Young cougar among red desert rocks.
 
Book Cover of Cougar:The American Lion

COUGAR: THE AMERICAN LION, by Kevin Hansen

Chapter Three - Cougars at Home


WHERE COUGARS LIVE
Biologists marvel at the cougar's remarkable adaptability. The best example of this is the cat's enormous geographic range. Cougars seem equally at home in Alberta's alpine forests, Arizona's Sonoran Desert, or Mexico's tropical jungles. While the lions don't seem particular about where they live, studies show that the cats do prefer certain types of terrain and vegetation. Habitat, a space and an environment suited to a particular species, and geographic range, a broader term indicating the map area in which a species occurs, are important concepts in understanding cougar life.(1) One expects to find cougars only in suitable habitats within a geographic range, and suitable cougar habitat contains two elements: cover and large prey.

Biologists marvel at the cougar's remarkable adaptability.
Mountain lions are stalking predators that must get close to their prey before ambushing from a short distance. They will take advantage of terrain (steep canyons, rock outcroppings, boulders) or vegetation (dense brush, thickets) to remain hidden while stalking. Cover refers to this combination of terrain and vegetation that allows the cat to stay out of sight while hunting and stalking; habitats that have good stalking cover attract mountain lions. Cover also helps protect the female cougar's vulnerable kittens. (In The Cycle of Life Chapter I explained how a dense thicket or pile of boulders is used as a den. This is what we mean by protective cover.)

Even the best stalking cover is of no value, however, if there is nothing to stalk. Deer are the lion's primary prey-mule deer in western North America and white tailed deer in eastern North America(2)-and deer must be present in sufficient numbers in the lion's habitat for the cat to survive.(3) While lions will take a variety of prey, killing one deer is more energy-efficient than killing several rabbits or squirrels, allowing the cat to procure a lot of fresh meat at once. This is particularly important to females with hungry kittens to feed. Since the presence of cougars in an area is largely dependent on the presence of deer, it is important to understand what makes up good deer habitat.

Like cougar habitat, suitable deer habitat must contain a combination of cover and food. But because deer are ungulates (hoofed, plant-eating mammals) they use both resources differently. Where the puma requires stalking cover, deer need escape cover, usually a dense tangle of vegetation into which predators cannot easily follow.(1) In the case of desert bighorn sheep (Ovis canadensis), on which cougars occasionally prey in New Mexico, Nevada and California, escape cover may not be "cover" in the normal sense, but a sheer rock face. The steep incline and exposure discourages a cougar from following and allows the sheep to escape.

Other types of cover are required. Deer need cover for giving birth to fawns and for resting (protective cover and resting cover). They cope with seasonal temperature extremes by taking refuge in timber stands (thermal cover); tall trees with dense foliage reduce heat loss through radiation on cold, clear winter nights, provide shade on hot sunny days, and serve as shields from strong winds for deer and cougars alike.(4)

Being herbivores (plant-eaters), deer will gravitate to habitats that have adequate forage. Mule deer (Odocoileus hemionus), the most common species of deer in the western United States and Canada, require a mix of food types(5) and are known to eat 788 plant species.(6) In southern Utah, mule deer feed extensively on bitterbrush and Gambel oak, two plants that also provide excellent cover. Not surprisingly, areas dominated by these two plants are also frequented by mountain lions.(7)

Maurice Hornocker(8) and later John Seidensticker and his coworkers studied cougars in the remote wilderness of the Idaho Primitive Area (now the Frank Church River of No Return Wilderness). There, they found that cougars referred steep, rocks areas covered with dense stands of Douglas fir and ponderosa pine, with sagebrush and grasslands mixed among the bluffs and talus slopes. The big cats avoided crossing large open areas with insufficient cover, preferring to travel around the perimeters.

Researchers Kenny Logan and Larry Irwin(10) studied cougars in the Bighorn Mountains of northern Wyoming; their work provided the first quantified evaluation of cougar habitat use, and their findings were similar to those of Seidensticker and Hornocker in Idaho. The cats frequented canyonland habitats with steep, rugged slopes (greater than 45 degrees) containing mixed conifer and brushy mountain mahogany cover. Grasslands and sagebrush areas with gentle slopes (less than 20 degrees) were generally avoided.

