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'{{Redirect|Mountain lion||Cougar (disambiguation)|and|Mountain lion (disambiguation)}} {{pp-pc1}}{{Use mdy dates|date=January 2015}} {{speciesbox | name = Cougar<ref name=MSW3/> | fossil_range = {{Fossil range|0.3|0}}<small>Middle [[Pleistocene]]–[[Holocene]]</small> | status = LC | status_system = iucn3.1 | status_ref =<ref name="iucn">{{IUCN2014.3|assessor=Caso, A.|assessor2=Lopez-Gonzalez, C.|assessor3=Payan, E.|assessor4=Eizirik, E.|assessor5=de Oliveira, T.|assessor6=Leite-Pitman, R.|assessor7=Kelly, M.|assessor8=Valderrama, C.|assessor9=Lucherini, M.|last-assessor-amp=yes|year=2008|id=18868|title=Puma concolor|downloaded=December 10, 2014}}</ref> | image = Cougar 25.jpg | image_caption = Captive cougar at the Cincinnati Zoo | genus = Puma | species = concolor | authority = ([[Carl Linnaeus|Linnaeus]], 1771) | range_map = Cougar_range_map_2010.png | range_map_caption = Cougar range | subdivision_ranks = [[Subspecies]] | subdivision = * ''P. c. anthonyi'' – eastern South America * ''[[Argentine cougar|P. c. cabrerae]]'' – central South America * ''[[Northern South American cougar|P. c. concolor]]'' – northern South America * ''[[Florida panther|P. c. coryi]]''? – south Florida * ''[[Costa Rican cougar|P. c. costaricensis]]'' – Central America * ''[[North American cougar|P. c. cougar]]'' – North America * ''P. c. puma'' – southern South America Also see [[#Subspecies|text]] | synonyms = * ''[[Felis]] concolor'' }} The '''cougar''' (''Puma concolor''), also commonly known as the '''mountain lion''', '''puma''', '''panther''', or '''catamount''', is a large [[Felidae|felid]] of the subfamily [[Felinae]] native to the Americas. Its [[range (biology)|range]], from the Canadian [[Yukon]] to the southern [[Andes]] of South America, is the greatest of any large wild terrestrial mammal in the [[Western Hemisphere]].<ref name="diet" /> An adaptable, [[Generalist and specialist species|generalist]] species, the cougar is found in most [[Americas|American]] [[habitat (ecology)|habitat]] types. It is the second-heaviest cat in the [[New World]], after the [[jaguar]]. Secretive and largely solitary by nature, the cougar is properly considered both [[nocturnal]] and [[crepuscular]], although there are daytime sightings.<ref name="cougar">[https://web.archive.org/web/20070224075326/http://www.nps.gov/ccso/cougar.htm Cougars]. US National Park Service.</ref><ref>Hansen, Kevin. (1992) ''Cougar: The American Lion.'' Northland. Flagstaff, AZ, ch. 4, ISBN 0873585445.</ref><ref>[https://web.archive.org/web/20110220144246/http://www.wildlife.state.nm.us/conservation/cougar/CougarEducationBooklet_1-31-11.pdf Cougar Education & Identification Course]. New Mexico Department of Game & Fish</ref><ref>[http://sagehen.ucnrs.org/Documents/visitors/wildlife/lion.pdf Living With California Mountain Lions]. California Department of Game & Fish</ref> The cougar is more closely related to smaller felines, including the [[Cat|domestic cat]] (subfamily Felinae), than to any species of subfamily [[Pantherinae]],<ref name=MSW3/><ref>Hartwell, Sarah. [http://www.messybeast.com/cathistory.htm The Domestication of the Cat]. messybeast.com</ref><ref>[http://www.messybeast.com/small-wildcats.htm Small Wild Cat Species]. messybeast.com</ref> of which only the jaguar is native to the Americas. The cougar is an [[ambush predator]] and pursues a wide variety of prey. Primary food sources are [[ungulate]]s, particularly [[deer]], but also [[livestock]]. It also hunts species as small as insects and [[rodents]]. This cat prefers habitats with dense underbrush and rocky areas for stalking, but can also live in open areas. The cougar is [[Territory (animal)|territorial]] and survives at low population densities. Individual territory sizes depend on terrain, vegetation, and abundance of prey. While large, it is not always the [[apex predator]] in its range, yielding to the jaguar, [[gray wolf]], [[American black bear]], and [[grizzly bear]]. It is reclusive and mostly avoids people. [[List of fatal cougar attacks in North America by decade|Fatal attacks on humans]] are rare, but have recently been increasing in North America as more people enter their territories.<ref name="Med"/> Intensive hunting following [[European colonization of the Americas]] and the ongoing human development of cougar [[habitat]] has caused populations to drop in most parts of its historical range. In particular, the cougar was [[Local extinction|extirpated]] in eastern North America in the beginning of the 20th century, except for an isolated [[Florida panther]] subpopulation. Transient males have been verified in Minnesota,<ref>[http://www.startribune.com/local/134835873.html Shot fells elusive cougar chased out of a culvert]. StarTribune.com (December 1, 2011). Retrieved on April 29, 2013.</ref> [[Missouri]],<ref>[http://mdc.mo.gov/discover-nature/wildlife-reporting/mountain-lion-reports/confirmed-mountain-lion-reports Confirmed Mountain Lion Reports]. Missouri Department of Conversation, mdc.mo.gov (March 20, 2014)</ref> [[Wisconsin]],<ref>{{cite news| url=http://www.dailymail.co.uk/news/article-2090026/Extinct-Sightings-cougars-rise-eastern-states.html |date=January 22, 2012 | location=London | work=Daily Mail | title=Extinct? Cougar sightings on the rise in eastern United States}}</ref> Iowa,<ref>[http://www.iowadnr.gov/home/ctl/detail/mid/2822/itemid/581 Mountain Lion Shot Early Friday Morning in Monona County]. Iowa Department of Natural Resources, iowadnr.gov (December 23, 2011)</ref><ref>[http://www.kwwl.com/story/16385438/mountain-lion-shot-by-police-in-monona-county Mountain lion shot by police in Monona County – KWWL.com – News & Weather for Waterloo, Dubuque, Cedar Rapids & Iowa City, Iowa |]. KWWL.com. Retrieved on April 29, 2013.</ref> the [[Upper Peninsula]] of Michigan, and Illinois, where a cougar was shot in the city limits of Chicago<ref>{{cite web|url=http://www.cougarnet.org/centralmidwest.html |title=The Cougar Network – Using Science to Understand Cougar Ecology |publisher=Cougarnet.org |accessdate=February 16, 2011}}</ref><ref>{{cite web|url=http://twitpic.com/od2y4 |title=Trail cam photo of cougar in the eastern Upper Peninsula|publisher=Twitpic.com |date=November 5, 2009 |accessdate=February 16, 2011}}</ref><ref>{{cite web|url=http://www.poorboysoutdoors.com/2010/05/indiana-confirms-green-county-mountain-lion/ |archiveurl=https://web.archive.org/web/20100711000325/http://www.poorboysoutdoors.com/2010/05/indiana-confirms-green-county-mountain-lion/ |archivedate=July 11, 2010 |title=Indiana confirms mountain lion in Green County |publisher=Poorboysoutdoors.com |date=May 10, 2010 |accessdate=February 16, 2011}}</ref> and, in at least one instance, observed as far east as coastal [[Connecticut]].<ref>{{cite news|url=http://content.usatoday.com/communities/ondeadline/post/2011/07/mountain-lion-killed-in-conn-had-walked-from-south-dakota/1|title=Mountain lion killed in Conn. had walked from S. Dakota|publisher=USA Today| date=July 26, 2011| accessdate=October 27, 2012}}</ref><ref>{{cite web|title=Connecticut Mountain Lion Likely Came From The Black Hills|url=http://www.npr.org/blogs/thetwo-way/2011/07/27/138748682/connecticut-mountain-lion-likely-came-from-the-black-hills|work=[[NPR]] News|accessdate=July 28, 2011}}</ref> Reports of [[eastern cougar]]s ''(P. c. cougar)'' still surface, although it was declared extirpated in 2011.<ref name="Eastern Cougar Is Declared Extinct">{{cite news|title=Eastern Cougar Is Declared Extinct|url=http://www.nytimes.com/2011/03/03/science/earth/03cougar.html?_r=0|publisher=The New York Times|accessdate=March 2, 2011|first=Felicity|last=Barringer|date=March 2, 2011}}</ref> ==Naming and etymology== With its vast range across the length of the Americas, ''P. concolor'' has dozens of names and various references in the [[mythology]] of the [[Indigenous peoples of the Americas|indigenous Americans]] and in contemporary culture. Currently, it is referred to as "puma" by most scientists<ref>{{cite web |title=Mountain Lion (Puma, Cougar) |publisher=San Diego Zoo |url=http://animals.sandiegozoo.org/animals/mountain-lion-puma-cougar |accessdate=December 14, 2014 }}</ref> and by the populations in 21 of the 23 countries in the Americas where "puma" is the common name in Spanish or Portuguese.<ref name=EOL>{{cite book |title=Encyclopedia of Life |url=http://eol.org/pages/311910/names/common_names |chapter=''Puma concolor''|accessdate=December 14, 2014 }}</ref> The cat has many local or regional names in the United States and Canada, of which cougar, puma, mountain lion, and panther are popular.<ref>{{Cite journal | last1 = Martinez Guerrero | first1 = J. H. | last2 = Pereda Solis | first2 = M. E. P| last3 = Zarate Ramos | first3 = J. J. | last4 = Rosales Alferez | first4 = F. | last5 = Herrera Casio| first5 = H. | title = Report of ''Physaloptera praeputialis'' (Von Linstow 1889, Nematoda) in Mountain Lion (''Puma concolor'', Linneaus 1771) | doi = 10.3923/javaa.2010.601.603 | journal = Journal of Animal and Veterinary Advances | volume = 9 | issue = 3 | pages = 601–603 | year = 2010 | pmid = | pmc = |type=free full text}}</ref> "Mountain lion" was a term first used in writing in 1858 from the diary of [[George A. Jackson]] of [[Colorado]].