Puma concolor

The cougar (''Puma concolor'') is a large cat native to the Americas. Its range spans from the Canadian Yukon to the southern Andes in South America and is the most widespread of any large wild terrestrial mammal in the Western Hemisphere. It is an adaptable, generalist species, occurring in most American habitat types. This wide range has brought it many common names, including puma, mountain lion, catamount and panther (for the Florida sub-population). It is the second-largest cat in the New World, after the jaguar (''Panthera onca''). Secretive and largely solitary by nature, the cougar is properly considered both nocturnal and crepuscular, although daytime sightings do occur. Despite its size, the cougar is more closely related to smaller felines, including the domestic cat (''Felis catus'') than to any species of the subfamily Pantherinae.

The cougar is an ambush predator that pursues a wide variety of prey. Primary food sources are ungulates, particularly deer, but it also hunts smaller prey such as rodents. It prefers habitats with dense underbrush and rocky areas for stalking, but also lives in open areas. Cougars are territorial and live at low population densities. Individual home ranges depend on terrain, vegetation and abundance of prey. While large, it is not always the apex predator in its range, yielding prey it has killed to American black bears, grizzly bears and packs of wolves. It is reclusive and mostly avoids people. Fatal attacks on humans are rare, but increased in North America as more people entered cougar habitat and built farms.

Intensive hunting following European colonization of the Americas and ongoing human development into cougar habitat has caused the cougar populations to drop in most parts of its historical range. In particular, the eastern cougar population is considered to have been mostly extirpated in eastern North America at the beginning of the 20th century, with the exception of the isolated Florida panther subpopulation.

Naming and etymology

The word ''cougar'' is borrowed from the Portuguese ''çuçuarana'', via French; it was originally derived from the Tupi language. A current form in Brazil is ''suçuarana''. In the 17th century, Georg Marcgrave named it ''cuguacu ara''. Marcgrave's rendering was reproduced in 1648 by his associate Willem Piso. ''Cuguacu ara'' was then adopted by John Ray in 1693. In 1774, Georges-Louis Leclerc, Comte de Buffon converted ''cuguacu ara'' to ''cuguar'', which was later modified to "cougar" in English.

The cougar holds the Guinness record for the animal with the greatest number of names, with over 40 in English alone. "Puma" is the common name used in Latin America and most parts of Europe. The term ''puma'' is also sometimes used in the United States. The first use of ''puma'' in English dates to 1777, introduced from Spanish from the Quechua language. In the western United States and Canada, it is also called "mountain lion", a name first used in writing in 1858. Other names include "panther" and "catamount" (meaning "cat of the mountains").

Taxonomy and evolution

''Felis concolor'' was the scientific name proposed by Carl Linnaeus in 1771 for a cat with a long tail from Brazil. The second half of the name, "concolor" is Latin for "of uniform color". It was placed in the genus ''Puma'' by William Jardine in 1834.
This genus is part of the Felinae. The cougar is most closely related to the jaguarundi and the cheetah.

Subspecies

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. Following this research, the authors of '' Mammal Species of the World'' recognized the following six subspecies in 2005:
* ''P. c. concolor'' includes the synonyms ''bangsi'', ''incarum'', ''osgoodi'', ''soasoaranna'', ''sussuarana'', ''soderstromii'', ''suçuaçuara'', and ''wavula''
* ''P. c. puma'' includes the synonyms ''araucanus'', ''concolor'', ''patagonica'', ''pearsoni'', and ''puma''
* ''P. c. couguar'' includes ''arundivaga'', ''aztecus'', ''browni'', ''californica'', ''floridana'', ''hippolestes'', ''improcera'', ''kaibabensis'', ''mayensis'', ''missoulensis'', ''olympus'', ''oregonensis'', ''schorgeri'', ''stanleyana'', ''vancouverensis'', and ''youngi''
* ''P. c. costaricensis''
* ''P. c. anthonyi'' includes ''acrocodia'', ''borbensis'', ''capricornensis'', ''concolor'', ''greeni'', and ''nigra''
* ''P. c. cabrerae'' 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.

, the Cat Classification Taskforce of the Cat Specialist Group recognizes only two subspecies as valid:
*'' 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

Evolution

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.
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. North American felids then invaded South America 2–4Mya as part of the Great American Interchange, following the formation of the Isthmus of Panama.

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, while other research suggests the cheetah diverged in the Old World itself.
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.

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.

Characteristics

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

Cougars are slender and agile members of the Felidae. They are the fourth largest cat species worldwide; adults stand about tall at the shoulders. Adult males are around long from nose to tail tip, and females average , with overall ranges between nose to tail suggested for the species in general. Of this length, the tail typically accounts for .Shivaraju, A. (2003
''Puma concolor''
. Animal Diversity Web, University of Michigan Museum of Zoology. Retrieved on September 15, 2011. Males generally weigh , averaging . Females typically weigh between , averaging . Cougar size is smallest close to the equator and larger towards the poles. The largest recorded cougar, shot in 1901, weighed ; claims of and have been reported, though they were probably exaggerated. On average, adult male cougars in British Columbia weigh and adult females , though several male cougars in British Columbia weighed between .

