Old page wikitext, before the edit (old_wikitext ) | '{{redirect|Catamount|the ski area|Catamount Ski Area}}
{{about|the large cat species}}
{{Taxobox
| name = Cougar<ref name=MSW3/>
| fossil_range = [[Middle Pleistocene]] to Recent
| status = LC
| trend = down
| status_system = iucn3.1
| status_ref =<ref name="iucn">{{IUCN2008|assessors=Caso, A., Lopez-Gonzalez, C., Payan, E., Eizirik, E., de Oliveira, T., Leite-Pitman, R., Kelly, M., Valderrama, C. & Lucherini, M.|year=2008|id=18868|title=Puma concolor|downloaded=22 March 2009}} Database entry includes justification for why this species is least concern</ref>
| image = Mountain lion.jpg
| image_width=250px
| regnum = [[Animal]]ia
| phylum = [[Chordate|Chordata]]
| classis = [[Mammal]]ia
| ordo = [[Carnivora]]
| familia = [[Felidae]]
| genus = ''[[Puma (genus)|Puma]]''
| species = '''''P. concolor'''''
| binomial = ''Puma concolor''
| binomial_authority = ([[Carolus Linnaeus|Linnaeus]], 1771)
| range_map = Wiki-Puma concolor.png
| range_map_caption = Cougar range
}}
The '''cougar''' (''Puma concolor''), also known as '''puma''', '''mountain lion''', '''catamount''', or '''panther''', depending on the region, is a [[mammal]] of the [[Felidae]] family, native to the [[Americas]]. This large, solitary cat has the greatest [[range (biology)|range]] of any wild terrestrial mammal in the [[Western Hemisphere]],<ref name="diet"/> extending from [[Yukon]] in [[Canada]] to the southern [[Andes]] of [[South America]]. An adaptable, [[Generalist and specialist species|generalist]] species, the cougar is found in every major [[the Americas|American]] [[habitat (ecology)|habitat]] type. It is the second heaviest cat in the American continents after the [[jaguar]], and the fourth heaviest in the world, along with the [[leopard]], 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 [[ungulate]]s such as [[deer]], [[elk]], and [[bighorn sheep]], as well as domestic [[cattle]], [[horse]]s, and [[Domestic sheep|sheep]], particularly in the northern part of its range, but it also hunts species as small as [[insect]]s and [[rodents]]. Moreover, it prefers habitats with dense underbrush and rocky areas for stalking, but it can live in open areas. The cougar is [[Territory (animal)|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 [[Apex predator|dominant species]] in its range, as when it competes for prey with other predators such as the [[jaguar]], [[gray wolf]], [[American Black Bear]], and the [[grizzly bear]]. It is a reclusive cat and usually avoids people. [[List of fatal cougar attacks in North America by decade|Attacks on humans]] remain rare, despite a recent increase in frequency.<ref name="Med"/>
Due to [[persecution]] following the [[European colonization of the Americas]], and continuing human development of cougar habitat, populations have dropped in most parts of its historical range. In particular, the cougar was [[Local extinction|extirpated]] in eastern [[North America]], except an isolated [[Florida panther|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|indigenous Americans]] and in contemporary culture. The Cougar has recently made a comeback in the state of [[Wyoming]], where it presently has the largest population in North America.
==Naming and etymology==
The cougar has numerous names in English, of which ''puma'' and ''mountain lion'' are popular. Other names include ''catamount'', ''panther'', ''mountain screamer'', and painter. Lexicographers regard ''painter'' as a primarily upper-Southern U.S. regional variant on "panther",<ref>{{cite web|url=http://www.bartleby.com/61/61/P0016100.html |title=transcription of ''American Heritage Dictionary'' |publisher=Bartleby.com |date= |accessdate=2009-03-08}}</ref> but a [[folk etymology]], fancying a resemblance between the typically dark tip of its tail and a paintbrush dipped in dark paint, has some currency.
The cougar holds the [[Guinness World Records|Guinness]] record for the animal with the highest number of names, presumably due to its wide distribution across North and South America. It has over 40 names in English alone.<ref>{{cite book | title = The Guinness Book of World Records | year = 2004 | page = 49}}</ref>
"Cougar" is borrowed from the [[Portuguese language|Portuguese]] ''çuçuarana'', via [[French language|French]]; the term was originally derived from the [[Tupi language]]. A current form in Brazil is ''suçuarana''. "Puma" comes, via [[Spanish language|Spanish]], from the [[Quechua language]].<ref>[http://www.etymonline.com/index.php?term=cougar ''Cougar''], [http://www.etymonline.com/index.php?term=puma ''Puma''] and
{{cite web | url = http://www.etymonline.com/index.php?term=jaguar | title = ''Jaguar'' at the Online Etymology Dictionary | year = 2001 | publisher = Douglas Harper | accessdate = 2006-08-06}}</ref>
==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 cat]]s in the subfamily [[Pantherinae]].<ref name=MSW3/> The [[family (biology)|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,<ref>{{cite news |first=Nicholas |last=Wade |authorlink= |coauthors= |title=DNA Offers New Insight Concerning Cat Evolution |url=http://www.nytimes.com/2006/01/06/science/06cats.html?ex=1294203600&en=4b75c4da1cdc2167&ei=5090 |work=[[New York Times]] |publisher= |date= 2006-01-06 |accessdate=2007-06-03}}</ref> as cats are poorly represented in the [[fossil record]],<ref name="Johnson2006"/> and there are significant [[confidence intervals]] with suggested dates.
[[Image:Cougar pounce.jpg|thumb|Although large, the cougar is closely related to small felines.]]
In the latest [[genomic]] study of Felidae, the common ancestor of today's ''[[Leopardus]]'', ''[[Lynx]]'', ''[[Puma (genus)|Puma]]'', ''[[Prionailurus]]'', and ''[[Felis]]'' lineages migrated across the [[Beringia|Bering land bridge]] into the [[Americas]] approximately 8 to 8.5 million years ([[annum|Ma]]) ago. The lineages subsequently diverged in that order.<ref name="Johnson2006">{{cite journal |author=Johnson, W.E., Eizirik, E., Pecon-Slattery, J., Murphy, W.J., Antunes, A., Teeling, E. & O'Brien, S.J. |date=2006-01-06 |number=5757 |url=http://www.sciencemag.org/cgi/content/abstract/311/5757/73 |title=The Late Miocene radiation of modern Felidae: A genetic assessment |journal=[[Science (journal)|Science]] |volume=311 |issue=5757 |pages=73–77 |doi=10.1126/science.1122277 |accessdate=2007-06-04 |pmid=16400146}}</ref> 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,<ref name="Johnson2006"/><ref name="Culver">{{cite journal |author=Culver, M. |coauthors=Johnson, W.E., Pecon-Slattery, J., O'Brein, S.J. |year=2000 |title=Genomic Ancestry of the American Puma |journal=Journal of Heredity |volume=91 |issue=3 |pages=186–97 |url=http://www.coryi.org/Florida_panther/Miscellaneous_Panther_Material/Genomic%20ancestry%20of%20the%20American%20puma.pdf |format = PDF | doi = 10.1093/jhered/91.3.186 |pmid=10833043}}</ref> 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,<ref name="Johnson2006"/><ref name="Culver"/> while other research suggests the cheetah diverged in the [[Old World]] itself.<ref>{{cite journal |first=Ross |last=Barnett |coauthors=Ian Barnes, Matthew J. Phillips, Larry D. Martin, C. Richard Harington, Jennifer A. Leonard, and Alan Cooper |date=2005-08-09 |url=http://linkinghub.elsevier.com/retrieve/pii/S0960982205008365 |title=Evolution of the extinct Sabretooths and the American cheetah-like cat |journal=Current Biology |volume=15 |issue=15 |pages=R589–R590 |doi=10.1016/j.cub.2005.07.052 |accessdate=2007-06-04}}</ref> The outline of [[Felinae|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 [[local extinction|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.<ref name="Culver"/>
===Subspecies===
Until the late 1990s, as many as 32 [[subspecies]] were recorded; however, a recent [[Genetics|genetic]] study of [[mitochondrial DNA]]<ref name="Culver"/> 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]]:<ref name=MSW3>{{MSW3 Wozencraft |pages=544–45}}</ref>
; Argentine puma {{nobold|(''Puma concolor cabrerae'')}} : includes the previous subspecies and synonyms ''hudsonii'' and ''puma'' (Marcelli, 1922);
; [[Costa Rican Cougar]] {{nobold|(''Puma concolor costaricensis'')}}
; Eastern South American cougar {{nobold|(''Puma concolor anthonyi'')}} : includes the previous subspecies and synonyms ''acrocodia'', ''borbensis'', ''capricornensis'', ''concolor'' (Pelzeln, 1883), ''greeni'' and ''nigra'';
; [[North American Cougar]] {{nobold|(''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 {{nobold|(''Puma concolor concolor'')}} : includes the previous subspecies and synonyms ''bangsi'', ''incarum'', ''osgoodi'', ''soasoaranna'', ''soderstromii'', ''sucuacuara'' and ''wavula'';
; Southern South American puma {{nobold|(''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.<ref name=improving>{{cite journal |first=Michael J. |last=Conroy |coauthors=Paul Beier; Howard Quigley; Michael R. Vaughan |year=2006 |month=January |url=http://www.wildlifejournals.org/perlserv/?request=get-pdf&file=i0022-541X-70-1-1.pdf |title=Improving The Use Of Science In Conservation: Lessons From The Florida Panther |journal=Journal of Wildlife Management |volume=70 |issue=1 |pages=1–7 |doi=10.2193/0022-541X(2006)70[1:ITUOSI]2.0.CO;2 |format=PDF |accessdate=2007-06-11}}</ref> Culver ''et al.'' themselves noted low [[microsatellite]] variation in the Florida panther, possibly due to [[inbreeding]];<ref name=Culver/> 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."<ref name=FloridaRecovery>{{cite web |url=http://www.fws.gov/verobeach/Florida%20panther%20files/Panther%20Recovery%20Plan%202006_01_31%20-%20no%20figures.pdf |title=Florida Panther Recovery Program (Draft) |accessdate=2007-06-11 |author=The Florida Panther Recovery Team |date=2006-01-31 |format=PDF |publisher=U.S. Fish and Wildlife Service}}</ref>
==Biology and behavior==
===Physical characteristics===
Cougars are slender and agile cats. Adults stand about 60 to 76 cm (2.0 to 2.5 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.<ref name="Texas">{{cite web |url=http://www.tpwd.state.tx.us/huntwild/wild/species/mlion/ |title=Mountain Lion (''Puma concolor'') |accessdate=2007-03-30 |publisher=Texas Parks and Wildlife}}</ref><ref name="NY">{{cite web |url=http://www.dec.state.ny.us/website/dfwmr/wildlife/endspec/eacofs.html |title=Eastern Cougar Fact Sheet |accessdate=2007-03-30 |publisher=New York State Department of Environmental Conservation}}</ref> Males have an average weight of about 53 to 72 kilograms (115 to 160 pounds). In rare cases, some may reach over 118 kg (260 lb). Female average weight is between 34 and 48 kg (75 and 105 lb).<ref name="CAP">{{cite paper |author=Nowell, K. and Jackson, P |title=Wild Cats. Status Survey and Conservation Action Plan. |publisher=IUCN/SSC Cat Specialist Group. IUCN, Gland, Switzerland |year=2006 |url=http://carnivoractionplans1.free.fr/wildcats.pdf |format=PDF |accessdate=2007-07-27 }}</ref> Cougar size is smallest close to the [[equator]], and larger towards the [[Polar region|poles]].<ref name="diet">{{cite journal |author=J. Agustin Iriarte, William L. Franklin, Warren E. Johnson, and Kent H. Redford |year=1990 |title=Biogeographic variation of food habits and body size of the America puma |journal=[[Oecologia]] |volume=85 |issue=2 |page=185 |url=http://www.springerlink.com/content/nvk62r701822qq17/ |accessdate=2007-04-04 |doi=10.1007/BF00319400}}</ref>
[[Image:Cougar sitting.jpg|thumb|left|Although cougars resemble the [[domestic cat]], they are about the same size as an adult human.]]
