Cougar range map
Cougar (Puma concolor) is a very large, New World wild cat (family Felidae), characterized by a slender body, long hind legs, retractable claws, and typically an unpatterned tawny, grayish, or reddish coat. Also known as puma, mountain lion, or panther, depending on region, this large, solitary cat has the greatest range of any wild terrestrial mammal in the Western Hemisphere (Iriarte et al. 1990), extending from Yukon in Canada to the southern Andes of South America. It also is the second heaviest cat in the New World, after the jaguar, and the fourth heaviest in the world, after the tiger, lion, and jaguar, although it is most closely related to smaller felines.
An adaptable, generalist species, the cougar is found in every major New World habitat type. A capable stalk-and-ambush predator, the cougar pursues a wide variety of prey. Primary food sources include ungulates such as deer and bighorn sheep, as well as domestic cattle, horses, and sheep, particularly in the northern part of its range, but it hunts species as small as insects and rodents. As a predator at the top of its food chain, the cougar helps keep populations of prey species in balance.
The cougar prefers habitats with dense underbrush and rocky areas for stalking, but it can live in open areas. It is a reclusive cat and usually avoids people. Attacks on humans remain rare.
Due to persecution following the European colonization of the Americas, and continuing human development of cougar habitat, populations have dropped in many parts of its historical range. In particular, the cougar was extirpated in eastern North America, except an isolated sub-population in Florida; the animal may be recolonizing parts of its former eastern territory. With its vast range, the cougar has dozens of common names and various references in the mythology of the indigenous peoples of the Americas and in contemporary culture.
Cougars are slender cats, with somewhat round heads and with erect ears. They have powerful forequarters, neck, and jaw, which allows them to grasp and hold large prey. As with all cats, they walk on four feet, in a digitigrade manner—that is on their toes. Their hind legs are longer and stronger than their fore legs, which gives them strength for sprinting and leaping, but not stamina for long distance running. They have large paws, with five retractable claws on their forepaws (one a dewclaw) and four on their hind paws. The larger front feet and claws are adaptations to clutching prey (Wright and Amirault-Langlais 2007).
Cougars have proportionally the largest hind legs in the cat family (Nowell and Jackson 2006), allowing great leaping and short-sprint ability. An exceptional vertical leap of 5.4 meters (18 feet) is reported for the cougar (SDZ 2008). Horizontal jumping capability is suggested anywhere from 6 to 12 meters (20 to 40 feet). The cougar can run as fast as 55 kilometers per hour (35 miles per hour)(CanGeo 2008), but is best adapted for short, powerful sprints rather than long chases.
Cougar coloring is plain (hence the Latin concolor) but can vary greatly between individuals and even between siblings. The coat is typically tawny, but ranges to silvery-gray or reddish, with lighter patches on the under body including the jaws, chin, and throat. Infants are spotted and born with blue eyes and rings on their tails (Nowell and Jackson 2006); juveniles are pale, and dark spots remain on their flanks (DEC 2008). Despite anecdotes to the contrary, and even reference works citing that melanistic (black) forms of cougars are common (Grzimek et al. 2004), other authorities maintain that all-black coloring (melanism) has never been documented in cougars (Glenn 2006). The term "black panther" is used colloquially to refer to melanistic individuals of other species, particularly jaguars and leopards (MB).
Cougars are large cats. While the cat family (Felidae) is usually divided into the "big cats" of the subfamily Pantherinae and the "small cats" of the subfamily Felinae, the cougar is part of the Felinae subfamily and yet can weigh as much or more as some of the "big cats." Cougars can be as large as jaguars, but are less muscled and powerful; where their ranges overlap, the cougar tends to be smaller than average. The cougar is on average heavier than the leopard, but smaller than the tiger or lion. Adult cougars stand about 60 to 80 centimeters (2.0 to 2.7 feet) tall at the shoulders. The length of adult males is around 2.4 meters (8 feet) long, nose to tail, with overall ranges between 1.5 and 2.75 meters (5 and 9 feet) nose to tail suggested for the species in general (TPW; DEC 2008). Males have an average weight of about 53 to 72 kilograms (115 to 160 pounds). In rare cases, some may reach over 120 kilograms (260 pounds). Female average weight is between 34 and 48 kilograms (75 and 105 pounds (Nowell and Jackson 2006). Cougar size is smallest close to the equator, and larger towards the poles (Iriarte et al. 1990).