<ref>{{cite book |title=George A. Jackson's Diary of 1858–1859 |author=Jackson, George A. |editor=LeHafen, Roy |journal=Colorado Magazine |year=1935 |volume=6 |pages=201–214 }}</ref> Other names include catamount (probably a contraction from "cat of the mountain"), mountain screamer, and painter. Lexicographers regard painter as a primarily upper-Southern US regional variant on panther.<ref>{{cite web|url=http://www.bartleby.com/61/61/P0016100.html|archiveurl=https://web.archive.org/web/20010313081943/http://www.bartleby.com/61/61/P0016100.html|archivedate=March 13, 2001|title=painter|work=transcription of the ''American Heritage Dictionary'', Bartleby.com|accessdate=March 8, 2009}}</ref> The word ''panther'' is commonly used to specifically designate the [[black panther]], a [[melanistic]] jaguar or leopard, and the [[Florida panther]], a subspecies of cougar (''P.{{nbsp}}c. coryi''). ''P. concolor'' holds the [[Guinness World Records|Guinness]] record for the animal with the greatest number of names, with over 40 in English alone.<ref>{{cite book| title = The Guinness Book of World Records| year = 2004| page = 49}}</ref> "Cougar" may be borrowed from the archaic [[Portuguese language|Portuguese]] ''çuçuarana''; the term was originally derived from the [[Tupi language]] ''susua'rana'', meaning "similar to deer (in hair color)". A current form in [[Brazil]] is ''[[wikt:suçuarana|suçuarana]]''. It may also be borrowed from the [[Guaraní language]] term ''guaçu&nbsp;ara'' or ''guazu&nbsp;ara''. Less common Portuguese terms are ''[[wikt:onça-parda|onça-parda]]'' (brown ''onça'', in distinction of the black-spotted [yellow] one, ''[[wikt:onça-pintada|onça-pintada]]'', the [[jaguar]]) or ''[[wikt:leão-baio|leão-baio]]'' (lit. [[Chestnut (color)|chestnut]] lion), or unusually non-native ''puma'' or ''leão-da-montanha'', more common names for the animal when native to a region other than South America (especially for those who do not know that ''suçuaranas'' are found elsewhere but with a different name). People in rural regions often refer to both the cougar and the jaguar as simply ''[[wikt:gata|gata]]'' (she-cat), and outside of the Amazon, both are colloquially referred to as simply ''[[wikt:onça|onça]]'' by many people (that is also a name for the leopard in [[Angola]]). In the 17th century, German naturalist [[Georg Marcgrave]] named the cat the ''cuguacu ara''. Marcgrave's rendering was reproduced by his associate, Dutch naturalist [[Willem Piso]], in 1648. ''Cuguacu ara'' was then adopted by English naturalist [[John Ray]] in 1693.<ref>{{cite journal|title=Words to the Wise|journal=Take Our Word for it|issue=205 |page=2 |url=http://www.takeourword.com/TOW205/page2.html |accessdate=July 31, 2012 }}</ref> The French naturalist [[Georges-Louis Leclerc, Comte de Buffon]] in 1774 (probably influenced by the word "jaguar") converted the ''cuguacu ara'' to ''cuguar'', which was later modified to "cougar" in English.<ref>{{OEtymD|jaguar}}</ref><ref>{{cite web| url= http://www.merriam-webster.com/dictionary/cougar| title = cougar | publisher=Merriam-Webster Dictionary Online}}</ref><ref>{{cite web| url = http://oxforddictionaries.com/definition/english/cougar| year = 1989| title = cougar | work=Oxford Dictionaries Online, Oxford University Press}}</ref> The first English record of "puma" was in 1777, where it had come from the Spanish, who in turn borrowed it from the Peruvian [[Quechua language]] in the 16th century, where it means "powerful".<ref>{{cite web |title=The Puma |publisher=Projeto Puma |url=http://uniplac.net/~puma/puma.htm |archiveurl=https://web.archive.org/web/20070706131806/http://uniplac.net/~puma/puma.htm |archivedate=July 6, 2007 |accessdate=July 31, 2012 }}</ref> ==Taxonomy and evolution== [[File:Puma face.jpg|thumb|Although large, the cougar is more closely related to smaller felines than to other big cats.]] [[File:Cougar closeup.jpg|thumb|Close-up of face]] Cougars are the largest of the small cats. They are placed in the subfamily [[Felinae]], although their 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 about 11 million years ago. Taxonomic research on felids remains partial, and much of what is known about their evolutionary history is based on [[mitochondrial DNA]] analysis,<ref>{{cite news |first=Nicholas |last=Wade |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 |date= January 6, 2006 |accessdate=June 3, 2007}}</ref> as cats are poorly represented in the [[fossil record]],<ref name="Johnson2006"/> and significant [[confidence intervals]] exist with suggested dates. In the latest [[genomic]] study of the 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 8.0 to 8.5&nbsp;million years ago (Mya). The lineages subsequently diverged in that order.<ref name="Johnson2006">{{cite journal |author1=Johnson, W.E.|author2= Eizirik, E.|author3= Pecon-Slattery, J.|author4= Murphy, W.J.|author5= Antunes, A.|author6= Teeling, E. |author7= O'Brien, S.J. |date=January 6, 2006 |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 |pmid=16400146|bibcode= 2006Sci...311...73J}}</ref> North American felids then invaded South America 3{{nbsp}}Mya as part of the [[Great American Interchange]], following formation of the [[Isthmus of Panama]]. The cougar was originally thought to belong in ''Felis'' (''Felis concolor''), the genus which includes the [[cat|domestic cat]]. As of 1993, it is now placed in ''Puma'' along with the [[jaguarundi]], a cat just a little more than a tenth its weight. 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. |author2=Johnson, W.E. |author3=Pecon-Slattery, J. |author4=O'Brien, S.J. |year=2000 |title=Genomic Ancestry of the American Puma |journal=Journal of Heredity |volume=91 |issue=3 |pages=186–97 | doi = 10.1093/jhered/91.3.186 |pmid=10833043}}</ref> but the relationship is unresolved. The cheetah lineage is suggested by some studies to have 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 |author2=Barnes, Ian |author3=Phillips, Matthew J. |author4=Martin, Larry D. |author5=Harington, C. Richard |author6=Leonard, Jennifer A. |author7=Cooper, Alan |date= 2005 |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 |pmid=16085477}}</ref> The outline of [[Felinae|small feline]] migration to the Americas is thus unclear. A high level of genetic similarity has been found among North American cougar populations, suggesting they are all fairly recent descendants of a small ancestral group. Culver ''et al.'' propose the original North American population of ''P.{{nbsp}}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 1980s, as many as 32 [[subspecies]] were recorded; [[Genetics|genetic]] study of mitochondrial DNA<ref name="Culver" /> found 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{{nbsp}}ed.) recognizes six subspecies, five of which are solely found in Latin America:<ref name=MSW3>{{MSW3 Wozencraft |id = 14000204|pages=544–45}}</ref> * [[Argentine cougar]] {{nobold|(''Puma concolor cabrerae'')}} Pocock, 1940:<br> includes the previous subspecies and synonyms ''hudsonii'' and ''puma'' * [[Costa Rican cougar]] {{nobold|(''P. c. costaricensis'')}} Merriam, 1901 * [[Eastern South American cougar]] {{nobold|(''P. c. anthonyi'')}} Nelson and Goldman, 1931:<br> includes the previous subspecies and synonyms ''acrocodia'', ''borbensis'', ''capricornensis'', ''concolor'', ''greeni'', and ''nigra'' * [[North American cougar]] {{nobold|(''P. c. couguar'')}} Kerr, 1792:<br> includes the previous subspecies and synonyms ''arundivaga'', ''aztecus'', ''browni'', ''californica'', ''floridana'', ''hippolestes'', ''improcera'', ''kaibabensis'', ''mayensis'', ''missoulensis'', ''olympus'', ''oregonensis'', ''schorgeri'', ''stanleyana'', ''vancouverensis'', and ''youngi'' * [[Northern South American cougar]] {{nobold|(''P. c. concolor'')}} Linnaeus, 1771:<br> includes the previous subspecies and synonyms ''bangsi'', ''incarum'', ''osgoodi'', ''soasoaranna, sussuarana'', ''soderstromii'', ''suçuaçuara'', and ''wavula'' * [[Southern South American cougar]] {{nobold|(''P. c. puma'')}} Molina, 1782:<br> includes the previous subspecies and synonyms ''araucanus'', ''concolor'', ''patagonica'', ''pearsoni'', and ''puma'' '''''[[Incertae sedis|Incerta sedis]]''''' * [[Florida panther]] {{nobold|(''P. c. coryi'')}} The status of the Florida panther remains uncertain. It is still regularly listed as subspecies ''P. c. coryi'' in research works, including those directly concerned with its conservation.<ref name=improving>{{cite journal |first=Michael J. |last=Conroy |author2=Beier, Paul|author3=Quigley, Howard|author4=Vaughan, Michael R. |year=2006 |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 }}</ref> Culver et al. noted low [[Microsatellite (genetics)|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 use of genetics in puma taxonomy is not resolved at this time."<ref name=FloridaRecovery>{{cite web |url=https://www.fws.gov/uploadedFiles/Panther%20Recovery%20Plan.pdf |title=Florida Panther Recovery Plan |accessdate=June 15, 2016 |author=The Florida Panther Recovery Team |date=November 1, 2008 |format=PDF |publisher=U.S. Fish and Wildlife Service}}</ref> ==Biology and behavior== ===Physical characteristics=== [[File:Cougarskull.jpg|thumb|Cougar skull and jawbone]] Cougars are slender and agile members of the cat family. They are the fourth-largest cat;<ref>[https://web.archive.org/web/20101121100620/http://cougarnet.org/cougarfacts.html Expanding Cougar Population]. The Cougar Net.org</ref> adults stand about {{convert|60|to|90|cm|in|abbr=on}} tall at the shoulders.