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.<ref name="WhosWho">{{cite web |url=http://www.hww.ca/hww2.asp?pid=1&id=87&cid=8 |title=Cougar |work=Hinterland Who's Who |accessdate=2007-05-22 |publisher=[[Canadian Wildlife Service]] and [[Canadian Wildlife Federation]]}}</ref>
Cougars can be almost 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 as heavy as 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''.<ref>{{cite web |last=Weissengruber |first=GE |coauthors=G Forstenpointner, G Peters, A Kübber-Heiss, and WT Fitch |title=Hyoid apparatus and pharynx in the lion (Panthera leo), jaguar (Panthera onca), tiger (Panthera tigris), cheetah (Acinonyx jubatus) and domestic cat (Felis silvestris f. catus) |work=Journal of Anatomy |publisher=Anatomical Society of Great Britain and Ireland |pages=195–209 |volume=201 |issue=201 |year=2002 |month=September |url=http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=1570911 |doi=10.1046/j.1469-7580.2002.00088.x |accessdate=2007-05-20}}</ref> 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.<ref>{{cite web |url=http://www.easterncougar.org/pages/abouteasterncougars.htm |title=About Eastern Cougars |accessdate=2007-06-03 |publisher=Eastern Cougar Foundation}}</ref>
[[Image:Puma concolor paw.jpg|thumb|Rear 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;<ref name="CAP"/> juveniles are pale, and dark spots remain on their flanks.<ref name="NY"/> Despite anecdotes to the contrary, all-black coloring ([[melanism]]) has never been documented in cougars.<ref>{{cite web |url=http://www.news-tribune.net/features/cnhinsoutdoors_story_032111454.html |title=Black cougar more talk than fact |date=2006-02-01|accessdate=2007-05-20 |publisher=Tahlequah Daily Press|quote=[[Game Warden]]: Never in the history of the United States has there ever been, in captivity or in the wild, a documented black mountain lion}}</ref> The term "[[black panther]]" is used colloquially to refer to melanistic individuals of other species, particularly jaguars and leopards.<ref>{{cite web |url=http://www.messybeast.com/genetics/mutant-pumas.html |title=Mutant Pumas}}</ref>
Cougars have large paws and proportionally the largest hind legs in the cat family.<ref name="CAP"/> 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.<ref>{{cite web |url=http://www.sandiegozoo.org/animalbytes/t-puma.html |title=Mountain Lion (Puma, Cougar) |accessdate=2007-04-02 |work=San Diego Zoo.org |publisher=[[San Diego Zoo|Zoological Society of San Diego]]}}</ref> Horizontal jumping capability from standing position is suggested anywhere from 6 to 12 m (20 to 40 ft). The cougar can run as fast as 55–72 km/h (35-45 mph),<ref name="CanGeo">{{cite web |url=http://www.canadiangeographic.ca/Magazine/mj04/indepth/justthefacts.asp |title=Cougars in Canada (Just the Facts) |accessdate=2007-04-02 |publisher=[[Canadian Geographic Magazine]]}}</ref> but is best adapted for short, powerful sprints rather than long chases. It is adept at climbing, which allows it to evade [[Canidae|canine]] competitors. Although it is not strongly associated with water, it can swim.<ref name="Sierra">{{cite web |url=http://arizona.sierraclub.org/conservation/mt-lion/index.asp |title= Mountain Lion, Puma concolor |accessdate=2007-05-20 |publisher=[[Sierra Club]]}}</ref>
===Hunting and diet===
A successful [[Generalist and specialist species|generalist]] [[predator]], the cougar will eat any animal it can catch, from [[insect]]s to large [[ungulate]]s. Like all cats, it is an [[obligate carnivore]], feeding only on meat. The Mean weight of vertebrate prey (MWVP) was positively correlated (r=0.875) with puma body weight and inversely correlated (r=-0.836) with food niche breadth in all [[the Americas|America]]. In general, MWVP was lower in areas closer to the Equator.<ref name="diet"/> 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, [[Mustang (horse)|wild horses of Arizona]], domestic horses, and domestic livestock such as cattle and sheep are also primary food bases in many areas.<ref name="SW">{{cite web |url=http://wf2dnvr2.webfeat.org/|title=Influence of Predation by Mountain Lions on Numbers and Survivorship of a Feral Horse Population|author=John W. Turner, Jr. and Michael L. Morrison|accessdate=2008-08-29 |year=2008 |publisher=[[The Southwestern Naturalist]] Vol. 46, No.2 pp. 183-190 }}</ref> A survey of North America research found 68% of prey items were ungulates, especially deer. Only the Florida Panther showed variation, often preferring feral [[Hog (swine)|hogs]] and [[armadillo]]s.<ref name="diet"/>
[[Image:CMM MountainLion.jpg|thumb|Cougars are ambush predators, feeding mostly on deer and other mammals.]]
Investigation in [[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 Wolf|gray wolves]], with whom the cougar competes for resources.<ref name="Yellowstone">{{cite web |title=Wildlife: Wolves |publisher=[[Yellowstone National Park]] |url=http://www.yellowstonenationalpark.com/wolves.htm |accessdate=2007-04-08}}<br />* {{cite web |author = Holly Akenson, James Akenson, Howard Quigley |title=Winter Predation and Interactions of Wolves and Cougars on Panther Creek in Central Idaho |url=http://www.yellowstonenationalpark.com/wolves.htm}}<br />* {{cite web |author=John K. Oakleaf, Curt Mack, Dennis L. Murray |title=Winter Predation and Interactions of Cougars and Wolves in the Central Idaho Wilderness |url=http://www.yellowstonenationalpark.com/wolves.htm}}</ref> 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.<ref>{{cite journal |last=Ross |first=R. |coauthors=Jalkotzy, MG., Festa-Bianchet, M. |month=May | year=1993 |title=Cougar predation on bighorn sheep in southwestern Alberta during winter |journal=Canadian Journal of Zoology |volume=75 |issue=5 |pages=771–75 |url=http://md1.csa.com/partners/viewrecord.php?requester=gs&collection=ENV&recid=4321651 |accessdate=2007-04-08 |doi=10.1139/z97-098}}</ref>
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.<ref name="diet"/> Other listed prey species of the cougar include [[mice]], [[porcupine]], and [[hare]]s. Birds and small reptiles are sometimes preyed upon in the south, but this is rarely recorded in North America.<ref name="diet"/>
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.<ref name="WhosWho"/> 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.<ref name="CAP"/> The cat 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.<ref>{{cite journal |last=Bauer |first=Jim W. |coauthors=Kenneth A. Logan, Linda L. Sweanor, Walter M. Boyce |month=December | year=2005 |title=Scavenging behavior in Puma |journal=The Southwestern Naturalist |volume=50 |issue=4 |pages=466–471 |doi= 10.1894/0038-4909(2005)050[0466:SBIP]2.0.CO;2 |accessdate=2007-05-09}}</ref>
===Reproduction and lifecycle===
Females reach sexual maturity between one-and-a-half and three years of age. They typically average one [[litter (animal)|litter]] every two to three years throughout their reproductive life;<ref name="Utah">{{cite web |url=http://www.wildlife.utah.gov/pdf/cmgtplan.pdf |title=Utah Cougar Management Plan (Draft) |accessdate=2007-05-02 |author=Cougar Discussion Group |date=1999-01-27 |year= |month= |format=PDF |work= |publisher=Utah Division of Wildlife Resources}}</ref> the period can be as short as one year.<ref name="CAP"/> Females are in [[estrus]] for approximately 8 days of a 23-day cycle; the [[gestation period]] is approximately 91 days.<ref name="CAP"/> Females are sometimes reported as [[monogamous]],<ref name="CanGeo"/> but this is uncertain and [[polygyny]] may be more common.<ref name="UWSP">{{cite web |author=Matthew Hamilton |coauthors= Peter Hundt, Ryan Piorkowski |url=http://www.uwsp.edu/wildlife/carnivore/Mountain%20Lion%20Natural%20History_files/Mountain%20Lion%20Natural%20History_copy(1).htm |title=Mountain Lions |accessdate=2007-05-10 |publisher=[[University of Wisconsin, Stevens Point]]}}</ref> Copulation is brief but frequent.