Unlike members of the "big cat" genus Panthera—tiger (Panthera tigris), lion (Panthera leo), jaguar (Panthera onca), and leopard (Panthera pardus—the cougar cannot roar, lacking the specialized larynx and hyoid apparatus of Panthera (Weissengruber et al. 2002). Like domestic cats, cougars vocalize low-pitched hisses, growls, and purrs, as well as chirps and whistles. They are well known for their screams, referenced in some of its common names, although these may often be the misinterpreted calls of other animals (ECF 2006)
Like all cats, the cougar is an obligate carnivore, requiring meat in its diet. (Some cat species, such as bobcats, supplement their meat diet with fruit). As with other cats, the teeth of cougars are well suited to their diet, with long canines for gripping prey and blade-like molars for cutting flesh (Voelker 1986).
A successful generalist predator, the cougar will eat any animal it can catch, from insects to large ungulates. Its most important prey species are various deer species, particularly in North America; mule deer, white-tailed deer, elk, and even the large moose are taken by the cat. Other species such as bighorn sheep, horses, and domestic livestock such as cattle and sheep are also primary food bases in many areas. A survey of North America research found 68 percent of prey items were ungulates, especially deer. Only the Florida panther (a subspecies of cougar) showed variation, often preferring feral hogs and armadillos (Iriarte et al. 1990). Investigation in Yellowstone National Park showed elk followed by mule deer were the cougar's primary targets; the prey base is shared with the park's gray wolves, with whom the cougar competes for resources (Akenson et al. 2007; Oakleaf et al. 2007). Another study on winter kills (November–April) in Alberta showed that ungulates accounted for greater than 99% of the cougar diet. Learned, individual prey recognition was observed, as some cougars rarely killed bighorn sheep, while others relied heavily on the species (Ross et al. 1993).
In the Central and South American cougar range, the ratio of deer in the diet declines. Small to mid-size mammals are preferred, including large rodents such as the capybara. Ungulates accounted for only 35 percent of prey items in one survey, approximately half that of North America. Competition with the larger jaguar has been suggested for the decline in the size of prey items (Iriarte et al. 1990). Other listed prey species of the cougar include mice, porcupine, and hares. Birds and small reptiles are sometimes preyed upon in the south, but this is rarely recorded in North America (Iriarte et al. 1990).
Though capable of sprinting, the cougar is typically an ambush predator. It stalks through brush and trees, across ledges, or other covered spots, before delivering a powerful leap onto the back of its prey and a suffocating neck bite. The cougar is capable of breaking the neck of some of its smaller prey with a strong bite and momentum bearing the animal to the ground (Wrightv and Amirault-Langlais. 2007). It has a flexible spine which aids its killing technique.
Kills are generally estimated at around one large ungulate every two weeks. The period shrinks for females raising young, and may be as short as one kill every three days when cubs are nearly mature at around 15 months (Nowell and Jackson 2006). The cat commonly drags a kill to a preferred spot, covers it with brush, and returns to feed over a period of days. It is generally reported that the cougar is a non-scavenger and will rarely consume prey it has not killed; but deer carcasses left exposed for study were scavenged by cougars in California, suggesting more opportunistic behavior (Bauer et al. 2005).
The cougar is adept at climbing, which allows it to evade canine competitors. Although it is not strongly associated with water, it can swim (Sierra 2006).
Females reach sexual maturity between one-and-a-half and three years of age. They typically average one litter every two to three years throughout their reproductive life (UDWR 1999); the period can be as short as one year (Nowell and Jackson 2006). Females are in estrus for approximately 8 days of a 23-day cycle; the gestation period is approximately 91 days (Nowell and Jackson 2006). Females are sometimes reported as monogamous (CanGeo 2008), but this is uncertain and polygyny may be more common (Hamilton et al. 2007). Copulation is brief but frequent.