<ref>[https://web.archive.org/web/20120323144937/http://www.floridapanther.org/panther_facts.html Florida Panther Facts]. Florida Panther Refuge</ref> Adult males are around {{convert|2.4|m|ft|abbr=on}} long from nose to tail tip, and females average {{convert|2.05|m|ft|abbr=on}}, with overall ranges between {{convert|1.5|to|2.75|m|ft|abbr=on}} 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=March 30, 2007 |publisher=Texas Parks and Wildlife}}</ref><ref name="NY">{{cite web |url=http://www.dec.ny.gov/animals/6974.html |title=Eastern Cougar Fact Sheet |accessdate=March 30, 2007 |publisher=New York State Department of Environmental Conservat ion}}</ref> Of this length, {{convert|63|to|95|cm|in|abbr=on}} is comprised by the tail.<ref name=ADW>Shivaraju, A. (2003) [http://animaldiversity.ummz.umich.edu/site/accounts/information/Puma_concolor.html ''Puma concolor'']. Animal Diversity Web, University of Michigan Museum of Zoology. Retrieved on September 15, 2011.</ref> Males typically weigh 53 to 100&nbsp;kg (115 to 220&nbsp;lb), averaging 62&nbsp;kg (137&nbsp;lb). Females typically weigh between 29 and 64&nbsp;kg (64 and 141&nbsp;lb), averaging 42&nbsp;kg (93&nbsp;lb).<ref name=ADW/><ref name="CAP">{{Cite journal |author1=Nowell, K. |author2=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=July 27, 2007 }}</ref><ref>{{cite web|url=http://pick4.pick.uga.edu/mp/20q?search=Puma+concolor |title=Puma concolor – Mountain Lion – Discover Life |publisher=Pick4.pick.uga.edu |accessdate=February 16, 2011}}</ref> Cougar size is smallest close to the [[equator]], and larger towards the [[Polar region|poles]].<ref name="diet">{{cite journal |author1=Iriarte, J. Agustin|author2= Franklin, William L.|author3= Johnson, Warren E. |author4=Redford, Kent H. |year=1990 |title=Biogeographic variation of food habits and body size of the America puma |journal=[[Oecologia]] |volume=85 |issue=2 |pages=185–190 |doi=10.1007/BF00319400}}</ref> The largest recorded cougar, shot in 1901, weighed 105.2&nbsp;kg (232&nbsp;lb); claims of 125.2&nbsp;kg (276&nbsp;lb) and 118&nbsp;kg (260&nbsp;lb) have been reported, though they were most likely exaggerated.<ref>{{cite book |last= Hornocker|first= Maurice|date= 2010|title= Cougar : ecology and conservation|url= http://library.wur.nl/WebQuery/clc/1941110|location= |publisher= Chicago [etc.] : University of Chicago Press|page= |isbn= 0226353443|access-date= }}</ref> On average, adult male cougars in British Columbia weigh 56.7&nbsp;kg (125&nbsp;lb) and adult females 45.4&nbsp;kg (100&nbsp;lb), though several male cougars in British Columbia weighed between 86.4 and 95.5&nbsp;kg (190 to 210&nbsp;lb).<ref>{{cite web|last=Spalding|first=D. J.|title=Cougar in British Columbia|url=http://www.env.gov.bc.ca/wld/documents/cougar.htm|work=British Columbia Fish and Wildlife Branch}}</ref> [[File:Cougar sitting.jpg|thumb|left|upright|Although cougars somewhat resemble the [[domestic cat]], they are about the same size as an adult human.]] The head of the cat is round and the ears are 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 |archiveurl=https://web.archive.org/web/20070518235841/http://www.hww.ca/hww2.asp?pid=1&id=87&cid=8 |archivedate=May 18, 2007 |title=Cougar |work=Hinterland Who's Who |accessdate=May 22, 2007 |publisher=[[Canadian Wildlife Service]] and [[Canadian Wildlife Federation]]}}</ref> Cougars can be almost as large as [[jaguar]]s, but are less muscular and not as powerfully built; where their ranges overlap, the cougar tends to be smaller on average. Besides the jaguar, the cougar is on average larger than all felids apart from lions and tigers. 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 journal |last=Weissengruber |first=GE |author2=Forstenpointner G |author3=Peters G |author4=Kübber-Heiss A |author5=Fitch WT |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'')|publisher=Anatomical Society of Great Britain and Ireland |pages=195–209 |volume=201 |issue=3 |year=2002 |doi=10.1046/j.1469-7580.2002.00088.x|pmc=1570911 |pmid=12363272 |journal=Journal of Anatomy}}</ref> Compared to "big cats", cougars are often silent with minimal communication through vocalizations outside of the mother-offspring relationship.<ref>{{cite book|author1=Hornocker, Maurice G. |author2=Negri, Sharon |title=Cougar: ecology and conservation|url=https://books.google.com/books?id=ZSDUxT3fSYEC&pg=PA114|accessdate=September 15, 2011|date=December 15, 2009|publisher=University of Chicago Press|isbn=978-0-226-35344-9|pages=114–}}</ref> Cougars sometimes voice low-pitched hisses, growls, and purrs, as well as chirps and whistles, many of which are comparable to those of domestic cats. They are well known for their screams, as referenced in some of their common names, although these screams are often misinterpreted to be the calls of other animals or humans.<ref>{{cite web |url=http://www.easterncougar.org/pages/abouteasterncougars.htm |title=About Eastern Cougars |accessdate=June 3, 2007 |publisher=Eastern Cougar Foundation}}</ref> [[File: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 underbody, 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|author=Glenn, Eddie|url=http://www.tahlequahdailypress.com/black-mountain-lion-more-rumor-than-fact/article_db970b99-4383-5306-a277-10beadd92364.html |title=Black cougar more talk than fact |date=February 2, 2006 |accessdate=June 15, 2016 |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|work=messybeast.com}}</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. The cougar's top running speed ranges between {{convert|64|and|80|km/h|abbr=on}},<ref name=ZoologicalWildLifeFoundation>{{cite web |url=http://zoologicalwildlifefoundation.com/animals/mammals/felines/cougar/|title=Cougar|publisher=Zoological Wildlife Foundation |accessdate=October 31, 2012}}</ref><ref>[http://mountainlion.org/FAQfrequentlyaskedquestions.asp Mountain Lion FAQ and Facts]. Mountainlion.org. Retrieved on April 29, 2013.</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=May 20, 2007 |publisher=[[Sierra Club]] |deadurl=yes |archiveurl=https://web.archive.org/web/20150123034652/http://arizona.sierraclub.org/conservation/mt-lion/index.asp |archivedate=January 23, 2015 }}</ref> ===Hunting and diet=== A successful [[Generalist and specialist species|generalist]] [[predator]], the cougar will eat any animal it can catch, from insects to large [[ungulate]]s (over 500&nbsp;kg). Like all cats, it is an [[obligate carnivore]], meaning it needs to feed exclusively on meat to survive. The mean weight of vertebrate prey (MWVP) that pumas attack increases with the puma's body weight; in general, MWVP is lower in areas closer to the [[equator]].<ref name="diet"/> Its most important prey species are various deer species, particularly in North America; [[mule deer]], [[white-tailed deer]], [[elk]] and even bull [[moose]] are taken. Other species such as the [[bighorn sheep|bighorn]] and [[Dall's sheep]], [[horse]], [[fallow deer]], [[caribou]], [[mountain goat]], [[coyote]], [[pronghorn]], and domestic livestock such as cattle and sheep are also primary food bases in many areas.<ref name="SW">{{cite journal |doi=10.2307/3672527 |title=Influence of predation by mountain lions on numbers and survivorship of a feral horse population |jstor=3672527 |first=John W. |last=Turner |first2=Michael L. |last2=Morrison |year=2008 |journal=The Southwestern Naturalist |volume=46 |issue=2 |pages=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"/> [[File:Mountain Lion441.jpg|thumb|A captive cougar feeding. 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 which 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=April 8, 2007}}<br />* {{cite web |author1=Akenson, Holly |author2=Akenson, James |author3=Quigley, Howard |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 |author1=Oakleaf, John K. |author2=Mack, Curt |author3=Murray, Dennis L. |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. |author2=Jalkotzy, M. G.|author3=Festa-Bianchet, M. | 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 |doi=10.1139/z97-098}}</ref> In [[Pacific Rim National Park Reserve]], [[Feces#Animal feces|scat]] samples showed [[raccoon]]s to make up 28% of the cougar's diet, [[harbor seal]]s and [[blacktail deer]] 24% each, [[North American river otter]]s 10%, [[California sea lion]] 7%, and [[American mink]] 4%; the remaining 3% were unidentified.<ref>{{cite web |url= http://www.petethomasoutdoors.com/2012/02/british-columbia-cougars-found-to-prey-on-seals-sea-lions.html|title= British Columbia cougars found to prey on seals, sea lions|author=<!--Staff writer(s); no by-line.--> |date= February 23, 2012|website= |publisher= |access-date= August 5, 2016}}</ref> In the Central and South American cougar range, the ratio of deer in the diet declines. Small to mid-sized mammals are preferred, including large rodents such as the [[capybara]]. Ungulates accounted for only 35% of prey items in one survey, about half that of North America. Competition with the larger jaguar 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]]s, [[American beaver|beaver]]s, raccoons, [[hare]]s, [[guanaco]], [[peccary]], [[vicuna]], [[rhea (bird)|rhea]], and [[wild turkey]].<ref>Whitaker, John O. ''The Audubon Society Field Guide to North American Mammals.'' Chanticleer Press, New York, 1980, pg. 598. ISBN 0-394-50762-2.