[[Image:Mountain lion kittens.jpg|thumb|left|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.<ref name="Utah"/> Kitten survival rates are just over one per litter.<ref name="CAP"/>
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).<ref name="Utah"/> 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."<ref name="Dispersal">{{cite journal |last=Sweanor |first=Linda |coauthors=Kenneth A. Logan, Maurice G. Hornocker |year=2000 |month=June |title=Cougar Dispersal Patterns, Metapopulation Dynamics, and Conservation |journal=Conservation Biology |volume=14 |issue=3 |pages=798–808 |doi=10.1046/j.1523-1739.2000.99079.x |url=http://www.blackwell-synergy.com/links/doi/10.1046/j.1523-1739.2000.99079.x |format=PDF |accessdate=2007-05-02}}</ref>
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]].<ref name="CAP"/> 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.<ref>{{cite journal |last=Biek |first=Roman |coauthors=Allen G. Rodrigo, David Holley, Alexei Drummond, Charles R. Anderson Jr., Howard A. Ross, and Mary Poss |year=2003 |month=September |title=Epidemiology, Genetic Diversity, and Evolution of Endemic Feline Immunodeficiency Virus in a Population of Wild Cougars |journal=Journal of Virology |volume=77 |issue=17 |pages=9578–89 |doi=10.1128/JVI.77.17.9578-9589.2003 |url=http://jvi.asm.org/cgi/content/abstract/77/17/9578 |accessdate=2007-05-22 |pmid=12915571}}</ref>
===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 kilometer]]s (58 to 386 [[square mile|sq mi]]) with female ranges half the size.<ref name="CanGeo"/> Other research suggests a much smaller lower limit of 25 km<sup>2</sup> (10 sq mi) but an even greater upper limit of 1300 km<sup>2</sup> (500 sq mi) for males.<ref name="Utah"/> 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 km<sup>2</sup> (300 sq mi).<ref name="Dordt">{{cite web |url=http://homepages.dordt.edu/~mahaffy/mtlion/mtlionshort_behaviour.html |title=Behavior of cougar in Iowa and the Midwest |accessdate=2007-05-11 |last=Mahaffy |first=James |year=2004 |month=December |publisher=[[Dordt College]]}}</ref> 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.<ref name="Sierra"/>
Home range sizes and overall cougar abundance depend on terrain, vegetation, and prey abundance.<ref name="Utah"/> One female adjacent to the [[San Andres Mountains]], for instance, was found with a large range of 215 km<sup>2</sup> (83 sq mi), necessitated by poor prey abundance.<ref name="Dispersal"/> Research has shown cougar abundances from 0.5 animals to as much as 7 (in one study in South America) per 100 km<sup>2</sup> (38 sq mi).<ref name="CAP"/>
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.<ref name="Dordt"/> When males encounter each other, they hiss and spit, and may engage in violent conflict if neither backs down.<ref name="UWSP"/> Hunting or relocation of the cougar may increase aggressive encounters by disrupting territories and bringing young, transient animals into conflict with established individuals.<ref>{{cite web |url=http://www.sinapu.org/PDF/Front%20Range%20lion%20study.pdf |format=PDF |title=Mountain Lion (Puma concolor) study on Boulder Open Space |accessdate=2007-05-11 |date=2007-03-22 |work=Letter to the Parks and Open Space Advisory Committee, Boulder, Colorado |publisher=Sinapu}}</ref>
==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.<ref name="WhosWho"/> 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.<ref name="iucn"/> Its preferred habitats include precipitous canyons, escarpments, rim rocks, and dense brush.<ref name="Sierra"/>
More recently<!-- More recently than what? -->, reports from the Michigan Department Of Natural Resources have stated that cougars may be in Michigan's Upper Peninsula.{{Fact|date=May 2009}}
[[Image:MountainLion.jpg|thumb|left|Cougar, photographed in the [[Arizona-Sonora Desert Museum]], Tucson, Arizona.]]
The cougar was [[Local extinction|extirpated]] across much of its eastern North American range with the exception of [[Florida]] in the two centuries after [[European colonization of the Americas|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.<ref>{{cite web |url=http://www.naturalhistorymag.com/master.html?http://www.naturalhistorymag.com/0305/0305_selections.html |title=Bookshelf |last=Marschall |first=Laurence A. |date= |year=2005 |month=March |work=Natural Selections |publisher=[[Natural History Magazine]] |accessdate=2007-05-06}}</ref> [[DNA]] evidence has suggested its presence in eastern North America,<ref>{{cite web |url=http://lfpress.ca/newsstand/News/Local/2007/05/24/4205568.html |first=Joe |last=Belanger |title=DNA evidence of cougars found in southern Ontario |date=2007-05-24 |publisher=[[London Free Press]] |accessdate=2007-06-05}}</ref> while a consolidated map of cougar sightings shows numerous reports, from the mid-western [[Great Plains]] through to [[Eastern Canada]].<ref>{{cite web |url=http://www.cougarnet.org/bigpicture.html |title=The "Big" Picture |accessdate=2007-05-20 |author=[http://www.cougarnet.org/network.html Board of Directors] | year=2004| publisher=The Cougar Network}} ''The Cougar Network methodology is recognized by the [http://www.fws.gov/northeast/ECougar/ U.S. Fish and Wildlife Service].''</ref> The only unequivocally known eastern population is the [[Florida panther]], which is critically endangered. There have also been sightings in [[Elliotsville, Maine]] in the central part of the [[Maine|state]] and in [[New Hampshire]] there have been recent sightings as early as 1997<ref> {{cite web | url = http://www.rick-davidson.com/PDF/Mountain%20Lion%20Sightings%20Open%20Office%20092008%20photo%20final.pdf | format = PDF | title = NH Sightings Catamount | accessdate = 2009-03-20 | publisher=[[Beech River Books]]}}</ref>
On April 14, 2008 police shot and killed a cougar on the north side of Chicago, Illinois. DNA tests were consistent with cougars from the Black Hills. Less than one year later, on March 5, 2009, a cougar was photographed and unsuccessfully tranquilized by state wildlife biologists in a tree near Spooner, Wisconsin in the northwestern part of the state.<ref> {{cite web | url = http://www.stltoday.com/stltoday/lifestyle/stories.nsf/pets/story/5F51AA5F3CCF8DAD862575A00066E78F?OpenDocument | format = HTML | title = Sightings show cougars expanding into central US | accessdate = 2009-04-22 | publisher=St.Louis Today}}</ref>
South of the [[Rio Grande]], the [[World Conservation Union|International Union for the Conservation of Nature and Natural Resources]] (IUCN) lists the cat in every [[Central America|Central]] and [[South American]] country except [[Costa Rica]] and [[Panama]].<ref name="iucn"/> While specific state and provincial statistics are often available in North America, much less is known about the cat in its southern range.<ref> {{cite web | url = http://www.nwf.org/cats/pdfs/cougarfacts.pdf | format = PDF | title = Cougar facts | accessdate = 2007-05-20 | publisher=[[National Wildlife Federation]]}}</ref>
The cougar's total breeding population is estimated at less than 50,000 by the IUCN, with a declining trend.<ref name="iucn"/> 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.<ref>{{cite web |url=http://www.dfw.state.or.us/wildlife/cougar/ |title=Cougar Management Plan |accessdate=2007-05-20 |year=2006 |work=Wildlife Division: Wildlife Management Plans |publisher=Oregon Department of Fish and Wildlife}}</ref> [[California]] has actively sought to protect the cat and a similar number of cougars has been suggested, between 4,000 and 6,000.<ref name="California">{{cite web |url=http://www.dfg.ca.gov/news/issues/lion.html |title=Mountain Lions in California |accessdate=2007-05-20 |year=2004 |publisher=California Department of Fish and Game}}</ref>
===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]].
[[Image:Cougar track.jpg|thumb|Front paw print of a cougar. An adult paw print is approximately 10 cm (4 [[inches]]) long.<ref name="GovBC"/>]]
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 [[American Black Bear]]s visited 24% of cougar kills in Yellowstone and [[Glacier National Park]]s, usurping 10% of carcasses.<ref>{{cite paper |author=COSEWIC. Canadian Wildlife Service |title=Assessment and Update Status Report on the Grizzly Bear (Ursus arctos) |publisher=[[Environment Canada]] |year=2002 |url=http://dsp-psd.pwgsc.gc.ca/Collection/CW69-14-166-2002E.pdf |format=PDF |accessdate=2007-04-08 }}</ref>
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.<ref>{{cite web |url=http://www.washingtonpost.com/ac2/wp-dyn?pagename=article&node=&contentId=A2158-2003May17¬Found=true |title=In Yellowstone, it's Carnivore Competition |last=Gugliotta |first=Guy |date=2003-05-19 |publisher=Washington Post |accessdate=2007-04-09}}</ref> 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.<ref>{{cite web |url=http://www.greateryellowstonescience.org/topic/wolves/wolvesoverview.html |title=Overview: Gray Wolves |accessdate=2007-04-09 |publisher=Greater Yellowstone Learning Center}}</ref> 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."<ref>{{cite web |url=http://www.oregonwild.org/press-room/press-clips/turf-wars-in-idaho-s-wilderness |title=Turf wars in Idaho's wilderness |last=Cockle |first=Richard |date=2006-10-29 |publisher=[[The Oregonian]] |accessdate=2007-04-09}}</ref> Both species, meanwhile, are capable of killing mid-sized predators such as [[bobcat]]s and [[coyote]]s and tend to suppress their numbers.<ref name="Yellowstone"/>
In the southern portion of its range, the cougar and jaguar share overlapping territory.<ref name="HAMDIG">{{cite web |url=http://www.ecology.info/ecology-jaguar-puma.htm |first=Paul |last=Hamdig |title=Sympatric Jaguar and Puma |publisher=Ecology Online Sweden |accessdate=August 30 2006 |dateformat=mdy}}</ref> The jaguar tends to take larger prey and the cougar smaller where they overlap, reducing the cougar's size.<ref name="diet"/> Of the two felines, the cougar appears best able to exploit a broader prey niche and smaller prey.<ref name="foodhabits">{{cite journal |author=Rodrigo Nuanaez, Brian Miller, and Fred Lindzey |year=2000 |title=Food habits of jaguars and pumas in Jalisco, Mexico |journal=Journal of Zoology |volume=252 |issue=3 |page=373 |url=http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=58851 |accessdate=2006-08-08}}</ref>
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.<ref>{{cite journal |last=Robinson |first=Hugh S. |coauthors=Robert B. Wielgus, and John C. Gwilliam |year=2002 |title=Cougar predation and population growth of sympatric mule deer and white-tailed deer |journal=Canadian Journal of Zoology |volume=80 |issue=3 |pages=556–68 |doi=10.1139/z02-025 |url=http://rparticle.web-p.cisti.nrc.ca/rparticle/AbstractTemplateServlet?journal=cjz&volume=80&year=&issue=&msno=z02-025&calyLang=fra |accessdate=2007-05-20 |quote=}}</ref> The [[Vancouver Island Marmot]], an endangered species [[Endemism|endemic]] to one region of dense cougar population, has seen decreased numbers due to cougar and gray wolf predation.<ref>{{cite journal |last=Bryant |first=Andrew A. |coauthors=Page, Rick E. |year=2005 |title=Cougar predation and population growth of sympatric mule deer and white-tailed deer |journal=Canadian Journal of Zoology |month=May |volume=83 |issue=5 |pages=674–82 |url=http://www.ingentaconnect.com/content/nrc/cjz/2005/00000083/00000005/art00006?crawler=true |accessdate=2007-05-20 |doi=10.1139/z05-055}}</ref>
In the southern part of [[South America]] the Puma is a top level predator that has controlled the population of [[Guanaco]] and other species since prehistoric times.
==Hybrids==
[[Image:Pumapard-1904.jpg|thumb|upright|Pumapard, taken in 1904]]
{{Main|Pumapard}}
A pumapard is a [[hybrid (biology)|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 Zoological Garden|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.<ref name="Geo">{{cite web|url=http://www.geocities.com/pride_lands/Liger_Tigon.html|title=Geocites - Liger & Tigon Info|accessdate=9 June 2008|dateformat=dmy}}</ref>
==Conservation status==
The [[World Conservation Union]] (IUCN) currently lists the cougar as a "[[least concern]]" species. The cougar is regulated under [[CITES#Appendix I|Appendix I]] of the [[CITES|Convention on International Trade in Endangered Species of Wild Fauna and Flora]] (CITES),<ref>{{cite web |url=http://www.cites.org/eng/app/appendices.shtml |title=Appendices I, II and III |accessdate=2007-05-24 |publisher=[[Convention on International Trade in Endangered Species of Wild Fauna and Flora]]}}</ref> rendering illegal international trade in specimens or parts.
[[Image:Cougar snow.jpg|thumb|left|Cougar conservation depends on preservation of their habitat.]]