The late Steven Laing was a member of a team of researchers who studied cougars in the Boulder-Escalante region of south-central Utah. This 10-year study, completed in 1989, is considered to be the most thorough examination of the cats yet completed. Laing, being responsible for examining cougar habitat use, found that cougars selected pinyon-juniper woodlands with lava boulders scattered in the understory cover (ground vegetation). The cats also frequented ponderosa pine/oakbrush, mixed aspen/spruce-fir, and spruce-fir habitats. They avoided sagebrush bottomlands, agricultural and pasture lands, slickrock sandstone canyons, and open meadows. These areas were typically at higher elevations with steeper slopes and denser understory cover. This combination of terrain and vegetation seems to enhance the cat's ability to survey and move through the landscape unseen.(7)

Harley Shaw observed a similar pattern in Arizona. Forested areas such as those found on the Mogollon Rim and Kaihab Plateau have little understory cover and hold relatively, low mountain lion densities, while chaparral and pinyon-juniper vegetations, which have dense understory cover, have higher densities of mountain lions. Shaw suspects understory cover for stalking is the key to habitat suitability.(11)

Cougars cannot successfully take prey if there is too much or too little cover.
Seidensticker and his colleagues(9) described suitable cougar habitat as a combination of vegetation, topography, prey numbers, and prey vulnerability. Prey vulnerability refers to the ease a prey species may be captured and killed by cougars and depends on the availability of cover (both stalking cover and escape cover) and the behavior of the prey. The relationship between vegetation and prey vulnerability is particularly important: cougars cannot successfully take prey if there is too much or too little cover. The best vegetation for stalking cover is moderately dense-thick enough for the lion to remain hidden, sparse enough for the cat to see its prey.(12) Thus, the presence of moderately dense stalking cover in a habitat increases the vulnerability of the prey found there. This also explains why dense vegetation is attractive to deer as escape cover-it reduces their vulnerability to cougars.

Writer Barry Lopez (a MLF Honorary Boardmember) calls the cougar "a dweller on the edge." Edges, or ecotones, are transitional borders between different habitat types-the places where forest meets clearing, where rocky ledge meets bush, and where willow thicket meets streamside banks.(2,13) Such areas provide good forage and cover for deer, which in turn attracts cougars. Florida panthers make frequent use of ecotones in Everglades National Park;(14) much of the panthers' habitat in that part of south Florida is slash pine woodland s with dense understory cover of saw palmetto. This woodland forms the eastern boundary of a flat, grassy wetland (the Everglades) dotted with islands of hardwood trees called hammocks. Panthers do most of their hunting and make most of their kills along the edges of these hammocks and in these woodlands. Laing found a similar pattern in the Boulder-Escalante study: "Riparian zones [streamside habitats] and rock ledges were the two ecotones most associated with highly used areas suggesting habitat selection based on prey densities, cover diversity, and possibly water availability." (7)

To a cougar, vegetation shape and density seems to be more important than vegetation type in determining habitat suitability. This partially explains the cat's ability to occupy such an extensive range and variety of habitats.(12)

So the best deer and cougar habitats appear to be forested areas that contain good deer forage and cover as well as a diversity of terrain and sufficient stalking cover.  Cougars seem consciously to select cover and terrain that allow them find prey, observe it while stalking, and approach close enough to make a kill.(4) Seidensticker's study in Idaho(9) showed that, over a span of years, kill sites were clustered in certain areas, which suggested that these areas offered advantages in taking prey.

HOME RANGE
Within mountain lion habitat, adult cougars space themselves out and confine their movement to individual fixed areas known as home ranges. Home range should not be confused with geographic range, which is a broader term indicating the entire map area in which cougars occur.(1) Cougar home ranges include hunting areas, water sources, resting areas, lookout positions, and denning sites where kittens or cubs can be safely reared. Cougars that occupy home ranges are called residents, and possession of a home range enhances a resident' lion's chances of more consistently finding prey, locating mates, and successfully rearing young.(15)

Possession of a home range is fundamental to the cougar's survival as a solitary predator. By having a fixed area of land to hunt in, the cougar is better able to consistently locate prey. It roams its home range constantly, learning the terrain, where the best cover is, where the deer most likely can be found. This is why survival for a transient cougar is more precarious than that of a resident cougar.  Transients are constantly moving through unfamiliar territory and have not yet perfected their hunting skills.

