Cougar

Following Linnaeus' first scientific description of the cougar, 32 cougar zoological specimens were described and proposed as subspecies until the late 1980s. Genetic analysis of cougar mitochondrial DNA indicate that many of these are too similar to be recognized as distinct at a molecular level, but that only six phylogeographic groups exist. The Florida panther samples showed a low microsatellite variation, possibly due to inbreeding.[20] Following this research, the authors of Mammal Species of the World recognized the following six subspecies in 2005:[2]

• P. c. concolor (Linnaeus, 1771) includes the synonyms bangsi, incarum, osgoodi, soasoaranna, sussuarana, soderstromii, suçuaçuara, and wavula
• P. c. puma (Molina, 1782) includes the synonyms araucanus, concolor, patagonica, pearsoni, and puma (Trouessart, 1904)
• P. c. couguar (Kerr, 1792) includes arundivaga, aztecus, browni, californica, floridana, hippolestes, improcera, kaibabensis, mayensis, missoulensis, olympus, oregonensis, schorgeri, stanleyana, vancouverensis, and youngi
• P. c. costaricensis (Merriam, 1901)
• P. c. anthonyi (Nelson and Goldman, 1931) includes acrocodia, borbensis, capricornensis, concolor, greeni, and nigra
• P. c. cabrerae Pocock, 1940 includes hudsonii and puma proposed by Marcelli in 1922

In 2006, the Florida panther was still referred to as a distinct subspecies P. c. coryi in research works.[21]

As of 2017, the Cat Classification Taskforce of the Cat Specialist Group recognizes only two subspecies as valid:[22]

• P. c. concolor in South America, possibly excluding the region northwest of the Andes
• P. c. couguar in North and Central America and possibly northwestern South America

The family Felidae is believed to have originated in Asia about 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.[19] Significant confidence intervals exist with suggested dates. In the latest genomic study of the Felidae, the common ancestor of today's Leopardus, Lynx, Puma, Prionailurus, and Felis lineages migrated across the Bering land bridge into the Americas 8.0 to 8.5 million years ago (Mya). The lineages subsequently diverged in that order.[19] North American felids then invaded South America 2–4 Mya as part of the Great American Interchange, following the formation of the Isthmus of Panama.[20] but the relationship is unresolved. The cheetah lineage is suggested by some studies to have diverged from the Puma lineage in the Americas and migrated back to Asia and Africa,[19][20] while other research suggests the cheetah diverged in the Old World itself.[23] A high level of genetic similarity has been found among North American cougar populations, suggesting they are all fairly recent descendants of a small ancestral group. Culver et al. propose the original North American cougar population 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 South American cougars.[20]

A coprolite identified as from a cougar was excavated in Argentina's Catamarca Province and dated to 17,002–16,573 years old. It contained Toxascaris leonina eggs. This finding indicates that the cougar and the parasite existed in South America since at least the Late Pleistocene.[24]

With the increase of human development and infrastructure growth in California, the cougar populations in the state are becoming more isolated from one another.[71]

A successful generalist predator, the cougar will eat any animal it can catch, from insects to large ungulates (over 500 kg (1,100 lb)). Like other cats, it is an obligate carnivore, meaning it must feed on meat to survive. The mean weight of vertebrate prey (MWVP) that pumas attack increases with the puma's body weight; in general, MWVP is lower in areas closer to the equator.[35] Its most important prey species are various deer species, particularly in North America; mule deer, white-tailed deer, elk, and even bull moose are taken. Other species taken include the bighorn and Dall's sheep, horses, fallow deer, caribou, mountain goat, coyote, American badger and pronghorn.[97] 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.[35]

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

Investigations at Yellowstone National Park showed that elk, followed by mule deer, were the cougar's primary targets; the prey base is shared with the park's gray wolves, with which the cougar competes for resources.[84] 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.[98]

In Pacific Rim National Park Reserve, scat samples showed raccoons to make up 28% of the cougar's diet, harbor seals and blacktail deer 24% each, North American river otters 10%, California sea lion 7%, and American mink 4%; the remaining 3% were unidentified.[99]

