Cougar

For other uses, see Cougar (disambiguation), Puma (disambiguation), or Panther.

Cougar Temporal range: Middle Pleistocene to Recent
File:Mountain lion.jpg
Conservation status
Near Threatened  (IUCN 3.1)
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Family:	Felidae
Genus:	Puma
Species:	P. concolor
Binomial name
Puma concolor (Linnaeus, 1771)
Cougar range map

The cougar (Puma concolor), also puma, mountain lion, or panther depending on region, is a mammal of the Felidae family, native to the Americas. This large, solitary cat has the greatest range of any wild terrestrial mammal in the Western Hemisphere, extending from Yukon in Canada to the southern Andes of South America. An adaptable, generalist species, the cougar is found in every major New World habitat type. It 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.

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. It prefers habitats with dense underbrush and rocky areas for stalking, but it can live in open areas. The cougar is territorial and persists at low population densities. Individual territory sizes depend on terrain, vegetation, and abundance of prey. While it is a large predator, it is not always the dominant species in its range, as when it competes for prey with animals such as the gray wolf, black bear, and the grizzly bear. It is a reclusive cat and usually avoids people. Attacks on humans remain rare, despite a recent increase in frequency.

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 names and various references in the mythology of the indigenous peoples of the Americas and in contemporary culture.

Naming and etymology

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 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.

"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.

Taxonomy and evolution

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. 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, as cats are poorly represented in the fossil record, and there are significant confidence intervals with suggested dates.

Although large, the cool cat is closely related to small felines.

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 (Ma) ago. The lineages subsequently diverged in that order. North American felids then invaded South America 3 Ma ago 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, but the relationship is unresolved. It has been suggested that the cheetah lineage diverged from the Puma lineage in the Americas (see American cheetah) and migrated back to Asia and Africa, while other research suggests the cheetah diverged in the Old World itself. 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. 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.

Subspecies

Until the late 1990s, as many as 32 subspecies were recorded; however, a recent genetic study of mitochondrial DNA 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:

Argentine puma (Puma concolor cabrerae)

includes the previous subspecies and synonyms hudsonii and puma (Marcelli, 1922);

Costa Rican Cougar (Puma concolor costaricensis)

Eastern South American cougar (Puma concolor anthonyi)

includes the previous subspecies and synonyms acrocodia, borbensis, capricornensis, concolor (Pelzeln, 1883), greeni and nigra;

North American Cougar (Puma concolor couguar)

includes the previous subspecies and synonyms arundivaga, aztecus, browni, californica, coryi, floridana, hippolestes, improcera, kaibabensis, mayensis, missoulensis, olympus, oregonensis, schorgeri, stanleyana, vancouverensis and youngi;

Northern South American cougar (Puma concolor concolor)

includes the previous subspecies and synonyms bangsi, incarum, osgoodi, soasoaranna, soderstromii, sucuacuara and wavula;

Southern South American puma (Puma concolor puma)

includes the previous subspecies and synonyms araucanus, concolor (Gay, 1847), patagonica, pearsoni and puma (Trouessart, 1904)

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. Culver et al. 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."

Biology and behavior

Physical characteristics

Although cougars resemble the domestic cat, they are about the same size as an adult human.

Cougars are slender and agile cats. Adults stand about 60 to 80 cm (2.0 to 2.7 ft) tall at the shoulders. The length of adult males is around 2.4 m (8 ft) long nose to tail, with overall ranges between 1.5 and 2.75 meters (5 and 9 ft) nose to tail suggested for the species in general. Males have an average weight of about 53 to 72 kilograms (115 to 160 pounds). In rare cases, some may reach over 120 kg (260 lb). Female average weight is between 34 and 48 kg (75 and 105 lb). Cougar size is smallest close to the equator, and larger towards the poles.

The head of the cat is round and the ears erect. Its powerful forequarters, neck, and jaw serve to grasp and hold large prey. It has five retractable claws on its forepaws (one a dewclaw) and four on its hind paws. The larger front feet and claws are adaptations to clutching prey.

