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{{Short description|Large species of cat native to the Americas}} | |||
{{Redirect|Mountain lion||Cougar (disambiguation)|and|Mountain lion (disambiguation)}} | |||
{{ |
{{Otheruses}} | ||
{{Redirect|Mountain lion|other uses|Mountain lion (disambiguation)}} | |||
{{Distinguish|Lion}} | |||
{{Use mdy dates|date=January 2015}} | |||
{{Redirect|Catamount}} | |||
{{speciesbox | |||
{{Featured article}} | |||
| name = Cougar<ref name=MSW3/> | |||
{{Pp-semi-indef|small=yes}} | |||
| fossil_range = {{Fossil range|0.3|0}}<small>Middle ]–]</small> | |||
{{Use mdy dates|date=May 2024}} | |||
{{Use American English|date=May 2024}} | |||
{{Speciesbox | |||
| name = Cougar | |||
| fossil_range = {{Fossil range|1.2|0}}<small>Early ] – ]</small> | |||
| image = Mountain Lion in Glacier National Park.jpg | |||
| image_caption = A ] in ], United States | |||
| status = LC | | status = LC | ||
| status_system = |
| status_system = IUCN3.1 | ||
| status_ref =<ref name= |
| status_ref =<ref name=iucn>{{cite iucn |title=''Puma concolor'' |name-list-style=amp |author1=Nielsen, C. |author2=Thompson, D. |author3=Kelly, M. |author4=Lopez-Gonzalez, C. A. |page=e.T18868A97216466 |year=2015 |errata=2016 |doi=10.2305/IUCN.UK.2015-4.RLTS.T18868A50663436.en |access-date=January 16, 2022}}</ref> | ||
| |
| status2 = CITES_A2 | ||
| status2_system = CITES | |||
| image_caption = Captive cougar at the Cincinnati Zoo | |||
| status2_ref =<ref name=iucn/>{{efn|The populations of Costa Rica and Panama are included in Appendix I.}} | |||
| genus = Puma | | genus = Puma | ||
| species = concolor | | species = concolor | ||
| authority = (], 1771) | | authority = (], 1771)<ref name="MSW3" /> | ||
| range_map = Cougar_range_map_2010.png | | range_map = Cougar_range_map_2010.png | ||
| range_map_caption = Cougar range | | range_map_caption = Cougar range (without recent confirmations across northern Canadian territories, eastern U.S. states, and Alaska) | ||
| subdivision_ranks = |
| subdivision_ranks = Subspecies | ||
| subdivision = | | subdivision = | ||
* '']'' | |||
* ''P. c. anthonyi'' – eastern South America | |||
* '']'' |
* '']'' | ||
* '']'' – northern South America | |||
* '']''? – south Florida | |||
* '']'' – Central America | |||
* '']'' – North America | |||
* ''P. c. puma'' – southern South America | |||
Also see ] | Also see ] | ||
| synonyms = | |||
* ''] concolor'' | |||
}} | }} | ||
The '''cougar''' ('''''Puma concolor''''') ({{IPAc-en|ˈ|k|uː|g|ər}}, '']''), also known as the '''panther''', '''mountain lion''', '''catamount''' and '''puma''', is a large ] native to the Americas. It inhabits ], ] and ], making it the most widely distributed wild, terrestrial ] in the ], and one of the most widespread in the world. Its range spans the ], ] and ] provinces of ], the ] and areas in the western ]. Further south, its range extends through Mexico to the ] and the southern ] in ]. It is an adaptable ], occurring in most American ] types. It prefers habitats with dense underbrush and rocky areas for stalking but also lives in open areas. | |||
The '''cougar''' (''Puma concolor''), also commonly known as the '''mountain lion''', '''puma''', '''panther''', or '''catamount''', is a large ] of the subfamily ] native to the Americas. Its ], from the Canadian ] to the southern ] of South America, is the greatest of any large wild terrestrial mammal in the ].<ref name="diet" /> An adaptable, ] species, the cougar is found in most ] ] types. It is the second-heaviest cat in the ], after the ]. Secretive and largely solitary by nature, the cougar is properly considered both ] and ], although there are daytime sightings.<ref name="cougar">. US National Park Service.</ref><ref>Hansen, Kevin. (1992) ''Cougar: The American Lion.'' Northland. Flagstaff, AZ, ch. 4, ISBN 0873585445.</ref><ref>. New Mexico Department of Game & Fish</ref><ref>. California Department of Game & Fish</ref> The cougar is more closely related to smaller felines, including the ] (subfamily Felinae), than to any species of subfamily ],<ref name=MSW3/><ref>Hartwell, Sarah. . messybeast.com</ref><ref>. messybeast.com</ref> of which only the jaguar is native to the Americas. | |||
The cougar is |
The cougar is largely solitary. Its activity pattern varies from ] and ] to ] and ] between protected and non-protected areas, and is apparently correlated with the presence of other predators, prey species, livestock and humans. It is an ] that pursues a wide variety of prey. ]s, particularly ], are its primary prey, but it also hunts ]s. It is ] and lives at low population densities. Individual ]s depend on terrain, vegetation and abundance of prey. While large, it is not always the dominant ] in its range, yielding prey to other predators. It is reclusive and mostly avoids people. ] are rare but increased in North America as more people entered cougar habitat and built farms. | ||
The cougar is listed as ] on the ]. Intensive hunting following ] and ongoing human development into cougar habitat has caused populations to decline in most parts of its historical range. In particular, the ] population is considered to be mostly ] in eastern North America since the early 20th century, with the exception of the isolated ] subpopulation. | |||
Intensive hunting following ] and the ongoing human development of cougar ] has caused populations to drop in most parts of its historical range. In particular, the cougar was ] in eastern North America in the beginning of the 20th century, except for an isolated ] subpopulation. Breeding populations have moved east into the far western parts of the Dakotas, Nebraska, and Oklahoma. Transient males have been verified in Minnesota,<ref>. StarTribune.com (December 1, 2011). Retrieved on April 29, 2013.</ref> ],<ref>. Missouri Department of Conversation, mdc.mo.gov (March 20, 2014)</ref> ],<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>. Iowa Department of Natural Resources, iowadnr.gov (December 23, 2011)</ref><ref>. KWWL.com. Retrieved on April 29, 2013.</ref> the ] 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 ].<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=] News|accessdate=July 28, 2011}}</ref> Reports of ]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== | ==Naming and etymology== | ||
{{Wiktionary}} | |||
With its vast range across the length of the Americas, ''P. concolor'' has dozens of names and various references in the ] of the ] 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 ] of ].<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 ], a ] jaguar or leopard, and the ], a subspecies of cougar (''P.{{nbsp}}c. coryi''). | |||
The word ''cougar'' is borrowed from the ] ''çuçuarana'', via French; it was originally derived from the ]. A current form in Brazil is ''suçuarana''.<ref>{{cite web |url=http://www.etymonline.com/index.php?term=jaguar |title=Online Etymology Dictionary |date=2001 |publisher=] |access-date=August 12, 2006 |archive-date=June 28, 2017 |archive-url=https://web.archive.org/web/20170628080622/http://www.etymonline.com/index.php?term=jaguar |url-status=live}}<br /> & corresponding entry for {{Webarchive|url=https://web.archive.org/web/20151202104804/http://www.etymonline.com/index.php?l=c&p=27 |date=December 2, 2015 }}</ref> In the 17th century, ] named it ''cuguacu ara''. Marcgrave's rendering was reproduced in 1648 by his associate ]. ''Cuguacu ara'' was then adopted by ] in 1693.<ref>{{cite magazine |date=October 14, 2006 |title=Words to the Wise |magazine=Take Our Word for It |issue=205 |page=2 |url=http://www.takeourword.com/TOW205/page2.html |access-date=July 31, 2012 |url-status=live |archive-url=https://web.archive.org/web/20120812212012/http://www.takeourword.com/TOW205/page2.html |archive-date=August 12, 2012}}</ref> In 1774, ] converted ''cuguacu ara'' to ''cuguar'', which was later modified to "cougar" in English.<ref>{{OEtymD|jaguar}}</ref><ref>{{cite web |year=1989 |url=http://oxforddictionaries.com/definition/english/cougar |title=cougar |work=Oxford Dictionaries Online, Oxford University Press| url-status = dead |archive-url=https://web.archive.org/web/20130601145645/http://oxforddictionaries.com/definition/english/cougar| archive-date=June 1, 2013}}</ref> | |||
''P. concolor'' holds the ] 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> | |||
The cougar holds the ] record for the animal with the greatest number of names, with over 40 in English alone.<ref>{{cite book |author=Folkard, C. |title=The Guinness Book of World Records |year=2004 |page=49 |publisher=Bt Bound}}</ref> "Puma" is the ] used in Latin America and most parts of Europe. The term ''puma'' is also sometimes used in the United States.<ref>{{cite journal |author1=Rau, J. R. |author2=Jiménez, J. E. |name-list-style=amp |year=2002 |title=Diet of puma (''Puma concolor'', Carnivora: Felidae) in coastal and Andean ranges of southern Chile |journal=Studies on Neotropical Fauna and Environment |volume=37 |issue=3 |pages=201–205 |doi=10.1076/snfe.37.3.201.8567 |bibcode=2002SNFE...37..201R |s2cid=84264487 |issn = 0165-0521}}</ref><ref>{{cite journal |author1=Scognamillo, D. |author2=Maxit, I. E. |author3=Sunquist, M. |author4=Polisar, J. |name-list-style=amp |year=2003 |title=Coexistence of jaguar (''Panthera onca'') and puma (''Puma concolor'') in a mosaic landscape in the Venezuelan llanos |journal=Journal of Zoology |volume=259 |issue=3 |pages=269–279 |doi=10.1017/S0952836902003230}}</ref><ref>{{cite journal |author1=Holmes, B. R. |author2=Laundré, J. W. |name-list-style=amp |year=2006 |title=Use of open, edge and forest areas by pumas Puma concolor in winter: are pumas foraging optimally? |journal=Wildlife Biology |volume=12 |issue=2 |pages=201–210 |doi=10.2981/0909-6396(2006)122.0.CO;2 |doi-access=free}}</ref><ref>{{cite journal |author1=Dickson, B. G. |author2=Roemer, G. W. |author3=McRae, B. H. |author4=Rundall, J. M. |name-list-style=amp |year=2013 |title=Models of regional habitat quality and connectivity for Pumas (''Puma concolor'') in the southwestern United States |journal=PLOS ONE |volume=8 |issue=12 |page=e81898 |doi=10.1371/journal.pone.0081898 |pmid=24367495 |pmc=3867332 |bibcode=2013PLoSO...881898D |doi-access=free}}</ref> The first use of ''puma'' in English dates to 1777, introduced from Spanish from the ].<ref>{{cite web |title=Puma |website=] |access-date=April 16, 2022 |url=https://www.etymonline.com/word/puma |archive-date=May 16, 2022 |archive-url=https://web.archive.org/web/20220516211512/https://www.etymonline.com/word/puma |url-status=live}}</ref> In the western United States and Canada, it is also called "mountain lion", a name first used in writing in 1858.<ref>{{cite magazine |title=George A. Jackson's Diary of 1858–1859 |author=Jackson, G. A. |journal=Colorado Magazine |year=1935 |volume=6 |pages=201–214}}</ref> Other names include "panther" (although it does not belong to the ] '']'') and "catamount" (meaning "cat of the mountains").<ref>{{cite book |author=Wild, Paula |year=2013 |title=The Cougar: Beautiful, Wild and Dangerous |publisher=Douglas and McIntyre Limited |page=25 |isbn=9781771620031}}</ref> | |||
"Cougar" may be borrowed from the archaic ] ''çuçuarana''; the term was originally derived from the ] ''susua'rana'', meaning "similar to deer (in hair color)". A current form in ] is '']''. It may also be borrowed from the ] term ''guaçu ara'' or ''guazu ara''. Less common Portuguese terms are '']'' (brown ''onça'', in distinction of the black-spotted one, '']'', the ]) or '']'' (lit. ] 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 '']'' (she-cat), and outside of the Amazon, both are colloquially referred to as simply '']'' by many people (that is also a name for the leopard in ]). | |||
In the 17th century, German naturalist ] named the cat the ''cuguacu ara''. Marcgrave's rendering was reproduced by his associate, Dutch naturalist ], in 1648. ''Cuguacu ara'' was then adopted by English naturalist ] 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 ] 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 ] 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== | ==Taxonomy and evolution== | ||
''Felis concolor'' was the ] proposed by ] in 1771 for a cat with a long tail from Brazil.<ref>{{cite book |author=Linnaeus, C. |year=1771 |title=Mantissa plantarum altera |location=Holmiae |publisher=Impensis Direct. Laurentii Salvii |language=la |page=522 |chapter=''Felis concolor'' |chapter-url=https://bibdigital.rjb.csic.es/viewer/15082/?offset=#page=529&viewer=picture&o=bookmarks&n=0&q= |access-date=August 29, 2019 |archive-date=December 26, 2019 |archive-url=https://web.archive.org/web/20191226233854/https://bibdigital.rjb.csic.es/viewer/15082/?offset=#page=529&viewer=picture&o=bookmarks&n=0&q= |url-status=live}}</ref> The specific epithet of the name, "concolor", is ] for "of uniform color". It was placed in the genus '']'' by ] in 1834.<ref>{{cite book |last1=Jardine |first1=W. |year=1834 |title=Naturalists' library, Mammalia, volume 2 |location=Edinburgh |publisher=Lizars, Stirling and Kenney |pages=266–267 |chapter=Genus II. ''Puma'' |chapter-url=https://archive.org/details/naturalistslibra16jardrich/page/266}}</ref> This genus is part of the ].<ref name="MSW3" /> The cougar is most closely related to the ] and the cheetah.<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|doi-access=free}}</ref> | |||
] | |||
] | |||
Cougars are the largest of the small cats. They are placed in the subfamily ], although their bulk characteristics are similar to those of the ]s in the subfamily ].<ref name=MSW3/> The ] ] 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 ] 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 ],<ref name="Johnson2006"/> and significant ] exist with suggested dates. | |||
In the latest ] study of the Felidae, the common ancestor of today's '']'', '']'', '']'', '']'', and '']'' lineages migrated across the ] into the Americas 8.0 to 8.5 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=] |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 ], following formation of the ]. The cougar was originally thought to belong in ''Felis'' (''Felis concolor''), the genus which includes the ]. As of 1993, it is now placed in ''Puma'' along with the ], a cat just a little more than a tenth its weight. | |||
The cougar and jaguarundi are most closely related to the modern ] 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 ]) and migrated back to Asia and Africa,<ref name="Johnson2006" /><ref name="Culver" /> while other research suggests the cheetah diverged in the ] 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 ] 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 ] during the ] some 10,000 years ago, when other large mammals, such as '']'', also disappeared. North America was then repopulated by a group of South American cougars.<ref name="Culver" /> | |||
===Subspecies=== | ===Subspecies=== | ||
] at ], in the Chilean part of ]]] | |||
Until the late 1980s, as many as 32 ] were recorded; ] 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 '']'' (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> | |||
Following Linnaeus's first scientific description of the cougar, 32 cougar ]s were described and proposed as ] until the late 1980s. ] analysis of cougar ] indicates that many of these are too similar to be recognized as distinct at a molecular level but that only six ] groups exist. The ] samples showed a low ] variation, possibly due to ].<ref name="Culver" /> Following this research, the authors of '']'' recognized the following six subspecies in 2005:<ref name="MSW3">{{MSW3 Wozencraft |id=14000204 |pages=544–545 |heading=Species ''Puma concolor''}}</ref> | |||
* ] {{nobold|(''Puma concolor cabrerae'')}} Pocock, 1940:<br> includes the previous subspecies and synonyms ''hudsonii'' and ''puma'' | |||
* ] {{nobold|(''P. c. costaricensis'')}} Merriam, 1901 | |||
* ] {{nobold|(''P. c. anthonyi'')}} Nelson and Goldman, 1931:<br> includes the previous subspecies and synonyms ''acrocodia'', ''borbensis'', ''capricornensis'', ''concolor'', ''greeni'', and ''nigra'' | |||
* ] {{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'' | |||
* ] {{nobold|(''P. c. concolor'')}} Linnaeus, 1771:<br> includes the previous subspecies and synonyms ''bangsi'', ''incarum'', ''osgoodi'', ''soasoaranna, sussuarana'', ''soderstromii'', ''suçuaçuara'', and ''wavula'' | |||
* ] {{nobold|(''P. c. puma'')}} Molina, 1782:<br> includes the previous subspecies and synonyms ''araucanus'', ''concolor'', ''patagonica'', ''pearsoni'', and ''puma'' | |||
* ''P. c. concolor'' {{small|(Linnaeus, 1771)}} includes the ] ''bangsi'', ''incarum'', ''osgoodi'', ''soasoaranna'', ''sussuarana'', ''soderstromii'', ''suçuaçuara'', and ''wavula'' | |||
''''']''''' | |||
* ''P. c. puma'' {{small|(], 1782)}} includes the synonyms ''araucanus'', ''concolor'', ''patagonica'', ''pearsoni'', and ''puma'' {{small|(], 1904)}} | |||
* ] {{nobold|(''P. c. coryi'')}} | |||
* ''P. c. couguar'' {{small|(], 1792)}} includes ''arundivaga'', ''aztecus'', ''browni'', ''californica'', ''floridana'', ''hippolestes'', ''improcera'', ''kaibabensis'', ''mayensis'', ''missoulensis'', ''olympus'', ''oregonensis'', ''schorgeri'', ''stanleyana'', ''vancouverensis'', and ''youngi'' | |||
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)702.0.CO;2 }}</ref> Culver et al. noted low ] variation in the Florida panther, possibly due to ];<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> | |||
* ''P. c. costaricensis'' {{small|(], 1901)}} | |||
* ''P. c. anthonyi'' {{small|(] and ], 1931)}} includes ''acrocodia'', ''borbensis'', ''capricornensis'', ''concolor'', ''greeni'', and ''nigra'' | |||
* ''P. c. cabrerae'' {{small|], 1940}} includes ''hudsonii'' and ''puma'' proposed by Marcelli in 1922 | |||
In 2006, the Florida panther was still referred to as a distinct subspecies ''P. c. coryi'' in research works.<ref name="improving">{{cite journal |first=M. J. |last=Conroy |author2=Beier, P. |author3=Quigley, H. |author4=Vaughan, M. 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)702.0.CO;2 |s2cid=85920592}}</ref> | |||