Only females are involved in parenting. Female cougars are fiercely protective of their kittens and have been seen to successfully fight off animals as large as grizzly bears in their defense. Litter size is between one and six kittens, typically two or three. Caves and other alcoves that offer protection are used as litter dens. Born blind, kittens are completely dependent on their mother at first, and begin to be weaned at around three months of age. As they grow, they begin to go out on forays with their mother, first visiting kill sites, and after six months beginning to hunt small prey on their own (UDWR 1999). Kitten survival rates are just over one per litter (Nowell and Jackson 2006).
Sub-adults leave their mother to attempt to establish their own territory at around two years of age and sometimes earlier; males tend to leave sooner. One study has shown high morbidity among cougars that travel farthest from the maternal range, often due to conflicts with other cougars ("intraspecific" conflict) (UDWR 1999). Research in New Mexico has shown that "males dispersed significantly farther than females, were more likely to traverse large expanses of non-cougar habitat, and were probably most responsible for nuclear gene flow between habitat patches" (Sweanor et al. 2000).
Life expectancy in the wild is reported at between 8 to 13 years, and probably averages 8 to 10; a female of at least 18 years was reported killed by hunters on Vancouver Island (Novell and Jackson 2006). Cougars may live as long as 20 years in captivity. Causes of death in the wild include disability and disease, competition with other cougars, starvation, accidents, and, where allowed, human hunting. Feline immunodeficiency virus, an endemic AIDS-like disease in cats, is well-adapted to the cougar (Biek et al. 2003).
Like almost all cats, the cougar is a solitary animal. Only mothers and kittens live in groups, with adults meeting only to mate. It is secretive and crepuscular, being most active around dawn and dusk.
The cougar is territorial and persists at low population densities.
Estimates of territory sizes vary greatly. Grzimek et al. (2004) report that the male range is at least 260 square kilometers (100 square miles) and overall the home range for the species varies from 32 to 1,031 square kilometers. A report in Canadian Geographic notes large male territories of 150 to 1000 square kilometers (58 to 386 sq mi) with female ranges half the size (CanGeo 2008). Other research suggests a lower limit of 25 km² (10 sq mi) for the species, but an even greater upper limit of 1300 km² (500 sq mi) for males (UDWR 1999). In the United States, very large ranges have been reported in Texas and the Black Hills of the northern Great Plains, in excess of 775 km² (300 sq mi) (Mahaffy 2004). Male ranges may include or overlap with those of females but, at least where studied, not with those of other males, which serves to reduce conflict between cougars. Ranges of females may overlap slightly with each other.
Scrape marks, urine, and feces are used to mark territory and attract mates. Males may scrape together a small pile of leaves and grasses and then urinate on it as a way of marking territory (Sierra 2006).
Home range sizes and overall cougar abundance depend on terrain, vegetation, and prey abundance (UDWR 1999). One female adjacent to the San Andres Mountains, for instance, was found with a large range of 215 km² (83 sq mi), necessitated by poor prey abundance (Sweanor et al. 2000). Research has shown cougar abundances from 0.5 animals to as much as 7 (in one study in South America) per 100 km² (38 sq mi) (Nowell and Jackson 2006).
Because males disperse further than females and compete more directly for mates and territory, they are most likely to be involved in conflict. Where a sub-adult fails to leave his maternal range, for example, he may be killed by his father (Mahaffy 2004). When males encounter each other, they hiss and spit, and may engage in violent conflict if neither backs down (Hamilton et al. 2007). Hunting or relocation of the cougar may increase aggressive encounters by disrupting territories and bringing young, transient animals into conflict with established individuals (WEG 2007).