</ref> Birds and small reptiles are sometimes preyed upon in the south, but this is rarely recorded in North America.<ref name="diet"/> Not all of their prey is listed here due to their large range. 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"/> Kills are generally estimated around one large ungulate every two weeks. The period shrinks for females raising young, and may be as short as one kill every three days when cubs are nearly mature around 15 months.<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. The cougar is generally reported to not be a [[scavenger]], and rarely consumes 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. |author2=Logan, Kenneth A. |author3=Sweanor, Linda L. |author4=Boyce, Walter M. |date=December 2005 |editor1-last=Jones |editor1-first=Cheri A. |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}}</ref> ===Reproduction and life cycle=== Females reach sexual maturity between one-and-a-half to three years of age. They typically average one [[litter (animal)|litter]] every two to three years throughout their reproductive lives,<ref name="Utah">{{cite web |url=http://www.wildlife.utah.gov/pdf/cmgtplan.pdf |title=Utah Cougar Management Plan (Draft) |accessdate=May 2, 2007 |author=Cougar Discussion Group |date=January 27, 1999 |format=PDF |publisher=Utah Division of Wildlife Resources}}</ref> though the period can be as short as one year.<ref name="CAP"/> Females are in [[estrus]] for about 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">{{cite web |url=http://www.canadiangeographic.ca/Magazine/mj04/indepth/justthefacts.asp |title=Cougars in Canada (Just the Facts) |accessdate=April 2, 2007 |publisher=[[Canadian Geographic Magazine]]}}</ref> but this is uncertain and [[polygyny]] may be more common.<ref name="UWSP">{{cite web |author1=Hamilton, Matthew |author2=Hundt, Peter |author3=Piorkowski, Ryan |url=http://www.uwsp.edu/wildlife/carnivore/Mountain%20Lion%20Natural%20History_files/Mountain%20Lion%20Natural%20History_copy(1).htm |title=Mountain Lions |accessdate=May 10, 2007 |publisher=[[University of Wisconsin, Stevens Point]]}}</ref> Copulation is brief but frequent. Chronic stress can result in low reproductive rates when in captivity as well as in the field.<ref>{{cite journal|author1=Bonier, F.|author2= Quigley, H. |author3=Austad, S. |year= 2004|title= A technique for non-invasively detecting stress response in cougars|journal= Wildlife Society Bulletin|volume= 32|issue=3|pages=711–717|doi=10.2193/0091-7648(2004)032[0711:ATFNDS]2.0.CO;2|url=http://post.queensu.ca/~bonierf/pubs/pub1.pdf}}</ref> [[File:Mountain lion kittens.jpg|thumb|left|Cougar cubs]] [[File:Puma cub Malibu Springs area National Park Service December 2013.jpg|thumb|left|Cougar cub]] Only females are involved in parenting. Female cougars are fiercely protective of their cubs, and have been seen to successfully fight off animals as large as Grizzly bears in their defense. Litter size is between one and six cubs; typically two. Caves and other alcoves that offer protection are used as litter dens. Born blind, cubs are completely dependent on their mother at first, and begin to be weaned at around three months of age. As they grow, they begin to go out on forays with their mother, first visiting kill sites, and after six months beginning to hunt small prey on their own.<ref name="Utah"/> Kitten survival rates are just over one per litter.<ref name="CAP"/> When cougars are born, they have spots, but they lose them as they grow, and by the age of 2{{nbsp}}1/2 years, they will be completely gone<ref>{{cite web|url=http://wildlife.utah.gov/dwr/learn-more/living-with-cougars.html |title=Staying safe in cougar country |publisher=Wildlife.utah.gov |accessdate=October 6, 2011}}</ref> Young 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 [[mortality rate|mortality]] amongst cougars that travel farthest from the maternal range, often due to conflicts with other cougars ([[intraspecific competition]]).<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 |author2=Logan, Kenneth A. |author3=Hornocker, Maurice G. |year=2000 |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}}</ref> Life expectancy in the wild is reported at eight to 13 years, and probably averages eight to 10; a female of at least 18 years was reported killed by hunters on [[Vancouver Island]].<ref name="CAP"/> Cougars may live as long as 20 years in captivity. One male North American cougar (''P. c. couguar''), named Scratch, was two months short of his 30th birthday when he died in 2007.<ref name="Scratch">{{cite web |url=http://www.bigcatrescue.org/forever/scratchcougar.htm |archiveurl=https://web.archive.org/web/20081119080302/http://www.bigcatrescue.org/forever/scratchcougar.htm |archivedate=November 19, 2008 |title=Scratch |accessdate=August 21, 2009|work=bigcatrescue.org}}</ref> Causes of death in the wild include disability and disease, competition with other cougars, starvation, accidents, and, where allowed, human hunting. [[Feline immunodeficiency virus]], an endemic HIV-like virus in cats, is well-adapted to the cougar.<ref>{{cite journal |last=Biek |first=Roman |author2=Rodrigo, Allen G. |author3=Holley, David |author4=Drummond, Alexei |author5=Anderson Jr., Charles R. |author6=Ross, Howard A. |author7=Poss, Mary |date= 2003 |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 |pmid=12915571 |pmc=187433}}</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&nbsp;km<sup>2</sup> (58 to 386&nbsp;sq&nbsp;mi) with female ranges half the size.<ref name="CanGeo"/> Other research suggests a much smaller lower limit of 25&nbsp;km<sup>2</sup> (10&nbsp;sq&nbsp;mi), but an even greater upper limit of 1300&nbsp;km<sup>2</sup> (500&nbsp;sq&nbsp;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&nbsp;km<sup>2</sup> (300&nbsp;sq&nbsp;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=May 11, 2007 |last=Mahaffy |first=James |date=December 2004 |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 [[territorial marking|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&nbsp;km<sup>2</sup> (83&nbsp;sq&nbsp;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&nbsp;km<sup>2</sup> (38&nbsp;sq&nbsp;mi).<ref name="CAP"/> Because males disperse farther than females and compete more directly for mates and territory, they are most likely to be involved in conflict. Where a subadult 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, 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 |archiveurl=https://web.archive.org/web/20070616200443/http://www.sinapu.org/PDF/Front%20Range%20lion%20study.pdf |archivedate=June 16, 2007 |format=PDF |title=Mountain Lion (''Puma concolor'') study on Boulder Open Space |accessdate=May 11, 2007 |date=March 22, 2007 |work=Letter to the Parks and Open Space Advisory Committee, Boulder, Colorado |publisher=Sinapu}}</ref> ==Ecology== ===Distribution and habitat=== [[File:Mountain-lion-01623.jpg|thumb|A cougar in Yellowstone National Park]] [[File:Puma concolor camera trap Arizona 2.jpg|thumb|A [[camera trap]] image of a cougar in Saguaro National Park, Arizona]] 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]]. 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. 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"/> 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 of its territory. Currently, it ranges across most western American states, the Canadian provinces of [[Alberta]], [[Saskatchewan]] and [[British Columbia]], and the Canadian territory of [[Yukon]]. 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=March 2005 |work=Natural Selections |publisher=[[Natural History Magazine]] |accessdate=May 6, 2007}}</ref> [[DNA]] evidence has suggested its presence in eastern North America,<ref>{{cite web|url=http://pbdba.lfpress.com/cgi-bin/publish.cgi?p=183996&s=pets_nature |first=Joe |last=Belanger |title=DNA tests reveal cougars roam region |date=May 25, 2007 |publisher=[[London Free Press]] |accessdate=June 5, 2007 |deadurl=yes |archiveurl=https://web.archive.org/web/20130921061306/http://pbdba.lfpress.com/cgi-bin/publish.cgi?p=183996&s=pets_nature |archivedate=September 21, 2013 }}</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=May 20, 2007 |author=Board of Directors |year=2004 |publisher=The Cougar Network |deadurl=yes |archiveurl=https://web.archive.org/web/20141120173547/http://www.cougarnet.org/bigpicture.html |archivedate=November 20, 2014 }} ''The Cougar Network methodology is recognized by the [http://www.fws.gov/northeast/ECougar/ U.S. Fish and Wildlife Service].''</ref> The [[Quebec]] wildlife services (known locally as MRNF) also considers cougar to be present in the province as a threatened species after multiple DNA tests confirmed cougar hair in lynx mating sites.<ref>{{cite web|url=http://www.mrnf.gouv.qc.ca/faune/especes/menacees/faire.jsp |title=Your part in helping endangered species |accessdate=January 7, 2010 |year=2010 |publisher=Ministry of Wildlife and Natural Resources, Quebec, Canada |deadurl=yes |archiveurl=https://web.archive.org/web/20110203082852/http://www.mrnf.gouv.qc.ca/faune/especes/menacees/faire.jsp |archivedate=February 3, 2011 }}</ref> The only unequivocally known eastern population is the [[Florida panther]], which is critically endangered. There have been unconfirmed sightings in [[Elliotsville Plantation, Maine]] (north of [[Monson, Maine|Monson]]); and in [[New Hampshire]], there have been unconfirmed 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 =March 20, 2009 | publisher=Beech River Books|author=Davidson, Rick|year=2009}}</ref> In 2009, the [[Michigan]] Department of Natural Resources confirmed a cougar sighting in Michigan's [[Upper Peninsula]].