{{as of|2017}}, the Cat Classification Taskforce of the Cat Specialist Group recognizes only two subspecies as ]:<ref name="Catsg2017">{{cite journal |author1=Kitchener, A. C. |author2=Breitenmoser-Würsten, C. |author3=Eizirik, E. |author4=Gentry, A. |author5=Werdelin, L. |author6=Wilting, A. |author7=Yamaguchi, N. |author8=Abramov, A. V. |author9=Christiansen, P. |author10=Driscoll, C. |author11=Duckworth, J. W. |author12=Johnson, W. |author13=Luo, S.-J. |author14=Meijaard, E. |author15=O'Donoghue, P. |author16=Sanderson, J. |author17=Seymour, K. |author18=Bruford, M. |author19=Groves, C. |author20=Hoffmann, M. |author21=Nowell, K. |author22=Timmons, Z. |author23=Tobe, S. |year=2017 |title=A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group |journal=Cat News |issue=Special Issue 11 |pages=33–34 |url=https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y#page=33 |access-date=September 3, 2020 |archive-date=July 30, 2018 |archive-url=https://web.archive.org/web/20180730142355/https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y#page=33 |url-status=live}}</ref> | |||
==Biology and behavior== | |||
*'']'' in South America, possibly excluding the region northwest of the Andes | |||
*'']'' in North and Central America and possibly northwestern South America | |||
===Evolution=== | |||
===Physical characteristics=== | |||
{{cladogram|title= | |||
] | |||
|caption=The ''Puma'' lineage of the family ], depicted along with closely related genera<ref name="bcw2">{{cite book |last1=Werdelin |first1=L. |last2=Yamaguchi |first2=N. |last3=Johnson |first3=W. E. |last4=O'Brien |first4=S. J. |chapter=Phylogeny and evolution of cats (Felidae) |name-list-style=amp |year=2010 |pages=59–82 |chapter-url=https://www.researchgate.net/publication/266755142 |editor1-last=Macdonald |editor1-first=D. W. |editor2-last=Loveridge |editor2-first=A. J. |title=Biology and Conservation of Wild Felids |publisher=Oxford University Press |location=Oxford, UK |isbn=978-0-19-923445-5 |access-date=April 4, 2022 |archive-date=September 25, 2018 |archive-url=https://web.archive.org/web/20180925141956/https://www.researchgate.net/publication/266755142 |url-status=live}}</ref> | |||
Cougars are slender and agile members of the cat family. They are the fourth-largest cat;<ref>. The Cougar Net.org</ref> adults stand about {{convert|60|to|90|cm|in|abbr=on}} tall at the shoulders.<ref>. 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) . Animal Diversity Web, University of Michigan Museum of Zoology. Retrieved on September 15, 2011.</ref> Males typically weigh 53 to 100 kg (115 to 220 lb), averaging 62 kg (137 lb). Females typically weigh between 29 and 64 kg (64 and 141 lb), averaging 42 kg (93 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 ], and larger towards the ].<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=] |volume=85 |issue=2 |pages=185–190 |doi=10.1007/BF00319400}}</ref> The largest recorded cougar, shot in 1901, weighed 105.2 kg (232 lb); claims of 125.2 kg (276 lb) and 118 kg (260 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 : University of Chicago Press|page= |isbn= 0226353443|access-date= }}</ref> On average, adult male cougars in British Columbia weigh 56.7 kg (125 lb) and adult females 45.4 kg (100 lb), though several male cougars in British Columbia weighed between 86.4 and 95.5 kg (190 to 210 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> | |||
|1={{clade | style=font-size:90%;line-height:100%;width:475px; | |||
|1={{clade | |||
|1={{clade | |||
|1={{clade | |||
|label1=''Lynx'' lineage | |||
|1='']'' | |||
}} | |||
|2={{clade | |||
|1={{clade | |||
|label1=''Puma'' lineage | |||
|1={{clade | |||
|label1='']'' | |||
|1=Cheetah ''A. jubatus'' ] | |||
|2={{clade | |||
|label1='']'' | |||
|1='''Cougar''' ] | |||
|label2='']'' | |||
|2=] ''H. yagouaroundi'' ] | |||
}} | |||
}} | |||
}} | |||
|2={{clade | |||
|label1=Domestic cat lineage | |||
|1='']'' | |||
|label2=Leopard cat lineage | |||
|2={{clade | |||
|1='']'' | |||
|2='']'' | |||
}} | |||
}} | |||
}} | |||
}} | |||
}} | |||
}} | |||
}} | |||
The ] ] 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 ] analysis.<ref name="Johnson2006" /> Significant ] exist with suggested dates. In the latest ] study of the Felidae, the common ancestor of today's '']'', '']'', '']'', '']'', and '']'' lineages migrated across the ] into the Americas 8.0 to 8.5 million years ago. 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. |name-list-style=amp |date=2006 |title=The Late Miocene radiation of modern Felidae: A genetic assessment |journal=] |volume=311 |issue=5757 |pages=73–77 |doi=10.1126/science.1122277 |pmid=16400146 |bibcode=2006Sci...311...73J |s2cid=41672825 |url=https://zenodo.org/record/1230866 |access-date=July 12, 2019 |url-access= |archive-date=October 4, 2020 |archive-url=https://web.archive.org/web/20201004075725/https://zenodo.org/record/1230866 |url-status=live}}</ref> North American felids then invaded South America 2–4{{nbsp}}Mya as part of the ], following the formation of the ].<ref name="Culver" /> | |||
], 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 ]) 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=] and ]}}</ref> | |||
The cheetah lineage is suggested by some studies to have diverged from the ''Puma'' lineage in the Americas and migrated back to Asia and Africa,<ref name="Johnson2006" /><ref name="Culver" /> while other research suggests the cheetah diverged in the ] itself.<ref>{{cite journal |author1=Ross, B. |author2=Barnes, I. |author3=Phillips, M. J. |author4=Martin, L D. |author5=Harington, C. R. |author6=Leonard, J. A. |author7=Cooper, A. |name-list-style=amp |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 |s2cid=17665121|doi-access=free |bibcode=2005CBio...15.R589B}}</ref> A high level of genetic similarity has been found among North American cougar populations, suggesting they are all fairly recent descendants of a small ancestral group. Culver et al. propose the original North American cougar population was ] during the ] some 10,000 years ago, when other large mammals, such as '']'', also disappeared. North America was then repopulated by ]s.<ref name="Culver" /> | |||
Cougars can be almost as large as ]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 "]", as it cannot roar, lacking the specialized ] and ] apparatus of '']''.<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> | |||
A ] identified as from a cougar was ] in Argentina's ] and dated to 17,002–16,573 years old. It contained '']'' eggs. This finding indicates that the cougar and the parasite have existed in South America since at least the ].<ref>{{cite journal |author1=Petrigh, R. S. |author2=Martínez, J. G. |author3=Mondini, M. |author4=Fugassa, M. H. |name-list-style=amp |year=2019 |title=Ancient parasitic DNA reveals ''Toxascaris leonina'' presence in Final Pleistocene of South America |journal=Parasitology |volume=146 |issue=10 |pages=1284–1288 |doi=10.1017/S0031182019000787 |pmid=31196226 |doi-access=free |hdl=11336/177873 |hdl-access=free}}</ref> The oldest fossil record of a cougar (''Puma concolor'') in South America (Argentina) is a partial skull from the late ] (]) age.<ref>{{cite journal |last1=Chimento |first1=N.R. |last2=Dondas |first2=A. |title=First record of ''Puma concolor'' (Mammalia, Felidae) in the Early-Middle Pleistocene of South America |journal=Journal of Mammalian Evolution |date=2018 |volume=25 |issue=3 |pages=381–389 |doi=10.1007/s10914-017-9385-x |s2cid=16249074}}</ref> | |||
] | |||
Cougar coloring is plain (hence the ] ''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 (]) 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=]: 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 "]" 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> | |||
==Characteristics== | |||
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>. 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 ] 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=] |deadurl=yes |archiveurl=https://web.archive.org/web/20150123034652/http://arizona.sierraclub.org/conservation/mt-lion/index.asp |archivedate=January 23, 2015 }}</ref> | |||
{{multiple image |align=right |direction=vertical |image1=Cougarskull.jpg |caption1=Cougar skull and jawbone |image2=Cougar track.jpg |caption2=Front paw print of a cougar}} | |||
The head of the cougar is round, and the ears are erect. Its powerful forequarters, neck, and jaw serve to grasp and hold large prey. It has four retractile claws on its hind paws and five on its forepaws, of which one is a ]. The larger front feet and claws are adaptations for clutching prey.<ref name="WhosWho">{{cite web |url=http://www.hww.ca/hww2.asp?pid=1&id=87&cid=8 |archive-url=https://web.archive.org/web/20070518235841/http://www.hww.ca/hww2.asp?pid=1&id=87&cid=8 |archive-date=May 18, 2007 |title=Cougar |work=Hinterland Who's Who |access-date=May 22, 2007 |publisher=] and ]}}</ref> | |||
===Hunting and diet=== | |||
A successful ] ], the cougar will eat any animal it can catch, from insects to large ]s (over 500 kg). Like all cats, it is an ], 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 ].<ref name="diet"/> Its most important prey species are various deer species, particularly in North America; ], ], ] and even bull ] are taken. Other species such as the ] and ], ], ], ], ], ], ], 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 ] and ]s.<ref name="diet"/> | |||
Cougars are slender and agile members of the ]. They are the fourth largest cat species worldwide;<ref>. The Cougar Net.org</ref> adults stand about {{cvt|60|to|90|cm}} tall at the shoulders.<ref>. Florida Panther Refuge</ref> Adult males are around {{cvt|2.4|m}} long from nose to tail tip, and females average {{cvt|2.05|m}}, with overall ranges between {{cvt|1.50|to|2.75|m}} 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'') |access-date=March 30, 2007 |publisher=Texas Parks and Wildlife |url-status=live |archive-url=https://web.archive.org/web/20070403213038/http://www.tpwd.state.tx.us/huntwild/wild/species/mlion/ |archive-date=April 3, 2007}}</ref><ref name="NY">{{cite web |url=http://www.dec.ny.gov/animals/6974.html |title=Eastern Cougar Fact Sheet |access-date=March 30, 2007 |publisher=New York State Department of Environmental Conservat ion |url-status=live |archive-url=https://web.archive.org/web/20071229075135/http://www.dec.ny.gov/animals/6974.html |archive-date=December 29, 2007}}</ref> Of this length, the tail typically accounts for {{cvt|63|to|95|cm}}.<ref name="ADW">Shivaraju, A. (2003) {{webarchive|url=https://web.archive.org/web/20090606152055/http://animaldiversity.ummz.umich.edu/site/accounts/information/Puma_concolor.html |date=June 6, 2009}}. Animal Diversity Web, University of Michigan Museum of Zoology. Retrieved on September 15, 2011.</ref> Males generally weigh {{cvt|53|to|72|kg}}. Females typically weigh between {{cvt|34|and|48|kg}}.<ref name="ADW" /><ref name="CAP">{{Cite web |author1=Nowell, K. |author2=Jackson, P. |name-list-style=amp |title=Wild Cats. Status Survey and Conservation Action Plan |publisher=IUCN/SSC Cat Specialist Group. IUCN, Gland, Switzerland |year=1996 |url=http://carnivoractionplans1.free.fr/wildcats.pdf |access-date=July 27, 2007 |url-status=live |archive-url=https://web.archive.org/web/20070807215533/http://carnivoractionplans1.free.fr/wildcats.pdf |archive-date=August 7, 2007}}</ref> Cougar size is smallest close to the ] and larger towards the ].<ref name="diet">{{cite journal |author1=Iriarte, J. A. |author2=Franklin, W. L. |author3=Johnson, W. E. |author4=Redford, K. H. |year=1990 |title=Biogeographic variation of food habits and body size of the America puma |journal=] |volume=85 |issue=2 |pages=185–190 |doi=10.1007/BF00319400 |pmid=28312554 |bibcode=1990Oecol..85..185I |s2cid=10134066}}</ref> The largest recorded cougar, shot in 1901, weighed {{cvt|105.2|kg}}; claims of {{cvt|125.2|kg}} and {{cvt|118|kg}} have been reported, though they were probably exaggerated.<ref>{{cite book |last=Hornocker |first=Maurice |date=2010 |title=Cougar : ecology and conservation |url=http://library.wur.nl/WebQuery/clc/1941110 |publisher=Chicago : University of Chicago Press |isbn=978-0226353449}}</ref> Male cougars in North America average {{cvt|62|kg}}, while the average female in the same region averages about {{cvt|42|kg}}.<ref>{{cite encyclopedia |last=Beler |first=Paul |title=puma |url=https://www.britannica.com/animal/puma-mammal-species |encyclopedia=Britannica|access-date=October 2, 2023|archive-date=October 6, 2023|archive-url=https://web.archive.org/web/20231006055338/https://www.britannica.com/animal/puma-mammal-species|url-status=live}}</ref> On average, adult male cougars in British Columbia weigh {{cvt|56.7|kg}} and adult females {{cvt|45.4|kg}}, though several male cougars in British Columbia weighed between {{cvt|86.4|and|95.5|kg}}.<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|url-status=dead|archive-url=https://web.archive.org/web/20110824110816/http://www.env.gov.bc.ca/wld/documents/cougar.htm|archive-date=August 24, 2011|access-date=June 5, 2011}}</ref> | |||
] | |||
Investigation in ] showed that ], followed by ], were the cougar's primary targets; the prey base is shared with the park's ], with which the cougar competes for resources.<ref name="Yellowstone">{{cite web |title=Wildlife: Wolves |publisher=] |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 ] 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> | |||
Depending on the locality, cougars can be smaller or bigger than jaguars but are less muscular and not as powerfully built, so on average, their weight is less. Whereas the size of cougars tends to increase as much as distance from the equator increases,<ref name="diet" /> which crosses the northern portion of South America, jaguars are generally smaller north of the ] in South America and larger south of it. For example, while South American jaguars are comparatively large, and may exceed {{cvt|90|kg}},<ref name="FrancisIserson2015">{{cite journal |author1=Francis, Adama M. |author2=Iserson, K. V. |year=2015 |title=Jaguar Attack on a Child: Case Report and Literature Review |journal=Western Journal of Emergency Medicine |volume=16 |issue=2 |pages=303–309 |doi=10.5811/westjem.2015.1.24043 |pmc=4380383 |pmid=25834674}}</ref> North American jaguars in Mexico's ] weigh approximately {{cvt|50|kg}}, about the same as female cougars.<ref name="Foodhabits">{{cite journal |author1=Nuanaez R. |author2=Miller, B. |author3=Lindzey F. |name-list-style=amp |year=2000 |title=Food habits of jaguars and pumas in Jalisco, Mexico |journal=Journal of Zoology |volume=252 |issue=3 |pages=373–379 |url=http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=58851 |doi=10.1111/j.1469-7998.2000.tb00632.x |access-date=May 20, 2007 |archive-date=August 10, 2016 |archive-url=https://web.archive.org/web/20160810211518/http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=58851 |url-status=live}}</ref> | |||
In ], ] samples showed ]s to make up 28% of the cougar's diet, ]s and ] 24% each, ]s 10%, ] 7%, and ] 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> | |||
Cougar coloring is plain (hence the Latin ''concolor'' in the scientific name) but can vary greatly across individuals and even siblings. The coat is typically tawny, but it ranges from silvery-grey to 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" /> A ] individual was seen in ] in Rio de Janeiro in 2013 when it was recorded by a camera trap, indicating that pure white individuals do exist within the species, though they are extremely rare.<ref name="Branco">{{cite web |url=https://revistapesquisa.fapesp.br/quando-o-pardo-e-branco/ |title=Quando o pardo é branco |date=2019 |work=revistapesquisa.fapesp.br |location=Brazil |language=pt |archive-url=https://web.archive.org/web/20200807222632/https://revistapesquisa.fapesp.br/quando-o-pardo-e-branco/ |archive-date=August 7, 2020 |access-date=August 18, 2020}}</ref><ref>{{Cite web |last=Dell'amore |first=Christine |date=December 3, 2020 |title=Extremely rare white cougar highlights a quirk of the species |url=https://www.nationalgeographic.com/animals/article/extrremely-rare-white-cougar-highlights-quirk-of-species?loggedin=true&rnd=1690918501116 |archive-url=https://web.archive.org/web/20230801194030/https://www.nationalgeographic.com/animals/article/extrremely-rare-white-cougar-highlights-quirk-of-species?loggedin=true&rnd=1690918501116 |url-status=dead |archive-date=August 1, 2023 |access-date=August 1, 2023 |website=National Geographic}}</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 ]. 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, ]s, ]s, raccoons, ]s, ], ], ], ], and ].<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. | |||
The cougar has large paws and proportionally the largest hind legs in the Felidae,<ref name="CAP" /> allowing for great leaping and powerful short sprints. It can leap from the ground up to {{cvt|5.5|m}} high into a tree.<ref name="Nowak, 1999">{{cite book |title=Walker's Mammals of the World |volume=1 |last=Nowak |first=R. M. |year=1999 |publisher=Johns Hopkins University Press |location=Baltimore |isbn=0-8018-5789-9 |page=818}}</ref> | |||
Though capable of sprinting, the cougar is typically an ]. 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"/> | |||
==Distribution and habitat== | |||
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 ], 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)0502.0.CO;2}}</ref> | |||
] image of a cougar in ], ]]] | |||
The cougar has the most extensive range of any wild land animal in the Americas, spanning 110 degrees of ] from the Yukon in Canada to the southern Andes in Chile.<ref name="iucn" /> The species was extirpated from eastern North America, aside from Florida, but they may be recolonizing their former range and isolated populations have been documented east of their contemporary ranges in both the Midwestern US and Canada.<ref>{{cite journal |author1=Morrison, C. D. |author2=Boyce, M. S. |author3=Nielsen, S. E. |year=2015 |title=Space-use, movement and dispersal of sub-adult cougars in a geographically isolated population |journal=PeerJ |volume=3 |pages=e1118 |doi=10.7717/peerj.1118 |pmid=26290786 |pmc=4540023 |doi-access=free}}</ref> | |||
===Reproduction and life cycle=== | |||
Females reach sexual maturity between one-and-a-half to three years of age. They typically average one ] 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 ] for about 8 days of a 23-day cycle; the ] is approximately 91 days.<ref name="CAP"/> Females are sometimes reported as ],<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=]}}</ref> but this is uncertain and ] 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=]}}</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)0322.0.CO;2|url=http://post.queensu.ca/~bonierf/pubs/pub1.pdf}}</ref> | |||