East of the [[Mississippi]], the only unequivocally known cougar population is the [[Florida panther]]. The [[United States Fish and Wildlife Service]] recognizes both an Eastern cougar and the Florida panther, affording protection under the [[Endangered Species Act]].<ref>{{cite web |url=http://www.fws.gov/endangered/i/a/saa48.html |title=Eastern Cougar |accessdate=2007-05-20 | work=Endangered and Threatened Species of the Southeastern United States (The Red Book)| year=1991 |publisher=[[U.S. Fish and Wildlife Service]]}}</ref><ref>{{cite web |url=http://www.fws.gov/endangered/i/a/saa05.html |title=Florida Panther |accessdate=2007-06-07 | work=Endangered and Threatened Species of the Southeastern United States (The Red Book)| year=1993 |publisher=[[United States Fish and Wildlife Service]]}}</ref> Certain taxonomic authorities have collapsed both designations into the [[North American Cougar]], with Eastern or Florida subspecies not recognized,<ref name="MSW3"/> while a subspecies designation remains recognized by some conservation scientists.<ref name=improving/> The most recent documented count for the Florida sub-population is 87 individuals, reported by recovery agencies in 2003.<ref>{{cite web |url=http://myfwc.com/panther/news/pdf/FWC2002-2003PantherGeneticRestorationAnnualReport.pdf |format=PDF |title=Florida Fish and Wildlife Conservation Commission. 2002–2003 Panther Genetfic Restoration Annual Report |accessdate=2007-06-05}}</ref>
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]].<ref name="CAP"/> 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 [[Hunting dog|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.<ref name="California"/> However statistics from the [[California Department of Fish and Game|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 six per year in the 1970s. The [[Bay Area Puma Project]] aims to obtain information on cougar populations in the San Francisco Bay area and the animals' interactions with habitat, prey, humans, and residential communities.<ref name=felidae1>{{cite web|url=http://www.felidaefund.org/research/bay_puma.html |title="Bay Area Puma Project" information page, Felidae Conservation Fund, accessed 18 February 2009 |publisher=Felidaefund.org |date= |accessdate=2009-03-08}}</ref>
Conservation threats to the species include persecution as a pest animal, degradation and fragmentation of their habitat, and depletion of their prey base. [[Wildlife corridor]] 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 km<sup>2</sup> (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.<ref>{{cite journal |last=Beier |first=Paul |year=1993 |month=March |title=Determining Minimum Habitat Areas and Habitat Corridors for Cougars |journal=Conservation Biology|volume=7|issue=1|pages=94–108 |url=http://links.jstor.org/sici?sici=0888-8892%28199303%297%3A1%3C94%3ADMHAAH%3E2.0.CO%3B2-I&size=LARGE&origin=JSTOR-enlargePage |accessdate=2007-05-20|doi=10.1046/j.1523-1739.1993.07010094.x}}</ref>
==Relationships with humans==
===In mythology and culture===
[[Image:MochePuma.jpg|thumb|[[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.<ref>Berrin, Katherine & Larco Museum. ''The Spirit of Ancient Peru:Treasures from the [[Larco Museum|Museo Arqueológico Rafael Larco Herrera]].'' New York: [[Thames and Hudson]], 1997.</ref> The sky and thunder god of the Inca, [[Viracocha]], has been associated with the animal.<ref>{{cite web |first=Kulmar |last=Tarmo |url=http://folklore.ee/folklore/vol12/inca.htm |coauthors=Kait Realo (translator) |title=On the role of Creation and Origin Myths in the Development of Inca State and Religion |accessdate=2007-05-22 |work=Electronic Journal of Folklore|publisher=Estonian Folklore Institute}}</ref>
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]] {{Fact|date=October 2008}} and the [[Cheyenne]], amongst others. To the [[Apache]] and [[Walapai]] of [[Arizona]], the wail of the Cougar was a harbinger of death.<ref>{{cite web |url=http://www.aphis.usda.gov/wildlife_damage/nwrc/publications/living/cougar.pdf |title=Living with Wildlife: Cougars |accessdate=2009-04-11 |format=PDF |publisher=[[USDA Wildlife Services]]}}</ref>
The cougar continues to be a symbol of strength and stealth. From [[Eurocopter Cougar|combat helicopters]], motor vehicles (see [[Mercury Cougar|Ford/Mercury Cougar]] and [[Ford Puma]]) to [[Puma AG|athletic shoes]], both "Cougar" and "Puma" are widely used as [[Brand|brand names]]. Various sports teams have also adopted the names, including the [[Argentina national rugby union team]], the [[National Autonomous University of Mexico]] soccer club as well as US universities, The [[Florida Panthers]], The [[College of Charleston]], [[Brigham Young University]], The [[University of Houston]], [[Washington State University]], and The [[University of Vermont]]. Many places, such as [[Cougar Mountain]], are also named after their association with cougars.<!--we don't need a massive list here, please-->
===Livestock predation===
Cougars can cause severe economic hardship on those whose livelihoods depend on livestock. During the early years of ranching, cougars were considered on par with wolves in destructiveness. According to figures in Texas in 1990 for example, 86 calves, 253 Mohair goats, 302 Mohair kids, 445 sheep and 562 lambs were confirmed to have been killed by cougars that year. In Nevada in 1992, cougars were confirmed to have killed 9 calves, 1 horse, 4 colts, 5 goats, 318 sheep and 400 lambs. In both cases, sheep were the most frequently attacked. Some instances of [[surplus killing]] have resulted in the deaths of 20 sheep in one attack.<ref name="Livestock">{{cite web | url=http://www.aws.vcn.com/mountain_lion_fact_sheet.html | title = Mountain Lion Fact Sheet | publisher = Abundant Wildlife Society of North America | accessdate = 2008-07-10}}</ref> Cougars frequently kill calves, sheep and goats by biting the top of the neck or head, differing greatly from the throat bite used by coyotes and indiscriminate mutilation by feral dogs. The size of the tooth puncture marks also helps distinguish kills made by cougars from those made by smaller predators.<ref name="Predation">{{cite web | url=http://texnat.tamu.edu/ranchref/predator//cougars/t-cougar.htm | title = Cougar Predation - Description | publisher = Procedures for Evaluating Predation on Livestock and Wildlife | accessdate = 2008-08-03}}</ref>
===Attacks on humans===
{{seealso|List of fatal cougar attacks in North America by decade}}
[[Image:MountainLionAttackProtocol.jpg|thumb|right|Mountain Lion warning sign]]
Due to the [[overpopulation|expanding human population]], 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.<ref name="Med">{{cite journal |last=McKee |first=Denise |year=2003 |title=Cougar Attacks on Humans: A Case Report |journal=Wilderness and Environmental Medicine |volume=14 |issue=3 |pages=169–73 |publisher=Wilderness Medical Society |url=http://www.wemjournal.org/wmsonline/?request=get-document&issn=1080-6032&volume=014&issue=03&page=0169 |accessdate=2007-05-20}}</ref> Attacks on people, livestock, and pets may occur when the cat [[Habituation|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.<ref name="GovBC"/>
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).<ref name="Beier">{{cite web |first=Paul |last=Beier |url=http://users.frii.com/mytymyk/lions/beier.htm |title=Cougar attacks on humans in United States and Canada |work=Wildlife Society Bulletin |year=1991 |accessdate=2007-05-20 |publisher=Northern Arizon University}}</ref> By 2004, the count had climbed to 88 attacks and 20 deaths.<ref>{{cite web |url=http://www.azgfd.gov/w_c/mtn_lion_attacks.shtml |title=Confirmed mountain lion attacks in the United States and Canada 1890 — Present |accessdate=2007-05-20 |publisher=Arizona Game and Fish}}</ref>
Within North America, the distribution of attacks is not uniform. The heavily populated state of [[California]] has seen a dozen attacks since 1986 (after just three from 1890 to 1985), including three fatalities.<ref name="California"/> Lightly populated [[New Mexico]] reported an attack in 2008, the first there since 1974.<ref>[http://www.wildlife.state.nm.us/publications/press_releases/documents/2008/062308pinosaltoslion.html Search continues for mountain lion that killed Pinos Altos man], New Mexico Department of Game and Fish, press release June 23, 2008]; [http://www.wildlife.state.nm.us/publications/press_releases/documents/2008/062508pinosaltoslion.html Wounded mountain lion captured, killed near Pinos Altos], New Mexico Department of Game and Fish, press release June 25, 2008]; [http://www.wildlife.state.nm.us/publications/press_releases/documents/2008/070108pinosaltoslion2.html Second mountain lion captured near Pinos Altos], New Mexico Department of Game and Fish, press release July 1, 2008]</ref>
As with many predators, a cougar may attack if cornered, if a fleeing human stimulates their instinct to chase, or if a person "[[wikt:play dead|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 persuading an attacking cougar to disengage.<ref name="Med"/><ref name="GovBC">{{cite web |url=http://www.env.gov.bc.ca/wld/documents/cougsf.htm |title=Safety Guide to Cougars |work=Environmental Stewardship Division |year=1991 |accessdate=2007-05-28 |publisher=[[Government of British Columbia]], Ministry of Environment}}</ref>
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.<ref name="Med"/> 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{{ndash}} and almost all fatalities{{ndash}} 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.<ref name="Beier"/>
==References==
{{reflist|2}}
==Other resources==
* {{cite book |last=Baron |first=David |title=The Beast in the Garden: A Modern Parable of Man and Nature |year=2004 |publisher=W. W. Norton and Company |location=New York |isbn=0393058077}}
* {{cite book |last=Kobalenko |first=Jerry |title=Forest Cats of North America |year=1997 |publisher=Island Press |location=Buffalo, New York |isbn=1-55209-174-4}}
* {{cite book |last=Logan |first=Ken |coauthors=Linda Sweanor |title=Desert Puma: Evolutionary Ecology and Conservation of an Enduring Carnivore |year=2001 |publisher=Island Press |isbn=1-55963-866-4}}
==External links==
{{portal|Cats|Cat03.jpg}}
{{portal|Mammals|Okapi2.jpg}}
{{wiktionary}}
{{commons|Puma concolor}}
{{wikispecies|Puma concolor}}
{{Spoken Wikipedia|Cougar1.ogg|2008-11-06}}
* [http://www.theanimalfiles.com/mammals/carnivores/mountain_lion.html Video, Pictures and Information on Mountain Lions]
* [http://www.cougarfund.org/ A Definitive Resource About Cougar] Comprehensive non-profit site with extensive information about cougars, from how to live safely in cougar country, to science abstracts, hunting regulations, state-by-state cougar management/policy info, to (rare) photos and videos of wild cougars.