Cougars are not territorial in the sense that they defend their home ranges to exclude all other cougars. Rather, the big cats have evolved a land tenure system(9) in which home ranges are maintained by resident lions but not transient lions. Male home ranges are typically larger than female home ranges, usually overlapping or encompassing several of the female ranges, but only occasionally overlapping those of other resident males; however, female home ranges commonly overlap. Exceptions to this pattern do exist. Studies in the Diablo Mountains of California(16) and the San Andres Mountains of New Mexico(17) showed overlap between male home ranges, while those of females did not.

In areas where home ranges overlap, cougars seem to avoid each other. This mutual avoidance is thought to be accomplished primarily through sight and smell.(2, 12) Smell is employed through the use of scrapes. In the "Cycle of Life" chapter it was explained how scrapes function as biological traffic signals within home ranges. By either making a scrape or sniffing the scrape of other individuals, cougars send and receive a variety of messages. Male residents can announce their presence, transient lions or females with dependent kittens can avoid male residents, and females can find males when they are ready to mate.(18) Two cougars in overlapping home ranges can both use the common area because scrapes allow them to use the area at different times.

After sniffing a scrape impregnated with the urine of another cougar, a lion will display a lip-curling grimace known as a flehman. This action is thought to allow them to use a special olfactory organ in the roof of their mouth to evaluate, or get a better "look" at the scent. Biologists speculate that males may use the flehman to determine from a female's urine whether she is ready to mate.(9,19,20) Both Seidensticker(9) and Lindzey(18) have observed this behavior in captive mountain lions.

Experts speculate that land tenure and mutual avoidance allow cougars too maintain home ranges in a number of beneficial ways. First they seem to reduce conflict. A large, powerful carnivore like the cougar depends exclusively on its good health to capture prey. Frequent fighting could lead to serious injury and starvation. This is not to say fighting between cats never occurs; as has already been pointed out, in some cougar populations, such as in southern New Mexico, fighting is common and even a major source of mortality.(17) Secondly, maintaining a matrix of adjacent and overlapping home ranges seems to limit cougar population density,(8,21,22) which in turn increases the cats' chances of finding prey. Finally, because home ranges are frequently hundreds of square miles in size, it would be impossible for a cougar to actively defend the entire area against all intruders.(15) As a result, a more flexible system of coexistence has evolved.

Researchers have learned much of what they know about home ranges through radio telemetry. This involves capturing a wild cougar and attaching a collar containing a small radio transmitter. By plotting the cat's locations on a topographic map, biologists can learn the size of the area a cougar uses throughout the year, what the density of cats in an area is, and the social structure of the population.(23) What they have learned is that home ranges vary widely in size, depending on local vegetation, prey density, and the time of the year. Male home range size can vary from 25 to 500 square miles, while females usually occupy smaller areas of from 8 to over 400 square miles. Sitton and Wallen documented some of the smallest home ranges in the Big Sur region of coastal California, where the average home ranges were 25 to 35 square miles for males and 18 to 25 square miles for females.(24) A warmer climate and abundant forage probably make it unnecessary for the deer to migrate between winter and summer range and the herds thus concentrate in specific areas. Consequently, the cougar population concentrates around the denser prey population. The presence of good stalking cover is likely a factor as well. Hemker and his colleagues found some of the largest home ranges during the Boulder-Escalante study in southern Utah, where males occupied areas of up to 513 square miles and females up to 426 square miles.(25)

As might be expected, male and female cougars use their home ranges differently. Besides a place to hunt, males require in area where they can mate with as many females as possible without interference from surrounding males. That is why male home ranges overlap two, three, or more female home ranges and why male home ranges usually do not overlap. Female home ranges are usually smaller and are used to provide sufficient prey and denning sites for rearing kittens, even in years of low prey density.(15)

Life is tough for a female cougar-much tougher than for a male. Seidensticker explains why: "Adult females are subject to more stress and hazards than are males. The female must hunt and kill large, potentially dangerous prey more frequently than males and must do so at regular, predictable intervals if she is to succeed in rearing her kittens, thus increasing the likelihood of accidental death."(9) On occasion, males have even been known to abandon their home range and move to a new area;(26) older males can also be displaced from their home ranges by prime males.(17) Females show much more attachment to their home ranges and tend to remain in the same area for their entire lives.