The cougar lives in all forest types, lowland and mountainous deserts, and in open areas with little vegetation up to an elevation of {{cvt|5800|m}}.<ref name="iucn" /> In the ], it prefers steep canyons, escarpments, rim rocks and dense brush.<ref>{{cite journal |author1=Dickson, B.G. |name-list-style=amp |author2=Beier, P. |year=2007 |title=Quantifying the influence of topographic position on cougar (''Puma concolor'') movement in southern California, USA |journal=Journal of Zoology |volume=271 |issue=3 |pages=270–277 |doi=10.1111/j.1469-7998.2006.00215.x |citeseerx=10.1.1.571.8947}}</ref> In Mexico, it was recorded in the ].<ref>{{cite journal |author1=Caso, A. |name-list-style=amp |author2=Domínguez, E. F. |year=2018 |title=Confirmed presence of jaguar, ocelot and jaguarundi in the Sierra of San Carlos, Mexico |journal=Cat News |issue=68 |pages=31–32}}</ref> In the ], it inhabits ] and semi-] forests in ].<ref>{{cite journal |author1=Ávila-Nájera, D. M. |name-list-style=amp |author2=Chávez, C. |author3=Pérez-Elizalde, S. |author4=Guzmán-Plazola, R. A. |author5=Mendoza, G. D. |author6=Lazcano-Barrero, M. A. |year=2018 |title=Ecology of ''Puma concolor'' (Carnivora: Felidae) in a Mexican tropical forest: adaptation to environmental disturbances |journal=Revista de Biología Tropical |volume=66 |issue=1 |pages=78–90 |doi=10.15517/rbt.v66i1.27862 |doi-access=free}}</ref> In ], it was recorded in the lower montane forest in ] and in a river basin in the ] above {{cvt|700|m}} in 2019.<ref>{{cite journal |author1=Morales-Rivas, A. |name-list-style=amp |author2=Álvarez, F. S. |author3=Pocasangre-Orellana, X. |author4=Girón, L. |author5=Guerra, G. N. |author6=Martínez, R. |author7=Pablo Domínguez, J. |author8=Leibl, F. |author9=Heibl, C. |year=2020 |title=Big cats are still walking in El Salvador: first photographic records of ''Puma concolor'' (Linnaeus, 1771) and an overview of historical records in the country |journal=Check List |volume=16 |issue=4 |pages=563–570 |doi=10.15560/16.3.563 |doi-access=free}}</ref> | |||
] | |||
In ], it was recorded in a ] ] close to a ] in the ], and close to water bodies in the ].<ref>{{cite journal |author1=Olarte-González, G. |name-list-style=amp |author2=Escovar-Fadul, T. |author3=Balaguera-Reina, S.A. |year=2015 |title=First record of ''Puma concolor'' Linneus, 1771 (Carnivora: Felidae) preying ''Odocoileus virginianus'' (Zimmermann, 1780) on a palm-oil plantation in the Meta department, Colombia |journal=Mammalogy Notes |volume=2 |issue=1 |pages=8–10 |doi=10.47603/manovol2n1.8-10 |doi-access=free}}</ref><ref>{{cite journal |author1=Boron, V. |name-list-style=amp |author2=Xofis, P. |author3=Link, A. |author4=Payan, E. |author5=Tzanopoulos, J. |year=2020 |title=Conserving predators across agricultural landscapes in Colombia: habitat use and space partitioning by jaguars, pumas, ocelots and jaguarundis |journal=Oryx |volume=54 |issue=4 |pages=554–563 |doi=10.1017/S0030605318000327 |doi-access=free}}</ref> | |||
] | |||
In the human-modified landscape of central Argentina, it inhabits ] with abundant vegetation cover and prey species.<ref name=Guerisoli2019>{{cite journal |author1=Guerisoli, M. D. L. M. |author2=Caruso, N. |author3=Luengos Vidal, E. M. |name-list-style=amp |author4=Lucherini, M. |year=2019 |title=Habitat use and activity patterns of ''Puma concolor'' in a human-dominated landscape of central Argentina |journal=Journal of Mammalogy |volume=100 |issue=1 |pages=202–211 |doi=10.1093/jmammal/gyz005 |doi-access=free |hdl=11336/104140 |hdl-access=free}}</ref> | |||
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> | |||
==Behavior and ecology== | |||
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 ] amongst cougars that travel farthest from the maternal range, often due to conflicts with other cougars (]).<ref name="Utah"/> Research in ] 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> | |||
Cougars are an important keystone species in Western Hemisphere ecosystems, linking numerous species at many trophic levels. In a comprehensive literature review of more than 160 studies on cougar ecology, ecological interactions with 485 other species in cougar-inhabited ecosystems have been shown to involve different areas of interaction, ranging from the use of other species as food sources and prey, fear effects on potential prey, effects from carcass remains left behind, to competitive effects on other predator species in shared habitat. The most common research topic in the literature used here was the cougar's diet and its prey's regulation.<ref name="LaBarge et al. 2022">{{Cite journal |last1=LaBarge |first1=L. R. |last2=Evans |first2=M. J. |last3=Miller |first3=J. R. B. |last4=Cannataro |first4=G. |last5=Hunt |first5=C. |last6=Elbroch |first6=L. M. |date=2022 |title=Pumas ''Puma concolor'' as ecological brokers: a review of their biotic relationships |journal=Mammal Review |volume=52 |issue=3 |pages=360–376 |doi=10.1111/mam.12281 |doi-access=free}}</ref> | |||
===Hunting and diet=== | |||
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 ].<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. ], 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> | |||
] northwest of Los Angeles]] | |||
] | |||
The cougar is a ] ]. It prefers large mammals such as ], ], ], ], ] and ]. It opportunistically takes smaller prey such as ]s, ]s, smaller carnivores, birds, and even domestic animals, including pets.<ref name="Naughton" /> The mean weight of cougar vertebrate prey increases with its body weight and is lower in areas closer to the ]. A survey of North America research found 68% of prey items were ungulates, especially deer. Only the Florida panther showed variation, often preferring feral ]s and ]s.<ref name="diet" /> Cougars have been known to prey on introduced ] populations in ]. One individual cougar was recorded as hunting 29 gemsbok, which made up 58% of its recorded kills. Most gemsbok kills were neonates, but some adults were also known to have been taken.<ref name="TWS">{{cite web |last1=Kobilinsky |first1=Dana |date=March 9, 2023 |title=Rising oryx numbers may distress New Mexico ecosystem |url=https://wildlife.org/rising-oryx-numbers-may-distress-new-mexico-ecosystem/ |access-date=March 12, 2023 |website=The Wildlife Society |archive-date=March 11, 2023 |archive-url=https://web.archive.org/web/20230311220740/https://wildlife.org/rising-oryx-numbers-may-distress-new-mexico-ecosystem/ |url-status=live}}</ref> Elsewhere in the southwestern United States, they have been recorded to also prey on ]s in the ],<ref>{{Cite web |url=https://wildlife.org/jwm-cougars-prey-on-feral-horses-in-the-great-basin |title=JWM: Cougars prey on feral horses in the Great Basin |date=August 20, 2021|access-date=May 10, 2023|archive-date=May 10, 2023|archive-url=https://web.archive.org/web/20230510190218/https://wildlife.org/jwm-cougars-prey-on-feral-horses-in-the-great-basin/|url-status=live}}</ref> as well as ]s in the ] and ]s.<ref>{{Cite web |url=https://wildlife.org/do-cougars-affect-ecosystems-by-preying-on-feral-donkeys/ |title=Do cougars affect ecosystems by preying on feral donkeys? |date=May 10, 2023|access-date=May 10, 2023|archive-date=May 10, 2023|archive-url=https://web.archive.org/web/20230510184027/https://wildlife.org/do-cougars-affect-ecosystems-by-preying-on-feral-donkeys/|url-status=live}}</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 ], being most active around dawn and dusk. | |||
Investigations at ] showed that elk and mule deer were the cougar's primary prey; the prey base is shared with the park's ], with which the cougar competes for resources.<ref name="Yellowstone">{{cite web |title=Wildlife: Wolves |publisher=] |url=http://www.yellowstonenationalpark.com/wolves.htm |access-date=April 8, 2007 |url-status=live |archive-url=https://web.archive.org/web/20070420174741/http://www.yellowstonenationalpark.com/wolves.htm |archive-date=April 20, 2007}}<br />* {{cite web |author1=Akenson, H. |author2=Akenson, J. |author3=Quigley, H. |title=Winter predation and interactions of Wolves and Cougars on Panther Creek in Central Idaho |url=http://www.yellowstonenationalpark.com/wolves.htm |url-status=live |archive-url=https://web.archive.org/web/20070420174741/http://www.yellowstonenationalpark.com/wolves.htm |archive-date=April 20, 2007}}<br />* {{cite web |author1=Oakleaf, John K. |author2=Mack, C. |author3=Murray, D. L. |title=Winter predation and interactions of Cougars and Wolves in the Central Idaho Wilderness |url=http://www.yellowstonenationalpark.com/wolves.htm |url-status=live |archive-url=https://web.archive.org/web/20070420174741/http://www.yellowstonenationalpark.com/wolves.htm |archive-date=April 20, 2007}}</ref> A study on winter kills from November to April in ] 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=771–775 |doi=10.1139/z97-098 |name-list-style=amp}}</ref> | |||
Estimates of territory sizes vary greatly. '']'' reports large male territories of 150 to 1000 km<sup>2</sup> (58 to 386 sq mi) with female ranges half the size.<ref name="CanGeo"/> Other research suggests a much smaller lower limit of 25 km<sup>2</sup> (10 sq mi), but an even greater upper limit of 1300 km<sup>2</sup> (500 sq mi) for males.<ref name="Utah"/> In the United States, very large ranges have been reported in Texas and the ] of the northern ], in excess of 775 km<sup>2</sup> (300 sq mi).<ref name="Dordt">{{cite web |url=http://homepages.dordt.edu/~mahaffy/mtlion/mtlionshort_behaviour.html |title=Behavior of cougar in Iowa and the Midwest |accessdate=May 11, 2007 |last=Mahaffy |first=James |date=December 2004 |publisher=]}}</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, ], and ] are used to mark territory and attract mates. Males may scrape together a small pile of leaves and grasses and then ].<ref name="Sierra"/> | |||
In the Central and South American cougar range area, the ratio of deer in the diet declines. Small to mid-sized mammals, including large rodents such as the ], are preferred. Ungulates accounted for only 35% of prey items in one survey, about half that of North America. Competition with the larger jaguar in South America has been suggested for the decline in the size of prey items.<ref name="diet" /> In Central or North America, the cougar and jaguar share the same prey, depending on its abundance.<ref name="GGLG2017">{{cite journal |last1=Gutiérrez-González |first1=C. E. |last2=López-González |first2=C. A. |title=Jaguar interactions with pumas and prey at the northern edge of jaguars' range |journal=PeerJ |volume=5 |pages=e2886 |date=2017 |doi=10.7717/peerj.2886 |pmid=28133569 |pmc=5248577 |name-list-style=amp |doi-access=free}}</ref> Other listed prey species of the cougar include ], ]s, ]s, ]s, ]s, ]es, ], ]s, ]s and ]s.<ref>{{cite book |author=Whitaker, J. O. |year=1980 |title=The Audubon Society Field Guide to North American Mammals |location=New York |publisher=Chanticleer Press |chapter=|chapter-url= |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" /> ]s (''Spheniscus magellanicus'') constitute the majority of prey items in cougar diet in ]'s ] and ].<ref>{{cite journal |author1=Martínez, J.I.Z. |name-list-style=amp |author2=Travaini, A. |author3=Zapata, S. |author4=Procopio, D. |author5=Santillán, M.Á. |year=2012 |title=The ecological role of native and introduced species in the diet of the puma ''Puma concolor'' in southern Patagonia |journal=Oryx |volume=46 |issue=1 |pages=106–111 |doi=10.1017/S0030605310001821 |doi-access=free |hdl=11336/81623 |hdl-access=free}}</ref> | |||
Home range sizes and overall cougar abundance depend on terrain, vegetation, and prey abundance.<ref name="Utah"/> One female adjacent to the ], for instance, was found with a large range of 215 km<sup>2</sup> (83 sq mi), necessitated by poor prey abundance.<ref name="Dispersal"/> Research has shown cougar abundances from 0.5 animals to as much as 7 (in one study in South America) per 100 km<sup>2</sup> (38 sq mi).<ref name="CAP"/> | |||
Although capable of sprinting, the cougar is typically an ]. 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 can break 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 ], but deer carcasses left exposed for study were scavenged by cougars in California, suggesting more opportunistic behavior.<ref>{{cite journal |last=Bauer |first=J. W. |author2=Logan, K. A. |author3=Sweanor, L. L. |author4=Boyce, W. M. |date=2005 |title=Scavenging behavior in Puma |journal=The Southwestern Naturalist |volume=50 |issue=4 |pages=466–471 |name-list-style=amp |doi=10.1894/0038-4909(2005)0502.0.CO;2 |s2cid=85632179}}</ref> | |||
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> | |||
===Interactions with other predators=== | |||
==Ecology== | |||
]s at ], using a ] for refuge]] | |||
Aside from humans, no species preys upon mature cougars in the wild, although conflicts with other predators or scavengers occur. Of the large predators in ] – the ] and ]s, ] and cougar – the massive grizzly bear appears dominant, often (though not always) able to drive a gray wolf pack, black bear or cougar off their kills. One study found that grizzlies and ]s visited 24% of cougar kills in Yellowstone and ], usurping 10% of carcasses. Bears gained up to 113%, and cougars lost up to 26% of their daily energy requirements from these encounters.<ref>{{Cite web |author=COSEWIC. Canadian Wildlife Service |title=Assessment and Update Status Report on the Grizzly Bear (''Ursus arctos'') |website=] |year=2002 |url=http://dsp-psd.pwgsc.gc.ca/Collection/CW69-14-166-2002E.pdf |access-date=April 8, 2007 |archive-date=October 9, 2022 |archive-url=https://ghostarchive.org/archive/20221009/http://dsp-psd.pwgsc.gc.ca/Collection/CW69-14-166-2002E.pdf |url-status=live}}</ref> In ] 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 killing, and when bears are most active, the cats take prey more frequently and spend less time feeding on each kill. Unlike several subordinate predators from other ecosystems, cougars do not appear to exploit spatial or temporal refuges to avoid 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|doi-access=free}}</ref><ref>{{cite magazine |author=ELboch, M. |date=November 1, 2014 |title=Mountain Lions Versus Black Bears |magazine=National Geographic|access-date=September 24, 2016 |url=http://voices.nationalgeographic.com/2014/11/01/mountain-lions-versus-black-bears/|url-status=dead|archive-url=https://web.archive.org/web/20160925021131/http://voices.nationalgeographic.com/2014/11/01/mountain-lions-versus-black-bears/|archive-date=September 25, 2016}}</ref> | |||
===Distribution and habitat=== | |||
] | |||
] 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 ], from northern ] in Canada to the southern ]. 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 gray wolf and the cougar compete more directly for prey, mostly in winter. Packs of wolves can steal cougars' kills, and there are some documented cases of cougars being killed by them. One report describes a large pack of seven to 11 wolves killing a female cougar and her kittens,<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|access-date=April 12, 2013|url-status=live|archive-url=https://web.archive.org/web/20131102015747/http://www.forwolves.org/ralph/wolves-deadcougar.htm|archive-date=November 2, 2013}}</ref> while in nearby ], a 2-year-old male cougar was found dead, apparently killed by a wolf pack.<ref name="IME">{{cite web |title=Predators clash above Elkhorn |url=http://archives.mtexpress.com/index2.php?ID=2005125077 |last=Kauffman |first=J. |date=2009 |publisher=Idaho Mountain Express|access-date=August 21, 2013|url-status=live|archive-url=https://web.archive.org/web/20170807071659/http://archives.mtexpress.com/index2.php?ID=2005125077|archive-date=August 7, 2017}}</ref> Conversely, one-to-one confrontations tend to be dominated by the cat, and there are various documented accounts where wolves have been ambushed and killed,<ref name="Wolf B4 Killed by Mountain Lion?">{{cite web |url=http://www.forwolves.org/ralph/32596.html |title=Wolf B4 Killed by Mountain Lion? |date=March 25, 1996 |publisher=forwolves.org|archive-url=https://web.archive.org/web/20131102020159/http://www.forwolves.org/ralph/32596.html|archive-date=November 2, 2013|url-status=live}}</ref><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=G. |date=2003 |newspaper=] |access-date=April 9, 2007 |url-status=dead |archive-url=https://web.archive.org/web/20130723074822/http://forests.org/shared/reader/welcome.aspx?linkid=22714&keybold=wildlife%20AND%20%20cougar |archive-date=July 23, 2013}}</ref><ref name="Autopsy Indicates Cougar Killed Wolf">{{cite web |url=http://www.igorilla.com/gorilla/animal/1999/cougar_kills_wolf.html |title=Autopsy Indicates Cougar Killed Wolf |date=2000 |publisher=igorilla.com|archive-url=https://web.archive.org/web/20150924033237/http://www.igorilla.com/gorilla/animal/1999/cougar_kills_wolf.html|archive-date=September 24, 2015|url-status=live|access-date=May 22, 2015}}</ref><ref name="Mountain lions kill collared wolves in Bitterroot">{{cite web |url=http://missoulian.com/news/state-and-regional/mountain-lions-kill-collared-wolves-in-bitterroot/article_68c0c60c-d792-59e3-b736-5b10c17eb10a.html |title=Mountain lions kill collared wolves in Bitterroot |publisher=missoulian.com |archive-url=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|archive-date=May 17, 2013|url-status=dead|access-date=May 29, 2012}}</ref> including adult male specimens.<ref name="This lion doesn't run, instead kills, eats wolf">{{cite web |url=https://www.jhnewsandguide.com/news/environmental/this-lion-doesn-t-run-instead-kills-eats-wolf/article_bdf4e68b-49d2-52b7-af68-302a559a9361.html |title=This lion doesn't run, instead kills, eats wolf |date=December 11, 2013 |publisher=National Geographic|access-date=October 7, 2019|archive-date=October 7, 2019|archive-url=https://web.archive.org/web/20191007213926/https://www.jhnewsandguide.com/news/environmental/this-lion-doesn-t-run-instead-kills-eats-wolf/article_bdf4e68b-49d2-52b7-af68-302a559a9361.html|url-status=live}}</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 ], 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 |access-date=April 9, 2007 |publisher=Greater Yellowstone Learning Center| archive-url = https://web.archive.org/web/20070929115515/http://www.greateryellowstonescience.org/topic/wolves/wolvesoverview.html| archive-date=September 29, 2007}}</ref> One researcher in Oregon noted: "When there is a pack around, cougars are not comfortable around their kills or raising kittens A lot of times a big cougar will kill a wolf, but the pack phenomenon changes the table."<ref>{{cite news |url=http://www.oregonwild.org/press-room/press-clips/turf-wars-in-idaho-s-wilderness |title=Turf wars in Idaho's wilderness |last=Cockle |first=Richard |date=2006 |newspaper=] |access-date=April 9, 2007 |url-status=dead |archive-url=https://web.archive.org/web/20071213022857/http://www.oregonwild.org/press-room/press-clips/turf-wars-in-idaho-s-wilderness |archive-date=December 13, 2007}}</ref> Both species are capable of killing mid-sized predators, such as ]s, ]es, ]s and ]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 |date=2013 |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|access-date=April 8, 2013 |url-status=live|archive-url=https://web.archive.org/web/20130411040326/http://missoulian.com/news/state-and-regional/cougars-vs-coyotes-photos-draw-internet-crowd/article_f6ecdb1a-a051-11e2-85a3-001a4bcf887a.html|archive-date=April 11, 2013}}</ref> | |||