* [http://dir.salon.com/story/people/feature/2002/02/12/cougar/index.html?pn=2 Description of a Cougar attack]
* [http://depts.washington.edu/natmap/facts/cougar_712.html Cougar Facts and Photos] - NatureMapping Program
* [http://biology.plosjournals.org/perlserv/?request=get-document&doi=10.1371/journal.pbio.0060040 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
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{{North American Game}}
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New page wikitext, after the edit (new_wikitext ) | '{{redirect|Catamount|the ski area|Catamount Ski Area}}
{{about|the large cat species}}
{{Taxobox
| name = Cougar<ref name=MSW3/>
| fossil_range = [[Middle Pleistocene]] to Recent
| status = LC
| trend = down
| status_system = iucn3.1
| status_ref =<ref name="iucn">{{IUCN2008|assessors=Caso, A., Lopez-Gonzalez, C., Payan, E., Eizirik, E., de Oliveira, T., Leite-Pitman, R., Kelly, M., Valderrama, C. & Lucherini, M.|year=2008|id=18868|title=Puma concolor|downloaded=22 March 2009}} Database entry includes justification for why this species is least concern</ref>
| image = Mountain lion.jpg
| image_width=250px
| regnum = [[Animal]]ia
| phylum = [[Chordate|Chordata]]
| classis = [[Mammal]]ia
| ordo = [[Carnivora]]
| familia = [[Felidae]]
| genus = ''[[Puma (genus)|Puma]]''
| species = '''''P. concolor'''''
| binomial = ''Puma concolor''
| binomial_authority = ([[Carolus Linnaeus|Linnaeus]], 1771)
| range_map = Wiki-Puma concolor.png
| range_map_caption = Cougar range
}}
The '''cougar''' (''Puma concolor''), also known as '''puma''', '''mountain lion''', '''catamount''', or '''panther''', depending on the region, is a [[mammal]] of the [[Felidae]] family, native to the [[Americas]]. This large, solitary cat has the greatest [[range (biology)|range]] of any wild terrestrial mammal in the [[Western Hemisphere]],<ref name="diet"/> extending from [[Yukon]] in [[Canada]] to the southern [[Andes]] of [[South America]]. An adaptable, [[Generalist and specialist species|generalist]] species, the cougar is found in every major [[the Americas|American]] [[habitat (ecology)|habitat]] type. It is the second heaviest cat in the American continents after the [[jaguar]], and the fourth heaviest in the world, along with the [[leopard]], 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 [[ungulate]]s such as [[deer]], [[elk]], and [[bighorn sheep]], as well as domestic [[cattle]], [[horse]]s, and [[Domestic sheep|sheep]], particularly in the northern part of its range, but it also hunts species as small as [[insect]]s and [[rodents]]. Moreover, it prefers habitats with dense underbrush and rocky areas for stalking, but it can live in open areas. The cougar is [[Territory (animal)|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 [[Apex predator|dominant species]] in its range, as when it competes for prey with other predators such as the [[jaguar]], [[gray wolf]], [[American Black Bear]], and the [[grizzly bear]]. It is a reclusive cat and usually avoids people. [[List of fatal cougar attacks in North America by decade|Attacks on humans]] remain rare, despite a recent increase in frequency.<ref name="Med"/>
Due to [[persecution]] following the [[European colonization of the Americas]], and continuing human development of cougar habitat, populations have dropped in most parts of its historical range. In particular, the cougar was [[Local extinction|extirpated]] in eastern [[North America]], except an isolated [[Florida panther|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|indigenous Americans]] and in contemporary culture. The Cougar has recently made a comeback in the state of [[Wyoming]], where it presently has the largest population in North America.
==Naming and etymology==
The cougar has numerous names in English, of which ''puma'' and ''mountain lion'' are popular. Other names include ''catamount'', ''panther'', ''mountain screamer'', and painter. Lexicographers regard ''painter'' as a primarily upper-Southern U.S. regional variant on "panther",<ref>{{cite web|url=http://www.bartleby.com/61/61/P0016100.html |title=transcription of ''American Heritage Dictionary'' |publisher=Bartleby.com |date= |accessdate=2009-03-08}}</ref> but a [[folk etymology]], fancying a resemblance between the typically dark tip of its tail and a paintbrush dipped in dark paint, has some currency.
The cougar holds the [[Guinness World Records|Guinness]] record for the animal with the highest number of names, presumably due to its wide distribution across North and South America. It has over 40 names in English alone.<ref>{{cite book | title = The Guinness Book of World Records | year = 2004 | page = 49}}</ref>
"Cougar" is borrowed from the [[Portuguese language|Portuguese]] ''çuçuarana'', via [[French language|French]]; the term was originally derived from the [[Tupi language]]. A current form in Brazil is ''suçuarana''. "Puma" comes, via [[Spanish language|Spanish]], from the [[Quechua language]].<ref>[http://www.etymonline.com/index.php?term=cougar ''Cougar''], [http://www.etymonline.com/index.php?term=puma ''Puma''] and
{{cite web | url = http://www.etymonline.com/index.php?term=jaguar | title = ''Jaguar'' at the Online Etymology Dictionary | year = 2001 | publisher = Douglas Harper | accessdate = 2006-08-06}}</ref>
==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 cat]]s in the subfamily [[Pantherinae]].<ref name=MSW3/> The [[family (biology)|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,<ref>{{cite news |first=Nicholas |last=Wade |authorlink= |coauthors= |title=DNA Offers New Insight Concerning Cat Evolution |url=http://www.nytimes.com/2006/01/06/science/06cats.html?ex=1294203600&en=4b75c4da1cdc2167&ei=5090 |work=[[New York Times]] |publisher= |date= 2006-01-06 |accessdate=2007-06-03}}</ref> as cats are poorly represented in the [[fossil record]],<ref name="Johnson2006"/> and there are significant [[confidence intervals]] with suggested dates.
[[Image:Cougar pounce.jpg|thumb|Although large, the cougar is closely related to small felines.]]
In the latest [[genomic]] study of Felidae, the common ancestor of today's ''[[Leopardus]]'', ''[[Lynx]]'', ''[[Puma (genus)|Puma]]'', ''[[Prionailurus]]'', and ''[[Felis]]'' lineages migrated across the [[Beringia|Bering land bridge]] into the [[Americas]] approximately 8 to 8.5 million years ([[annum|Ma]]) ago. The lineages subsequently diverged in that order.<ref name="Johnson2006">{{cite journal |author=Johnson, W.E., Eizirik, E., Pecon-Slattery, J., Murphy, W.J., Antunes, A., Teeling, E. & O'Brien, S.J. |date=2006-01-06 |number=5757 |url=http://www.sciencemag.org/cgi/content/abstract/311/5757/73 |title=The Late Miocene radiation of modern Felidae: A genetic assessment |journal=[[Science (journal)|Science]] |volume=311 |issue=5757 |pages=73–77 |doi=10.1126/science.1122277 |accessdate=2007-06-04 |pmid=16400146}}</ref> 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,<ref name="Johnson2006"/><ref name="Culver">{{cite journal |author=Culver, M. |coauthors=Johnson, W.E., Pecon-Slattery, J., O'Brein, S.J. |year=2000 |title=Genomic Ancestry of the American Puma |journal=Journal of Heredity |volume=91 |issue=3 |pages=186–97 |url=http://www.coryi.org/Florida_panther/Miscellaneous_Panther_Material/Genomic%20ancestry%20of%20the%20American%20puma.pdf |format = PDF | doi = 10.1093/jhered/91.3.186 |pmid=10833043}}</ref> 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,<ref name="Johnson2006"/><ref name="Culver"/> while other research suggests the cheetah diverged in the [[Old World]] itself.<ref>{{cite journal |first=Ross |last=Barnett |coauthors=Ian Barnes, Matthew J. Phillips, Larry D. Martin, C. Richard Harington, Jennifer A. Leonard, and Alan Cooper |date=2005-08-09 |url=http://linkinghub.elsevier.com/retrieve/pii/S0960982205008365 |title=Evolution of the extinct Sabretooths and the American cheetah-like cat |journal=Current Biology |volume=15 |issue=15 |pages=R589–R590 |doi=10.1016/j.cub.2005.07.052 |accessdate=2007-06-04}}</ref> The outline of [[Felinae|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 [[local extinction|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.<ref name="Culver"/>
===Subspecies===
Until the late 1990s, as many as 32 [[subspecies]] were recorded; however, a recent [[Genetics|genetic]] study of [[mitochondrial DNA]]<ref name="Culver"/> 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]]:<ref name=MSW3>{{MSW3 Wozencraft |pages=544–45}}</ref>
; Argentine puma {{nobold|(''Puma concolor cabrerae'')}} : includes the previous subspecies and synonyms ''hudsonii'' and ''puma'' (Marcelli, 1922);
; [[Costa Rican Cougar]] {{nobold|(''Puma concolor costaricensis'')}}
; Eastern South American cougar {{nobold|(''Puma concolor anthonyi'')}} : includes the previous subspecies and synonyms ''acrocodia'', ''borbensis'', ''capricornensis'', ''concolor'' (Pelzeln, 1883), ''greeni'' and ''nigra'';
; [[North American Cougar]] {{nobold|(''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 {{nobold|(''Puma concolor concolor'')}} : includes the previous subspecies and synonyms ''bangsi'', ''incarum'', ''osgoodi'', ''soasoaranna'', ''soderstromii'', ''sucuacuara'' and ''wavula'';
; Southern South American puma {{nobold|(''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.<ref name=improving>{{cite journal |first=Michael J. |last=Conroy |coauthors=Paul Beier; Howard Quigley; Michael R. Vaughan |year=2006 |month=January |url=http://www.wildlifejournals.org/perlserv/?request=get-pdf&file=i0022-541X-70-1-1.pdf |title=Improving The Use Of Science In Conservation: Lessons From The Florida Panther |journal=Journal of Wildlife Management |volume=70 |issue=1 |pages=1–7 |doi=10.2193/0022-541X(2006)70[1:ITUOSI]2.0.CO;2 |format=PDF |accessdate=2007-06-11}}</ref> Culver ''et al.'' themselves noted low [[microsatellite]] variation in the Florida panther, possibly due to [[inbreeding]];<ref name=Culver/> 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."<ref name=FloridaRecovery>{{cite web |url=http://www.fws.gov/verobeach/Florida%20panther%20files/Panther%20Recovery%20Plan%202006_01_31%20-%20no%20figures.pdf |title=Florida Panther Recovery Program (Draft) |accessdate=2007-06-11 |author=The Florida Panther Recovery Team |date=2006-01-31 |format=PDF |publisher=U.S. Fish and Wildlife Service}}</ref>
==Biology and behavior==
===Physical characteristics===
Cougars are slender and agile cats. Adults stand about 60 to 76 cm (2.0 to 2.5 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.<ref name="Texas">{{cite web |url=http://www.tpwd.state.tx.us/huntwild/wild/species/mlion/ |title=Mountain Lion (''Puma concolor'') |accessdate=2007-03-30 |publisher=Texas Parks and Wildlife}}</ref><ref name="NY">{{cite web |url=http://www.dec.state.ny.us/website/dfwmr/wildlife/endspec/eacofs.html |title=Eastern Cougar Fact Sheet |accessdate=2007-03-30 |publisher=New York State Department of Environmental Conservation}}</ref> Males have an average weight of about 53 to 72 kilograms (115 to 160 pounds). In rare cases, some may reach over 118 kg (2600 lb). Female average weight is between 34 and 48 kg (200000 and 30000000 lb).<ref name="CAP">{{cite paper |author=Nowell, K. and Jackson, P |title=Wild Cats. Status Survey and Conservation Action Plan. |publisher=IUCN/SSC Cat Specialist Group. IUCN, Gland, Switzerland |year=2006 |url=http://carnivoractionplans1.free.fr/wildcats.pdf |format=PDF |accessdate=2007-07-27 }}</ref> Cougar size is smallest close to the [[equator]], and larger towards the [[Polar region|poles]].<ref name="diet">{{cite journal |author=J. Agustin Iriarte, William L. Franklin, Warren E. Johnson, and Kent H. Redford |year=1990 |title=Biogeographic variation of food habits and body size of the America puma |journal=[[Oecologia]] |volume=85 |issue=2 |page=185 |url=http://www.springerlink.com/content/nvk62r701822qq17/ |accessdate=2007-04-04 |doi=10.1007/BF00319400}}</ref>
[[Image:Cougar sitting.jpg|thumb|left|Although cougars resemble the [[domestic cat]], they are about the same size as an adult human.]]