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. Despite its size, it is not typically classified among the "big cats," as it cannot roar, lacking the specialized larynx and hyoid apparatus of Panthera. 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.

paw of a cougar

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-grey 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; juveniles are pale, and dark spots remain on their flanks. Despite anecdotes to the contrary, all-black coloring (melanism) has never been documented in cougars. The term "black panther" is used colloquially to refer to melanistic individuals of other species, particularly jaguars and leopards.

Cougars have large paws and proportionally the largest hind legs in the cat family. This physique allows it great leaping and short-sprint ability. An exceptional vertical leap of 5.4 m (18 ft) is reported for the cougar. Horizontal jumping capability is suggested anywhere from 6 to 12 m (20 to 40 ft). The cougar can run as fast as 55 km/h (35 mph), but is best adapted for short, powerful sprints rather than long chases. It is adept at climbing, which allows it to evade canine competitors. Although it is not strongly associated with water, it can swim.

Hunting and diet

Cougars are ambush predators, feeding mostly on deer and other mammals.

A successful generalist predator, the cougar will eat any animal it can catch, from insects to large ungulates. Like all cats, it is an obligate carnivore, feeding only on meat. 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% of prey items were ungulates, especially deer. Only the Florida Panther showed variation, often preferring feral hogs and armadillos. 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. 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.

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% 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. 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.

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.

Reproduction and lifecycle

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; the period can be as short as one year. Females are in estrus for approximately 8 days of a 23-day cycle; the gestation period is approximately 91 days. Females are sometimes reported as monogamous, but this is uncertain and polygyny may be more common. Copulation is brief but frequent.

Cougar kittens

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. Kitten survival rates are just over one per litter.

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 amongst cougars that travel farthest from the maternal range, often due to conflicts with other cougars ("intraspecific" conflict). 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."

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. 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.

Social structure and home range

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.

Estimates of territory sizes vary greatly. Canadian Geographic reports large male territories of 150 to 1000 square kilometers (58 to 386 sq mi) with female ranges half the size. Other research suggests a much smaller lower limit of 25 km² (10 sq mi) but an even greater upper limit of 1300 km² (500 sq mi) for males. 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). 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.

Home range sizes and overall cougar abundance depend on terrain, vegetation, and prey abundance. 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. 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).

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. When males encounter each other, they hiss and spit, and may engage in violent conflict if neither backs down. Hunting or relocation of the cougar may increase aggressive encounters by disrupting territories and bringing young, transient animals into conflict with established individuals.

Ecology

Distribution and habitat

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. 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. Its preferred habitats include precipitous canyons, escarpments, rim rocks, and dense brush.

Cougar, photographed in the Arizona-Sonora Desert Museum, Tucson, Arizona.

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, it ranges across most western American states, the Canadian provinces of Alberta and British Columbia, and the Canadian Yukon Territory. There have been widely-debated reports of possible recolonization of eastern North America. DNA evidence has suggested its presence in eastern North America, while a consolidated map of cougar sightings shows numerous reports, from the mid-western Great Plains through to Eastern Canada. 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. While specific state and provincial statistics are often available in North America, much less is known about the cat in its southern range.

The cougar's total breeding population is estimated at less than 50,000 by the IUCN, with a declining trend. 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. California has actively sought to protect the cat and a similar number of cougars has been suggested, between 4,000 and 6,000.

Ecological role

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 brown bear and gray wolf. In the south, the cougar must compete with the larger jaguar. In Florida it encounters the American alligator.

Front paw print of a cougar. An adult paw print is approximately 10 cm (4 inches) long.

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% of cougar kills in Yellowstone and Glacier National Parks, usurping 10% of carcasses.

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 fourteen wolves killing a female cougar and her kittens. Conversely, lone wolves are at a disadvantage, and have been reported killed by cougars. 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. 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." Both species, meanwhile, are capable of killing mid-sized predators such as bobcats and coyotes and tend to suppress their numbers.