Depending on the locality, cougars can be smaller or bigger than jaguars, but are less muscular and not as powerfully built, so their weight is, on average, less. Whereas cougars tend to be larger as distance increases from the equator, which crosses the northern portion of South America, jaguars are generally smaller north of the Amazon River in South America and larger south of it. For example, while South American jaguars are comparatively large, and may exceed , North American jaguars in Mexico's Chamela-Cuixmala Biosphere Reserve weigh approximately , about the same as female cougars.

Cougar coloring is plain (hence the Latin ''concolor'' one color"in the scientific name), but can vary greatly across individuals, and even siblings. The coat is typically tawny, but it otherwise ranges from silvery-grey or reddish with lighter patches on the underbody, 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. A leucistic individual was seen in Serra dos Órgãos National Park in Rio de Janeiro in 2013 when it was recorded by a camera trap, indicating that extremely rare, pure white individual cougars do exist in the species.

The cougar has large paws and proportionally the largest hind legs in the Felidae, allowing for its great leaping and short-sprint ability. It is capable of leaping from the ground up to high into a tree.

Distribution and habitat

The cougar has the largest range of any wild land animal in the Americas, spanning 110 degrees of latitude from the Yukon Territory in Canada to the southern Andes in Chile. The species was extirpated from eastern North America, aside from Florida, but they may be recolonizing their former range and isolated populations have been documented east of their contemporary ranges in both the US and Canada.

The cougar lives in all forest types, lowland and mountainous deserts and in open areas with little vegetation up to an elevation of . In the Santa Ana Mountains, it prefers steep canyons, escarpments, rim rocks and dense brush.
In Mexico, it was recorded in the Sierra de San Carlos. In the Yucatán Peninsula, it inhabits secondary and semi-deciduous forests in El Eden Ecological Reserve.
In El Salvador, it was recorded in lower montane forest in Montecristo National Park and in a river basin in the Morazán Department above in 2019.
In Colombia, it was recorded in a palm oil plantation close to a riparian forest in the Llanos Basin, and close to water bodies in the Magdalena River Valley.
In the human-modified landscape of central Argentina, it inhabits bushland with abundant vegetation cover and prey species.

Behavior and ecology

Cougars are important keystone species in Western Hemisphere ecosystems, linking numerous different species at many trophic levels. In a comprehensive literature review of more than 160 studies on cougar ecology, ecological interactions with 485 other species in cougar-inhabited ecosystems have been shown to involve different areas of interaction, ranging from the use of other species as food sources and prey, fear effects on potential prey, effects from carcass remains left behind, to competitive effects on other predator species in shared habitat. The most common research topic in the literature used here was the diet of the cougar and the regulation of its prey.Laura R. LaBarge, Michael J. Evans, Jennifer R. B. Miller, Gillian Cannataro, Christian Hunt, L. Mark Elbroch: ''Pumas ''Puma concolor'' as ecological brokers: a review of their biotic relationships.'' In: ''Mammal Review'' 25, Januar 2022, S. 304–312. DOI:10.1111/mam.12281.

Hunting and diet

The cougar is a generalist, hypercarnivore. They prefer large mammals such as mule deer, white-tailed deer, elk, moose, mountain goat and bighorn sheep. They will opportunistically take smaller prey such as rodents, lagomorphs, smaller carnivores, birds and even domestic animals including pets. The mean weight of cougar vertebrate prey increases with its body weight and is lower in areas closer to the equator. 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.

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 wolves, with which the cougar competes for resources.
*
* A study on winter kills from November to 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-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. 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. Other listed prey species of the cougar include mice, porcupines, beavers, raccoons, hares, guanacoes, peccaries, vicuna, rheas, and wild turkeys. Birds and small reptiles are sometimes preyed upon in the south, but this is rarely recorded in North America. Magellanic penguins (''Spheniscus magellanicus'') constitute the majority of prey items in cougar diet in Patagonia's Bosques Petrificados de Jaramillo National Park and Monte León National Park.

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. 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. 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.

Interactions with other predators

Aside from humans, no species preys upon mature cougars in the wild, although conflicts with other predators or scavengers occur. Of the large predators in Yellowstone National Park – the grizzly bear, the black bear, the gray wolf, and the cougar – the massive grizzly bear appears dominant, often (but not always) able to drive a gray wolf pack, an American black bear, and a cougar off their kills. One study found that grizzlies and American black bears visited 24% of cougar kills in Yellowstone and Glacier National Parks, usurping 10% of carcasses. Bears gained up to 113% and cougars lost up to 26% of their respective daily energy requirements from these encounters. In Colorado and California, American black bears were found to visit 48% and 77% of kills, respectively. In general, cougars are subordinate to American black bears when it comes to kills and when bears are most active, the cats take prey more frequently and spend less time feeding on each kill. Unlike several subordinate predators from other ecosystems, cougars do not appear to take advantage of spatial or temporal refuges to avoid their competitors.

The gray wolf and the cougar compete more directly for prey, mostly in winter. Packs of wolves can steal cougars' kills, and there are some documented cases of cougars being killed by them. One report describes a large pack of seven to 11 wolves killing a female cougar and her kittens, while in nearby Sun Valley, Idaho

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