In the Central and South American cougar range, the ratio of deer in the diet declines. Small to mid-sized mammals are preferred, including large rodents such as the capybara. Ungulates accounted for only 35% of prey items in one survey, about half that of North America. Competition with the larger jaguar in South America has been suggested for the decline in the size of prey items.[35] However, a study by Gutiérrez-González and López-González showed that the cougar and jaguar in Central or North America may share the same prey, depending on its abundance.[100] Other listed prey species of the cougar include mice, porcupines, beavers, raccoons, hares, guanaco, peccary, vicuna, rhea, and wild turkey.[101] Birds and small reptiles are sometimes preyed upon in the south, but this is rarely recorded in North America.[35]

In Patagonia's Monte León National Park, the local cougar population has specialized on hunting penguins.[102]

Although 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.[26] Kills are generally estimated 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 around 15 months.[33] The cat drags a kill to a preferred spot, covers it with brush, and returns to feed over a period of days. The cougar is generally reported to not be a scavenger, but deer carcasses left exposed for study were scavenged by cougars in California, suggesting more opportunistic behavior.[103]

North American cougar cub at Malibu Springs

Cubs

Females reach sexual maturity between one-and-a-half to three years of age. They typically average one litter every two to three years throughout their reproductive lives,[104] though the period can be as short as one year.[33] Females are in estrus for about 8 days of a 23-day cycle; the gestation period is approximately 91 days.[33] Females are sometimes reported as monogamous,[105] but this is uncertain and polygyny may be more common.[106] Copulation is brief but frequent. Chronic stress can result in low reproductive rates when in captivity as well as in the field.[107]

Only females are involved in parenting. Litter size is between one and six cubs; typically two. Caves and other alcoves that offer protection are used as litter dens. Born blind, cubs 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.[104] Kitten survival rates are just over one per litter.[33] Newborn cougars have spots that fade and eventually disappear by the age of 2 1/2 years.[108] Juveniles remain with their mothers at least for two years.[109]

Young adults leave their mother to attempt to establish their own territories at around two years of age and sometimes earlier; males tend to leave sooner. One study has shown high mortality amongst cougars that travel farthest from the maternal range, often due to conflicts with other cougars (intraspecific competition).[104] 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."[110]

Life expectancy in the wild is reported at eight to 13 years, and probably averages eight to 10; a female of at least 18 years was reported killed by hunters on Vancouver Island.[33] 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 HIV-like virus in cats, is well-adapted to the cougar.[111]

Like almost all cats, the cougar is a mostly solitary animal. Only mothers and kittens live in groups, with adults meeting rarely. While generally loners, cougars will reciprocally share kills with one another and seem to organize themselves into small communities defined by the territories of dominant males. Cats within these areas socialize more frequently with each other than with outsiders.[112] Estimates of territory sizes for cougars vary greatly. Canadian Geographic reports large male territories of 150 to 1000 km² (58 to 386 sq mi) with female ranges half that size.[105] 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.[104] 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).[113] Male ranges may include or overlap with those of females but, at least where studied, not with those of other males, which reduces 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.[114]

Home range sizes and overall cougar abundance depend on terrain, vegetation, and prey abundance.[104] One female adjacent to the San Andres Mountains was found with a large range of 215 km² (83 sq mi), necessitated by poor prey abundance.[110] Research has shown cougar abundances from 0.5 animals to as many as 7 (in one study in South America) per 100 km² (39 sq mi).[33]

Because males disperse farther than females and compete more directly for mates and territory, they are more likely to be involved in conflict. Where a juvenile fails to leave his maternal range, for example, he may be killed by his father.[113] When males encounter each other, they hiss, spit, and may engage in violent conflict if neither backs down.[106] Hunting or relocation of the cougar may increase aggressive encounters by disrupting territories and bringing young, transient animals into conflict with established individuals.[115]

Mountain Lion warning sign in California, US.

The pertinent North American subspecies is P. concolor couguar. Due to the expanding human population, cougar ranges increasingly overlap with areas inhabited by humans. Attacks on humans are very rare, as cougar prey recognition is a learned behavior and they do not generally recognize humans as prey.[129] In a 10-year study in New Mexico of wild cougars who were not habituated to humans, the animals did not exhibit threatening behavior to researchers who approached closely (median distance=18.5 m; 61 feet) except in 6% of cases; 14/16 of those were females with cubs.[130] Attacks on people, livestock, and pets may occur when a puma habituates to humans or is in a condition of severe starvation. Attacks are most frequent during late spring and summer, when juvenile cougars leave their mothers and search for new territory.[25]