The amount of stalking cover and prey numbers in cougar habitat obviously influence home range size. Home range size and the degree of overlap in turn influence the density of a cougar population in a given area.(27) Density estimates of 3 to 7 adult cougars per 100 square miles have been made in southern Alberta(28) and 5 to 8 adult cougars per 100 square miles in the Diablo Mountains of California.(16) Both habitats are characterized by good stalking cover and abundant prey. Lion densities in dryer desert climates seem to be lower. Linda Sweanor and Kenny Logan estimate only 2 adult cougars per 100 square miles in their New Mexico study area, which lies in the Chihuahuan Desert.(29) Densities in southern Utah were even lower, at .5 to .8 adult cougars per 100 square miles, which was 30 percent lower than densities estimated elsewhere.(25) When stalking cover, prey, and water are scarce, cougars expend more energy searching for and stalking prey in larger home ranges. As a result, the cougar population is scattered more thinly across the region. (Fred Lindzey cautions that estimating densities of a solitary and highly mobile predator like the mountain lion is difficult, and that a variety of estimation methods are used by biologists, so one must he careful in making comparisons.(12))

As a result of the functions of land tenure and mutual avoidance, cougars appear to "saturate" an area at a given density. Research indicates that the density of cougars in a particular area is socially regulated through home range size and overlap and does not increase above a level socially tolerable for cougars. In other words, healthy cougar populations appear to be self-regulating. Harley Shaw explains: "In all studies of lions where relatively good documentation of lion numbers has been made, lion densities have peaked and stabilized at points between ten and twenty square miles per adult resident. Evidence indicates that, if left alone, adult resident lions will probably not populate beyond such densities."(22)

POPULATION DYNAMICS
Wildlife biologists have long puzzled over the cougar's solitary life style and how it benefits the animal as a predator. "I find it curious that an animal starting in a litter becomes antisocial in adulthood," writes Harley Shaw. "The earliest moments of a lion's life involve touching its siblings. Its growth involves play, interaction, and cooperation during early efforts in hunting. Yet the animal ultimately comes to avoid other adults except at breeding time."(22)

The cooperative hunting methods of wolves have been extensively studied and their success is well known, as is their gregarious life style. One cannot help but wonder if in comparison the cougar's solitary life style, especially hunting, puts the cat at a distinct disadvantage. Seidensticker and his fellow researchers don't think so: "The mountain lion, too, kills large potentially dangerous prey, but unlike the wolf, a pursuit predator, the lion is a stalking predator whose success depends solely on the element of surprise. In the broken land where lions find sufficient cover to stalk and launch successful attacks, the prey usually are scattered and time-consuming to find. Under such conditions, a solitary social structure is apparently the most effective life style."(9)

While hunting alone may have its advantages, cougars may not be as solitary as once thought. Rick Hopkins points out that adult males are not solitary by choice and probably spend most of their time searching for receptive females.(27) Susan de Treville once monitored four radio collared adult resident males to within 50 yards of each other in the Big Sur region of California.(30) Andrew Kitchener writes: "...far from having a chaotic, random system of home ranges driven by the need for [solitary] cats to avoid each other at all costs, most wild cats maintain a predictable system of land tenure, which promotes social stability and maximizes the reproductive success of both males and females... far from being strangers, neighboring cats probably know each other very well from their own distinctive smells."(15)

Solitary or not, cougars that inhabit a common geographic area are referred to as a population,(1) and within a population not all cougars are created equal. This feline social hierarchy consists of three classes of animals: resident adult males and females, transient males and females, and dependent offspring of resident females. Resident adults maintain established home ranges and do most of the breeding in a population. Transients constantly move through the home ranges of residents in search of a vacant home range of their own; while female transients tend to delay breeding until they find and occupy a home range, this may not be true of males. Dependent offspring include kittens and juveniles that still rely on their mother to hunt for them.(23)

Determining whether a cougar is an adult, transient, or kitten is more complex than it sounds; finding a reliable way to determine age is something which has long eluded biologists. Age is usually established using a combination of tooth wear, body weight, color spotting, and behavior. Kittens are newborn to 16 months old and are still with their mother in her home range; they may still have spots which fade by the third of fourth month. Transients are 17 to 23 months old and have left their mother's home range but have not yet settled in a home range of their own; spotting may still be present on the insides of the front legs. Resident adults are mature cats, at least 24 months old, and occupy an established home range; spotting is absent or very faint,

and females may show evidence of nursing.(26)