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.<ref name="WhosWho">{{cite web |url=http://www.hww.ca/hww2.asp?pid=1&id=87&cid=8 |title=Cougar |work=Hinterland Who's Who |accessdate=2007-05-22 |publisher=[[Canadian Wildlife Service]] and [[Canadian Wildlife Federation]]}}</ref>
Cougars can be almost 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 as heavy as 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''.<ref>{{cite web |last=Weissengruber |first=GE |coauthors=G Forstenpointner, G Peters, A Kübber-Heiss, and WT Fitch |title=Hyoid apparatus and pharynx in the lion (Panthera leo), jaguar (Panthera onca), tiger (Panthera tigris), cheetah (Acinonyx jubatus) and domestic cat (Felis silvestris f. catus) |work=Journal of Anatomy |publisher=Anatomical Society of Great Britain and Ireland |pages=195–209 |volume=201 |issue=201 |year=2002 |month=September |url=http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=1570911 |doi=10.1046/j.1469-7580.2002.00088.x |accessdate=2007-05-20}}</ref> 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.<ref>{{cite web |url=http://www.easterncougar.org/pages/abouteasterncougars.htm |title=About Eastern Cougars |accessdate=2007-06-03 |publisher=Eastern Cougar Foundation}}</ref>
[[Image:Puma concolor paw.jpg|thumb|Rear 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;<ref name="CAP"/> juveniles are pale, and dark spots remain on their flanks.<ref name="NY"/> Despite anecdotes to the contrary, all-black coloring ([[melanism]]) has never been documented in cougars.<ref>{{cite web |url=http://www.news-tribune.net/features/cnhinsoutdoors_story_032111454.html |title=Black cougar more talk than fact |date=2006-02-01|accessdate=2007-05-20 |publisher=Tahlequah Daily Press|quote=[[Game Warden]]: Never in the history of the United States has there ever been, in captivity or in the wild, a documented black mountain lion}}</ref> The term "[[black panther]]" is used colloquially to refer to melanistic individuals of other species, particularly jaguars and leopards.<ref>{{cite web |url=http://www.messybeast.com/genetics/mutant-pumas.html |title=Mutant Pumas}}</ref>
Cougars have large paws and proportionally the largest hind legs in the cat family.<ref name="CAP"/> 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.<ref>{{cite web |url=http://www.sandiegozoo.org/animalbytes/t-puma.html |title=Mountain Lion (Puma, Cougar) |accessdate=2007-04-02 |work=San Diego Zoo.org |publisher=[[San Diego Zoo|Zoological Society of San Diego]]}}</ref> Horizontal jumping capability from standing position is suggested anywhere from 6 to 12 m (20 to 40 ft). The cougar can run as fast as 55–72 km/h (35-45 mph),<ref name="CanGeo">{{cite web |url=http://www.canadiangeographic.ca/Magazine/mj04/indepth/justthefacts.asp |title=Cougars in Canada (Just the Facts) |accessdate=2007-04-02 |publisher=[[Canadian Geographic Magazine]]}}</ref> but is best adapted for short, powerful sprints rather than long chases. It is adept at climbing, which allows it to evade [[Canidae|canine]] competitors. Although it is not strongly associated with water, it can swim.<ref name="Sierra">{{cite web |url=http://arizona.sierraclub.org/conservation/mt-lion/index.asp |title= Mountain Lion, Puma concolor |accessdate=2007-05-20 |publisher=[[Sierra Club]]}}</ref>
===Hunting and diet===
A successful [[Generalist and specialist species|generalist]] [[predator]], the cougar will eat any animal it can catch, from [[insect]]s to large [[ungulate]]s. Like all cats, it is an [[obligate carnivore]], feeding only on meat. The Mean weight of vertebrate prey (MWVP) was positively correlated (r=0.875) with puma body weight and inversely correlated (r=-0.836) with food niche breadth in all [[the Americas|America]]. In general, MWVP was lower in areas closer to the Equator.<ref name="diet"/> 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, [[Mustang (horse)|wild horses of Arizona]], domestic horses, and domestic livestock such as cattle and sheep are also primary food bases in many areas.<ref name="SW">{{cite web |url=http://wf2dnvr2.webfeat.org/|title=Influence of Predation by Mountain Lions on Numbers and Survivorship of a Feral Horse Population|author=John W. Turner, Jr. and Michael L. Morrison|accessdate=2008-08-29 |year=2008 |publisher=[[The Southwestern Naturalist]] Vol. 46, No.2 pp. 183-190 }}</ref> A survey of North America research found 68% of prey items were ungulates, especially deer. Only the Florida Panther showed variation, often preferring feral [[Hog (swine)|hogs]] and [[armadillo]]s.<ref name="diet"/>
[[Image:CMM MountainLion.jpg|thumb|Cougars are ambush predators, feeding mostly on deer and other mammals.]]
Investigation in [[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 Wolf|gray wolves]], with whom the cougar competes for resources.<ref name="Yellowstone">{{cite web |title=Wildlife: Wolves |publisher=[[Yellowstone National Park]] |url=http://www.yellowstonenationalpark.com/wolves.htm |accessdate=2007-04-08}}<br />* {{cite web |author = Holly Akenson, James Akenson, Howard Quigley |title=Winter Predation and Interactions of Wolves and Cougars on Panther Creek in Central Idaho |url=http://www.yellowstonenationalpark.com/wolves.htm}}<br />* {{cite web |author=John K. Oakleaf, Curt Mack, Dennis L. Murray |title=Winter Predation and Interactions of Cougars and Wolves in the Central Idaho Wilderness |url=http://www.yellowstonenationalpark.com/wolves.htm}}</ref> 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.<ref>{{cite journal |last=Ross |first=R. |coauthors=Jalkotzy, MG., Festa-Bianchet, M. |month=May | year=1993 |title=Cougar predation on bighorn sheep in southwestern Alberta during winter |journal=Canadian Journal of Zoology |volume=75 |issue=5 |pages=771–75 |url=http://md1.csa.com/partners/viewrecord.php?requester=gs&collection=ENV&recid=4321651 |accessdate=2007-04-08 |doi=10.1139/z97-098}}</ref>
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.<ref name="diet"/> Other listed prey species of the cougar include [[mice]], [[porcupine]], and [[hare]]s. Birds and small reptiles are sometimes preyed upon in the south, but this is rarely recorded in North America.<ref name="diet"/>
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.<ref name="WhosWho"/> 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.<ref name="CAP"/> The cat 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.<ref>{{cite journal |last=Bauer |first=Jim W. |coauthors=Kenneth A. Logan, Linda L. Sweanor, Walter M. Boyce |month=December | year=2005 |title=Scavenging behavior in Puma |journal=The Southwestern Naturalist |volume=50 |issue=4 |pages=466–471 |doi= 10.1894/0038-4909(2005)050[0466:SBIP]2.0.CO;2 |accessdate=2007-05-09}}</ref>
===Reproduction and lifecycle===
Females reach sexual maturity between one-and-a-half and three years of age. They typically average one [[litter (animal)|litter]] every two to three years throughout their reproductive life;<ref name="Utah">{{cite web |url=http://www.wildlife.utah.gov/pdf/cmgtplan.pdf |title=Utah Cougar Management Plan (Draft) |accessdate=2007-05-02 |author=Cougar Discussion Group |date=1999-01-27 |year= |month= |format=PDF |work= |publisher=Utah Division of Wildlife Resources}}</ref> the period can be as short as one year.<ref name="CAP"/> Females are in [[estrus]] for approximately 8 days of a 23-day cycle; the [[gestation period]] is approximately 91 days.<ref name="CAP"/> Females are sometimes reported as [[monogamous]],<ref name="CanGeo"/> but this is uncertain and [[polygyny]] may be more common.<ref name="UWSP">{{cite web |author=Matthew Hamilton |coauthors= Peter Hundt, Ryan Piorkowski |url=http://www.uwsp.edu/wildlife/carnivore/Mountain%20Lion%20Natural%20History_files/Mountain%20Lion%20Natural%20History_copy(1).htm |title=Mountain Lions |accessdate=2007-05-10 |publisher=[[University of Wisconsin, Stevens Point]]}}</ref> Copulation is brief but frequent.