Between 1890 and 1990, in North America there were 53 reported, confirmed attacks on humans, resulting in 48 nonfatal injuries and 10 deaths of humans (the total is greater than 53 because some attacks had more than one victim).[131] By 2004, the count had climbed to 88 attacks and 20 deaths.[132]

Within North America, the distribution of attacks is not uniform. The heavily populated state of California saw a dozen attacks 1986 to 2004 (after just three from 1890 to 1985), including three fatalities.[66] Washington state was the site of a fatal attack in 2018, its first since 1924.[133] Lightly populated New Mexico reported an attack in 2008, the first there since 1974.[134]

As with many predators, a cougar may attack if cornered, if a fleeing human stimulates their instinct to chase, or if a person "plays dead". Standing still may cause the cougar to consider a person easy prey.[135] Exaggerating the threat to the animal through intense eye contact, loud 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 persuading an attacking cougar to disengage.[129][25]

When cougars do attack, they usually employ 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.[129] 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.[131] Preceding attacks on humans, cougars display aberrant behavior, such as activity during daylight hours, a lack of fear of humans, and stalking humans. There have sometimes been incidents of pet cougars mauling people.[136][137]

Research on new wildlife collars may be able to reduce human-animal conflicts by predicting when and where predatory animals hunt. This may save the lives of humans, pets, and livestock as well as the lives of these large predatory mammals that are important to the balance of ecosystems.[138]

Pumas in the Southern cone of America – often called Argentine cougars by North Americans – are reputed to be extremely reluctant to attack man; in legend, they defended people against jaguars.[139] The nineteenth century naturalists Félix de Azara[140] and William Henry Hudson[141] thought that attacks on people, even children or sleeping adults, did not happen. Hudson, citing anecdotal evidence from hunters, claimed that pumas were positively inhibited from attacking people, even in self-defense. In fact, attacks on humans, although exceedingly rare, have occurred.[142][143]

An early, authenticated, non-fatal case occurred near Lake Viedma, Patagonia in 1877 when a female mauled the Argentine scientist Francisco P. Moreno; Moreno afterwards showed the scars to Theodore Roosevelt. In this instance, however, Moreno had been wearing a guanaco-hide poncho round his neck and head as protection against the cold;[144] in Patagonia the guanaco is the puma's chief prey animal.[145] Another authenticated case occurred In 1997 in Iguazú National Park, northeast Argentina when the 20-month son of a ranger was killed by a female puma. Forensic analysis found specimens of the child's hair and clothing fibers in the animal's stomach. In this area the coatí is the puma's chief prey. Despite prohibitory signs, coatis are hand-fed by tourists in the park, causing unnatural approximation between cougars and humans. This particular puma had been raised in captivity and released into the wild.[146] In 2012 a 23-year-old woman was found dead in a mountainous area in Salta Province in northwest Argentina. Claw incisions, which severed a jugular vein, indicated that the attacker was a felid; differential diagnosis ruled out other possible perpetrators.[lower-alpha 1] There were no bite marks on the victim, who had been herding goats.[147] In 2019 in Córdoba Province an elderly man was badly injured by a puma after he attempted to defend his dog from it.[148]

Fatal attacks by other carnivores such as feral dogs can be misattributed to pumas without appropriate forensic knowledge.[149]

During the early years of ranching, cougars were considered on par with wolves in destructiveness. According to figures in Texas in 1990, 86 calves (0.0006% of Texas's 13.4 million cattle and calves), 253 mohair goats, 302 mohair kids, 445 sheep (0.02% of Texas's 2 million sheep and lambs) and 562 lambs (0.04% of Texas's 1.2 million lambs) were confirmed to have been killed by cougars that year.[150][151] In Nevada in 1992, cougars were confirmed to have killed nine calves, one horse, four foals, five goats, 318 sheep, and 400 lambs. In both reports, sheep were the most frequently attacked. Some instances of surplus killing have resulted in the deaths of 20 sheep in one attack.[152] A cougar's killing bite is applied to the back of the neck, head, or throat and the cat inflicts puncture marks with its claws usually seen on the sides and underside of the prey, sometimes also shredding the prey as it holds on. Coyotes also typically bite the throat but the work of a cougar is generally clean, while bites inflicted by coyotes and dogs leave ragged edges. The size of the tooth puncture marks also helps distinguish kills made by cougars from those made by smaller predators.[153]