Harley Shaw uses a slightly different system of classifying lions. Resident lions are adult males and females that use established home ranges and are reproductively active. Immature lions are defined as offspring of resident adults that are still traveling with, or close to, the mother. Transient lions are young, newly independent adults searching for a home range.(31)

Researchers Kenny Logan and Linda Sweanor prefer the terms emigrant and disperser to transient. These refer to lions that have emigrated or dispersed from their birth area but have not set up a home range. This dispersal out of a previously occupied area is called emigration, while movement into a new area is called immigration.(1) Emigrants become immigrants when they enter a new population.(17) (While it is good science for researchers to refine their techniques and terms, the lack of consistency also makes it difficult for scientists to compare information. It also frustrates writers attempting to explain mountain lions.)

Because no reliable method yet exists for accurately aging cougars, determining how many resident adults, transients, and kittens are present in a cougar population at any given time is equally difficult. This is further complicated by the fact that young transient cats are constantly on the move.(12) These factors, if not accounted for, can distort population estimates made by wildlife biologists.

Resident females generally outnumber resident males in a population.(25,32,33) This may be because females require smaller home ranges that frequently overlap those of other female cougars. Females also seem generally more tolerant of adjacent females and, as a result, more females might concentrate in a smaller area. Resident males, on the other hand, require larger home ranges that rarely overlap, and they tend to be intolerant of other males and are more widely scattered than resident females.

Young transient cougars usually leave their mother's home range (disperse) sometime during their second year. The newly independent cats occasionally linger in the birth area, while others leave immediately. They may wander for more than a before establishing their own home range.(22) If sufficient space is available, however some females will remain in their birth areas; one young female in Nevada stayed in the mountain range of her birth, bred at 24 months, and established a home range adjacent to her mother's. Female transients have even been known to take over their mother's home range after her death.(18)

Males tend to wander farther than females,(26) although how far either gender travels seems to vary. Transient males in the Diablo Mountains of California traveled up to 35 miles from their areas of birth.(16) In one Nevada population, males covered an average of 31 miles, while females averaged 18 miles.(26) One young cougar marked in the Bighorn Mountains of Wyoming appeared in northern Colorado, 300 miles from the original location.(34)

Transient males and females seldom remain in one location longer than six months. How the cats "test" a population for vacant home ranges is poorly understood. It is generally thought that a transient either finds a vacant home range or displaces an older resident. If a vacant home range is found, the young male or female will settle down to the business of establishing its place in the population. If the population density is so high that no vacant home areas exist, the transient moves on.

Kenny Logan believes male transients enter a new population and for a period of time, make kills, avoid other lions, and avoid scrapings. If they think they can establish a home range they begin to scrape. Meanwhile, the resident male, closely monitoring its home range, is busy investigating the newcomer's scrapes and kills. If the resident and newcomer encounter each other they may fight. As a result, the resident may kill the newcomer, the newcomer may kill the resident, or the newcomer may drive off or displace the resident.(17)

The role of transients in populations is important because they are the primary source of replacements for resident cougars who die as a result of hunting, old age, or accidents. Transients also ensure genetic mixing between populations,(12) and appear to be a major factor in the recovery of hunted populations. Cougar populations in isolated mountain ranges, such as those common to the Great Basin, are particularly vulnerable because immigration of transients is low.(35)

For example, aggressive predator control efforts wiped out all cougars in and around Yellowstone National Park by the 1920s, and - except for a few transients passing through - no cougars lived there for almost 50 years. But transients seem to be slowly recolonizing the 2.2 million-acre park; research biologist Kerry Murphy directs a field study of cougars in the region and estimates 14 to 17 resident mountain lions are now present. Radio collars have been attached to 9 females and 7 males, and the population produced 21 kittens in 1991. The region has abundant deer and elk, and the cougar population is increasing.(36)

As mentioned earlier, the availability of good hunting sites and prey numbers limit the number of cougars in an area. But birth, death, emigration, and immigration are all factors that greatly influence cougar population maintenance and growth. In absence of hunting by humans, the cougar population will remain relatively constant.(12) Rick Hopkins found that the unhunted cougar population in the Diablo Mountains of California was relatively stable with a low turnover of residents.(16)

COUGARS ON THE MOVE
There is nothing more distinctive about cats than the way they move, and mountain lions are no different. The big cats are the epitome of graceful, lithe motion. Stealthy shadows that do not move so much as flow across the landscape.