The cougar was ] across much of its eastern North American range (with the exception of Florida) in the two centuries after ], and faced grave threats in the remainder of its territory. Currently, it ranges across most western American states, the Canadian provinces of ], ] and ], and the Canadian territory of ]. 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=] |accessdate=May 6, 2007}}</ref> ] 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=] |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 ] 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 .''</ref> The ] 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 ], which is critically endangered. There have been unconfirmed sightings in ] (north of ]); and in ], 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 ] Department of Natural Resources confirmed a cougar sighting in Michigan's ].<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 ] 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> | |||
The cougar and jaguar share overlapping territory in the southern portion of its range.<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 |access-date=August 30, 2006 |archive-url=https://web.archive.org/web/20060716064117/http://www.ecology.info/ecology-jaguar-puma.htm| archive-date = July 16, 2006}}</ref> The jaguar tends to take the larger prey where ranges overlap, reducing both the cougar's potential size and the likelihood of direct competition between the two cats.<ref name="diet" /> Cougars appear better than jaguars at exploiting a broader prey niche and smaller prey.<ref name="foodhabits">{{cite journal |author1=Nuanaez, R. |author2=Miller, B. |author3=Lindzey, F. |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> | |||
On April 14, 2008, police shot and killed a cougar on the north side of ], Illinois. DNA tests were consistent with cougars from the ] of ]. 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 ], 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> | |||
===Social spacing and interactions=== | |||
Other eastern sightings since 2010 have occurred in locations such as ],<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> ]<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 ],<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> ]<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> ],<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 ],<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 ].<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> | |||
The cougar is a mostly solitary animal. Only mothers and kittens live in groups, with adults meeting rarely. While generally loners, cougars will reciprocally share kills and seem to organize themselves into small communities defined by the territories of dominant males. Cats within these areas socialize more frequently with each other than with outsiders.<ref>{{cite journal |title=Adaptive social strategies in a solitary carnivore |first1=L. M. |last1=Elbroch |first2=M. |last2=Levy |first3=M. |last3=Lubell |first4=H. |last4=Quigley |first5=A. |last5=Caragiulo |date=2017 |journal=Science Advances |volume=3 |issue=10 |name-list-style=amp |page=e1701218 |doi=10.1126/sciadv.1701218 |pmid=29026880 |pmc=5636203 |bibcode=2017SciA....3E1218E}}</ref> | |||
In the vicinity of a cattle ranch in northern Mexico, cougars exhibited ] activity that overlapped foremost with the activity of ].<ref>{{cite journal |author1=Gutiérrez-González, C.E. |name-list-style=amp |author2=López-González, C.A. |year=2017 |title=Jaguar interactions with pumas and prey at the northern edge of jaguars' range |journal=PeerJ |volume=5 |pages=e2886 |doi=10.7717/peerj.2886 |doi-access=free |pmid=28133569 |pmc=5248577}}</ref> In a nature reserve in central Mexico, the activity of cougars was ] and nocturnal, overlapping largely with the activity of the ] (''Dasypus novemcinctus'').<ref>{{cite journal |author1=Soria-Díaz, L. |author2=Monroy-Vilchis, O. |author3=Zarco-González, Z. |year=2016 |title=Activity pattern of puma (''Puma concolor'') and its main prey in central Mexico |journal=Animal Biology |volume=66 |pages=13–20 |doi=10.1163/15707563-00002487 |name-list-style=amp |url=https://www.researchgate.net/publication/292946569 |access-date=April 24, 2024 |archive-date=May 5, 2024 |archive-url=https://web.archive.org/web/20240505151909/https://www.researchgate.net/publication/292946569_Activity_pattern_of_puma_Puma_concolor_and_its_main_prey_in_central_Mexico |url-status=live}}</ref> Cougars in the montane Abra-Tanchipa Biosphere Reserve in southeastern Mexico displayed a ] activity pattern.<ref>{{cite journal |author1=Hernández-Saintmartín, A.D. |author2=Rosas-Rosas, O.C. |author3=Palacio-Núñez, J. |author4=Tarango-Arámbula, L.A. |author5=Clemente-Sánchez, F. |author6=Hoogesteijn, A.L. |year=2013 |title=Activity patterns of jaguar, puma and their potential prey in San Luis Potosí, Mexico |journal=Acta Zoológica Mexicana |volume=29 |issue=3 |url=https://www.scielo.org.mx/scielo.php?pid=S0065-17372013000300005&script=sci_arttext |name-list-style=amp |access-date=April 20, 2024 |archive-date=April 20, 2024 |archive-url=https://web.archive.org/web/20240420154521/https://www.scielo.org.mx/scielo.php?pid=S0065-17372013000300005&script=sci_arttext |url-status=live}}</ref> Data from 12 years of camera trapping in the Pacific slope and Talamanca Cordillera of Costa Rica showed cougars as cathemeral.<ref>{{cite journal |author1=Botts, R.T. |author2=Eppert, A.A. |author3=Wiegman, T.J. |author4=Rodriguez, A. |author5=Blankenship, S.R. |author6=Asselin, E.M. |author7=Garley, W.M. |author8=Wagner, A.P. |author9=Ullrich, S.E. |author10=Allen, G.R. |author11=Mooring, M.S. |year=2020 |title=Circadian activity patterns of mammalian predators and prey in Costa Rica |journal=Journal of Mammalogy |volume=101 |issue=5 |pages=1313–1331 |doi=10.1093/jmammal/gyaa103 |doi-access=free |pmid=33343263 |pmc=7733402 |name-list-style=amp}}</ref> Both cougars and jaguars in the ] of Belize were nocturnal but avoided each other.<ref>{{cite journal |author1=Harmsen, B.J. |author2=Foster, R.J. |author3=Silver, S.C. |author4=Ostro, L.E.T. |author5=Doncaster, C.P. |year=2009 |title=Spatial and temporal interactions of sympatric Jaguars (''Panthera onca'') and Pumas (''Puma concolor'') in a Neotropical forest |journal=Journal of Mammalogy |volume=90 |issue=3 |pages=612–620 |doi=10.1644/08-MAMM-A-140R.1 |name-list-style=amp}}</ref> In a protected cloud forest in the central Andes of Colombia, cougars were active from late afternoon to shortly before sunrise and sometimes during noon and early afternoon.<ref>{{cite journal |author1=Cepeda-Duque, J.C. |author2=Gómez–Valencia, B. |author3=Alvarez, S. |author4=Gutiérrez–Sanabria, D.R. |author5=Lizcano, D.J. |year=2021 |title=Daily activity pattern of pumas (''Puma concolor'') and their potential prey in a tropical cloud forest of Colombia |journal=Animal Biodiversity and Conservation |volume=44 |issue=2 |pages=267–278 |doi=10.32800/abc.2021.44.0267 |name-list-style=amp|doi-access=free}}</ref> In protected areas of the ]-] Landscape in Bolivia and Peru, cougars were active throughout the day but with a tendency to nocturnal activity that overlapped with the activity of main prey species.<ref>{{cite journal |author1=Ayala, G.M. |name-list-style=amp |author2=Viscarra, M.E. |author3=Sarmento, P. |author4=Negrões, N. |author5=Fonseca, C. |author6=Wallace, R.B. |year=2021 |title=Activity patterns of jaguar and puma and their primary prey in the Greater Madidi-Tambopata Landscape (Bolivia, Peru) |journal=Mammalia |volume=85 |issue=3 |pages=208–219 |doi=10.1515/mammalia-2020-0058}}</ref> During an 8-year-long study in a modified landscape in southeastern Brazil, male cougars were primarily nocturnal, but females were active at night and day.<ref>{{cite journal |author1=Azevedo, F.C. |author2=Lemos, F.G. |author3=Freitas-Junior, M.C. |author4=Rocha, D.G. |author5=Azevedo, F.C.C. |year=2018 |title=Puma activity patterns and temporal overlap with prey in a human-modified landscape at southeastern Brazil |journal=Journal of Zoology |volume=305 |issue=4 |pages=246–255 |doi=10.1111/jzo.12558 |name-list-style=amp}}</ref> Cougars were diurnal in the Brazilian ], but crepuscular and nocturnal in protected areas in the ], ] and ] biomes.<ref>{{cite journal |author1=Foster, V.C. |author2=Sarmento, P. |author3=Sollmann, R. |author4=Tôrres, N. |author5=Jácomo, A.T. |author6=Negrões, N. |author7=Fonseca, C. |author8=Silveira, L. |year=2013 |title=Jaguar and Puma activity patterns and predator-prey interactions in four Brazilian Biomes |journal=Biotropica |volume=45 |issue=3 |pages=373–379 |doi=10.1111/btp.12021 |jstor=23525363 |bibcode=2013Biotr..45..373F |name-list-style=amp}}</ref> Cougars in the Atlantic Forest were active throughout the day but displayed peak activity during early mornings in protected areas and crepuscular and nocturnal activity in less protected areas.<ref>{{cite journal |author1=Paviolo, A. |name-list-style=amp |author2=Di Blanco, Y. E. |author3=De Angelo, C.D. |author4=Di Bitetti, M.S. |year=2009 |title=Protection affects the abundance and activity patterns of pumas in the Atlantic Forest |journal=Journal of Mammalogy |volume=90 |issue=4 |pages=926–934 |doi=10.1644/08-MAMM-A-128.1 |doi-access=free |hdl=11336/59545 |hdl-access=free}}</ref> In central Argentina, cougars were active day and night in protected areas but were active immediately after sunset and before sunrise outside protected areas.<ref>{{cite journal |author1=Zanón-Martínez, J.I. |name-list-style=amp |author2=Kelly, M.J. |author3=Mesa-Cruz, J.B. |author4=Sarasola, J.H. |author5=DeHart, C. |author6=Travaini, A. |year=2016 |title=Density and activity patterns of pumas in hunted and non-hunted areas in central Argentina |journal=Wildlife Research |volume=43 |issue=6 |pages=449–460 |doi=10.1071/WR16056|hdl=11336/44202 |hdl-access=free }}</ref> Cougars displayed a foremost crepuscular and nocturnal activity pattern in a ranching area in southern Argentina.<ref name=Guerisoli2019/> | |||
South of the ], the ] (IUCN) lists the cat in every ] and ] 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=]}}</ref> | |||
] sizes and overall cougar abundance depend on terrain, vegetation, and prey abundance.<ref name="Utah2">{{cite web |author=Cougar Discussion Group |date=1999 |title=Utah Cougar Management Plan (Draft) |url=http://www.wildlife.utah.gov/pdf/cmgtplan.pdf |url-status=live |archive-url=https://web.archive.org/web/20070616200443/http://www.wildlife.utah.gov/pdf/cmgtplan.pdf |archive-date=June 16, 2007 |access-date=May 2, 2007 |publisher=Utah Division of Wildlife Resources}}</ref> Research suggests a lower limit of {{cvt|25|km2}} and upper limit of {{cvt|1300|km2}} of home range for males.<ref name="Utah">{{cite web |url=http://www.wildlife.utah.gov/pdf/cmgtplan.pdf |title=Utah Cougar Management Plan (Draft) |access-date=May 2, 2007 |author=Cougar Discussion Group |date=1999 |publisher=Utah Division of Wildlife Resources |url-status=live |archive-url=https://web.archive.org/web/20070616200443/http://www.wildlife.utah.gov/pdf/cmgtplan.pdf |archive-date=June 16, 2007}}</ref> Large male home ranges of {{cvt|150|to|1000|km2}} with female ranges half that size.<ref name="CanGeo2">{{cite web |title=Cougars in Canada (Just the Facts) |url=http://www.canadiangeographic.ca/Magazine/mj04/indepth/justthefacts.asp |url-status=dead |archive-url=https://web.archive.org/web/20070820172927/http://www.canadiangeographic.ca/Magazine/mj04/indepth/justthefacts.asp |archive-date=August 20, 2007 |access-date=April 2, 2007 |publisher=]}}</ref> One female adjacent to the ] was found with a big range of {{cvt|215|km2}}, necessitated by poor prey abundance.<ref name="Dispersal2">{{cite journal |author1=Sweanor, L. |author2=Logan, K. A. |author3=Hornocker, M. G. |name-list-style=amp |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 |bibcode=2000ConBi..14..798S |s2cid=26735359}}</ref> Research has shown cougar abundances from 0.5 animals to as many as seven per {{cvt|100|km2}}.<ref name="CAP2">{{Cite web |author1=Nowell, K. |author2=Jackson, P. |name-list-style=amp |year=1996 |title=Wild Cats. Status Survey and Conservation Action Plan |url=http://carnivoractionplans1.free.fr/wildcats.pdf |url-status=live |archive-url=https://web.archive.org/web/20070807215533/http://carnivoractionplans1.free.fr/wildcats.pdf |archive-date=August 7, 2007 |access-date=July 27, 2007 |publisher=IUCN/SSC Cat Specialist Group. IUCN, Gland, Switzerland}}</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 ], 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> | |||
Male home ranges include or overlap with females but, at least where studied, not with those of other males. The home ranges of females overlap slightly. Males create scrapes composed of leaves and ] with their hind feet, and ] them with ] and sometimes ].<ref>{{cite journal |author1=Allen, M. L. |name-list-style=amp |author2=Wittmer, H. U. |author3=Wilmers, C. C. |year=2014 |title=Puma communication behaviours: understanding functional use and variation among sex and age classes |journal=Behaviour |volume=151 |issue=6 |pages=819–840 |doi=10.1163/1568539X-00003173}}</ref> When males encounter each other, they vocalize and may engage in violent conflict if neither backs down.<ref name="UWSP22">{{cite web |author1=Hamilton, M. |author2=Hundt, P. |author3=Piorkowski, R. |title=Mountain Lions |url=http://www.uwsp.edu/wildlife/carnivore/Mountain%20Lion%20Natural%20History_files/Mountain%20Lion%20Natural%20History_copy(1).htm |url-status=dead |archive-url=https://web.archive.org/web/20070613114602/http://www.uwsp.edu/wildlife/carnivore/Mountain%20Lion%20Natural%20History_files/Mountain%20Lion%20Natural%20History_copy%281%29.htm |archive-date=June 13, 2007 |access-date=May 10, 2007 |publisher=]}}</ref> | |||
In 2012 research in ], 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 km<sup>2</sup>, that is 0.63 cougars every 100 km<sup>2</sup>.<ref>Research of Nicolás Guarda, supported by ], ], and a private enterprise. See article in Chilean newspaper ], , retrieved on January 28, 2013, in Spanish Language.</ref> | |||
Cougars communicate with various vocalizations. Aggressive sounds include growls, spits, snarls, and hisses. During the mating season, estrus females produce ] or yowls to attract mates, and males respond with similar vocals. Mothers and offspring keep in contact with whistles, chirps, and mews.<ref name="Naughton">{{cite book |author=Naughton, D. |year=2014 |title=The Natural History of Canadian Mammals |publisher=University of Toronto Press |pages=368–373 |isbn=978-1-4426-4483-0}}</ref><ref>{{cite book |author1=Hornocker, M. G. |author2=Negri, S. |name-list-style=amp |title=Cougar: ecology and conservation |url=https://books.google.com/books?id=ZSDUxT3fSYEC&pg=PA114 |date=2009 |publisher=University of Chicago Press |isbn=978-0-226-35344-9 |pages=113–114}}</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> | |||
=== |
===Reproduction and life cycle=== | ||
{{multiple image |align=right |direction=vertical |image1=Puma cub Malibu Springs area National Park Service December 2013.jpg |caption1=North American cougar ] in the ] |image2=Mountain lion kittens.jpg |caption2=Cubs}} | |||
] | |||
] | |||
Aside from humans, no species preys upon mature cougars in the wild, although conflicts with other predators or scavengers occur. The ] ecosystem provides a fruitful microcosm to study inter-predator interaction in North America. Of the three large predators, the massive ] 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 ]s visited 24% of cougar kills in Yellowstone and ], 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=] |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> | |||
Females reach ] at the age of 18 months to three years and are in ] for about eight days of a 23-day cycle; the ] is approximately 91 days.<ref name="CAP" /> Both adult males and females may mate with multiple partners, and a female's litter can have multiple paternities.<ref name="Naughton" /> Copulation is brief but frequent. Chronic stress can result in low reproductive rates in captivity as well as in the field.<ref>{{cite journal |author1=Bonier, F. |author2=Quigley, H. |author3=Austad, S. |name-list-style=amp |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)0322.0.CO;2 |s2cid=85819163}}</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 ], 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 ], 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 ... 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=] |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> | |||
] is 82–103 days long.<ref name="Naughton" /> Only females are involved in parenting. Litter size is between one and six cubs, typically two. Caves and other alcoves that offer protection are used as litter dens. Born blind, cubs are completely dependent on their mother at first and begin to be weaned at around three months of age. As they grow, they 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" /><ref name="Naughton" /> Kitten survival rates are just over one per litter.<ref name="CAP" /> | |||
Both species, meanwhile, are capable of killing mid-sized predators, such as ]s and ]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> | |||