<ref>{{cite web|url=http://www.mlive.com/outdoors/index.ssf/2009/11/photo_shows_cougar_presence_in.html |title=Photo shows cougar presence in Michigan|author=Skinner, Victor|work= The Grand Rapids Press|date= November 15, 2009 |accessdate=February 16, 2011}}</ref> Typically, extreme-range sightings of cougars involve young males, which can travel great distances to establish ranges away from established males; all four confirmed cougar kills in [[Iowa]] since 2000 involved males.<ref>{{cite news|url=http://gazetteonline.com/breaking-news/2009/12/15/cedar-rapids-man-shoots-mountain-lion-in-iowa-county |title=Cedar Rapids man shoots mountain lion in Iowa County |date=December 15, 2009 |work=Cedar Rapids Gazette |accessdate=December 16, 2009 |deadurl=yes |archiveurl=https://web.archive.org/web/20100323181821/http://gazetteonline.com/breaking-news/2009/12/15/cedar-rapids-man-shoots-mountain-lion-in-iowa-county |archivedate=March 23, 2010 }}</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]] of [[South Dakota]]. Less than a year later, on March{{nbsp}}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 news|author=Carlson, James A. |url=http://seattletimes.nwsource.com/html/nationworld/2009099370_apuslivingwithlions.html |title=Sightings show cougars expanding into central US |accessdate=April 22, 2009 |date=April 22, 2009 |publisher=Seattle Times |deadurl=yes |archiveurl=https://web.archive.org/web/20120703095930/http://seattletimes.nwsource.com/html/nationworld/2009099370_apuslivingwithlions.html |archivedate=July 3, 2012 }}</ref> Other eastern sightings since 2010 have occurred in locations such as [[Greene County, Indiana]],<ref>{{cite web | url = http://www.in.gov/portal/news_events/53876.htm | title = Mountain Lion Confirmed in Rural Greene County | date =May 7, 2010 | publisher=Indiana Department of Natural Resources}}</ref> [[Greenwich, Connecticut|Greenwich]]<ref name="CT Greenwich lion">{{cite news|title=Mountain lion reportedly spotted roaming Connecticut town|url=http://www.foxnews.com/us/2011/06/10/mountain-lion-reportedly-spotted-roaming-connecticut-town/|publisher=Fox News|accessdate=June 12, 2011|date=June 10, 2011}}</ref> and [[Milford, Connecticut]],<ref>{{cite news|url=http://www.cnn.com/2011/US/06/11/connecticut.mountain.lion/index.html|title=Mountain Lion killed by car on Connecticut highway|publisher=CNN |date=June 11, 2011}}</ref> [[Morgan County, Illinois|Morgan County]]<ref>{{cite news|url=http://www.sj-r.com/top-stories/x1272748311/Cougar-photographed-in-Morgan-County|archiveurl=https://web.archive.org/web/20131102184210/http://www.sj-r.com/top-stories/x1272748311/Cougar-photographed-in-Morgan-County|archivedate=November 2, 2013|title=Cougar photographed in Morgan County|publisher=The State Journal-Register |date=October 29, 2012|accessdate=October 31, 2012}}</ref> [[Pike County, Illinois|Pike County]],<ref>{{cite news| url=http://www.prairiestateoutdoors.com/index.php?/pso/article/jurys_still_out_but_pike_county_cougar_sighting_could_be_states_third_in_tw/|title=Jury's still out, but Pike County cougar sighting could be state's third in two months|publisher=The State Journal-Register |date=November 11, 2012|accessdate=November 13, 2012}}</ref> and [[Whiteside County, Illinois]],<ref>{{cite news| url=https://www.facebook.com/photo.php?fbid=767940479898919&set=a.213944418631864.65384.211596162200023&type=1&theater|title=Timeline Photos|work=Facebook|date=November 21, 2013|accessdate=November 21, 2013}}</ref> and [[Bourbon County, Kentucky]].<ref>{{cite web |url= http://www.usatoday.com/story/news/nation-now/2014/12/17/puma-killed-in-kentucky/20536509/|title= First cougar seen in Kentucky since Civil War is killed|author=<!--Staff writer(s); no by-line.--> |date= December 17, 2014|website= |publisher= |access-date=}}</ref> South of the [[Rio Grande]], the [[International Union for Conservation of Nature]] (IUCN) lists the cat in every [[Central America|Central]] and [[South American]] country.<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 | archiveurl = https://web.archive.org/web/20070221182221/http://www.nwf.org/cats/pdfs/cougarfacts.pdf | archivedate = February 21, 2007 | format = PDF | title = Cougar facts | accessdate =May 20, 2007 | 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"/> US 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=May 20, 2007 |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=May 20, 2007 |year=2004 |publisher=California Department of Fish and Game| archiveurl = https://web.archive.org/web/20070430012010/http://www.dfg.ca.gov/news/issues/lion.html| archivedate = April 30, 2007}}</ref> In 2012 research in [[Río Los Cipreses National Reserve]], Chile, based in 18 motion-sensitive cameras counted a population of two males and two females, one of them with at least two cubs, in an area of 600&nbsp;km<sup>2</sup>, that is 0.63 cougars every 100&nbsp;km<sup>2</sup>.<ref>Research of Nicolás Guarda, supported by [[National Forest Corporation (Chile)|Conaf]], [[Pontifical Catholic University of Chile]], and a private enterprise. See article in Chilean newspaper [[La Tercera]], [http://www.latercera.com/noticia/tendencias/2013/01/659-506141-9-investigacion-midio-por-primera-vez-poblacion-de-pumas-en-zona-central.shtml Investigación midió por primera vez población de pumas en zona central], retrieved on January 28, 2013, in Spanish Language.</ref> === Habitat fragmentation === With the increase of human development and infrastructure growth in California, the North American Cougar populations are becoming more isolated from one another.<ref>{{Cite journal | last = Ernest | first = Holly B. | last2 = Vickers | first2 = T. Winston | last3 = Morrison | first3 = Scott A. | last4 = Buchalski | first4 = Michael R. | last5 = Boyce | first5 = Walter M. | title = Fractured Genetic Connectivity Threatens a Southern California Puma (Puma concolor) Population | url = http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0107985 | journal = PLoS ONE | volume = 9 | issue = 10 | pages = e107985 | doi = 10.1371/journal.pone.0107985 | pmc = 4189954 | pmid = 25295530 | year = 2014 }}</ref> ===Ecological role=== [[File:Cougar track.jpg|thumb|Front paw print of a cougar. An adult paw print is approximately 10&nbsp;cm (4&nbsp;inches) long.<ref name="GovBC"/>]] [[File:Feeling Unwelcome.jpg|thumb|Juvenile cougars conflict with coyotes]] Aside from humans, no species preys upon mature cougars in the wild, although conflicts with other predators or scavengers occur. The [[Yellowstone National Park]] ecosystem provides a fruitful microcosm to study inter-predator interaction in North America. Of the three large predators, the massive [[grizzly 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 grizzlies and [[American black bear]]s visited 24% of cougar kills in Yellowstone and [[Glacier National Park (U.S.)|Glacier National Parks]], usurping 10% of carcasses. Bears gained up to 113%, and cougars lost up to 26%, of their respective daily energy requirements from these encounters.<ref>{{Cite journal |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=April 8, 2007 }}</ref> In Colorado and California, black bears were found to visit 48% and 77% of kills respectively. In general, cougars are subordinate to black bears when it comes to kills and when bears are most active, the cats take prey more frequently and spend less time feeding on each kill. In addition, unlike several subordinate predators from other ecosystems, cougars do not appear to take advantage of spatial or temporal refuges to avoid their competitors.<ref>{{cite journal|author=Elbroch, L. M.|author2=Lendrum, P. E.|author3=Allen, M. L.|author4=Wittmer, H. U.|year=2014|title=Nowhere to hide: pumas, black bears, and competition refuges|journal=Behavioral Ecology|doi=10.1093/beheco/aru189|volume=26|pages=247–254}}</ref><ref>{{cite web|author=ELboch, M.|date=November 1, 2014|title=Mountain Lions Versus Black Bears|publisher=National Geographic|accessdate=September 24, 2016|url=http://voices.nationalgeographic.com/2014/11/01/mountain-lions-versus-black-bears/}}</ref> The gray wolf and the cougar compete more directly for prey, especially in winter. Wolves can steal kills and occasionally kill the cat. One report describes a large pack of 7 to 11 wolves killing a female cougar and her kittens.<ref name="Park wolf pack kills mother cougar">{{cite web|title=Park wolf pack kills mother cougar|url=http://www.forwolves.org/ralph/wolves-deadcougar.htm|publisher=forwolves.org|accessdate=April 12, 2013}}</ref> Conversely, lone female or young wolves are vulnerable to predation, and have been reported ambushed and killed by cougars.<ref>{{cite news |url=http://forests.org/shared/reader/welcome.aspx?linkid=22714&keybold=wildlife%20AND%20%20cougar |title=In Yellowstone, it's Carnivore Competition |last=Gugliotta |first=Guy |date=May 19, 2003 |work=Washington Post |accessdate=April 9, 2007}}</ref> Various accounts of cougars killing lone wolves, including a six-year-old female, have also been documented.<ref name="Wolf B4 Killed by Mountain Lion?">{{cite web|title=Wolf B4 Killed by Mountain Lion?