Remedial hunting appears to have the paradoxical effect of increased livestock predation and complaints of human-puma conflicts. In a 2013 study the most important predictor of puma problems were remedial hunting of puma the previous year. Each additional puma on the landscape increased predation and human-puma complaints by 5%, but each additional animal killed on the landscape during the previous year increased complaints by 50%. The effect had a dose-response relationship with very heavy (100% removal of adult puma) remedial hunting leading to a 150% – 340% increase in livestock and human conflicts.[154] This effect is attributed to the removal of older pumas that have learned to avoid people and their replacement by younger males that react differently to humans. Remedial hunting enables younger males to enter the former territories of the older animals.[155][156]

Predation by cougars on dogs "is widespread, but occurs at low frequencies"[157]

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 its name to both Inca regions and people. The Moche people represented the puma often in their ceramics.[158] The sky and thunder god of the Inca, Viracocha, has been associated with the animal.[159]

In North America, mythological descriptions of the cougar have appeared in the stories of the Hocąk language ("Ho-Chunk" or "Winnebago") of Wisconsin and Illinois[160] and the Cheyenne, amongst others. To the Apache and Walapai of Arizona, the wail of the cougar was a harbinger of death.[161] The Algonquins and Ojibwe believe that the cougar lived in the underworld and was wicked, whereas it was a sacred animal among the Cherokee.[162]

Several sports teams currently, or in the past, have used the cougar as the mascot or nickname for their team, although the name actually used has depended on the most popular regional name for the species. Among college sports teams, Brigham Young University in Utah and Washington State University in the northwestern United States, as well as the University of Houston in Texas use the cougar as their mascot. On Canada's prairies, Mount Royal University in Calgary and the University of Regina in Saskatchewan use the cougar as their mascot. The University of Vermont also uses the mascot, but uses the term "catamount" instead of cougar, as was traditional in the region where the school is located.

In professional hockey, the cougar was used by two Midwestern teams and one Northwestern team. The Chicago Cougars of the World Hockey Association operated from 1972 to 1975, while the Detroit Red Wings of the National Hockey League were called the Cougars from their inaugural season in 1926 until 1930. The Detroit Cougars were actually related to the Victoria Cougars of the Western Hockey League, which had won the Stanley Cup in 1925, in that when the Victoria Cougars, which had operated as the Aristocrats from 1918 to 1922 and as the Cougars from 1922 to 1926, disbanded in 1926, the owners of the newly formed Detroit club purchased the rights to many of the players of the Victoria club, and retained the Cougar nickname. The Florida Panthers of the National Hockey League use the name and image of the southeastern United States cougar subspecies, the Florida panther (the state animal), as its mascot.

The University of Pittsburgh in western Pennsylvania also uses the cougar as its sports mascot, and for many other clubs and organizations, but also uses the locally preferred name, "panther", corresponding to nearby geographic features (Panther Hollow and Panther Hollow Lake which got their names by 1885, 24 years before the University selected the name for its teams).[163] The University of Pittsburgh campus has about 20 physical representations of panthers or cougars at different locations around the campus, as well as four such statues on the Panther Hollow Bridge over Panther Hollow.

By contrast, Penn State University in central Pennsylvania uses the Nittany Lion as its mascot, "Nittany" being the name of a nearby mountain, as well as the valley in which the university is located, and "lion" being the abbreviated name of a stuffed mountain lion/cougar originally captioned as "Brush Lion" that was on display in a building at the university, and which is today housed in the Penn State All-Sports Museum.[164] The campus also has a Nittany Lion Shrine, featuring a large statue of a mountain lion on campus.

Many high schools also use the cougar as their sports team mascot.

The Sacramento Mountain Lions (2010–2012) of the United Football League used the California and southwestern United States regional name of the species. The Carolina Panthers of the National Football League have a black cat on their uniforms, as opposed to a brownish- or tawny-coloured cat, so it is most often presumed to represent a black panther, the black melanistic phase of either the jaguar (which actually only lived in the far Southwestern United States, and not the Southeastern United States where the Carolinas are located), or the leopard of Africa and Asia. A previously existing "Carolina" sports team, the Carolina Cougars, who operated in the American Basketball Association from 1969 to 1974, actually used the cougar, as opposed to the panther, as its nickname and mascot.

Most sports teams named the "Wildcats" use a logo similar to a bobcat, or in some cases in Canada, a Canada lynx, names which are often used for sports teams in their own right. By contrast, Wildcat Lager Beer brewed by Labatt, has always shown a picture of a cougar on its label.