There is nothing more distinctive about cats than the way they move.

Consider the effect of broken terrain on prey, explains Kenny Logan. "Mountain lions are ballerinas at getting across broken terrain-steep slopes, boulders, outcroppings, and undercut ledges." Mule deer and elk are not as good at negotiating such rugged landscapes.(17) Here the cougar uses both cover and superior agility to its advantage, and employs yet another weapon in its predatory arsenal-it hunts under the cover of darkness.

Cougars are not strictly nocturnal, as many once thought. Rather, they tend to be active at the same time as their prey, and deer tend to be active at dawn, dusk, and at night. Animals that are active during the twilight of dawn and dusk are said to be crepuscular. The big cat's excellent night vision makes it well suited for stalking during these low light periods. Florida panthers appear to be more active at sunrise and sunset,(37) and Paul Beier reports that in southern California mountain lions are active throughout the night.(39) During the winter in Idaho, radio-collared cougars were found to be more active at night (40 percent of the time they were located) than during daylight hours (14 percent of the time they were located). Daytime activity increased during the summer with an increased abundance of ground squirrels.(9) Bruce Ackerman studied cougar activity patterns in southern Utah and found the cats most active at sunrise and sunset, but less so during the winter. He also found that single adults were more active than females with young, but that the females became more active as their kittens grew.(3)

Most mule deer herds in mountainous areas of western North America migrate to lower elevations in the winter to avoid heavy snow and then come back to higher elevations in the summer. Predictably, cougars also shift their home range use to follow these seasonal movements.(25) This elevational movement of cats following deer herds has been observed in Arizona, Idaho, California, and Nevada; in Idaho, cougars' winter home ranges were smaller than summer home ranges.(9,25) Biologist Dave Ashman and his coworkers documented a case in Nevada where a male cougar occupied one mountain range in winter, then crossed 10 miles of flat, open desert to take up summer residence in another mountain range. Elsewhere in the state, cougars followed deer herds to lower elevations or traditional wintering grounds. During the cold Nevada winters, cougars avoided the north-facing slopes and frequented the south-facing slopes where there was less snow and more deer. When summer arrived, the cat shifted to the cooler north-facing slope, where there was more vegetation. Cougars also tended to restrict their movements to between 6,500 and 8,500 feet, where vegetation, deer, and other prey were most plentiful.(26)

Paul Beier has monitored some lions continuously for 24-hour periods to better understand their movements. He found that the lions in his California study area travel at a steady speed of approximately one-half mile per hour. Though there is a lot of variation, the cats seem to follow a cycle of traveling for a little less than an hour, followed by 30 minutes of rest.(38)

Mountain lions can cover a lot of ground in their nightly wanderings; a lion in the San Andres Mountains of New Mexico has been documented at 10 miles in a night.(17) Not surprisingly, Hemker and his coworkers found in their Utah study area that females with kittens less than six months old moved significantly shorter distances than females with older kittens, females without kittens, transients, or resident males. One resident male traveled eight miles in a one-day period, but this represents only the straight-line distance between locations on a map and does not reflect the actual distance covered.(25) Here is why.

Cougars have two basic patterns of movement: hunting and traveling. When the big cat is looking for prey, it traverses a zigzag course through it home range. An entire day searching for prey likely brings the cougar only a few linear miles from its starting point. The cat is basically searching and has no specific destination in mind. A traveling cat, with a specific destination in mind is a different matter. The route will be more direct, though it will still take advantage of the landscape, following ridges, major drainages, or using low passes.(22)

A common myth about the daily movements of cougars is that they follow a regular and predictable circuit around their home range. This assumption may stem partly from observations of domestic cats or vague attempt apply human behavior to cougars. A variety of studies that monitored cougar movements with radio telemetry found no such patterns. Cougars move freely and arbitrarily about their home ranges and have never been seen to pass a fixed location at set intervals.(22) As a stalking predator that depends on the element of surprise to capture its prey, regularity of movement would handicap the cougar's ability to survive.