|url=http://www.forwolves.org/ralph/32596.html|publisher=forwolves.org|date=March 25, 1996}}</ref><ref name="Autopsy Indicates Cougar Killed Wolf">{{cite web|title=Autopsy Indicates Cougar Killed Wolf|url=http://www.igorilla.com/gorilla/animal/1999/cougar_kills_wolf.html|publisher=igorilla.com|date=April–May 2000|accessdate=May 22, 2015}}</ref><ref name="Mountain lions kill collared wolves in Bitterroot">{{cite web|title=Mountain lions kill collared wolves in Bitteroot |url=http://missoulian.com/news/state-and-regional/mountain-lions-kill-collared-wolves-in-bitterroot/article_68c0c60c-d792-59e3-b736-5b10c17eb10a.html |publisher=missoulian.com |accessdate=May 29, 2012 |deadurl=yes |archiveurl=https://web.archive.org/web/20130517113554/http://missoulian.com/news/state-and-regional/mountain-lions-kill-collared-wolves-in-bitterroot/article_68c0c60c-d792-59e3-b736-5b10c17eb10a.html |archivedate=May 17, 2013 }}</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=April 9, 2007 |publisher=Greater Yellowstone Learning Center| archiveurl = https://web.archive.org/web/20070929115515/http://www.greateryellowstonescience.org/topic/wolves/wolvesoverview.html| archivedate = September 29, 2007}}</ref> In nearby [[Sun Valley, Idaho]], a cougar/wolf encounter that resulted in the death of the cougar was documented.<ref name="IME">{{cite web|title=Predators clash above Elkhorn|url=http://www.mtexpress.com/index2.php?ID=2005125077|publisher=Idaho Mountain Express|accessdate=August 21, 2013}}</ref> One researcher in Oregon noted: "When there is a pack around, cougars are not comfortable around their kills or raising kittens&nbsp;... 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=October 29, 2006 |publisher=[[The Oregonian]] |accessdate=April 9, 2007 |deadurl=yes |archiveurl=https://web.archive.org/web/20071213022857/http://www.oregonwild.org/press-room/press-clips/turf-wars-in-idaho-s-wilderness |archivedate=December 13, 2007 }}</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"/> Although cougars can kill coyotes, the latter have been documented attempting to prey on cougar cubs.<ref name="Cougars vs. coyotes photos draw Internet crowd">{{cite web|title=Cougars vs. coyotes photos draw Internet crowd|url=http://missoulian.com/news/state-and-regional/cougars-vs-coyotes-photos-draw-internet-crowd/article_f6ecdb1a-a051-11e2-85a3-001a4bcf887a.html|publisher=missoulian.com|accessdate=April 8, 2013}}</ref> 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 | archiveurl = https://web.archive.org/web/20060716064117/http://www.ecology.info/ecology-jaguar-puma.htm| archivedate = July 16, 2006}}</ref> The jaguar tends to take larger prey and the cougar smaller where they overlap, reducing the cougar's size and also further reducing the likelihood of direct competition.<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 |author1=Nuanaez, Rodrigo|author2= Miller, Brian |author3=Lindzey, Fred |year=2000 |title=Food habits of jaguars and pumas in Jalisco, Mexico |journal=Journal of Zoology |volume=252 |issue=3 |pages=373–379 |doi=10.1111/j.1469-7998.2000.tb00632.x }}</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. |author2=Wielgus, Robert B. |author3=Gwilliam, John C. |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 }}</ref> The [[Vancouver Island marmot]], an endangered species [[endemic]] to one region of dense cougar population, has seen decreased numbers due to cougar and gray wolf predation.<ref>{{cite journal |last=Bryant |first=Andrew A. |author2=Page, Rick E. |date=May 2005 |title=Timing and causes of mortality in the endangered Vancouver Island marmot (Marmota vancouverensis) |journal=Canadian Journal of Zoology |volume=83 |issue=5 |pages=674–82 |doi=10.1139/z05-055}}</ref> Nevertheless, there is a measurable effect on the quality of deer populations by puma predation.<ref>Fountain, Henry (November 16, 2009) [http://www.nytimes.com/2009/11/17/science/17obprey.html "Observatory: When Mountain Lions Hunt, They Prey on the Weak"]. ''[[New York Times]]''.</ref><ref>{{cite journal|doi=10.1046/j.1523-1739.1996.10040964.x|title=Resilience and Conservation of Large Carnivores in the Rocky Mountains|year=1996|last1=Weaver|first1=John L.|last2=Paquet|first2=Paul C.|last3=Ruggiero|first3=Leonard F.|journal=Conservation Biology|volume=10|issue=4|pages=964–976}}</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.<ref>Busch, Robert H. ''The Cougar Almanac.'' New York, 2000, pg 94. ISBN 1592282954.</ref> ==Hybrids== [[File:Pumapard-1904.jpg|thumb|upright|Pumapard, photographed in 1904]] {{Main article|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=June 9, 2008|archiveurl=https://web.archive.org/web/20071015110856/http://www.geocities.com/pride_lands/Liger_Tigon.html|archivedate=October 15, 2007}}</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&nbsp;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.php|title=Appendices I, II and III |accessdate=May 24, 2007 |publisher=[[Convention on International Trade in Endangered Species of Wild Fauna and Flora]]}}</ref> rendering illegal international trade in specimens or parts. [[File:Cougar snow.jpg|thumb|left|Cougar conservation depends on preservation of its habitat]] In the United States east of the [[Mississippi River]], the only unequivocally known cougar population is the [[Florida panther]]. Until 2011, the [[United States Fish and Wildlife Service]] (USFWS) recognized both an [[Eastern cougar]] (claimed to be a subspecies by some, denied by others)<ref>{{cite book |last1=Bolgiano |first1=Chris |title=Mountain Lion:An Unnatural History of Pumas and People |place=[[Mechanicsburg, PA]] |publisher=Stackpole Books |date=August 1995 |edition=Hardcover |isbn=978-0-8117-1044-2}}</ref><ref>{{cite book |last1=Eberhart |first1=George M. |title=Mysterious Creatures: A Guide to Cryptozoology |url=http://www.ebook3000.com/animals/Mysterious-Creatures--A-Guide-to-Cryptozoology_64383.html |volume=Volume 2 |location=Santa Barbara, CA |publisher=ABC-CLIO |year=2002 |pages= 153–161 |isbn=1-57607-283-5 }}</ref> 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=May 20, 2007 | work=Endangered and Threatened Species of the Southeastern United States (The Red Book)| year=1991 |publisher=[[U.S. Fish and Wildlife Service]]| archiveurl = https://web.archive.org/web/20070403000340/http://www.fws.gov/endangered/i/a/saa48.html| archivedate = April 3, 2007}}</ref><ref>{{cite web |url=http://www.fws.gov/endangered/i/a/saa05.html |title=Florida Panther |accessdate=June 7, 2007 | work=Endangered and Threatened Species of the Southeastern United States (The Red Book)| year=1993 |publisher=[[United States Fish and Wildlife Service]] |archiveurl = https://web.archive.org/web/20070604163840/http://www.fws.gov/endangered/i/a/saa05.html|archivedate = June 4, 2007}}</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/> In 2003 the documented count for the Florida sub-population was 87 individuals.<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=June 5, 2007| archiveurl = https://web.archive.org/web/20070616200443/http://myfwc.com/panther/news/pdf/FWC2002-2003PantherGeneticRestorationAnnualReport.pdf| archivedate = June 16, 2007}}</ref> In March 2011, the USFWS declared the Eastern cougar extinct. With the taxonomic uncertainty about its existence as a subspecies as well as the possibility of eastward migration of cougars from the western range, the subject remains open.<ref name="barringer">{{cite news |last1=Barringer |first1=Felicity |url=http://www.nytimes.com/2011/03/01/us/03cougar.html?hpw |title=U.S. Declares Eastern Cougar Extinct, With an Asterisk |work=The New York Times |accessdate=March 2, 2011 |date=March 2, 2011}}</ref> This uncertainty has been recognized by Canadian authorities. The Canadian federal agency called [[Committee on the Status of Endangered Wildlife in Canada]] rates its current data as "insufficient" to draw conclusions regarding the eastern cougar's survival, and says on its Web site "Despite many sightings in the past two decades from eastern Canada, there are insufficient data to evaluate the taxonomy or assign a status to this cougar." Notwithstanding numerous reported sightings in [[Ontario]], Quebec, [[New Brunswick]] and [[Nova Scotia]], it has been said that the evidence is inconclusive: ". . . there may not be a distinct 'eastern' subspecies, and some sightings may be of escaped pets."<ref>[http://www.cosewic.gc.ca/eng/sct1/searchdetail_e.cfm?id=132 Committee on Status of Endangered Wildlife in Canada]. Cosewic.gc.ca. Retrieved on September 15, 2011.</ref><ref>[http://www.naturecanada.ca/endangered_know_our_species_ecougar.asp Eastern Cougar, Nature Canada]. Naturecanada.ca. Retrieved on September 15, 2011.</ref> The cougar is also protected across much of the rest of its 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]]. 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. Texas is the only state in the United States with a viable population of cougars that does not protect that population in some way. In Texas, cougars are listed as nuisance wildlife and any person holding a hunting or a trapping permit can kill a cougar regardless of the season, number killed, sex or age of the animal.<ref>[http://www.tpwd.state.tx.us/huntwild/wild/nuisance/mountain_lion/ TPWD: Mountain Lions]. Tpwd.state.tx.us (July 16, 2007). Retrieved on September 15, 2011.