[[Image:Mountain lion kittens.jpg|thumb|left|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.<ref name="Utah"/> Kitten survival rates are just over one per litter.<ref name="CAP"/>
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).<ref name="Utah"/> 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."<ref name="Dispersal">{{cite journal |last=Sweanor |first=Linda |coauthors=Kenneth A. Logan, Maurice G. Hornocker |year=2000 |month=June |title=Cougar Dispersal Patterns, Metapopulation Dynamics, and Conservation |journal=Conservation Biology |volume=14 |issue=3 |pages=798–808 |doi=10.1046/j.1523-1739.2000.99079.x |url=http://www.blackwell-synergy.com/links/doi/10.1046/j.1523-1739.2000.99079.x |format=PDF |accessdate=2007-05-02}}</ref>
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]].<ref name="CAP"/> 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.<ref>{{cite journal |last=Biek |first=Roman |coauthors=Allen G. Rodrigo, David Holley, Alexei Drummond, Charles R. Anderson Jr., Howard A. Ross, and Mary Poss |year=2003 |month=September |title=Epidemiology, Genetic Diversity, and Evolution of Endemic Feline Immunodeficiency Virus in a Population of Wild Cougars |journal=Journal of Virology |volume=77 |issue=17 |pages=9578–89 |doi=10.1128/JVI.77.17.9578-9589.2003 |url=http://jvi.asm.org/cgi/content/abstract/77/17/9578 |accessdate=2007-05-22 |pmid=12915571}}</ref>
===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 kilometer]]s (58 to 386 [[square mile|sq mi]]) with female ranges half the size.<ref name="CanGeo"/> Other research suggests a much smaller lower limit of 25 km<sup>2</sup> (10 sq mi) but an even greater upper limit of 1300 km<sup>2</sup> (500 sq mi) for males.<ref name="Utah"/> 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 km<sup>2</sup> (300 sq mi).<ref name="Dordt">{{cite web |url=http://homepages.dordt.edu/~mahaffy/mtlion/mtlionshort_behaviour.html |title=Behavior of cougar in Iowa and the Midwest |accessdate=2007-05-11 |last=Mahaffy |first=James |year=2004 |month=December |publisher=[[Dordt College]]}}</ref> 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.<ref name="Sierra"/>
Home range sizes and overall cougar abundance depend on terrain, vegetation, and prey abundance.<ref name="Utah"/> One female adjacent to the [[San Andres Mountains]], for instance, was found with a large range of 215 km<sup>2</sup> (83 sq mi), necessitated by poor prey abundance.<ref name="Dispersal"/> Research has shown cougar abundances from 0.5 animals to as much as 7 (in one study in South America) per 100 km<sup>2</sup> (38 sq mi).<ref name="CAP"/>
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.<ref name="Dordt"/> When males encounter each other, they hiss and spit, and may engage in violent conflict if neither backs down.<ref name="UWSP"/> Hunting or relocation of the cougar may increase aggressive encounters by disrupting territories and bringing young, transient animals into conflict with established individuals.<ref>{{cite web |url=http://www.sinapu.org/PDF/Front%20Range%20lion%20study.pdf |format=PDF |title=Mountain Lion (Puma concolor) study on Boulder Open Space |accessdate=2007-05-11 |date=2007-03-22 |work=Letter to the Parks and Open Space Advisory Committee, Boulder, Colorado |publisher=Sinapu}}</ref>
==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.<ref name="WhosWho"/> 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.<ref name="iucn"/> Its preferred habitats include precipitous canyons, escarpments, rim rocks, and dense brush.<ref name="Sierra"/>
More recently<!-- More recently than what? -->, reports from the Michigan Department Of Natural Resources have stated that cougars may be in Michigan's Upper Peninsula.{{Fact|date=May 2009}}
[[Image:MountainLion.jpg|thumb|left|Cougar, photographed in the [[Arizona-Sonora Desert Museum]], Tucson, Arizona.]]
The cougar was [[Local extinction|extirpated]] across much of its eastern North American range with the exception of [[Florida]] in the two centuries after [[European colonization of the Americas|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.<ref>{{cite web |url=http://www.naturalhistorymag.com/master.html?http://www.naturalhistorymag.com/0305/0305_selections.html |title=Bookshelf |last=Marschall |first=Laurence A. |date= |year=2005 |month=March |work=Natural Selections |publisher=[[Natural History Magazine]] |accessdate=2007-05-06}}</ref> [[DNA]] evidence has suggested its presence in eastern North America,<ref>{{cite web |url=http://lfpress.ca/newsstand/News/Local/2007/05/24/4205568.html |first=Joe |last=Belanger |title=DNA evidence of cougars found in southern Ontario |date=2007-05-24 |publisher=[[London Free Press]] |accessdate=2007-06-05}}</ref> while a consolidated map of cougar sightings shows numerous reports, from the mid-western [[Great Plains]] through to [[Eastern Canada]].<ref>{{cite web |url=http://www.cougarnet.org/bigpicture.html |title=The "Big" Picture |accessdate=2007-05-20 |author=[http://www.cougarnet.org/network.html Board of Directors] | year=2004| publisher=The Cougar Network}} ''The Cougar Network methodology is recognized by the [http://www.fws.gov/northeast/ECougar/ U.S. Fish and Wildlife Service].''</ref> The only unequivocally known eastern population is the [[Florida panther]], which is critically endangered. There have also been sightings in [[Elliotsville, Maine]] in the central part of the [[Maine|state]] and in [[New Hampshire]] there have been recent sightings as early as 1997<ref> {{cite web | url = http://www.rick-davidson.com/PDF/Mountain%20Lion%20Sightings%20Open%20Office%20092008%20photo%20final.pdf | format = PDF | title = NH Sightings Catamount | accessdate = 2009-03-20 | publisher=[[Beech River Books]]}}</ref>
On April 14, 2008 police shot and killed a cougar on the north side of Chicago, Illinois. DNA tests were consistent with cougars from the Black Hills. Less than one year later, on March 5, 2009, a cougar was photographed and unsuccessfully tranquilized by state wildlife biologists in a tree near Spooner, Wisconsin in the northwestern part of the state.<ref> {{cite web | url = http://www.stltoday.com/stltoday/lifestyle/stories.nsf/pets/story/5F51AA5F3CCF8DAD862575A00066E78F?OpenDocument | format = HTML | title = Sightings show cougars expanding into central US | accessdate = 2009-04-22 | publisher=St.Louis Today}}</ref>
South of the [[Rio Grande]], the [[World Conservation Union|International Union for the Conservation of Nature and Natural Resources]] (IUCN) lists the cat in every [[Central America|Central]] and [[South American]] country except [[Costa Rica]] and [[Panama]].<ref name="iucn"/> While specific state and provincial statistics are often available in North America, much less is known about the cat in its southern range.<ref> {{cite web | url = http://www.nwf.org/cats/pdfs/cougarfacts.pdf | format = PDF | title = Cougar facts | accessdate = 2007-05-20 | publisher=[[National Wildlife Federation]]}}</ref>
The cougar's total breeding population is estimated at less than 50,000 by the IUCN, with a declining trend.<ref name="iucn"/> 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.<ref>{{cite web |url=http://www.dfw.state.or.us/wildlife/cougar/ |title=Cougar Management Plan |accessdate=2007-05-20 |year=2006 |work=Wildlife Division: Wildlife Management Plans |publisher=Oregon Department of Fish and Wildlife}}</ref> [[California]] has actively sought to protect the cat and a similar number of cougars has been suggested, between 4,000 and 6,000.<ref name="California">{{cite web |url=http://www.dfg.ca.gov/news/issues/lion.html |title=Mountain Lions in California |accessdate=2007-05-20 |year=2004 |publisher=California Department of Fish and Game}}</ref>
===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]].
[[Image:Cougar track.jpg|thumb|Front paw print of a cougar. An adult paw print is approximately 10 cm (4 [[inches]]) long.<ref name="GovBC"/>]]
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 [[American Black Bear]]s visited 24% of cougar kills in Yellowstone and [[Glacier National Park]]s, usurping 10% of carcasses.<ref>{{cite paper |author=COSEWIC. Canadian Wildlife Service |title=Assessment and Update Status Report on the Grizzly Bear (Ursus arctos) |publisher=[[Environment Canada]] |year=2002 |url=http://dsp-psd.pwgsc.gc.ca/Collection/CW69-14-166-2002E.pdf |format=PDF |accessdate=2007-04-08 }}</ref>
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.<ref>{{cite web |url=http://www.washingtonpost.com/ac2/wp-dyn?pagename=article&node=&contentId=A2158-2003May17¬Found=true |title=In Yellowstone, it's Carnivore Competition |last=Gugliotta |first=Guy |date=2003-05-19 |publisher=Washington Post |accessdate=2007-04-09}}</ref> 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.<ref>{{cite web |url=http://www.greateryellowstonescience.org/topic/wolves/wolvesoverview.html |title=Overview: Gray Wolves |accessdate=2007-04-09 |publisher=Greater Yellowstone Learning Center}}</ref> 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."<ref>{{cite web |url=http://www.oregonwild.org/press-room/press-clips/turf-wars-in-idaho-s-wilderness |title=Turf wars in Idaho's wilderness |last=Cockle |first=Richard |date=2006-10-29 |publisher=[[The Oregonian]] |accessdate=2007-04-09}}</ref> Both species, meanwhile, are capable of killing mid-sized predators such as [[bobcat]]s and [[coyote]]s and tend to suppress their numbers.<ref name="Yellowstone"/>
In the southern portion of its range, the cougar and jaguar share overlapping territory.<ref name="HAMDIG">{{cite web |url=http://www.ecology.info/ecology-jaguar-puma.htm |first=Paul |last=Hamdig |title=Sympatric Jaguar and Puma |publisher=Ecology Online Sweden |accessdate=August 30 2006 |dateformat=mdy}}</ref> The jaguar tends to take larger prey and the cougar smaller where they overlap, reducing the cougar's size.<ref name="diet"/> Of the two felines, the cougar appears best able to exploit a broader prey niche and smaller prey.<ref name="foodhabits">{{cite journal |author=Rodrigo Nuanaez, Brian Miller, and Fred Lindzey |year=2000 |title=Food habits of jaguars and pumas in Jalisco, Mexico |journal=Journal of Zoology |volume=252 |issue=3 |page=373 |url=http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=58851 |accessdate=2006-08-08}}</ref>
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.<ref>{{cite journal |last=Robinson |first=Hugh S. |coauthors=Robert B. Wielgus, and John C. Gwilliam |year=2002 |title=Cougar predation and population growth of sympatric mule deer and white-tailed deer |journal=Canadian Journal of Zoology |volume=80 |issue=3 |pages=556–68 |doi=10.1139/z02-025 |url=http://rparticle.web-p.cisti.nrc.ca/rparticle/AbstractTemplateServlet?journal=cjz&volume=80&year=&issue=&msno=z02-025&calyLang=fra |accessdate=2007-05-20 |quote=}}</ref> The [[Vancouver Island Marmot]], an endangered species [[Endemism|endemic]] to one region of dense cougar population, has seen decreased numbers due to cougar and gray wolf predation.<ref>{{cite journal |last=Bryant |first=Andrew A. |coauthors=Page, Rick E. |year=2005 |title=Cougar predation and population growth of sympatric mule deer and white-tailed deer |journal=Canadian Journal of Zoology |month=May |volume=83 |issue=5 |pages=674–82 |url=http://www.ingentaconnect.com/content/nrc/cjz/2005/00000083/00000005/art00006?crawler=true |accessdate=2007-05-20 |doi=10.1139/z05-055}}</ref>
In the southern part of [[South America]] the Puma is a top level predator that has controlled the population of [[Guanaco]] and other species since prehistoric times.
==Hybrids==
[[Image:Pumapard-1904.jpg|thumb|upright|Pumapard, taken in 1904]]
{{Main|Pumapard}}
A pumapard is a [[hybrid (biology)|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 Zoological Garden|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.<ref name="Geo">{{cite web|url=http://www.geocities.com/pride_lands/Liger_Tigon.html|title=Geocites - Liger & Tigon Info|accessdate=9 June 2008|dateformat=dmy}}</ref>
==Conservation status==
The [[World Conservation Union]] (IUCN) currently lists the cougar as a "[[least concern]]" species. The cougar is regulated under [[CITES#Appendix I|Appendix I]] of the [[CITES|Convention on International Trade in Endangered Species of Wild Fauna and Flora]] (CITES),<ref>{{cite web |url=http://www.cites.org/eng/app/appendices.shtml |title=Appendices I, II and III |accessdate=2007-05-24 |publisher=[[Convention on International Trade in Endangered Species of Wild Fauna and Flora]]}}</ref> rendering illegal international trade in specimens or parts.
[[Image:Cougar snow.jpg|thumb|left|Cougar conservation depends on preservation of their habitat.]]