CHAPTER NOTES

  1. Dassman, R.F. 1981. Wildlife biology, 2nd edition. John Wiley and Sons, New York.

  2. Dixon, K.R. 1982. Mountain lion. Pages 711-727 in J.A. Chapman and G.A. Feldhamer, eds. Wild mammals of North America. John Hopkins University Press. Baltimore.

  3. Ackerman, B.B. 1982. Cougar predation and ecological energetics in southern Utah. M.S. thesis, Utah State University, Logan.

  4. Lindzey, F.G., B.B. Ackerman, D. Barnhurst, T. Becker, T.P. Hemker, S.P. Laing, C. Mecham, and W.D. Van Sickle. 1989. Boulder-Escalante cougar project final report. Utah Division of Wildlife Resources, Salt Lake City, Utah.

  5. Wallmo, O.C. 1978. Mule and black-tailed deer. Pages 30-41 in J.L. Schmidt and D.L. Gilbert, eds. Big game of North America: Ecology and management. Wildlife Management Institute. Stackpole Books.

  6. Mills, J. 1990. Deer old game. National Wildlife, October-November 1990: 59.

  7. Laing, S.P. 1988. Cougar habitat selection and spatial use patterns in southern Utah. M.S. thesis. University of Wyoming, Laramie.

  8. Hornocker, M.G. 1969a. Winter territoriality in mountain lions. Journal of Wildlife Management, 33:457-464.

  9. Seidensticker, J.C., IV, M.G. Hornocker, W.V. Wiles, and J.P. Messick. 1973. Mountain lion social organization in the Idaho Primative Area. Wildlife Monographs, 35.

  10.  Logan, K.A. and L.L. Irwin. 1985. Mountain lion habitats in the Big Horn Mountains, Wyoming. Wildlife Society Bulletin 13: 257-262.

  11.  Shaw, H.G. 1991. Wildlife Biologist, General Wildlife Services, Chino Valley, Arizona. (Personal communication)

  12. Lindzey, F. 1987. Mountain lion. Pages 656-668 in M. Novak, J.A. Baker, M.E. Obbard, and B. Malloch, eds. Wild furbearer management and conservation in North America. Ministry of Natural Resources, Ontario, Canada.

  13. Lopez, B. 1981. The elusive mountain lion. GEO June: 98-116. 

  14. Bass, O.L. 1991. Wildlife Biologist, Everglades National Park Research Center, Homestead, Florida. (Personal communication)

  15. Kitchener, A. 1991. The natural history of the wild cats.  Cornell University Press. Ithaca, New York. 

  16. Hopkins, R.A. 1989. Ecology of the puma in the Diablo Range, California. Ph.D. dissertation, University of California at Berkeley.

  17. Logan, K.A. 1991. Wildlife Research Institute, Inc., Moscow, Idaho. (Personal communication) 

  18. Lindzey, F.G. 1991. Wildlife Biologist, U.S. Fish and Wildlife Service, Wyoming Cooperative Fish and Wildlife Research Unit, University of Wyoming, Laramie, Wyoming. (Personal communication) 

  19. Kiltie, R.A. 1991. How cats work. Pages 54-67 in J. Seidensticker and S. Lumpkin, eds. Great cats: Majestic creatures of the wild. Rodale Press, Emmaus, Pennsylvania. 

  20. Mellen, J. 1991. Cat behavior. Pages 68-75 in J. Seidensticker and S. Lumpkin, eds. Great cats: Majestic creatures of the wild. Rodale  Press, Emmaus, Pennsylvania. 

  21. Hornocker, M.G. 1970. An analysis of mountain lion predation upon mule deer and elk in the Idaho Primative Area. Wildlife  Monographs. 21:1-39. 

  22. Shaw, H.G. 1989. Soul among lions. Johnson Books. Boulder, Colorado. 

  23. Lynch, W. 1989. The elusive cougar. Canadian Geographic August/September: 24-31. 

  24.  Sitton, L.W. and S. Wallen. 1976. California mountain lion study. California Department of Fish and Game. Sacramento. 

  25. Hemker, T.P., F.G. Lindzey, and B.B. Ackerman. 1984.  Population characteristics and movement patterns of cougars in southern Utah. Journal of Wildlife Management, 48(4):1275-1284. 