</ref> Killed animals are not required to be reported to [[Texas Parks and Wildlife Department]]. Conservation work in Texas is the effort of a non-profit organization, [[Balanced Ecology Inc. (BEI)]], as part of their Texas Mountain Lion Conservation Project.<ref>[http://balancedecology.org/MountainLionWebSite/Mountain_Lion_Conservation_Project.html Texas Mountain Lion Conservation Project]. Balanced Ecology Inc. balancedecology.org</ref> 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 without a permit in California except under very specific circumstances, such as when a cougar is in act of pursuing livestock or domestic animals, or is declared a threat to public safety.<ref name="California"/> Permits are issued when owners can prove property damage on their livestock or pets. For example, multiple dogs have been attacked and killed, sometimes while with the owner. Many attribute this to the protection cougars have from being hunted and are now becoming desensitized to humans; most are removed from the population after the attacks have already occurred. 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 the 1970s with an average of over 112 cats killed per year from 2000 to 2006 compared to six per year in the 1970s. They also state on their website that there is a healthy number of cougars in California. 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/bapp.html |archiveurl=https://web.archive.org/web/20100323040005/http://www.felidaefund.org/research/bapp.html |archivedate=March 23, 2010 |title=Bay Area Puma Project (BAPP)|publisher=Felidae Conservation Fund |accessdate=March 8, 2009}}</ref> Conservation threats to the species include persecution as a pest animal, [[environmental degradation]] and [[habitat fragmentation]], and depletion of their prey base. [[Wildlife corridor]]s 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&nbsp;km<sup>2</sup> (850&nbsp;sq&nbsp;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 |date=March 1993 |title=Determining Minimum Habitat Areas and Habitat Corridors for Cougars |journal=Conservation Biology|volume=7|issue=1|pages=94–108|doi=10.1046/j.1523-1739.1993.07010094.x |jstor=2386646}}</ref> On March 2, 2011, the [[United States Fish and Wildlife Service]] declared the Eastern cougar (''Puma concolor couguar'') officially extinct.<ref>[http://www.fws.gov/northeast/ECougar/newsreleasefinal.html Northeast Region, U.S. Fish and Wildlife Service]. Fws.gov. Retrieved on September 15, 2011.</ref> ==Relationships with humans== ===In mythology=== The grace and power of the cougar have been widely admired in the cultures of the [[indigenous peoples of the Americas]]. The Inca city of [[Cusco]] is reported to have been designed in the shape of a cougar, and the animal also gave its name to both Inca regions and people. The [[Moche (culture)|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 |others=Kait Realo (translator) |title=On the role of Creation and Origin Myths in the Development of Inca State and Religion |accessdate=May 22, 2007 |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 [[Hocąk language]] ("Ho-Chunk" or "Winnebago") of [[Wisconsin]] and [[Illinois]]<ref>[http://hotcakencyclopedia.com/ho.Cougars.html '' Cougars,''] The Encyclopedia of Hočąk (Winnebago) Mythology. Retrieved: 2009/12/08.</ref> 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=April 11, 2009 |format=PDF |publisher=[[United States Department of Agriculture|USDA]] [[Wildlife Services]]}}</ref> The Algonquins and Ojibwe believe that the cougar lived in the underworld and was wicked, whereas it was a sacred animal among the Cherokee.<ref>{{cite book|title=The Element Encyclopedia of Magical Creatures |author1=Matthews, John |author2=Matthews, Caitlín |year=2005 |publisher=HarperElement |isbn=978-1-4351-1086-1 |page=364 }}</ref> ===Livestock predation=== During the early years of ranching, cougars were considered on par with wolves in destructiveness. According to figures in Texas in 1990, 86 calves (0.0006% of a total of 13.4&nbsp;million cattle & calves in Texas), 253 Mohair goats, 302 Mohair kids, 445 sheep (0.02% of a total of 2.0&nbsp;million sheep & lambs in Texas) and 562 lambs (0.04% of 1.2&nbsp;million lambs in Texas) were confirmed to have been killed by cougars that year.<ref name="NASS-cattle">{{cite web | url=http://usda.mannlib.cornell.edu/usda/nass/Catt//1990s/1990/Catt-07-27-1990.pdf | title = Cattle report 1990 |publisher = National Agricultural Statistics Service| accessdate=September 11, 2009}}</ref><ref name="NASS-sheep-goats">{{cite web | url=http://usda.mannlib.cornell.edu/usda/nass/SheeGoat//1990s/1990/SheeGoat-02-02-1990.pdf | title = Sheep and Goats report 1990| publisher=National Agricultural Statistics Service |accessdate =September 11, 2009}}</ref> In Nevada in 1992, cougars were confirmed to have killed 9 calves, 1 horse, 4 foals, 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 =July 10, 2008}}</ref> A cougar's killing bite is applied to the back of the neck, head, or throat and they inflict puncture marks with their claws usually seen on the sides and underside of the prey, sometimes also shredding the prey as they hold on. Coyotes also typically bite the throat region but do not inflict the claw marks and farmers will normally see the signature zig-zag pattern that coyotes create as they feed on the prey whereas cougars typically drag in a straight line. The work of a cougar is generally clean, differing greatly from the indiscriminate mutilation by coyotes and 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://agrilife.org/texnatwildlife/predators-and-predation/predator-species/cougars/| title = Cougar Predation – Description | publisher=Procedures for Evaluating Predation on Livestock and Wildlife | accessdate =August 3, 2008}}</ref> Remedial hunting appears to have the paradoxical effect of increased livestock predation and complaints of human-puma conflicts. In a 2013 study the most important predictor of puma problems were remedial hunting of puma the previous year. Each additional puma on the landscape increased predation and human-puma complaints by 5% but each additional animal killed on the landscape the previous year increased complaints by 50%, an order of magnitude higher. The effect had a dose-response relationship with very heavy (100% removal of adult puma) remedial hunting leading to a 150% – 340% increase in livestock and human conflicts.<ref>{{cite journal |title=Effects of Remedial Sport Hunting on Cougar Complaints and Livestock Depredations |author1=Peebles, Kaylie A.|author2= Wielgus, Robert B.|author3= Maletzke, Benjamin T.|author4= Swanson, Mark E. |journal=PLoS ONE |date=November 2013 |volume=8 |issue=11 |pages=e79713|doi=10.1371/journal.pone.0079713|pmid=24260291 |pmc=3834330}}</ref> This effect is attributed to the fact that inexperienced younger male pumas are most likely to approach human developments, whereas remedial hunting removes older pumas who have learned to avoid people in their established territories. Remedial hunting enables younger males to enter the former territories of the older animals.<ref>{{cite journal |title=Cougar attacks on humans in the United States and Canada |author=Beier, Paul |journal=Wildlife Society Bulletin |volume=19 |pages=403–412 |year=1991 |jstor=3782149 }}</ref><ref>{{cite journal |title=Mountain lion and human activity in California: testing speculations |author1=Torres SG|author2= Mansfield TM|author3= Foley JE|author4= Lupo T|author5= Brinkhaus A |year=1996 |journal=Wildlife Society Bulletin |volume=24 |issue=3 |pages=451–460 |jstor=3783326 }}</ref> ===Attacks on humans=== ====North American subspecies==== {{See also|List of fatal cougar attacks in North America}} [[File:MountainLionAttackProtocol.jpg|thumb|right|Mountain Lion warning sign]] The pertinent North American subspecies is ''P. concolor couguar''. Due to the [[Human overpopulation|expanding human population]], cougar [[range (biology)|range]]s increasingly overlap with areas inhabited by [[humans]]. Attacks on humans are very rare, as cougar prey recognition is a learned behavior and they do not generally recognize humans as prey.<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|pmid=14518628 |doi=10.1580/1080-6032(2003)14[169:CAOHAC]2.0.CO;2}}</ref> Attacks on people, livestock, and pets may occur when a puma [[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=May 20, 2007 |publisher=Northern Arizon University |deadurl=yes |archiveurl=https://web.archive.org/web/20120622033418/http://users.frii.com/mytymyk/lions/beier.htm |archivedate=June 22, 2012 }}</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=May 20, 2007 |publisher=Arizona Game and Fish Department}}</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>[[New Mexico Department of Game and Fish]]: [https://web.archive.org/web/20120620114802/http://www.wildlife.state.nm.us/publications/press_releases/documents/2008/062308pinosaltoslion.html Search continues for mountain lion that killed Pinos Altos man], June 23, 2008; [https://web.archive.org/web/20120620114802/http://www.wildlife.state.nm.us/publications/press_releases/documents/2008/062508pinosaltoslion.html Wounded mountain lion captured, killed near Pinos Altos], June 25, 2008; [https://web.archive.org/web/20120620114802/http://www.wildlife.state.nm.us/publications/press_releases/documents/2008/070108pinosaltoslion2.html Second mountain lion captured near Pinos Altos], 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]]". Standing still may cause the cougar to consider a person easy prey.