Juveniles remain with their mothers for one to two years.<ref name="Naughton" /> When a female reaches estrous again, her offspring must ] or the male will kill them. Males tend to disperse further than females.<ref name="UWSP">{{cite web |author1=Hamilton, M. |author2=Hundt, P. |author3=Piorkowski, R. |title=Mountain Lions |url=http://www.uwsp.edu/wildlife/carnivore/Mountain%20Lion%20Natural%20History_files/Mountain%20Lion%20Natural%20History_copy(1).htm |url-status=dead |archive-url=https://web.archive.org/web/20070613114602/http://www.uwsp.edu/wildlife/carnivore/Mountain%20Lion%20Natural%20History_files/Mountain%20Lion%20Natural%20History_copy%281%29.htm |archive-date=June 13, 2007 |access-date=May 10, 2007 |publisher=]}}</ref> One study has shown a high ] among cougars that travel farthest from their maternal range, often due to conflicts with other cougars.<ref name="Utah" /> In a study area in ], males dispersed farther than females, traversed large expanses of non-cougar habitat and were probably most responsible for nuclear gene flow between habitat patches.<ref name="Dispersal">{{cite journal |author1=Sweanor, L. |author2=Logan, K. A. |author3=Hornocker, M. G. |s2cid=26735359 |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 |bibcode=2000ConBi..14..798S |name-list-style=amp}}</ref> | |||
Life expectancy in the wild is reported at 8 to 13 years and probably averages 8 to 10; a female of at least 18 years was reported killed by ]s on ].<ref name="CAP" /> Cougars may live as long as 20 years in captivity. Causes of death in the wild include disability and disease, competition with other cougars, starvation, accidents, and, where allowed, hunting. The ] is well-adapted to the cougar.<ref>{{cite journal |author1=Biek, R. |author2=Rodrigo, A. G. |author3=Holley, D. |author4=Drummond, A. |author5=Anderson Jr., C. R. |author6=Ross, H. A. |author7=Poss, M. |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=9578–9589 |doi=10.1128/JVI.77.17.9578-9589.2003 |pmid=12915571 |pmc=187433 |name-list-style=amp}}</ref> | |||
As with any predator at or near the top of its ], 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 ] observed that the population of ], a favored cougar prey, was declining while the population of the less frequently preyed-upon ] 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 ], an endangered species ] 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) . '']''.</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> | |||
==Conservation== | |||
In the southern part of South America, the puma is a top level predator that has controlled the population of ] and other species since prehistoric times.<ref>Busch, Robert H. ''The Cougar Almanac.'' New York, 2000, pg 94. ISBN 1592282954.</ref> | |||
]]] | |||
The cougar has been listed as ] on the ] since 2008. However, it is also listed on ].<ref name="iucn" /> Hunting it is prohibited in California, ], ], ], ], ], ], Colombia, ], ], ], Brazil, Chile, ], ] and most of Argentina. Hunting is regulated in Canada, Mexico, ], and the United States.<ref name="CAP" /> Establishing ]s and protecting sufficient range areas are critical for the sustainability of cougar populations. Research simulations showed that it faces a low extinction risk in areas larger than {{cvt|2200|km2}}. Between one and four new individuals entering a population per decade markedly increases persistence, thus highlighting the importance of habitat corridors.<ref>{{cite journal |last=Beier |first=P. |s2cid=55580710 |date=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 |bibcode=1993ConBi...7...94B}}</ref> | |||
==Hybrids== | |||
] | |||
{{Main article|Pumapard}} | |||
A pumapard is a ] animal resulting from a union between a cougar and a ]. Three sets of these hybrids were bred in the late 1890s and early 1900s by ] at his animal park in ], Germany. Most did not reach adulthood. One of these was purchased in 1898 by ]. 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. | |||
The ] population is afforded protection under the ].<ref>{{cite web |url=http://www.fws.gov/endangered/i/a/saa48.html |title=Eastern Cougar |access-date=May 20, 2007 |work=Endangered and Threatened Species of the Southeastern United States (The Red Book) |year=1991 |publisher=]| archive-url = https://web.archive.org/web/20070403000340/http://www.fws.gov/endangered/i/a/saa48.html| archive-date=April 3, 2007}}</ref><ref>{{cite web |url=http://www.fws.gov/endangered/i/a/saa05.html |title=Florida Panther |access-date=June 7, 2007 |work=Endangered and Threatened Species of the Southeastern United States (The Red Book) |year=1993 |publisher=] |archive-url = https://web.archive.org/web/20070604163840/http://www.fws.gov/endangered/i/a/saa05.html|archive-date = June 4, 2007}}</ref> The Texas Mountain Lion Conservation Project was launched in 2009 and aimed at raising local people's awareness of the status and ecological role of the cougar and mitigating conflict between landowners and cougars.<ref>{{cite web |date=2007–2013 |title=Texas Mountain Lion Conservation Project |website=Balanced Ecology Inc. |url=http://balancedecology.org/MountainLionWebSite/Mountain_Lion_Conservation_Project.html |access-date=January 18, 2010 |archive-date=July 29, 2020 |archive-url=https://web.archive.org/web/20200729010958/http://balancedecology.org/MountainLionWebSite/Mountain_Lion_Conservation_Project.html |url-status=live}}</ref> | |||
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 ]. 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> | |||
The cougar is threatened by ], ], and depletion of its prey base due to ]. Hunting is legal in the western United States. In Florida, heavy traffic causes frequent accidents involving cougars. Highways are a major barrier to the dispersal of cougars.<ref name="iucn" /> The cougar populations in California are becoming fragmented with the increase in human population and infrastructure growth in the state.<ref>{{Cite journal |last1=Ernest |first1=H. B. |last2=Vickers |first2=T. W. |last3=Morrison |first3=S. A. |last4=Buchalski |first4=M. R. |last5=Boyce |first5=W. M. |title=Fractured genetic connectivity threatens a southern California Puma (''Puma concolor'') population |journal=PLOS ONE |volume=9 |issue=10 |pages=e107985 |doi=10.1371/journal.pone.0107985 |pmc=4189954 |pmid=25295530 |year=2014 |bibcode=2014PLoSO...9j7985E |doi-access=free}}</ref> | |||
==Conservation status== | |||
The ] (IUCN) currently lists the cougar as a "]" species. The cougar is regulated under ] of the ] (CITES),<ref>{{cite web |url=http://www.cites.org/eng/app/appendices.php|title=Appendices I, II and III |accessdate=May 24, 2007 |publisher=]}}</ref> rendering illegal international trade in specimens or parts. | |||
] in proximity of {{cvt|5|km2}} of cougar habitat is pronounced in areas with a median human density of {{cvt|32.48|/km2|/sqmi|disp=preunit|inhabitants|inhabitants}} and a median livestock population density of {{cvt|5.3|/km2|/sqmi|disp=preunit|heads|heads}}. Conflict is generally lower in areas more than {{cvt|16.1|km}} away from roads and {{cvt|27.8|km}} away from settlements.<ref>{{cite journal |author1=Guerisoli, M. D. L. M. |name-list-style=amp |author2=Luengos Vidal, E. |author3=Caruso, N. |author4=Giordano, A. J. |author5=Lucherini, M. |year=2021 |title=Puma–livestock conflicts in the Americas: A review of the evidence |journal=Mammal Review |volume=51 |issue=2 |pages=228–246 |doi=10.1111/mam.12224 |s2cid=226336427}}</ref> | |||
] | |||
In the United States east of the ], the only unequivocally known cougar population is the ]. Until 2011, the ] (USFWS) recognized both an ] (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=] |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 ].<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=]| 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=] |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> | |||
==Relationships with humans== | |||
This uncertainty has been recognized by Canadian authorities. The Canadian federal agency called ] 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 ], Quebec, ] and ], 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>. Cosewic.gc.ca. Retrieved on September 15, 2011.</ref><ref>. Naturecanada.ca. Retrieved on September 15, 2011.</ref> | |||
===Attacks on humans=== | |||
====In North America==== | |||
{{See also|List of fatal cougar attacks in North America}} | |||
] | |||
Due to the ], cougar ]s increasingly overlap with areas inhabited by humans.<ref>{{Cite web |date=November 20, 2023 |title=Mountain lion caught on camera in Tesoro Viejo in Madera County |url=https://abc30.com/tesoro-viejo-madera-county-mountain-lion-camera-spotting/14085089/ |access-date=November 21, 2023 |website=ABC30 Fresno |language=en |archive-date=November 21, 2023 |archive-url=https://web.archive.org/web/20231121004735/https://abc30.com/tesoro-viejo-madera-county-mountain-lion-camera-spotting/14085089/ |url-status=live}}</ref> 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 |pmid=14518628 |doi=10.1580/1080-6032(2003)142.0.CO;2|doi-access=free}}</ref> In a 10-year study in New Mexico of wild cougars who were not habituated to humans, the animals did not exhibit threatening behavior to researchers who approached closely (median distance=18.5 m; 61 feet) except in 6% of cases; {{frac|14|16}} of those were females with cubs.<ref>{{cite journal |title=Puma responses to close approaches by researchers |first1=Linda L. |last1=Sweanor |first2=Kenneth A. |last2=Logan |first3=Maurice G. |last3=Hornocker |journal=Wildlife Society Bulletin |volume=33 |issue=3 |pages=905–913 |year=2005 |doi=10.2193/0091-7648(2005)332.0.CO;2 |s2cid=86209378 |doi-access=free}}</ref> Attacks on people, livestock, and pets may occur when a puma ] 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" /> | |||
The cougar is also protected across much of the rest of its range. As of 1996, cougar hunting was prohibited in ], Brazil, ], Chile, ], Costa Rica, ], Guatemala, ], Nicaragua, ], Paraguay, ], Venezuela, and ]. The cat had no reported legal protection in ], El Salvador, and ].<ref name="CAP"/> Regulated cougar hunting is still common in the United States and Canada, although they are protected from all hunting in the ]; it is permitted in every U.S. state from the ] 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>. Tpwd.state.tx.us (July 16, 2007). Retrieved on September 15, 2011.</ref> Killed animals are not required to be reported to ]. Conservation work in Texas is the effort of a non-profit organization, ], as part of their Texas Mountain Lion Conservation Project.<ref>. Balanced Ecology Inc. balancedecology.org</ref> Cougars are generally hunted with packs of ], 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 ] 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 ] 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> | |||
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 the United States and Canada |work=Wildlife Society Bulletin |year=1991 |access-date=May 20, 2007 |publisher=Northern Arizona University |url-status=dead |archive-url=https://web.archive.org/web/20120622033418/http://users.frii.com/mytymyk/lions/beier.htm |archive-date=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 |access-date=May 20, 2007 |publisher=Arizona Game and Fish Department |url-status=dead |archive-url=https://web.archive.org/web/20070518181449/http://www.azgfd.gov/w_c/mtn_lion_attacks.shtml |archive-date=May 18, 2007}}</ref> | |||
Conservation threats to the species include persecution as a pest animal, ] and ], and depletion of their prey base. ]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 km<sup>2</sup> (850 sq mi) or more. As few as one to four new animals entering a population per decade markedly increases persistence, foregrounding the importance of habitat corridors.<ref>{{cite journal |last=Beier |first=Paul |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> | |||
Within North America, the distribution of attacks is not uniform. The heavily populated state of California saw a dozen attacks from 1986 to 2004 (after just three from 1890 to 1985), including three fatalities.<ref name="California">{{cite web |url=http://www.dfg.ca.gov/news/issues/lion.html |title=Mountain Lions in California |access-date=May 20, 2007 |year=2004 |publisher=California Department of Fish and Game| archive-url = https://web.archive.org/web/20070430012010/http://www.dfg.ca.gov/news/issues/lion.html| archive-date = April 30, 2007}}</ref> In March 2024, two brothers in California were attacked by a male cougar, with one being fatally wounded; it was the state's first fatal attack in 20 years.<ref>{{Cite news |last1=Razek |first1=Raja |last2=Flynn |first2=Jessica |date=March 25, 2024 |title=First fatal mountain lion attack in California in 20 years leaves one man dead, brother injured, authorities say |url=https://www.ctvnews.ca/climate-and-environment/first-fatal-mountain-lion-attack-in-california-in-20-years-leaves-one-man-dead-brother-injured-authorities-say-1.6821205 |access-date=March 25, 2024 |work=CTV News |archive-date=March 25, 2024 |archive-url=https://web.archive.org/web/20240325170316/https://www.ctvnews.ca/climate-and-environment/first-fatal-mountain-lion-attack-in-california-in-20-years-leaves-one-man-dead-brother-injured-authorities-say-1.6821205 |url-status=live}}</ref><ref>{{Cite news |last1=Dowd |first1=Katie |last2=Bartlett |first2=Amanda |title=Mountain lion kills man in Northern California for first time in 30 years |url=https://www.sfgate.com/bayarea/article/mountain-lion-kills-man-northern-california-19365657.php |access-date=March 28, 2024 |work=SFGATE |language=en |archive-date=March 28, 2024 |archive-url=https://web.archive.org/web/20240328003539/https://www.sfgate.com/bayarea/article/mountain-lion-kills-man-northern-california-19365657.php |url-status=live}}</ref> Washington state was the site of a fatal attack in 2018, its first since 1924.<ref>{{cite news |last1=Fisher |first1=Michelle |title=Victim in Deadly Washington State Cougar Attack Had Boston Ties |url=http://boston.cbslocal.com/2018/05/21/cougar-attack-victim-sj-brooks-washington-seattle-boston/|access-date=May 23, 2018 |publisher=CBS BOston |date=May 21, 2018|archive-date=May 23, 2018|archive-url=https://web.archive.org/web/20180523173303/http://boston.cbslocal.com/2018/05/21/cougar-attack-victim-sj-brooks-washington-seattle-boston/|url-status=live}}</ref> Lightly populated New Mexico reported an attack in 2008, the first there since 1974.<ref>]: , June 23, 2008; , June 25, 2008; , July 1, 2008</ref> | |||
On March 2, 2011, the ] declared the Eastern cougar (''Puma concolor couguar'') officially extinct.<ref>. Fws.gov. Retrieved on September 15, 2011.</ref> | |||
As with many predators, a cougar may attack if cornered, if a fleeing human stimulates their instinct to chase, or if a person "]". Standing still may cause the cougar to consider a person easy prey.<ref>{{cite magazine |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 |magazine=Scientific American|access-date=March 10, 2012 |date=April 14, 2009|url-status=live|archive-url=https://web.archive.org/web/20110319160359/https://www.scientificamerican.com/article.cfm?id=should-you-run-or-freeze-when-you-see-a-mountain-lion|archive-date=March 19, 2011}}</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 |access-date=May 28, 2007 |publisher=], Ministry of Environment |url-status=dead |archive-url=https://web.archive.org/web/20070823061650/http://www.env.gov.bc.ca/wld/documents/cougsf.htm |archive-date=August 23, 2007}}</ref> | |||
==Relationships with humans== | |||
When cougars do attack, they usually employ their characteristic neck bite, attempting to position their teeth between the ] and into the ]. 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 before 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 ], particularly on ], 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=https://www.nbcnews.com/id/wbna27756765 |agency=Associated Press |work=NBC News |date=November 16, 2008 |title=Neighbor saves Miami teen from cougar|access-date=February 11, 2012|archive-date=September 23, 2020|archive-url=https://web.archive.org/web/20200923224535/http://www.nbcnews.com/id/27756765|url-status=live}}</ref><ref>{{Cite news |title=2-Year-Old Boy Hurt In Pet Cougar Attack |url=https://www.nytimes.com/1995/06/04/nyregion/2-year-old-boy-hurt-in-pet-cougar-attack.html |date=June 4, 1995 |work=The New York Times|url-status=live|archive-url=https://web.archive.org/web/20170625060218/http://www.nytimes.com/1995/06/04/nyregion/2-year-old-boy-hurt-in-pet-cougar-attack.html|archive-date=June 25, 2017}}</ref> | |||
===In mythology=== | |||
The grace and power of the cougar have been widely admired in the cultures of the ]. The Inca city of ] 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 ] people represented the puma often in their ceramics.<ref>Berrin, Katherine & Larco Museum. ''The Spirit of Ancient Peru:Treasures from the ].'' New York: ], 1997.</ref> The sky and thunder god of the Inca, ], 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> | |||
Research on new wildlife collars may reduce human-animal conflicts by predicting when and where predatory animals hunt. This may save the lives of humans, pets, and livestock, as well as the lives of these large predatory mammals that are important to the balance of ecosystems.<ref>Williams, Terrie M. (November 6, 2014) {{webarchive|url=https://web.archive.org/web/20141109194703/http://www.latimes.com/opinion/op-ed/la-oe-williams-wildlife-coburn-wastebook-20141107-story.html |date=November 9, 2014 }} '']''</ref> | |||
In North America, mythological descriptions of the cougar have appeared in the stories of the ] ("Ho-Chunk" or "Winnebago") of ] and ]<ref> The Encyclopedia of Hočąk (Winnebago) Mythology. Retrieved: 2009/12/08.</ref> and the ], amongst others. To the ] and ] of ], 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=] ]}}</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> | |||
=== |
====In South America==== | ||