The cougar has the largest range of any wild land animal in the Americas. Its range spans 110 degrees of latitude, from northern Yukon in Canada to the southern Andes. It is one of only three cat species, along with the bobcat and Canadian lynx, native to Canada (Wright and Amirault-Langlais. 2007). Its wide distribution stems from its adaptability to virtually every habitat type: It is found in all forest types as well as in lowland and mountainous deserts. Studies show that the cougar prefers regions with dense underbrush, but can live with little vegetation in open areas (IUCN 2002). Its preferred habitats include precipitous canyons, escarpments, rim rocks, and dense brush (Sierra 2006).>
The cougar was extirpated across much of its eastern North American range with the exception of Florida in the two centuries after European colonization and faced grave threats in the remainder. Currently, the cougar ranges across most western American states, the Canadian provinces of Alberta and British Columbia, and the Canadian Yukon Territory. Some believe that small relict populations (around 50 individuals) may exist, especially in the Appalachian Mountains and eastern Canada. There have been widely-debated reports of possible recolonization of eastern North America (Marschall 2005), DNA evidence has suggested its presence in eastern North America (Belanger 2007), while a consolidated map of cougar sightings shows numerous reports, from the mid-western Great Plains through to Eastern Canada(CN 2004). Sightings of cougars in the eastern United States continue as cougars with offspring have been sighted in Maine, Vermont, New Hampshire, and Michigan. However, the only unequivocally known eastern population is the Florida panther, which is critically endangered.
South of the Rio Grande, the International Union for the Conservation of Nature and Natural Resources (IUCN) lists the cat in every Central and South American country except Costa Rica and Panama (IUCN 2002). While specific state and provincial statistics are often available in North America, much less is known about the cat in its southern range (NWF 2007).
The cougar's total breeding population is estimated at less than 50,000 by the IUCN, with a declining trend (IUCN 2002). U.S. state-level statistics are often more optimistic, suggesting cougar populations have rebounded. In Oregon, a healthy population of 5,000 was reported in 2006, exceeding a target of 3,000 (ODFW 2006). California has actively sought to protect the cat and a similar number of cougars has been suggested, between 4,000 and 6,000 (CDFG 2007).
Aside from humans, no species preys upon mature cougars in the wild. The cat is not, however, the apex predator throughout much of its range. In its northern range, the cougar interacts with other powerful predators such as the gray wolf, black bear, and the grizzly bear. In the south, the cougar must compete with the larger jaguar. In Florida, it encounters the American alligator.
The Yellowstone National Park ecosystem provides a fruitful microcosm to study inter-predator interaction in North America. Of the three large predators, the massive brown bear appears dominant, often, although not always, able to drive both the gray wolf pack and the cougar off their kills. One study found that brown or black bears visited 24 percent of cougar kills in Yellowstone and Glacier National Parks, usurping 10 percent of carcasses (COSEWIC 2002).
The gray wolf and the cougar compete more directly for prey, especially in winter. While individually more powerful than the gray wolf, a solitary cougar may be dominated by the pack structure of the canines. Wolves can steal kills and occasionally kill the cat. One report describes a large pack of 14 wolves killing a female cougar and her kittens. Conversely, lone wolves are at a disadvantage, and have been reported killed by cougars (Gugliotta 2003). Wolves more broadly affect cougar population dynamics and distribution by dominating territory and prey opportunities, and disrupting the feline's behavior. Preliminary research in Yellowstone, for instance, has shown displacement of the cougar by wolves (GYLC 2007). One researcher in Oregon notes: "When there is a pack around, cougars are not comfortable around their kills or raising kittens A lot of times a big cougar will kill a wolf, but the pack phenomenon changes the table" (Cockle 2006). Both species, meanwhile, are capable of killing mid-sized predators such as bobcats and coyotes and tend to suppress their numbers (Akenson 2007).
In the southern portion of its range, the cougar and jaguar share overlapping territory (Hamdig 2006). The jaguar tends to take larger prey and the cougar smaller where they overlap, reducing the cougar's size (Iriarte et al. 1990). Of the two felines, the cougar appears best able to exploit a broader prey niche and smaller prey (Nuanaez et al. 2000).
As with any predator at or near the top of its food chain, the cougar impacts the population of prey species. Predation by cougars has been linked to changes in the species mix of deer in a region. For example, a study in British Columbia observed that the population of mule deer, a favored cougar prey, was declining while the population of the less frequently preyed-upon white-tailed deer was increasing (Robinson et al. 2002). The Vancouver Island marmot, an endangered species endemic to one region of dense cougar population, has seen decreased numbers due to cougar and gray wolf predation (Bryant and Page 2005).