<ref>{{cite web|last=Subramanian|first=Sushma|title=Should You Run or Freeze When You See a Mountain Lion?|url=http://www.scientificamerican.com/article.cfm?id=should-you-run-or-freeze-when-you-see-a-mountain-lion|publisher=Scientific American|accessdate=March 10, 2012|date=April 14, 2009}}</ref> Exaggerating the threat to the animal through intense eye contact, loud shouting, and any other action to appear larger and more menacing, may make the animal retreat. Fighting back with sticks and rocks, or even bare hands, is often effective in persuading an attacking cougar to disengage.<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=May 28, 2007 |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 – and almost all fatalities – were children. The same study showed the highest proportion of attacks to have occurred in [[British Columbia]], particularly on [[Vancouver Island]] where cougar populations are especially dense.<ref name="Beier" /> Preceding attacks on humans, cougars display aberrant behavior, such as activity during daylight hours, a lack of fear of humans, and stalking humans. There have sometimes been incidents of pet cougars mauling people.<ref>{{cite news|url=http://www.msnbc.msn.com/id/27756765/ns/us_news-life/t/neighbor-saves-miami-teen-cougar/|agency=Associated Press|publisher=MSNBC|date=November 16, 2008|title=Neighbor saves Miami teen from cougar|accessdate=February 11, 2012}}</ref><ref>{{Cite journal|title=2-Year-Old Boy Hurt In Pet Cougar Attack|url=http://www.nytimes.com/1995/06/04/nyregion/2-year-old-boy-hurt-in-pet-cougar-attack.html|date=June 4, 1995|work=New York Times}}</ref> Research on new wildlife collars may be able to reduce human-animal conflicts by predicting when and where predatory animals hunt. This can not only save human lives and the lives of their pets and livestock but also save these large predatory mammals that are important to the balance of ecosystems.<ref>Williams, Terrie M. (November 6, 2014) [http://www.latimes.com/opinion/op-ed/la-oe-williams-wildlife-coburn-wastebook-20141107-story.html "As species decline, so does research funding"] ''[[Los Angeles Times]]''</ref> ====Other subspecies==== The puma of the plains of South America (''P. concolor cabrerae'',<ref>Wozencraft, W. Christopher (2005) [http://www.vertebrates.si.edu/msw/mswcfapp/msw/taxon_browser.cfm?msw_id=10843 "''Puma concolor cabrerae''"] in Don E. Wilson, DeeAnn M. Reeder (eds.) ''Mammal Species of the World'', 3rd ed. JHU Press. ISBN 0801882214.</ref> called the [[Argentine cougar]] by North Americans) is noted for its extreme unwillingness to attack humans. According to the Anglo-Argentine naturalist [[William Henry Hudson]]<ref name="gutenberg.org">Hudson, W. H. (1892) [http://www.gutenberg.org/files/7446/7446-h/7446-h.htm ''The Naturalist In La Plata''], chapter 2.</ref> "It does not attack man, and Azara is perfectly correct when he affirms that it never hurts, or threatens to hurt, man or child, even when it finds them sleeping. This, however, is not a full statement of the facts; the puma will not even defend itself against man". And: "All who have killed or witnessed the killing of the puma—and I have questioned scores of hunters on this point—agree that it resigns itself in this unresisting, pathetic manner to death at the hands of man. Claudio Gay, in his Natural History of Chili, says, "When attacked by man its energy and daring at once forsake it, and it becomes a weak, inoffensive animal, and trembling, and uttering piteous moans, and shedding abundant tears, it seems to implore compassion from a generous enemy". Hudson adds that, except in relation to man, the puma is a ferocious animal, attacking even the much stronger jaguar.<ref name="gutenberg.org"/> ==See also== {{Portal bar|Cats|Mammals}} ==References== {{Reflist|30em}} ==Further reading== {{Refbegin|30em}} * {{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 |url=https://books.google.com/books?id=8WRnVkgi0z4C&lpg=PP1|isbn=0-393-05807-7}} * {{cite book |last1=Bolgiano |first1=Chris |title=Mountain Lion:An Unnatural History of Pumas and People |place=[[Mechanicsburg, PA]] |publisher=Stackpole Books |year=2001 |edition=Paperback |url=http://www.washingtonpost.com/wp-srv/style/longterm/books/chap1/mountain.htm|isbn=0-8117-2867-6}} * {{cite book |last1=Eberhart |first1=George M. |title=Mysterious Creatures: A Guide to Cryptozoology |url=http://www.ebook3000.com/animals/Mysterious-Creatures--A-Guide-to-Cryptozoology_64383.html |volume=Volume 2 |location=Santa Barbara, CA |publisher=ABC-CLIO |year=2002 |pages= 153–161 |isbn=1-57607-283-5 }} * {{cite book |editor1-last=Hornocker |editor1-first=Maurice |editor2-last=Negri |editor2-first=Sharon |year=2010 |editor3-first=Fred |editor3-last=Lindzey|title=Cougar: Ecology and Conservation |edition=Hardcover |location=Chicago |publisher=University of Chicago Press |url=https://books.google.com/books?id=ZSDUxT3fSYEC&lpg=PP1 |isbn=978-0-226-35344-9}} * {{cite book |last1=Kobalenko |first1=Jerry |year=2005 |title=Forest Cats of North America |location=Hove |publisher=Firefly Books Ltd. |isbn=1-55209-172-4}} * {{cite journal|first1=Todd |last1=Lester |title=Search for Cougars in the East North America |format=PDF |work=North American BioFortean Review. |pages=15–17 |url=http://www.strangeark.com/nabr/NABR7.pdf |volume=3 |issue=7 |publisher=Zoological Miscellania website |date=October 2001}} * {{cite book |last=Logan |first=Ken |author2=Linda Sweanor |title=Desert Puma: Evolutionary Ecology and Conservation of an Enduring Carnivore |year=2001 |url=https://books.google.com/books?id=04joHfHvpsEC&lpg=PP1&dq=Desert%20Puma%3A%20Evolutionary%20Ecology%20and%20Conservation%20of%20an%20Enduring%20Carnivore&pg=PP1#v=onepage&q&f=true|publisher=Island Press |isbn=1-55963-866-4}} * {{cite web |url=http://www.mountainlion.org/publications.asp |title=Publications |publisher=Mountain Lion Foundation}} * {{cite book |first1=Gerry |last1=Parker |year=1994 |title=The Eastern Panther – Mystery Cat of the Appalachians |publisher=Nimbus Publishing (CN) |edition=Softcover |isbn=1-55109-268-9}} * {{cite book |last=Wright |first=Bruce S |title=The Eastern Panther: A Question of Survival |location=Toronto |publisher=Clark, Irwin, and Company|year= 1972 |isbn=0772005281}} * {{cite web |url=http://www.easterncougar.org/pages/bibliography.html |title=Annotated Bibliography |publisher=easterncougar.org – Cougar Rewilding Foundation}} {{Refend}} ==External links== {{Sister project links | wikt=Cougar | commons=Puma concolor | b=no | n=no | q=no | s=no | v=no | voy=no | species=Puma concolor | d=no | mw=no }} {{NIE Poster|year=1905|Puma}} {{Spoken Wikipedia|Cougar1.ogg|November 6, 2008}} * [http://www.catsg.org/index.php?id=94 Species portrait ''Puma concolor''; IUCN/SSC Cat Specialist Group] * [http://northernbushcraft.com/animalTracks/cougar/notes.htm Cougar Tracks]: How to identify cougar tracks in the wild * [http://animals.nationalgeographic.com/animals/mammals/mountain-lion/?rptregcta=reg_free_np&rptregcampaign=20130924_rw_membership_r3p_c1#close-modal Puma sounds] (they growl, hiss and scream but cannot roar like true lions of the genus ''[[Panthera]]'') at [[National Geographic Society]] * [http://santacruzpumas.org/ Santa Cruz Puma Project] * [http://www.eprn.homestead.com/ Eastern Puma Research Network] * [http://felidaefund.org/ Felidae Conservation Fund] * [http://www.easterncougar.org/ Cougar Rewilding Foundation, formerly "Eastern Cougar Foundation"] * [http://www.cougarnet.org/ The Cougar Network --Using Science to Understand Cougar Ecology] * [http://www.mountainlion.org/about_the_foundation.asp Mountain Lion Foundation – Saving America's Lion] * [http://www.savethecougar.org/ SaveTheCougar.org]: Sightings of cougars in Michigan * [http://www.amazonarium.com.br/docs/peopleandjaguarcomplete.pdf People and Cougar/Jaguars A Guide for Coexistence] * [http://www.cougarfund.org/ The Cougar Fund – Protecting America's Greatest Cat.] A Definitive Resource About Cougars] Comprehensive, non-profit [[501(c)(3)]] 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, and rare photos and videos of wild cougars. * [http://sagehen.ucnrs.org/Documents/visitors/wildlife/lion.pdf Living with California Mountain Lions] {{Carnivora|Fe.}} {{North American Game}} {{Featured article}} {{taxonbar}} {{Authority control}} [[Category:Puma (genus)]] [[Category:Felids of Central America]] [[Category:Felids of North America]] [[Category:Felids of South America]] [[Category:Fauna of the California chaparral and woodlands]] [[Category:Fauna of the Sierra Nevada (U.S.)]] [[Category:Mammals of North America]] [[Category:Mammals of South America]] [[Category:Megafauna of North America]] [[Category:Megafauna of South America]] [[Category:Mammals of Argentina]] [[Category:Mammals of Bolivia]] [[Category:Mammals of Brazil]] [[Category:Mammals of Canada]] [[Category:Mammals of Chile]] [[Category:Mammals of Colombia]] [[Category:Mammals of Guatemala]] [[Category:Mammals of Guyana]] [[Category:Mammals of Peru]] [[Category:Animals described in 1771]] [[Category:Taxa named by Carl Linnaeus]] [[Category:Pleistocene carnivorans]] [[Category:Quaternary carnivorans]] [[Category:Pleistocene mammals of North America]] [[Category:Pleistocene mammals of South America]] [[Category:Extant Middle Pleistocene first appearances]]'
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