In the southern portion of its range, the cougar and jaguar share overlapping territory. The jaguar tends to take larger prey and the cougar smaller where they overlap, reducing the cougar's size. Of the two felines, the cougar appears best able to exploit a broader prey niche and smaller prey.

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. 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.

Hybrids

Pumapard, taken in 1904

Main article: Pumapard

A pumapard is a hybrid animal resulting from a union between a cougar and a leopard. Three sets of these hybrids were bred in the late 1890s and early 1900s by Carl Hagenbeck at his animal park in Hamburg, Germany. Most did not reach adulthood. One of these was purchased in 1898 by Berlin Zoo. A similar hybrid in Berlin Zoo purchased from Hagenbeck was a cross between a male leopard and a female puma. Hamburg Zoo's specimen was the reverse pairing, the one in the black and white photo, fathered by a puma bred to an Indian leopardess. Whether born to a female puma mated to a male leopard, or to a male puma mated to a female leopard, pumapards inherit a form of dwarfism. Those reported grew to only half the size of the parents. They have a puma-like long body (proportional to the limbs, but nevertheless shorter than either parent), but short legs. The coat is variously described as sandy, tawny or greyish with brown, chestnut or "faded" rosettes.

Conservation status

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. The cougar is regulated under Appendix I of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), rendering illegal international trade in specimens or parts.

Cougar conservation depends on preservation of their habitat.

East of the Mississippi, the only unequivocally known cougar population is the Florida panther. The U.S. Fish and Wildlife Service recognizes both an Eastern cougar and the Florida panther, affording protection under the Endangered Species Act. Certain taxonomic authorities have collapsed both designations into the North American Cougar, with Eastern or Florida subspecies not recognized, while a subspecies designation remains recognized by some conservation scientists. The most recent documented count for the Florida sub-population is 87 individuals, reported by recovery agencies in 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. 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. 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.

Attacks on humans

See also: List of fatal cougar attacks in North America by decade

Cougar warning message

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. 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. The heavily populated state of California has seen a dozen attacks since 1986 (after just three from 1890 to 1985), including three fatalities. Attacks are most frequent during late spring and summer, when juvenile cougars leave their mothers and search for new territory.

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.

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. Children are at greatest risk of attack, and least likely to survive an encounter. Detailed research into attacks prior to 1991 showed that 64% 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.

In mythology and culture

File:MochePuma.jpg

Moche puma, Larco Museum collection

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. The sky and thunder god of the Inca, Viracocha, has been associated with the animal.

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 and the Cheyenne, amongst others. To the Apache and Walapai of Arizona, the wail of the Cougar was harbinger of death.

The cougar continues to be a symbol of strength and stealth. From combat helicopters, motor vehicles (see Ford/Mercury Cougar and Ford Puma) to athletic shoes, both "Cougar" and "Puma" are widely used as brand names. Various sports teams have also adopted the names, including the Argentina national rugby union team as well as US universities, Brigham Young University, The University of Houston, and Washington State University. Many places, such as Cougar Mountain, are also named after their association with cougars.

Other resources

• Baron, David (2004). The Beast in the Garden: A Modern Parable of Man and Nature. New York: W. W. Norton and Company. ISBN 0393058077.
• Kobalenko, Jerry (1997). Forest Cats of North America. Buffalo, New York: Island Press. ISBN 1-55209-174-4. {{cite book}}: Check |isbn= value: checksum (help)
• Logan, Ken (2001). Desert Puma: Evolutionary Ecology and Conservation of an Enduring Carnivore. Island Press. ISBN 1-55963-866-4. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

External links

• Description of a Cougar attack
• Cougar Facts and Photos - NatureMapping Program
• No Place for Predators? Liza Gross, PLoS Biology, explains how Washington State wildlife officials implemented a hunting policy, in response to a state measure passed to protect wildlife, that led to the highest rates of human-caused cougar mortality since the height of the bounty era