The cougar has numerous names in English, of which puma and mountain lion are popular. Other names include catamount, panther, painter because of its black tail tip, and mountain screamer. In North America, "panther" is used most often to refer to the Florida panther sub-population. In South America, "panther" refers to both the spotted and black color morphs of the jaguar, while it is also broadly used to refer to the Old World leopard.
The cougar holds the world record for the animal with the highest number of names due to its wide distribution across North and South America. It has over 40 names in English alone (Guinness 2003, 49).
"Cougar" is borrowed from the Portuguese çuçuarana, via French; the term was originally derived from the Tupi language. A current form in Brazil is suçuarana. "Puma" comes, via Spanish, from the Quechua language of Peru (Harper 2001a, 2001b).
The cougar is the largest of the "small cats." It is placed in the subfamily Felinae, although its bulk characteristics are similar to those of the big cats in the subfamily Pantherinae (Wozencraft 2005). The family Felidae is believed to have originated in Asia approximately 11 million years ago. Taxonomic research on felids remains partial and much of what is known about their evolutionary history is based on mitochondrial DNA analysis (Nicholas 2006), as cats are poorly represented in the fossil record (Johnson et al. 2006), and there are significant confidence intervals with suggested dates.
In the latest genomic study of Felidae, the common ancestor of today's Leopardus, Lynx, Puma, Prionailurus, and Felis lineages migrated across the Bering land bridge into the Americas approximately 8 to 8.5 million years ago (mya). The lineages subsequently diverged in that order (Johnson et al. 2006). North American felids then invaded South America 3 mya as part of the Great American Interchange, following formation of the Isthmus of Panama. The cougar was originally thought to belong in Felis, the genus which includes the domestic cat, but it is now placed in Puma along with the jaguarundi, a cat just a little more than a tenth its weight.
Studies have indicated that the cougar and jaguarundi are most closely related to the modern cheetah of Africa and western Asia (Johnson et al. 2006; Culver et al. 2000), but the relationship is unresolved. It has been suggested that the cheetah lineage diverged from the Puma lineage in the Americas and migrated back to Asia and Africa (Johnson et al. 2006; Culver et al. 2000), while other research suggests the cheetah diverged in the Old World itself (Ross et al. 2005). The outline of small feline migration to the Americas is thus unclear.
Recent studies have demonstrated a high level of genetic similarity among the North American cougar populations, suggesting that they are all fairly recent descendants of a small ancestral group. Culver et al. (2000) suggest that the original North American population of Puma concolor was extirpated during the Pleistocene extinctions some 10,000 years ago, when other large mammals, such as Smilodon, also disappeared. North America was then repopulated by a group of South American cougars (Culver et al. 2000).
Until the late 1990s, as many as 32 subspecies were recorded; however, a recent genetic study of mitochondrial DNA (Culver et al. 2000) found that many of these are too similar to be recognized as distinct at a molecular level. Following the research, the canonical Mammal Species of the World (3rd edition) recognizes six subspecies, five of which are solely found in Latin America (Wozencraft 2005):
The status of the Florida panther, here collapsed into the North American cougar, remains uncertain. It is still regularly listed as subspecies Puma concolor coryi in research works, including those directly concerned with its conservation (Conroy et al. 2006). Culver et al. (2000) themselves noted microsatellite variation in the Florida panther, possibly due to inbreeding ; responding to the research, one conservation team suggests "the degree to which the scientific community has accepted the results of Culver et al. and the proposed change in taxonomy is not resolved at this time" (FPRT 2006).
The World Conservation Union (IUCN) currently lists the cougar as a "near threatened" species. It has shifted the cougar's status from "least concern," while leaving open the possibility that it may be raised to "vulnerable" when greater data on the cat's distribution becomes available (IUCN 2002). The cougar is regulated under Appendix I of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES 2008) rendering illegal international trade in specimens or parts.
East of the Mississippi, the only unequivocally known cougar population in the United States is the Florida panther. The U.S. Fish and Wildlife Service historically recognized both an Eastern cougar and the Florida panther, affording protection under the Endangered Species Act (FWS 1991; FWS 1993; FWS 1996). Certain taxonomic authorities have collapsed both designations into the North American cougar, with Eastern or Florida subspecies not recognized (Wozencraft 2005), while a subspecies designation remains recognized by some conservation scientists (Conroy et al. 2006). The most recent documented count for the Florida sub-population is 87 individuals, reported by recovery agencies in 2003 (FFWCC 2003).