Cougars in the ] of South America are reputed to be extremely reluctant to attack people; in legend, they defended people against jaguars.<ref>{{cite web |last1=Chébez |first1=J. C. |last2=Nigro |first2=R. Á. |title=Aportes preliminaares para un plan de conservación y manejo del Puma (''Puma concolor'') en la República Argentina |language=es |url=http://maaz.ihmc.us/rid=1PP8CK25F-1FYP3P-2VPC/chebez-y-nigro-aportes-para-un-plan-de-conservacion-y-m.pdf |access-date=February 23, 2018 |archive-date=February 26, 2018 |archive-url=https://web.archive.org/web/20180226211612/http://maaz.ihmc.us/rid=1PP8CK25F-1FYP3P-2VPC/chebez-y-nigro-aportes-para-un-plan-de-conservacion-y-m.pdf |url-status=live}}</ref> The nineteenth-century naturalists ]<ref>{{cite book |last=Azara |first=F. d. |title=The Natural History of the Quadrupeds of Paraguay and the River la Plata |pages=207–208 |publisher=Adam and Charles Black |location=], ] |year=1838 |url=https://books.google.com/books?id=QNc0AQAAMAAJ&pg=PA207 |quote=I have not heard that they have assaulted or attempted to assault man, nor dogs and boys, even when it encounters them asleep |access-date=June 6, 2020 |archive-date=May 5, 2024 |archive-url=https://web.archive.org/web/20240505151936/https://books.google.com/books?id=QNc0AQAAMAAJ&pg=PA207#v=onepage&q&f=false |url-status=live}}</ref> and ]<ref>{{cite book |last=Hudson |first=W. H. |title=The Naturalist in La Plata |pages=31–49 |publisher=Chapman and Hall Ltd |year=1892 |location=London |url=https://archive.org/stream/naturalistinlapl1892huds#page/n43/mode/2up|access-date=February 15, 2018 |quote=This, however, is not a full statement of the facts; the puma will not even defend itself against man}}</ref> thought that attacks on people, even children or sleeping adults, did not happen. Hudson, citing anecdotal evidence from hunters, claimed that pumas were positively inhibited from attacking people, even in self-defense. Attacks on humans, although exceedingly rare, have occurred.<ref>{{cite book |last=Roosevelt |first=Theodore |title=Through the Brazilian Wilderness |pages=27–8 |publisher=Charles Scribner's Sons |year=1914 |location=New York |url=https://archive.org/stream/cu31924086561119#page/n53/mode/2up|access-date=February 15, 2018}}</ref><ref>{{cite book |last1=Young |first1=S. P. |last2=Goldman |first2=E. A. |title=The Puma: Mysterious American Cat |year=1964 |publisher=Dover Publications Inc |url=https://archive.org/details/pumamysteriousam00youn/page/99 |location=New York |pages=99, 103–105 |url-access=registration}}</ref> | |||
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 million cattle & calves in Texas), 253 Mohair goats, 302 Mohair kids, 445 sheep (0.02% of a total of 2.0 million sheep & lambs in Texas) and 562 lambs (0.04% of 1.2 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 ] 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> | |||
An early, authenticated, non-fatal case occurred near ], Patagonia, in 1877 when a female mauled the Argentine scientist ]; Moreno afterward showed the scars to ]. In this instance, however, Moreno had been wearing a ]-hide ] round his neck and head as protection against the cold;<ref>{{cite book |last=Roosevelt |first=Theodore |title=Through the Brazilian Wilderness |publisher=Charles Scribner's Sons |year=1914 |location=New York |url=https://archive.org/stream/cu31924086561119#page/n53/mode/2up|access-date=February 15, 2018 |pages=26–31}}</ref> in Patagonia the guanaco is the puma's chief prey animal.<ref>{{cite journal |title=Response of pumas (''Puma concolor'') to migration of their primary prey in Patagonia |last1=Gelin |first1=Maria L. |last2=Branch |first2=Lyn C. |last3=Thornton |first3=Daniel H. |last4=Novaro |first4=Andrés J. |last5=Gould |first5=Matthew J. |last6=Caragiulo |first6=Anthony |journal=PLOS ONE |volume=12 |issue=12 |pages=e0188877 |doi=10.1371/journal.pone.0188877 |pmid=29211753 |pmc=5718558 |year=2017 |bibcode=2017PLoSO..1288877G |doi-access=free}}</ref> Another authenticated case occurred in 1997 in ] in northeastern Argentina, when the 20-month-old son of a ranger was killed by a female puma. Forensic analysis found specimens of the child's hair and clothing fibers in the animal's stomach. The ] is the puma's chief prey in this area. Despite prohibitory signs, coatis are hand-fed by tourists in the park, causing unnatural approximation between cougars and humans. This particular puma had been raised in captivity and released into the wild.<ref>{{cite journal |last=Faletti |first=Dra. Alicia |title=Caso Ignacio Terán Luna |journal=Revista Química Viva |year=2013 |volume=12 |issue=2 |language=es |issn=1666-7948 |url=http://www.quimicaviva.qb.fcen.uba.ar/contratapa/ignacio.htm |access-date=February 16, 2018 |archive-date=February 20, 2018 |archive-url=https://web.archive.org/web/20180220024430/http://www.quimicaviva.qb.fcen.uba.ar/contratapa/ignacio.htm |url-status=live}}</ref> On March 13, 2012, Erica Cruz, a 23-year-old shepherdess was found dead in a mountainous area near ], Salta Province, in northwestern Argentina.<ref>{{Cite web |last=Redacción |date=2012 |title=Un puma mató a una pastora en Salta |url=https://www.rionegro.com.ar/un-puma-mato-a-una-pastora-en-salta-OBRN_835496/ |access-date=November 30, 2021 |website=Diario Río Negro |language=es |archive-date=November 30, 2021 |archive-url=https://web.archive.org/web/20211130020602/https://www.rionegro.com.ar/un-puma-mato-a-una-pastora-en-salta-OBRN_835496/ |url-status=live}}</ref> Claw incisions, which severed a jugular vein, indicated that the attacker was a felid; differential diagnosis ruled out other possible perpetrators.{{efn|There are no jaguars in the area; other felids were too small to kill humans.}} There were no bite marks on the victim, who had been herding goats.<ref>{{cite journal |title=Ataque fatal en humano, por puma (''Puma concolor'') |last1=Portelli |first1=C.M. |last2=Eveling |first2=C.R. |last3=Lamas |first3=J. |last4=Mamaní |first4=P.J. |journal=Cuadernos de Medicina Forense |volume=18 |issue=3–4 |year=2012 |pages=139–142 |doi=10.4321/S1135-76062012000300008 |language=es |url=http://scielo.isciii.es/scielo.php?script=sci_abstract&pid=S1135-76062012000300008 |access-date=February 26, 2018 |doi-access=free |archive-date=February 26, 2018 |archive-url=https://web.archive.org/web/20180226211656/http://scielo.isciii.es/scielo.php?script=sci_abstract&pid=S1135-76062012000300008 |url-status=live}}</ref> In 2019 in ] an elderly man was badly injured by a cougar after he attempted to defend his dog from it, while in neighboring Chile a 28-year-old woman was attacked and killed in ], in ], on October 20, 2020.<ref name="BioBioexp">{{Cite news |title=Expertos tras eventual ataque de un puma en Corral: 'Es inusual, no buscan enfrentar a los humanos' |url=https://www.biobiochile.cl/noticias/sociedad/animales/2020/10/21/expertos-tras-ataque-mortal-de-un-puma-en-corral-hay-que-tener-conciencia-es-un-animal-salvaje.shtml |last=Contreras |first=E. |date=2020|access-date=December 4, 2020 |work=] |language=es|archive-date=November 1, 2020|archive-url=https://web.archive.org/web/20201101014350/https://www.biobiochile.cl/noticias/sociedad/animales/2020/10/21/expertos-tras-ataque-mortal-de-un-puma-en-corral-hay-que-tener-conciencia-es-un-animal-salvaje.shtml|url-status=live}}</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> | |||
Fatal attacks by other carnivores, such as feral dogs, can be misattributed to cougars without appropriate forensic knowledge.<ref>{{cite journal |last1=Fonseca |first1=G. M. |last2=Palacios |first2=R. |title=An Unusual Case of Predation: Dog Pack or Cougar Attack? |journal=Journal of Forensic Sciences |volume=58 |issue=1 |year=2013 |pages=224–227 |doi=10.1111/j.1556-4029.2012.02281.x |pmid=22971181 |hdl=11336/10589 |s2cid=205771079|hdl-access=free}}</ref> | |||
===Attacks on humans=== | |||
===Predation on domestic animals=== | |||
====North American subspecies==== | |||
]'', a 1920s silent film created by the ] which explains the procedures to successfully hunt livestock-threatening cougars]] | |||
{{See also|List of fatal cougar attacks in North America}} | |||
] | |||
During the early years of ranching, cougars were considered on par with wolves in destructiveness. According to figures in ] in 1990, 86 calves (0.0006% of Texas's 13.4 million cattle and calves), 253 mohair goats, 302 mohair kids, 445 sheep (0.02% of Texas's 2 million sheep and lambs) and 562 lambs (0.04% of Texas's 1.2 million lambs) were confirmed to have been killed by cougars that year.<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 | access-date=September 11, 2009 | url-status=live | archive-url=https://web.archive.org/web/20110608154849/http://usda.mannlib.cornell.edu/usda/nass/Catt//1990s/1990/Catt-07-27-1990.pdf | archive-date=June 8, 2011 |df=mdy-all}}</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 | access-date=September 11, 2009 | url-status=live | archive-url=https://web.archive.org/web/20110608154900/http://usda.mannlib.cornell.edu/usda/nass/SheeGoat//1990s/1990/SheeGoat-02-02-1990.pdf | archive-date=June 8, 2011 |df=mdy-all}}</ref> In ] in 1992, cougars were confirmed to have killed nine calves, one horse, four foals, five goats, 318 sheep, and 400 lambs. In both reports, sheep were the most frequently attacked. Some instances of ] 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 |access-date=July 10, 2008 |archive-url=https://archive.today/20120720162250/http://www.aws.vcn.com/mountain_lion_fact_sheet.html |archive-date=July 20, 2012 |url-status=dead}}</ref> A cougar's killing bite is applied to the back of the neck, head, or ] and the cat inflicts puncture marks with its claws usually seen on the sides and underside of the prey, sometimes also shredding the prey as it holds on. Coyotes also typically bite the throat, but the work of a cougar is generally clean, while bites inflicted by coyotes and dogs leave ragged edges. The size of the tooth puncture marks also helps distinguish kills made by cougars from those made by smaller predators.<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 |access-date=August 3, 2008 |url-status=dead |archive-url=https://web.archive.org/web/20110111165333/http://agrilife.org/texnatwildlife/predators-and-predation/predator-species/cougars/ |archive-date=January 11, 2011}}</ref> | |||
The pertinent North American subspecies is ''P. concolor couguar''. | |||
Remedial hunting appears to have the paradoxical effect of increased livestock predation and complaints of human-cougar conflicts. In a 2013 study, the most important predictor of cougar problems was the remedial hunting of cougars the previous year. Each additional cougar on the landscape increased predation and human-cougar complaints by 5%, but each animal killed during the previous year increased complaints by 50%. The effect had a dose-response relationship with very heavy (100% removal of adult cougars) 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 |bibcode=2013PLoSO...879713P|doi-access=free}}</ref> This effect is attributed to the removal of older cougars that have learned to avoid people and their replacement by younger males that react differently to humans. Remedial hunting enables younger males to enter the former territories of the older animals.<ref>{{cite journal |title=Cougar attacks on humans in the United States and Canada |author=Beier, Paul |journal=Wildlife Society Bulletin |volume=19 |issue=4 |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> Predation by cougars on dogs "is widespread, but occurs at low frequencies".<ref>{{cite book |editor-last1=Gompper |editor-first=Matthew E. |author1=Butler, James R. A. |author2=Linnell, John D. C. |author3=Morrant, Damian |author4=Athreya, Vidya |author5=Lescureux, Nicolas |author6=McKeown, Adam |chapter=5: Dog eat dog, cat eat dog: social-ecological dimensions of dog predation by wild carnivores |title=Free-ranging dogs and wildlife conservation |date=2014 |publisher=Oxford University Press |page=127}}</ref> | |||
Due to the ], cougar ]s increasingly overlap with areas inhabited by ]. 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)142.0.CO;2}}</ref> Attacks on people, livestock, and pets may occur when a puma ] 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"/> | |||
===In mythology=== | |||
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> | |||
The grace and power of the cougar have been widely admired in the cultures of the ]. The ] city of ] 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 ] people often represented the cougar in their ceramics.<ref>Berrin, Katherine & Larco Museum. ''The Spirit of Ancient Peru: Treasures from the ]''. New York: Thames and Hudson, 1997.</ref> The sky and thunder god of the Inca, ], 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 |access-date=May 22, 2007 |work=Electronic Journal of Folklore |publisher=Estonian Folklore Institute |url-status=live |archive-url=https://web.archive.org/web/20070630045739/http://www.folklore.ee/folklore/vol12/inca.htm |archive-date=June 30, 2007}}</ref> | |||
In North America, mythological descriptions of the cougar have appeared in the stories of the ] ("Ho-Chunk" or "Winnebago") of ] and ]<ref>; {{webarchive|url=https://web.archive.org/web/20100421212756/http://hotcakencyclopedia.com/ho.Cougars.html |date=April 21, 2010 }}. ''The Encyclopedia of Hočąk (Winnebago) Mythology''. Retrieved: 2009/12/08.</ref> and the ], among others. To the ] and ] of the Southwestern United States, 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 |access-date=April 11, 2009 |publisher=] ] |url-status=live |archive-url=https://web.archive.org/web/20090420203146/http://www.aphis.usda.gov/wildlife_damage/nwrc/publications/living/cougar.pdf |archive-date=April 20, 2009}}</ref> The ] and ] believe that the cougar lived in the underworld and was wicked, whereas it was a sacred animal among the ].<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> | |||
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 ] reported an attack in 2008, the first there since 1974.<ref>]: , June 23, 2008; , June 25, 2008; , July 1, 2008</ref> | |||
==See also == | |||
As with many predators, a cougar may attack if cornered, if a fleeing human stimulates their instinct to chase, or if a person "]". 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=], Ministry of Environment}}</ref> | |||
{{Portal|Cats|Mammals}} | |||
* ]—hybrid of cougar and ] | |||
* ] | |||
* ]—hybrid of cougar and ] | |||
== Explanatory notes == | |||
When cougars do attack, they usually employ their characteristic neck bite, attempting to position their teeth between the ] and into the ]. 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 ], particularly on ] 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> | |||
{{Notelist}} | |||
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) '']''</ref> | |||
====Other subspecies==== | |||
The puma of the plains of South America (''P. concolor cabrerae'',<ref>Wozencraft, W. Christopher (2005) in Don E. Wilson, DeeAnn M. Reeder (eds.) ''Mammal Species of the World'', 3rd ed. JHU Press. ISBN 0801882214.</ref> called the ] by North Americans) is noted for its extreme unwillingness to attack humans. According to the Anglo-Argentine naturalist ]<ref name="gutenberg.org">Hudson, W. H. (1892) , 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== | ==References== | ||
{{Reflist|30em}} | {{Reflist|30em}} | ||
==Further reading== | == Further reading == | ||
* Mark Elbroch: ''The Cougar Conundrum: Sharing the World with a Successful Predator''. Island Press, 2020, {{ISBN|9781610919982}}. | |||
{{Refbegin|30em}} | |||
* Maurice Hornocker (ed.), Sharon Negri (ed.): ''Cougar: Ecology and Conservation''. University of Chicago Press, 2009, {{ISBN|9780226353470}}. | |||
* {{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}} | |||
*Kenneth A. Logan, Linda L. Sweanor: ''Desert Puma: Evolutionary Ecology And Conservation Of An Enduring Carnivore''. Island Press, 2001, {{ISBN|9781610910583}}. | |||
* {{cite book |last1=Bolgiano |first1=Chris |title=Mountain Lion:An Unnatural History of Pumas and People |place=] |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}} | |||
* Paula Wild: ''The Cougar: Beautiful, Wild and Dangerous''. Douglas and McIntyre, 2013, {{ISBN|9781771620031}}. | |||
* {{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== | ==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}} | {{NIE Poster|year=1905|Puma}} | ||
{{Commons and category|Puma concolor|Puma concolor}} | |||
{{Spoken Misplaced Pages|Cougar1.ogg|November 6, 2008}} | |||
{{Wikispecies|Puma concolor}} | |||
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{{Spoken Misplaced Pages|Cougar1.ogg|date=November 6, 2008}} | |||
* {{cite web |url=http://www.catsg.org/index.php?id=94 |website=IUCN/SSC Cat Specialist Group |title=Cougar ''Puma concolor''}} | |||
* : How to identify cougar tracks in the wild | * : How to identify cougar tracks in the wild | ||
* |
* {{cite web |url=http://animals.nationalgeographic.com/animals/mammals/mountain-lion/?rptregcta=reg_free_np&rptregcampaign=20130924_rw_membership_r3p_c1#close-modal |archive-url=https://web.archive.org/web/20131103143721/http://animals.nationalgeographic.com/animals/mammals/mountain-lion/?rptregcta=reg_free_np&rptregcampaign=20130924_rw_membership_r3p_c1#close-modal |url-status=dead |archive-date=November 3, 2013 |title=Puma sounds |publisher=National Geographic Society |date=September 10, 2010}} | ||
* | * | ||
* | * | ||
* | * | ||
* | * , formerly the "Eastern Cougar Foundation" | ||
* | * ; {{Webarchive|url=https://web.archive.org/web/20180731190549/http://www.cougarnet.org/ |date=July 31, 2018 }} | ||
* |
* {{cite web |url=http://www.mountainlion.org/about_the_foundation.asp |website=Mountain Lion Foundation |title=Saving America's Lion |access-date=December 11, 2010 |archive-date=July 31, 2018 |archive-url=https://web.archive.org/web/20180731153509/http://www.mountainlion.org/about_the_foundation.asp |url-status=dead}} | ||
* : Sightings of cougars in Michigan | * : Sightings of cougars in Michigan | ||
* . A Definitive Resource About Cougars: Comprehensive, non-profit ] site with extensive information about cougars, from how to live safely in cougar country, to science abstracts, hunting regulations, state-by-state cougar management/policy info, and rare photos and videos of wild cougars. | |||
* | |||
* A Definitive Resource About Cougars] Comprehensive, non-profit ] site with extensive information about cougars, from how to live safely in cougar country, to science abstracts, hunting regulations, state-by-state cougar management/policy info, and rare photos and videos of wild cougars. | |||
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Latest revision as of 21:26, 16 December 2024
Large species of cat native to the Americas For other uses, see Cougar (disambiguation). "Mountain lion" redirects here. For other uses, see Mountain lion (disambiguation). Not to be confused with Lion. "Catamount" redirects here. For other uses, see Catamount (disambiguation).