East of the [[Mississippi]], the only unequivocally known cougar population is the [[Florida panther]]. The [[United States Fish and Wildlife Service]] recognizes both an Eastern cougar and the Florida panther, affording protection under the [[Endangered Species Act]].<ref>{{cite web |url=http://www.fws.gov/endangered/i/a/saa48.html |title=Eastern Cougar |accessdate=2007-05-20 | work=Endangered and Threatened Species of the Southeastern United States (The Red Book)| year=1991 |publisher=[[U.S. Fish and Wildlife Service]]}}</ref><ref>{{cite web |url=http://www.fws.gov/endangered/i/a/saa05.html |title=Florida Panther |accessdate=2007-06-07 | work=Endangered and Threatened Species of the Southeastern United States (The Red Book)| year=1993 |publisher=[[United States Fish and Wildlife Service]]}}</ref> Certain taxonomic authorities have collapsed both designations into the [[North American Cougar]], with Eastern or Florida subspecies not recognized,<ref name="MSW3"/> while a subspecies designation remains recognized by some conservation scientists.<ref name=improving/> The most recent documented count for the Florida sub-population is 87 individuals, reported by recovery agencies in 2003.<ref>{{cite web |url=http://myfwc.com/panther/news/pdf/FWC2002-2003PantherGeneticRestorationAnnualReport.pdf |format=PDF |title=Florida Fish and Wildlife Conservation Commission. 2002–2003 Panther Genetfic Restoration Annual Report |accessdate=2007-06-05}}</ref>
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]].<ref name="CAP"/> 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 [[Hunting dog|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.<ref name="California"/> However statistics from the [[California Department of Fish and Game|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 six per year in the 1970s. The [[Bay Area Puma Project]] aims to obtain information on cougar populations in the San Francisco Bay area and the animals' interactions with habitat, prey, humans, and residential communities.<ref name=felidae1>{{cite web|url=http://www.felidaefund.org/research/bay_puma.html |title="Bay Area Puma Project" information page, Felidae Conservation Fund, accessed 18 February 2009 |publisher=Felidaefund.org |date= |accessdate=2009-03-08}}</ref>
Conservation threats to the species include persecution as a pest animal, degradation and fragmentation of their habitat, and depletion of their prey base. [[Wildlife corridor]] 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 km<sup>2</sup> (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.<ref>{{cite journal |last=Beier |first=Paul |year=1993 |month=March |title=Determining Minimum Habitat Areas and Habitat Corridors for Cougars |journal=Conservation Biology|volume=7|issue=1|pages=94–108 |url=http://links.jstor.org/sici?sici=0888-8892%28199303%297%3A1%3C94%3ADMHAAH%3E2.0.CO%3B2-I&size=LARGE&origin=JSTOR-enlargePage |accessdate=2007-05-20|doi=10.1046/j.1523-1739.1993.07010094.x}}</ref>
==Relationships with humans==
===In mythology and culture===
[[Image:MochePuma.jpg|thumb|[[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.<ref>Berrin, Katherine & Larco Museum. ''The Spirit of Ancient Peru:Treasures from the [[Larco Museum|Museo Arqueológico Rafael Larco Herrera]].'' New York: [[Thames and Hudson]], 1997.</ref> The sky and thunder god of the Inca, [[Viracocha]], has been associated with the animal.<ref>{{cite web |first=Kulmar |last=Tarmo |url=http://folklore.ee/folklore/vol12/inca.htm |coauthors=Kait Realo (translator) |title=On the role of Creation and Origin Myths in the Development of Inca State and Religion |accessdate=2007-05-22 |work=Electronic Journal of Folklore|publisher=Estonian Folklore Institute}}</ref>
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]] {{Fact|date=October 2008}} and the [[Cheyenne]], amongst others. To the [[Apache]] and [[Walapai]] of [[Arizona]], the wail of the Cougar was a harbinger of death.<ref>{{cite web |url=http://www.aphis.usda.gov/wildlife_damage/nwrc/publications/living/cougar.pdf |title=Living with Wildlife: Cougars |accessdate=2009-04-11 |format=PDF |publisher=[[USDA Wildlife Services]]}}</ref>
The cougar continues to be a symbol of strength and stealth. From [[Eurocopter Cougar|combat helicopters]], motor vehicles (see [[Mercury Cougar|Ford/Mercury Cougar]] and [[Ford Puma]]) to [[Puma AG|athletic shoes]], both "Cougar" and "Puma" are widely used as [[Brand|brand names]]. Various sports teams have also adopted the names, including the [[Argentina national rugby union team]], the [[National Autonomous University of Mexico]] soccer club as well as US universities, The [[Florida Panthers]], The [[College of Charleston]], [[Brigham Young University]], The [[University of Houston]], [[Washington State University]], and The [[University of Vermont]]. Many places, such as [[Cougar Mountain]], are also named after their association with cougars.<!--we don't need a massive list here, please-->
===Livestock predation===
Cougars can cause severe economic hardship on those whose livelihoods depend on livestock. During the early years of ranching, cougars were considered on par with wolves in destructiveness. According to figures in Texas in 1990 for example, 86 calves, 253 Mohair goats, 302 Mohair kids, 445 sheep and 562 lambs were confirmed to have been killed by cougars that year. In Nevada in 1992, cougars were confirmed to have killed 9 calves, 1 horse, 4 colts, 5 goats, 318 sheep and 400 lambs. In both cases, sheep were the most frequently attacked. Some instances of [[surplus killing]] have resulted in the deaths of 20 sheep in one attack.<ref name="Livestock">{{cite web | url=http://www.aws.vcn.com/mountain_lion_fact_sheet.html | title = Mountain Lion Fact Sheet | publisher = Abundant Wildlife Society of North America | accessdate = 2008-07-10}}</ref> Cougars frequently kill calves, sheep and goats by biting the top of the neck or head, differing greatly from the throat bite used by coyotes and indiscriminate mutilation by feral dogs. The size of the tooth puncture marks also helps distinguish kills made by cougars from those made by smaller predators.<ref name="Predation">{{cite web | url=http://texnat.tamu.edu/ranchref/predator//cougars/t-cougar.htm | title = Cougar Predation - Description | publisher = Procedures for Evaluating Predation on Livestock and Wildlife | accessdate = 2008-08-03}}</ref>
===Attacks on humans===
{{seealso|List of fatal cougar attacks in North America by decade}}
[[Image:MountainLionAttackProtocol.jpg|thumb|right|Mountain Lion warning sign]]
Due to the [[overpopulation|expanding human population]], 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.<ref name="Med">{{cite journal |last=McKee |first=Denise |year=2003 |title=Cougar Attacks on Humans: A Case Report |journal=Wilderness and Environmental Medicine |volume=14 |issue=3 |pages=169–73 |publisher=Wilderness Medical Society |url=http://www.wemjournal.org/wmsonline/?request=get-document&issn=1080-6032&volume=014&issue=03&page=0169 |accessdate=2007-05-20}}</ref> Attacks on people, livestock, and pets may occur when the cat [[Habituation|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.<ref name="GovBC"/>
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).<ref name="Beier">{{cite web |first=Paul |last=Beier |url=http://users.frii.com/mytymyk/lions/beier.htm |title=Cougar attacks on humans in United States and Canada |work=Wildlife Society Bulletin |year=1991 |accessdate=2007-05-20 |publisher=Northern Arizon University}}</ref> By 2004, the count had climbed to 88 attacks and 20 deaths.<ref>{{cite web |url=http://www.azgfd.gov/w_c/mtn_lion_attacks.shtml |title=Confirmed mountain lion attacks in the United States and Canada 1890 — Present |accessdate=2007-05-20 |publisher=Arizona Game and Fish}}</ref>
Within North America, the distribution of attacks is not uniform. The heavily populated state of [[California]] has seen a dozen attacks since 1986 (after just three from 1890 to 1985), including three fatalities.<ref name="California"/> Lightly populated [[New Mexico]] reported an attack in 2008, the first there since 1974.<ref>[http://www.wildlife.state.nm.us/publications/press_releases/documents/2008/062308pinosaltoslion.html Search continues for mountain lion that killed Pinos Altos man], New Mexico Department of Game and Fish, press release June 23, 2008]; [http://www.wildlife.state.nm.us/publications/press_releases/documents/2008/062508pinosaltoslion.html Wounded mountain lion captured, killed near Pinos Altos], New Mexico Department of Game and Fish, press release June 25, 2008]; [http://www.wildlife.state.nm.us/publications/press_releases/documents/2008/070108pinosaltoslion2.html Second mountain lion captured near Pinos Altos], New Mexico Department of Game and Fish, press release July 1, 2008]</ref>
As with many predators, a cougar may attack if cornered, if a fleeing human stimulates their instinct to chase, or if a person "[[wikt:play dead|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 persuading an attacking cougar to disengage.<ref name="Med"/><ref name="GovBC">{{cite web |url=http://www.env.gov.bc.ca/wld/documents/cougsf.htm |title=Safety Guide to Cougars |work=Environmental Stewardship Division |year=1991 |accessdate=2007-05-28 |publisher=[[Government of British Columbia]], Ministry of Environment}}</ref>
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.<ref name="Med"/> 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{{ndash}} and almost all fatalities{{ndash}} 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.<ref name="Beier"/>
==References==
{{reflist|2}}
==Other resources==
* {{cite book |last=Baron |first=David |title=The Beast in the Garden: A Modern Parable of Man and Nature |year=2004 |publisher=W. W. Norton and Company |location=New York |isbn=0393058077}}
* {{cite book |last=Kobalenko |first=Jerry |title=Forest Cats of North America |year=1997 |publisher=Island Press |location=Buffalo, New York |isbn=1-55209-174-4}}
* {{cite book |last=Logan |first=Ken |coauthors=Linda Sweanor |title=Desert Puma: Evolutionary Ecology and Conservation of an Enduring Carnivore |year=2001 |publisher=Island Press |isbn=1-55963-866-4}}
==External links==
{{portal|Cats|Cat03.jpg}}
{{portal|Mammals|Okapi2.jpg}}
{{wiktionary}}
{{commons|Puma concolor}}
{{wikispecies|Puma concolor}}
{{Spoken Wikipedia|Cougar1.ogg|2008-11-06}}
* [http://www.theanimalfiles.com/mammals/carnivores/mountain_lion.html Video, Pictures and Information on Mountain Lions]
* [http://www.cougarfund.org/ A Definitive Resource About Cougar] Comprehensive non-profit site with extensive information about cougars, from how to live safely in cougar country, to science abstracts, hunting regulations, state-by-state cougar management/policy info, to (rare) photos and videos of wild cougars.
* [http://dir.salon.com/story/people/feature/2002/02/12/cougar/index.html?pn=2 Description of a Cougar attack]
* [http://depts.washington.edu/natmap/facts/cougar_712.html Cougar Facts and Photos] - NatureMapping Program
* [http://biology.plosjournals.org/perlserv/?request=get-document&doi=10.1371/journal.pbio.0060040 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
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{{North American Game}}
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