  26. Ashman, D., G.C. Christensen, M.L. Hess, G .K. Tsukamoto, and M.S. Wickersham. 1983. The mountain lion in Nevada. Nevada Department of Wildlife, Reno.  

  27. Hopkins, R.A. 1991. Wildlife Biologist, H.T. Harvey and Associates, Alviso, California. (Personal communication) 

  28. Pall, O., M. Jalkotzy, and I. Ross. 1988. The cougar in Alberta. Fish and Wildlife Division. Alberta Forestry, Lands and Wildlife. Associated Resource Consultants. Calgary, Alberta. 

  29. Sweanor, L.L. 1990. Mountain lion social organization in a desert environment. M.S. thesis, University of Idaho, Moscow. 

  30. de Treville, S. 1991. Wildlife Biologist. de Treville Environmental Engineering. San Diego, California. (Personal communication) 

  31. Shaw, H. 1987. Mountain lion field guide. 3rd Edition. Special Report Number 9. Arizona Game and Fish Department. 

  32. Anderson, A.E. 1983. A critical review of literature on puma (Felis con color), Colorado Division of Wildlife. Special Report Number 54.

  33. Logan, K.A. 1983. Mountain lion population and habitat characteristics in the Big Horn Mountains of Wyoming. M.S. thesis, University of Wyoming, Laramie. 

  34. Parfit, M. 1985. Its days as a varmint are over, but the cougar is still on the run. Smithsonian, November: 68-79. 

  35. Ackerman, B.B., F.G. Lindzey, and T.P. Hemker. 1984. Cougar food habits in southern Utah. Journal of Wildlife Management, 48:147-155. 

  36. Murphy, K. 1991. Wildlife Biologist. Wildlife Research Intitute, Inc., Moscow, Idaho. (Personal communication) 

  37. Maehr, D.S., E.D. Land, J.C. Roof, and J.W. McCown. 1990c. Day beds, natal dens, and activity of Florida panthers. Proceedings of the annual conference of southeast fish and wildlife agencies, 44: In press. 

  38. Beier, P. 1992. Project Leader, Orange County Cooperative Mountain Lion Study, Department of Forestry and Resource Management,  University of California, Berkeley. (Personal communication) 

* Cougar The American Lion Line Illustrations Copyright (1992-2009) by Linnea Fronce



Chapter Index to Cougar: The American Lion

Written by Kevin Hansen in association with the Mountain Lion Foundation.


  • back of lion's head.

    Chapter 1: The Consummate Cat

    Start off by reading about the history of the cougar including the evolution of native cat species, the two dozen or so subspecies of cougars and their general appearance. Learn about their discovery in the western hemisphere by early explorers and the many names they have been given by different cultures.

  • Mother lion carrying kitten in mouth.

    Chapter 2: The Cycle of Life

    Beginning from birth, this chapter covers the life span of a cougar. A dependent kitten will mature in about two years, disperse off to establish its own home range, breed with others in neighboring ranges, and perhaps live to ten years of age. Cougars struggle mightily to survive in the face of active threats, which have greater or lesser impact depending upon their stage of life.

  • Lion peering out from overhanging cave entrance.

    Chapter 3: Cougars at Home

    Although cougars are adaptable and can survive any where that has cover and large prey, human hunting has limited them to the western portion in North America. The size and overlap of an individual's home range depends on its age and sex, and a cougar will use markings to define the borders. Get an in-depth look at their population dynamics and discover how far they will travel to find food.

  • Adult lion peering out from brush.

    Chapter 4: An Almost Perfect Predator

    A mountain lion's keen senses, muscular agility, and ability to adapt to almost any landscape and prey make it a successful hunter. Their walking stride, retractable claws and powerful jaw allow them to sneak through bushes undetected and quickly take down prey. Predators play an important role in the health of prey populations and studies have shown they do not significantly reduce the number of deer and elk in a region.

  • Kitten looking out from underneath a car.

    Chapter 5: Cougars and Humans

    Cougars were admired by many Native American cultures, and commonly found in their spiritual beliefs and folklore. But when early European explorers arrived, cougars were seen as a threat and competition. From the late 1600's to mid 1900's, bounties were often paid to anyone who killed a cougar. As ranching increased so did predator control, and then along with sport hunting, cougars were wiped out in most of the United States.