Cougar Temporal range: 1.2–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓ Early Pleistocene – Holocene | |
---|---|
A North American cougar in Glacier National Park, United States | |
Conservation status | |
Least Concern (IUCN 3.1) | |
CITES Appendix II (CITES) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Carnivora |
Suborder: | Feliformia |
Family: | Felidae |
Subfamily: | Felinae |
Genus: | Puma |
Species: | P. concolor |
Binomial name | |
Puma concolor (Linnaeus, 1771) | |
Subspecies | |
Also see text | |
Cougar range (without recent confirmations across northern Canadian territories, eastern U.S. states, and Alaska) |
The cougar (Puma concolor) (/ˈkuːɡər/, KOO-gər), also known as the panther, mountain lion, catamount and puma, is a large cat native to the Americas. It inhabits North, Central and South America, making it the most widely distributed wild, terrestrial mammal in the Western Hemisphere, and one of the most widespread in the world. Its range spans the Yukon, British Columbia and Alberta provinces of Canada, the Rocky Mountains and areas in the western United States. Further south, its range extends through Mexico to the Amazon Rainforest and the southern Andes Mountains in Patagonia. It is an adaptable generalist species, occurring in most American habitat types. It prefers habitats with dense underbrush and rocky areas for stalking but also lives in open areas.
The cougar is largely solitary. Its activity pattern varies from diurnality and cathemerality to crepuscularity and nocturnality between protected and non-protected areas, and is apparently correlated with the presence of other predators, prey species, livestock and humans. It is an ambush predator that pursues a wide variety of prey. Ungulates, particularly deer, are its primary prey, but it also hunts rodents. It is territorial and lives at low population densities. Individual home ranges depend on terrain, vegetation and abundance of prey. While large, it is not always the dominant apex predator in its range, yielding prey to other predators. It is reclusive and mostly avoids people. Fatal attacks on humans are rare but increased in North America as more people entered cougar habitat and built farms.
The cougar is listed as Least Concern on the IUCN Red List. Intensive hunting following European colonization of the Americas and ongoing human development into cougar habitat has caused populations to decline in most parts of its historical range. In particular, the eastern cougar population is considered to be mostly locally extinct in eastern North America since the early 20th century, with the exception of the isolated Florida panther subpopulation.
Naming and etymology
The word cougar is borrowed from the Portuguese çuçuarana, via French; it was originally derived from the Tupi language. A current form in Brazil is suçuarana. In the 17th century, Georg Marcgrave named it cuguacu ara. Marcgrave's rendering was reproduced in 1648 by his associate Willem Piso. Cuguacu ara was then adopted by John Ray in 1693. In 1774, Georges-Louis Leclerc, Comte de Buffon converted cuguacu ara to cuguar, which was later modified to "cougar" in English.
The cougar holds the Guinness record for the animal with the greatest number of names, with over 40 in English alone. "Puma" is the common name used in Latin America and most parts of Europe. The term puma is also sometimes used in the United States. The first use of puma in English dates to 1777, introduced from Spanish from the Quechua language. In the western United States and Canada, it is also called "mountain lion", a name first used in writing in 1858. Other names include "panther" (although it does not belong to the genus Panthera) and "catamount" (meaning "cat of the mountains").
Taxonomy and evolution
Felis concolor was the scientific name proposed by Carl Linnaeus in 1771 for a cat with a long tail from Brazil. The specific epithet of the name, "concolor", is Latin for "of uniform color". It was placed in the genus Puma by William Jardine in 1834. This genus is part of the Felinae. The cougar is most closely related to the jaguarundi and the cheetah.
Subspecies
Following Linnaeus's first scientific description of the cougar, 32 cougar zoological specimens were described and proposed as subspecies until the late 1980s. Genetic analysis of cougar mitochondrial DNA indicates that many of these are too similar to be recognized as distinct at a molecular level but that only six phylogeographic groups exist. The Florida panther samples showed a low microsatellite variation, possibly due to inbreeding. Following this research, the authors of Mammal Species of the World recognized the following six subspecies in 2005:
- P. c. concolor (Linnaeus, 1771) includes the synonyms bangsi, incarum, osgoodi, soasoaranna, sussuarana, soderstromii, suçuaçuara, and wavula
- P. c. puma (Molina, 1782) includes the synonyms araucanus, concolor, patagonica, pearsoni, and puma (Trouessart, 1904)
- P. c. couguar (Kerr, 1792) includes arundivaga, aztecus, browni, californica, floridana, hippolestes, improcera, kaibabensis, mayensis, missoulensis, olympus, oregonensis, schorgeri, stanleyana, vancouverensis, and youngi
- P. c. costaricensis (Merriam, 1901)
- P. c. anthonyi (Nelson and Goldman, 1931) includes acrocodia, borbensis, capricornensis, concolor, greeni, and nigra
- P. c. cabrerae Pocock, 1940 includes hudsonii and puma proposed by Marcelli in 1922
In 2006, the Florida panther was still referred to as a distinct subspecies P. c. coryi in research works.
As of 2017, the Cat Classification Taskforce of the Cat Specialist Group recognizes only two subspecies as valid:
- P. c. concolor in South America, possibly excluding the region northwest of the Andes
- P. c. couguar in North and Central America and possibly northwestern South America
Evolution
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The Puma lineage of the family Felidae, depicted along with closely related genera |
The family Felidae is believed to have originated in Asia about 11 million years ago (Mya). Taxonomic research on felids remains partial, and much of what is known about their evolutionary history is based on mitochondrial DNA analysis. Significant confidence intervals exist with suggested dates. In the latest genomic study of the Felidae, the common ancestor of today's Leopardus, Lynx, Puma, Prionailurus, and Felis lineages migrated across the Bering land bridge into the Americas 8.0 to 8.5 million years ago. The lineages subsequently diverged in that order. North American felids then invaded South America 2–4 Mya as part of the Great American Interchange, following the formation of the Isthmus of Panama.
The cheetah lineage is suggested by some studies to have diverged from the Puma lineage in the Americas and migrated back to Asia and Africa, while other research suggests the cheetah diverged in the Old World itself. A high level of genetic similarity has been found among North American cougar populations, suggesting they are all fairly recent descendants of a small ancestral group. Culver et al. propose the original North American cougar population was extirpated during the Pleistocene extinctions some 10,000 years ago, when other large mammals, such as Smilodon, also disappeared. North America was then repopulated by South American cougars.
A coprolite identified as from a cougar was excavated in Argentina's Catamarca Province and dated to 17,002–16,573 years old. It contained Toxascaris leonina eggs. This finding indicates that the cougar and the parasite have existed in South America since at least the Late Pleistocene. The oldest fossil record of a cougar (Puma concolor) in South America (Argentina) is a partial skull from the late Calabrian (Ensenadan) age.
Characteristics
Cougar skull and jawboneFront paw print of a cougarThe head of the cougar is round, and the ears are erect. Its powerful forequarters, neck, and jaw serve to grasp and hold large prey. It has four retractile claws on its hind paws and five on its forepaws, of which one is a dewclaw. The larger front feet and claws are adaptations for clutching prey.
Cougars are slender and agile members of the Felidae. They are the fourth largest cat species worldwide; adults stand about 60 to 90 cm (24 to 35 in) tall at the shoulders. Adult males are around 2.4 m (7 ft 10 in) long from nose to tail tip, and females average 2.05 m (6 ft 9 in), with overall ranges between 1.50 to 2.75 m (4 ft 11 in to 9 ft 0 in) nose to tail suggested for the species in general. Of this length, the tail typically accounts for 63 to 95 cm (25 to 37 in). Males generally weigh 53 to 72 kg (117 to 159 lb). Females typically weigh between 34 and 48 kg (75 and 106 lb). Cougar size is smallest close to the equator and larger towards the poles. The largest recorded cougar, shot in 1901, weighed 105.2 kg (232 lb); claims of 125.2 kg (276 lb) and 118 kg (260 lb) have been reported, though they were probably exaggerated. Male cougars in North America average 62 kg (137 lb), while the average female in the same region averages about 42 kg (93 lb). On average, adult male cougars in British Columbia weigh 56.7 kg (125 lb) and adult females 45.4 kg (100 lb), though several male cougars in British Columbia weighed between 86.4 and 95.5 kg (190 and 211 lb).
Depending on the locality, cougars can be smaller or bigger than jaguars but are less muscular and not as powerfully built, so on average, their weight is less. Whereas the size of cougars tends to increase as much as distance from the equator increases, which crosses the northern portion of South America, jaguars are generally smaller north of the Amazon River in South America and larger south of it. For example, while South American jaguars are comparatively large, and may exceed 90 kg (200 lb), North American jaguars in Mexico's Chamela-Cuixmala Biosphere Reserve weigh approximately 50 kg (110 lb), about the same as female cougars.
Cougar coloring is plain (hence the Latin concolor in the scientific name) but can vary greatly across individuals and even siblings. The coat is typically tawny, but it ranges from silvery-grey to reddish with lighter patches on the underbody, including the jaws, chin, and throat. Infants are spotted and born with blue eyes and rings on their tails; juveniles are pale, and dark spots remain on their flanks. A leucistic individual was seen in Serra dos Órgãos National Park in Rio de Janeiro in 2013 when it was recorded by a camera trap, indicating that pure white individuals do exist within the species, though they are extremely rare.
The cougar has large paws and proportionally the largest hind legs in the Felidae, allowing for great leaping and powerful short sprints. It can leap from the ground up to 5.5 m (18 ft) high into a tree.
Distribution and habitat
The cougar has the most extensive range of any wild land animal in the Americas, spanning 110 degrees of latitude from the Yukon in Canada to the southern Andes in Chile. The species was extirpated from eastern North America, aside from Florida, but they may be recolonizing their former range and isolated populations have been documented east of their contemporary ranges in both the Midwestern US and Canada.
The cougar lives in all forest types, lowland and mountainous deserts, and in open areas with little vegetation up to an elevation of 5,800 m (19,000 ft). In the Santa Ana Mountains, it prefers steep canyons, escarpments, rim rocks and dense brush. In Mexico, it was recorded in the Sierra de San Carlos. In the Yucatán Peninsula, it inhabits secondary and semi-deciduous forests in El Eden Ecological Reserve. In El Salvador, it was recorded in the lower montane forest in Montecristo National Park and in a river basin in the Morazán Department above 700 m (2,300 ft) in 2019. In Colombia, it was recorded in a palm oil plantation close to a riparian forest in the Llanos Basin, and close to water bodies in the Magdalena River Valley. In the human-modified landscape of central Argentina, it inhabits bushland with abundant vegetation cover and prey species.
Behavior and ecology
Cougars are an important keystone species in Western Hemisphere ecosystems, linking numerous species at many trophic levels. In a comprehensive literature review of more than 160 studies on cougar ecology, ecological interactions with 485 other species in cougar-inhabited ecosystems have been shown to involve different areas of interaction, ranging from the use of other species as food sources and prey, fear effects on potential prey, effects from carcass remains left behind, to competitive effects on other predator species in shared habitat. The most common research topic in the literature used here was the cougar's diet and its prey's regulation.
Hunting and diet
The cougar is a generalist hypercarnivore. It prefers large mammals such as mule deer, white-tailed deer, elk, moose, mountain goat and bighorn sheep. It opportunistically takes smaller prey such as rodents, lagomorphs, smaller carnivores, birds, and even domestic animals, including pets. The mean weight of cougar vertebrate prey increases with its body weight and is lower in areas closer to the equator. A survey of North America research found 68% of prey items were ungulates, especially deer. Only the Florida panther showed variation, often preferring feral hogs and armadillos. Cougars have been known to prey on introduced gemsbok populations in New Mexico. One individual cougar was recorded as hunting 29 gemsbok, which made up 58% of its recorded kills. Most gemsbok kills were neonates, but some adults were also known to have been taken. Elsewhere in the southwestern United States, they have been recorded to also prey on feral horses in the Great Basin, as well as feral donkeys in the Sonoran and Mojave Deserts.
Investigations at Yellowstone National Park showed that elk and mule deer were the cougar's primary prey; the prey base is shared with the park's wolves, with which the cougar competes for resources. A study on winter kills from November to April in Alberta showed that ungulates accounted for greater than 99% of the cougar diet. Learned, individual prey recognition was observed, as some cougars rarely killed bighorn sheep, while others relied heavily on the species.
In the Central and South American cougar range area, the ratio of deer in the diet declines. Small to mid-sized mammals, including large rodents such as the capybara, are preferred. Ungulates accounted for only 35% of prey items in one survey, about half that of North America. Competition with the larger jaguar in South America has been suggested for the decline in the size of prey items. In Central or North America, the cougar and jaguar share the same prey, depending on its abundance. Other listed prey species of the cougar include mice, porcupines, American beavers, raccoons, hares, guanacoes, peccaries, vicuñas, rheas and wild turkeys. Birds and small reptiles are sometimes preyed upon in the south, but this is rarely recorded in North America. Magellanic penguins (Spheniscus magellanicus) constitute the majority of prey items in cougar diet in Patagonia's Bosques Petrificados de Jaramillo National Park and Monte León National Park.
