock losses.

Tanner, 1. T. 1975. The stability and the intrinsic gro"1h rates of prey and predatorpopulations. Ecology 56:855-867.

Calculates survival rates of cougars.

'.--=:;-'_@l~~~

'·1

-38-

Thompson, B. C. 1984. Texas. Pp. 56-59 in J. Roberson and F. Lindzey, (eds.). Proc. 2ndMtn. Lion Workshop. Utah Div. WildJ. Res., Salt Lake City.

Provides mountain lion status, harvest information, and research programs in Texas.

, ToweiIJ, D. E., C. Maser, L. D. Bryant, and M. L. Johnson. 1988. Reproductivecharacteristics of eastern Oregon cougars. Northwest Sci. 62:147-150.

Reports reproductive characteristics of 46 male and 51 female cougars killed by hunters inOregon in December. Mean litter size based on placental scars is 2.4.

Toweill, D. E., C. Maser, M. L. Johnson, and L. D. Bryant. 1984. Size and reproductivecharacteristics of western Oregon cougars. Pp. 176-184 in J. Roberson and F. Lindzey,(eds.). Proc. 2nd Mtn. Lion Workshop. Utah Div. WildJ. Res., Salt Lake City.

Presents sizes, reproductive status, and reproductive characteristics of 87 cougars killed byhunters in Oregon's Cascade range.

Trainer, C. E., and G. Matson. 1989. Age determination in cougar from cementum annulicounts of tooth sections. P. 71 in J. H. Smith, ed. Trans. 3rd Mountain LionWorkshop, Arizona Chapter, The Wildlife Society and Arizona Game and Fish Dept.

Discusses accuracy of aging mountain lions using canine and premolar cementum annuli.

TruJio, L. A 1989. What mountain lion scientists say about their research: results of asurvey on mountain lion research methods. Mountain Lion Preservation Foundation.34pp.

Describes results of a survey of 30 mountain lion researchers onmountain lion research techniques and data analysis.

--------------..........-"'....- ...~m.i J J;·gi._~g;

-39-

Trulio, L. A. 1989. Preliminary results: Survey on mountain lion research methods. Pp. 44­46 in R. H. Smith, ed. Trans. 3rd mountain lion workshop, Arizona Chapter, TheWildlife Society and Arizona Game and Fish Dept.

Describes preliminary results of a survey of 30 mountain lion researchers on mountain lionresearch techniques and data analysis.

Tsukamoto, G. 1984. Nevada. Pp. 44-48 in J. Roberson and F. Lindzey, (eds.). Proc. 2ndMtn. Lion Workshop. Utah Div. Wildl. Res., Salt Lake City.

Describes status of mountain lions in Nevada. Harvest objectives are determinoo byestimating the lion population in a mountain range and setting a 25-30% harvest rate.Major management problems are population estimation refinement, lion depredationanalysis and research, and population monitoring.

Turbak, G. 1986. America's great cats. Northland Press, Flagstaff, Ariz. 77pp.

Popular literature including chapter on cougar biology, ecology, and research.

Van Dyke, F. G., and R. H. Brocke. 1987. Sighting and track reports as indices of mountainlion presence. Wildl. Soc. Bull. 15:251-256.

Evaluates reliability of mountain lion sightings reported by lion hunters, deer hunters andcampers, and concludes that sightings alone should never be used to describe mountain liondistribution.

Van Dyke, F. G., and R. H. Brocke. 1987. Searching technique for mountain lion sign atspecific locations. Wildl. Soc. Bull. 15:256-259.

Describes index developed to evaluate reliability of lion sightings submitted by public.

Van Dyke, F. G., H. B. Brocke, and H. G. Shaw. 1986. Use of road track counts as indicesof mountain lion presence. J. Wildl. Manage. 50:102-109.

Describes use of track counts to survey lion popUlations. Track frequency found to berelated to lion density.

-40-

Van Dyke, F. G., H. B. Rainer, H. G. Shaw, B. B. Ackerman, T. P. Hemker, and F. G.Lindzey. 1986. Reactions of mountain lions to logging and human activity. J. Wild!.Manage. 50:95-102.

Mountain lions avoid logged areas, select residence in areas with less roads than average,and avoid human residences. Transient lions most likely segment of population impactedby human activity.

Van Sickle, W. 1990. Methods for estimating cougar rumbers in southern Utah. M.S.Thesis. Univ. Wyo., Laramie. 73pp.

Describes results of census techniques including roadside track counts, capture/recapture,and aerial surveys for tracks in snow. Aerial surveys considered viable method for estimatingcougar numbers.

Van Sickle, W. D., and F. G. Lindzey. 1991. Evaluation of a cougar population estimatorbased on probability sampling. J. Wild!. Manage. 55:738-743.

Describes use of probability sampling of snow tracks located using helicopters as a methodof estimating cougar numbers.

Waid, D. D., D. B. P.ence, and R. J. Warren. 1985. The ecology of mountain lions in BigBend National Park. Research Highlights - 1985 Noxious Brush and Weed Control,Range and Wildlife, Tex. Tech. Univ. 16:35-36.

Weaver, R. A., and L. Sitton. 1978. Changing status of mountain lion in California andlivestock depredation problems. Proc. Vertebr. Pest Conf. 8:214-219.

Reviews mountain lion management in California and discusses livestock depredationproblem.

Weaver, D. 1976. Synopsis of California Mountain lion study. Pp 12-15 in G. C. Christensenand R. J. Fischer, co-chairmen. Trans. Mountain Lion Workshop. U.S. Fish Wildl.Serv., Portland, Oregon and Nev. Fish Game Dep., Reno.

-41-

Welch, B. 1979. Mountain lion research. Job Perform. Rep., P-R Proj. W-124-R-2, Job 1.New Mex. Game Fish Dep. Pp. 41-43.

Presents data on horne range, daily activities and movement patterns, dispersal, and speciescomposition of lion kills in New Mexico.

Wilson, R. 1984. Wyoming. Pp. 88-92 in J. Roberson and F. Lindzey, (eds.). Proc. 2nd Mtn.Lion Workshop. Utah Div. Wildl. Res., Salt Lake City.

Provides information on mountain lion distribution, population status, public attitude towardslions, harvest, and management programs in Wyoming.

Wrigley, R. E. 1982. Manitoba's big cat: the story of the cougar in Manitoba. ManitobaMuseum of Man and Nature, Winnipeg, Canada.

Popular book on history of cougar in Manitoba, cougar biology and .ecology.

-- ··_·_,~·__~_.~." __'C "·"·".·,>,_' ....,'¥..',,'~·.,,"",,",·O'-.