The cougar is also protected across much of the rest of their range. As of 1996, cougar hunting was prohibited in Argentina, Brazil, Bolivia, Chile, Colombia, Costa Rica, French Guiana, Guatemala, Honduras, Nicaragua, Panama, Paraguay, Suriname, Venezuela, and Uruguay. (Costa Rica and Panama are not listed as current range countries by the IUCN.) The cat had no reported legal protection in Ecuador, El Salvador, and Guyana (Nowell and Jackson 2006). Regulated cougar hunting is still common in the United States and Canada, although they are protected from all hunting in the Yukon; it is permitted in every U.S. state from the Rocky Mountains to the Pacific Ocean, with the exception of California. Cougars are generally hunted with packs of dogs, until the animal is "treed." When the hunter arrives on the scene, he shoots the cat from the tree at close range. The cougar cannot be legally killed in California except under very specific circumstances, such as when an individual is declared a public safety threat (CDFG 2007). However statistics from the Department of Fish and Game indicate that cougar killings in California have been on the rise since 1970s with an average of over 112 cats killed per year from 2000 to 2006 compared to 6 per year in the 1970s.
Conservation threats to the species include persecution as a pest animal, degradation and fragmentation of their habitat, and depletion of their prey base. Habitat corridors and sufficient range areas are critical to the sustainability of cougar populations. Research simulations have shown that the animal faces a low extinction risk in areas of 2200 km² (850 sq mi) or more. As few as one to four new animals entering a population per decade markedly increases persistence, foregrounding the importance of habitat corridors (Beier 1993).
Due to the growth of urban areas, cougar ranges increasingly overlap with areas inhabited by humans. Attacks on humans are rare, as cougar prey recognition is a learned behavior and they do not generally recognize humans as prey (McKee 2003). Attacks on people, livestock, and pets may occur when the cat habituates to humans. There have been 108 confirmed attacks on humans with twenty fatalities in North America since 1890, fifty of the incidents having occurred since 1991 (AZGFD 2007; Beier 1991). The heavily populated state of California has seen a dozen attacks since 1986 (after just three from 1890 to 1985), including three fatalities (CDFG 2007). Attacks are most frequent during late spring and summer, when juvenile cougars leave their mothers and search for new territory (GovBC 1991).
As with many predators, a cougar may attack if cornered, if a fleeing human being stimulates their instinct to chase, or if a person "plays dead." Exaggerating the threat to the animal through intense eye contact, loud but calm shouting, and any other action to appear larger and more menacing, may make the animal retreat. Fighting back with sticks and rocks, or even bare hands, is often effective in spurring an attacking cougar to disengage (McKee 2003; GovBC 1991).
When the cougar does attack, they usually employs their characteristic neck bite, attempting to position their teeth between the vertebrae and into the spinal cord. Neck, head, and spinal injuries are common and sometimes fatal (McKee 2003). Children are at greatest risk of attack, and least likely to survive an encounter. Detailed research into attacks prior to 1991 showed that 64 percent of all victims—and almost all fatalities—were children. The same study showed the highest proportion of attacks to have occurred in British Columbia, particularly on Vancouver Island where cougar populations are especially dense (Beier 1991).
The grace and power of the cougar have been widely admired in the cultures of the indigenous peoples of the Americas. The Inca city of Cusco is reported to have been designed in the shape of a cougar, and the animal also gave their name to both Inca regions and people. The Moche people represented the puma often in their ceramics (Berrin and Larco Museum 1997). The sky and thunder god of the Inca, Viracocha, has been associated with the animal (Kulmar 2007).
In North America, mythological descriptions of the cougar have appeared in the stories of the Hotcâk language ("Ho-Chunk" or "Winnebago") of Wisconsin and Illinois (Blowsnake) and the Cheyenne, among others. To the Apache and Walapai of Arizona, the wail of the cougar was harbinger of death (USDA 2007).
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