Although capable of sprinting, the cougar is typically an ambush predator. It stalks through brush and trees, across ledges, or other covered spots, before delivering a powerful leap onto the back of its prey and a suffocating neck bite. The cougar can break the neck of some of its smaller prey with a strong bite and momentum bearing the animal to the ground. Kills are generally estimated around one large ungulate every two weeks. The period shrinks for females raising young, and may be as short as one kill every three days when cubs are nearly mature around 15 months. The cat drags a kill to a preferred spot, covers it with brush, and returns to feed over a period of days. The cougar is generally reported to not be a scavenger, but deer carcasses left exposed for study were scavenged by cougars in California, suggesting more opportunistic behavior.
Interactions with other predators
Aside from humans, no species preys upon mature cougars in the wild, although conflicts with other predators or scavengers occur. Of the large predators in Yellowstone National Park – the grizzly and black bears, gray wolf and cougar – the massive grizzly bear appears dominant, often (though not always) able to drive a gray wolf pack, black bear or cougar off their kills. One study found that grizzlies and American black bears visited 24% of cougar kills in Yellowstone and Glacier National Parks, usurping 10% of carcasses. Bears gained up to 113%, and cougars lost up to 26% of their daily energy requirements from these encounters. 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 killing, and when bears are most active, the cats take prey more frequently and spend less time feeding on each kill. Unlike several subordinate predators from other ecosystems, cougars do not appear to exploit spatial or temporal refuges to avoid competitors.
The gray wolf and the cougar compete more directly for prey, mostly in winter. Packs of wolves can steal cougars' kills, and there are some documented cases of cougars being killed by them. One report describes a large pack of seven to 11 wolves killing a female cougar and her kittens, while in nearby Sun Valley, Idaho, a 2-year-old male cougar was found dead, apparently killed by a wolf pack. Conversely, one-to-one confrontations tend to be dominated by the cat, and there are various documented accounts where wolves have been ambushed and killed, including adult male specimens. Wolves more broadly affect cougar population dynamics and distribution by dominating territory and prey opportunities, and disrupting the feline's behavior. Preliminary research in Yellowstone, for instance, has shown displacement of the cougar by wolves. One researcher in Oregon noted: "When there is a pack around, cougars are not comfortable around their kills or raising kittens A lot of times a big cougar will kill a wolf, but the pack phenomenon changes the table." Both species are capable of killing mid-sized predators, such as bobcats, Canada lynxes, wolverines and coyotes, and tend to suppress their numbers. Although cougars can kill coyotes, the latter have been documented attempting to prey on cougar cubs.
The cougar and jaguar share overlapping territory in the southern portion of its range. The jaguar tends to take the larger prey where ranges overlap, reducing both the cougar's potential size and the likelihood of direct competition between the two cats. Cougars appear better than jaguars at exploiting a broader prey niche and smaller prey.
Social spacing and interactions
The cougar is a mostly solitary animal. Only mothers and kittens live in groups, with adults meeting rarely. While generally loners, cougars will reciprocally share kills and seem to organize themselves into small communities defined by the territories of dominant males. Cats within these areas socialize more frequently with each other than with outsiders.
In the vicinity of a cattle ranch in northern Mexico, cougars exhibited nocturnal activity that overlapped foremost with the activity of calves. In a nature reserve in central Mexico, the activity of cougars was crepuscular and nocturnal, overlapping largely with the activity of the nine-banded armadillo (Dasypus novemcinctus). Cougars in the montane Abra-Tanchipa Biosphere Reserve in southeastern Mexico displayed a cathemeral activity pattern. Data from 12 years of camera trapping in the Pacific slope and Talamanca Cordillera of Costa Rica showed cougars as cathemeral. Both cougars and jaguars in the Cockscomb Basin of Belize were nocturnal but avoided each other. In a protected cloud forest in the central Andes of Colombia, cougars were active from late afternoon to shortly before sunrise and sometimes during noon and early afternoon. In protected areas of the Madidi-Tambopata Landscape in Bolivia and Peru, cougars were active throughout the day but with a tendency to nocturnal activity that overlapped with the activity of main prey species. During an 8-year-long study in a modified landscape in southeastern Brazil, male cougars were primarily nocturnal, but females were active at night and day. Cougars were diurnal in the Brazilian Pantanal, but crepuscular and nocturnal in protected areas in the Cerrado, Caatinga and ecotone biomes. Cougars in the Atlantic Forest were active throughout the day but displayed peak activity during early mornings in protected areas and crepuscular and nocturnal activity in less protected areas. In central Argentina, cougars were active day and night in protected areas but were active immediately after sunset and before sunrise outside protected areas. Cougars displayed a foremost crepuscular and nocturnal activity pattern in a ranching area in southern Argentina.
Home range sizes and overall cougar abundance depend on terrain, vegetation, and prey abundance. Research suggests a lower limit of 25 km (9.7 sq mi) and upper limit of 1,300 km (500 sq mi) of home range for males. Large male home ranges of 150 to 1,000 km (58 to 386 sq mi) with female ranges half that size. One female adjacent to the San Andres Mountains was found with a big range of 215 km (83 sq mi), necessitated by poor prey abundance. Research has shown cougar abundances from 0.5 animals to as many as seven per 100 km (39 sq mi).
Male home ranges include or overlap with females but, at least where studied, not with those of other males. The home ranges of females overlap slightly. Males create scrapes composed of leaves and duff with their hind feet, and mark them with urine and sometimes feces. When males encounter each other, they vocalize and may engage in violent conflict if neither backs down.
Cougars communicate with various vocalizations. Aggressive sounds include growls, spits, snarls, and hisses. During the mating season, estrus females produce caterwauls or yowls to attract mates, and males respond with similar vocals. Mothers and offspring keep in contact with whistles, chirps, and mews.
Reproduction and life cycle
North American cougar cub in the Santa Monica MountainsCubsFemales reach sexual maturity at the age of 18 months to three years and are in estrus for about eight days of a 23-day cycle; the gestation period is approximately 91 days. Both adult males and females may mate with multiple partners, and a female's litter can have multiple paternities. Copulation is brief but frequent. Chronic stress can result in low reproductive rates in captivity as well as in the field.
Gestation is 82–103 days long. Only females are involved in parenting. Litter size is between one and six cubs, typically two. Caves and other alcoves that offer protection are used as litter dens. Born blind, cubs are completely dependent on their mother at first and begin to be weaned at around three months of age. As they grow, they go out on forays with their mother, first visiting kill sites and, after six months, beginning to hunt small prey on their own. Kitten survival rates are just over one per litter.
Juveniles remain with their mothers for one to two years. When a female reaches estrous again, her offspring must disperse or the male will kill them. Males tend to disperse further than females. One study has shown a high mortality rate among cougars that travel farthest from their maternal range, often due to conflicts with other cougars. In a study area in New Mexico, males dispersed farther than females, traversed large expanses of non-cougar habitat and were probably most responsible for nuclear gene flow between habitat patches.
Life expectancy in the wild is reported at 8 to 13 years and probably averages 8 to 10; a female of at least 18 years was reported killed by hunters on Vancouver Island. Cougars may live as long as 20 years in captivity. Causes of death in the wild include disability and disease, competition with other cougars, starvation, accidents, and, where allowed, hunting. The feline immunodeficiency virus is well-adapted to the cougar.
Conservation
The cougar has been listed as Least Concern on the IUCN Red List since 2008. However, it is also listed on CITES Appendix II. Hunting it is prohibited in California, Costa Rica, Honduras, Nicaragua, Guatemala, Panama, Venezuela, Colombia, French Guiana, Suriname, Bolivia, Brazil, Chile, Paraguay, Uruguay and most of Argentina. Hunting is regulated in Canada, Mexico, Peru, and the United States. Establishing wildlife corridors and protecting sufficient range areas are critical for the sustainability of cougar populations. Research simulations showed that it faces a low extinction risk in areas larger than 2,200 km (850 sq mi). Between one and four new individuals entering a population per decade markedly increases persistence, thus highlighting the importance of habitat corridors.
The Florida panther population is afforded protection under the Endangered Species Act. The Texas Mountain Lion Conservation Project was launched in 2009 and aimed at raising local people's awareness of the status and ecological role of the cougar and mitigating conflict between landowners and cougars.
The cougar is threatened by habitat loss, habitat fragmentation, and depletion of its prey base due to poaching. Hunting is legal in the western United States. In Florida, heavy traffic causes frequent accidents involving cougars. Highways are a major barrier to the dispersal of cougars. The cougar populations in California are becoming fragmented with the increase in human population and infrastructure growth in the state.
Human–wildlife conflict in proximity of 5 km (1.9 sq mi) of cougar habitat is pronounced in areas with a median human density of 32.48 inhabitants/km (84.1 inhabitants/sq mi) and a median livestock population density of 5.3 heads/km (14 heads/sq mi). Conflict is generally lower in areas more than 16.1 km (10.0 mi) away from roads and 27.8 km (17.3 mi) away from settlements.
Relationships with humans
Attacks on humans
In North America
See also: List of fatal cougar attacks in North AmericaDue to the expanding human population, cougar ranges increasingly overlap with areas inhabited by humans. Attacks on humans are very rare, as cougar prey recognition is a learned behavior and they do not generally recognize humans as prey. In a 10-year study in New Mexico of wild cougars who were not habituated to humans, the animals did not exhibit threatening behavior to researchers who approached closely (median distance=18.5 m; 61 feet) except in 6% of cases; 14⁄16 of those were females with cubs. Attacks on people, livestock, and pets may occur when a puma habituates to humans or is in a condition of severe starvation. Attacks are most frequent during late spring and summer when juvenile cougars leave their mothers and search for new territory.
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). By 2004, the count had climbed to 88 attacks and 20 deaths.
Within North America, the distribution of attacks is not uniform. The heavily populated state of California saw a dozen attacks from 1986 to 2004 (after just three from 1890 to 1985), including three fatalities. In March 2024, two brothers in California were attacked by a male cougar, with one being fatally wounded; it was the state's first fatal attack in 20 years. Washington state was the site of a fatal attack in 2018, its first since 1924. Lightly populated New Mexico reported an attack in 2008, the first there since 1974.
As with many predators, a cougar may attack if cornered, if a fleeing human stimulates their instinct to chase, or if a person "plays dead". Standing still may cause the cougar to consider a person easy prey. 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.
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. Children are at greatest risk of attack and least likely to survive an encounter. Detailed research into attacks before 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. 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.
Research on new wildlife collars may reduce human-animal conflicts by predicting when and where predatory animals hunt. This may save the lives of humans, pets, and livestock, as well as the lives of these large predatory mammals that are important to the balance of ecosystems.
In South America
Cougars in the southern cone of South America are reputed to be extremely reluctant to attack people; in legend, they defended people against jaguars. The nineteenth-century naturalists Félix de Azara and William Henry Hudson thought that attacks on people, even children or sleeping adults, did not happen. Hudson, citing anecdotal evidence from hunters, claimed that pumas were positively inhibited from attacking people, even in self-defense. Attacks on humans, although exceedingly rare, have occurred.
An early, authenticated, non-fatal case occurred near Lake Viedma, Patagonia, in 1877 when a female mauled the Argentine scientist Francisco P. Moreno; Moreno afterward showed the scars to Theodore Roosevelt. In this instance, however, Moreno had been wearing a guanaco-hide poncho round his neck and head as protection against the cold; in Patagonia the guanaco is the puma's chief prey animal. Another authenticated case occurred in 1997 in Iguazú National Park in northeastern Argentina, when the 20-month-old son of a ranger was killed by a female puma. Forensic analysis found specimens of the child's hair and clothing fibers in the animal's stomach. The coatí is the puma's chief prey in this area. Despite prohibitory signs, coatis are hand-fed by tourists in the park, causing unnatural approximation between cougars and humans. This particular puma had been raised in captivity and released into the wild. On March 13, 2012, Erica Cruz, a 23-year-old shepherdess was found dead in a mountainous area near Rosario de Lerma, Salta Province, in northwestern Argentina. Claw incisions, which severed a jugular vein, indicated that the attacker was a felid; differential diagnosis ruled out other possible perpetrators. There were no bite marks on the victim, who had been herding goats. In 2019 in Córdoba Province, Argentina an elderly man was badly injured by a cougar after he attempted to defend his dog from it, while in neighboring Chile a 28-year-old woman was attacked and killed in Corral, in Los Ríos Region, on October 20, 2020.
Fatal attacks by other carnivores, such as feral dogs, can be misattributed to cougars without appropriate forensic knowledge.
Predation on domestic animals
During the early years of ranching, cougars were considered on par with wolves in destructiveness. According to figures in Texas in 1990, 86 calves (0.0006% of Texas's 13.4 million cattle and calves), 253 mohair goats, 302 mohair kids, 445 sheep (0.02% of Texas's 2 million sheep and lambs) and 562 lambs (0.04% of Texas's 1.2 million lambs) were confirmed to have been killed by cougars that year. In Nevada in 1992, cougars were confirmed to have killed nine calves, one horse, four foals, five goats, 318 sheep, and 400 lambs. In both reports, sheep were the most frequently attacked. Some instances of surplus killing have resulted in the deaths of 20 sheep in one attack. A cougar's killing bite is applied to the back of the neck, head, or throat and the cat inflicts puncture marks with its claws usually seen on the sides and underside of the prey, sometimes also shredding the prey as it holds on. Coyotes also typically bite the throat, but the work of a cougar is generally clean, while bites inflicted by coyotes and dogs leave ragged edges. The size of the tooth puncture marks also helps distinguish kills made by cougars from those made by smaller predators.
Remedial hunting appears to have the paradoxical effect of increased livestock predation and complaints of human-cougar conflicts. In a 2013 study, the most important predictor of cougar problems was the remedial hunting of cougars the previous year. Each additional cougar on the landscape increased predation and human-cougar complaints by 5%, but each animal killed during the previous year increased complaints by 50%. The effect had a dose-response relationship with very heavy (100% removal of adult cougars) remedial hunting, leading to a 150–340% increase in livestock and human conflicts. This effect is attributed to the removal of older cougars that have learned to avoid people and their replacement by younger males that react differently to humans. Remedial hunting enables younger males to enter the former territories of the older animals. Predation by cougars on dogs "is widespread, but occurs at low frequencies".
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 people often represented the cougar in their ceramics. The sky and thunder god of the Inca, Viracocha, has been associated with the animal.
In North America, mythological descriptions of the cougar have appeared in the stories of the Hocąk language ("Ho-Chunk" or "Winnebago") of Wisconsin and Illinois and the Cheyenne, among others. To the Apache and Walapai of the Southwestern United States, the wail of the cougar was a harbinger of death. The Algonquins and Ojibwe believe that the cougar lived in the underworld and was wicked, whereas it was a sacred animal among the Cherokee.
See also
- Bougar—hybrid of cougar and bobcat
- List of largest cats
- Pumapard—hybrid of cougar and leopard
Explanatory notes
- The populations of Costa Rica and Panama are included in Appendix I.
- There are no jaguars in the area; other felids were too small to kill humans.
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This, however, is not a full statement of the facts; the puma will not even defend itself against man
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Further reading
- Mark Elbroch: The Cougar Conundrum: Sharing the World with a Successful Predator. Island Press, 2020, ISBN 9781610919982.
- Maurice Hornocker (ed.), Sharon Negri (ed.): Cougar: Ecology and Conservation. University of Chicago Press, 2009, ISBN 9780226353470.
- Kenneth A. Logan, Linda L. Sweanor: Desert Puma: Evolutionary Ecology And Conservation Of An Enduring Carnivore. Island Press, 2001, ISBN 9781610910583.
- Paula Wild: The Cougar: Beautiful, Wild and Dangerous. Douglas and McIntyre, 2013, ISBN 9781771620031.
External links
Listen to this article (43 minutes) This audio file was created from a revision of this article dated 6 November 2008 (2008-11-06), and does not reflect subsequent edits.(Audio help · More spoken articles)- "Cougar Puma concolor". IUCN/SSC Cat Specialist Group.
- Cougar Tracks: How to identify cougar tracks in the wild
- "Puma sounds". National Geographic Society. September 10, 2010. Archived from the original on November 3, 2013.
- Santa Cruz Puma Project
- Eastern Puma Research Network
- Felidae Conservation Fund
- Cougar Rewilding Foundation, formerly the "Eastern Cougar Foundation"
- The Cougar Network --Using Science to Understand Cougar Ecology; Archived July 31, 2018, at the Wayback Machine
- "Saving America's Lion". Mountain Lion Foundation. Archived from the original on July 31, 2018. Retrieved December 11, 2010.
- SaveTheCougar.org: Sightings of cougars in Michigan
- 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.
- Living with California Mountain Lions
- Oregon's first fatal cougar attack in the wild claims hiker near Mount Hood
Game animals and shooting in North America | ||
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Game birds | ||
Waterfowl | ||
Big game | ||
Other quarry | ||
See also |
Taxon identifiers | |
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Puma concolor |
|
Felis concolor |
- IUCN Red List least concern species
- Cougar
- Apex predators
- Big cats
- Carnivorans of Brazil
- Extant Middle Pleistocene first appearances
- Fauna of the Amazon
- Fauna of the California chaparral and woodlands
- Felids of Central America
- Felids of North America
- Felids of South America
- Mammals described in 1771
- Mammals of Patagonia
- Mammals of the Andes
- Pleistocene carnivorans
- Pleistocene mammals of North America
- Pleistocene mammals of South America
- Puma (genus)
- Quaternary carnivorans
- Taxa named by Carl Linnaeus