Editing Cougar (section)

==Behavior and ecology==
The cougar is a [[keystone species]] in Western Hemisphere ecosystems as it links numerous species at many [[trophic level]]s interacting with 485 other species as food source and prey, carcass remains left behind and competitive effects on other [[Predation|predator]]s in shared habitat.<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 |bibcode=2022MamRv..52..360L}}</ref>

===Hunting and diet===
[[File:P-35 and the Bears (25382928560).jpg|thumb|right|Camera trap image of cougar in the [[Santa Susana Mountains]] northwest of Los Angeles]]
[[File:P-35 and the Bears (25657563476).jpg|thumb|Cougar with deer kill]]

The cougar is a [[Generalist and specialist species|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 [[rodent]]s, [[lagomorph]]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 [[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 [[Hog (swine)|hog]]s and [[armadillo]]s.<ref name="diet" /> 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.<ref name="TWS">{{cite web |last1=Kobilinsky |first1=D. |date=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 |work=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 [[feral horse]]s in the [[Great Basin]],<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=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 [[feral donkey]]s in the [[Sonoran Desert|Sonoran]] and [[Mojave Desert]]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>

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 [[Wolf|wolves]], with which the cougar competes for resources.<ref name="Yellowstone">{{cite web |title=Wildlife: Wolves |publisher=[[Yellowstone National Park]] |url=http://www.yellowstonenationalpark.com/wolves.htm |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 [[Alberta]] showed that ungulates accounted for greater than 99% of the cougar diet. Learned, individual prey recognition was observed, as some cougars rarely killed bighorn sheep, while others relied heavily on the species.<ref>{{cite journal |last=Ross |first=R. |author2=Jalkotzy, M. G. |author3=Festa-Bianchet, M. |year=1993 |title=Cougar predation on bighorn sheep in southwestern Alberta during winter |journal=Canadian Journal of Zoology |volume=75 |issue=5 |pages=771–775 |doi=10.1139/z97-098 |name-list-style=amp}}</ref>

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.<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 [[Mouse|mice]], [[porcupine]]s, [[American beaver]]s, [[raccoon]]s, [[hare]]s, [[guanaco]]es, [[Peccary|peccaries]], [[vicuña]]s, [[Rhea (bird)|rhea]]s and [[wild turkey]]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" /> [[Magellanic penguin]]s (''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]].<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>

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.<ref name="WhosWho" /> Kills are generally estimated around one large ungulate every two weeks. The period shrinks for females raising young, and may be as short as one kill every three days when cubs are nearly mature around 15 months.<ref name="CAP" /> The cat drags a kill to a preferred spot, covers it with brush, and returns to feed over a period of days. The cougar is generally reported to not be a [[scavenger]], 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)050[0466:SBIP]2.0.CO;2 |s2cid=85632179}}</ref>

===Interactions with other predators===
[[File:Feeling Unwelcome.jpg|thumb|Juvenile cougar in conflict with [[coyote]]s at [[National Elk Refuge]], using a [[split-rail fence#Buck-and-rail fence|buck-and-rail fence]] 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 [[Yellowstone National Park]] – the [[grizzly bear|grizzly]] and [[American black bear|black bear]]s, [[Wolf|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 bear]]s visited 24% of cougar kills in Yellowstone and [[Glacier National Park (U.S.)|Glacier National Parks]], usurping 10% of carcasses. Bears gained up to 113%, and cougars lost up to 26% of their 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=[[Environment Canada]] |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 [[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.<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>

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 [[Sun Valley, Idaho]], 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=[[The Washington Post]] |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 [[Yellowstone]], for instance, has shown displacement of the cougar by wolves.<ref>{{cite web |url=http://www.greateryellowstonescience.org/topic/wolves/wolvesoverview.html |title=Overview: Gray Wolves |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&nbsp;[...] 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=[[The Oregonian]] |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 [[bobcat]]s, [[Canada lynx]]es, [[wolverine]]s and [[coyote]]s, and tend to suppress their numbers.<ref name="Yellowstone" /> Although cougars can kill coyotes, the latter have been documented attempting to prey on cougar cubs.<ref name="Cougars vs. coyotes photos draw Internet crowd">{{cite web |title=Cougars vs. coyotes photos draw Internet crowd |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 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>

===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.<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 [[Nocturnality|nocturnal]] activity that overlapped foremost with the activity of [[Calf (animal)|calves]].<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 [[Crepuscular animal|crepuscular]] and nocturnal, overlapping largely with the activity of the [[nine-banded armadillo]] (''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 [[Cathemerality|cathemeral]] 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 [[Cockscomb Basin Wildlife Sanctuary|Cockscomb Basin]] 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 [[Madidi National Park|Madidi]]-[[Tambopata National Reserve|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.<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 [[Pantanal]], but crepuscular and nocturnal in protected areas in the [[Cerrado]], [[Caatinga]] and [[ecotone]] 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|bibcode=2016WildR..43..449Z |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/>

[[Home range]] 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=[[Canadian Geographic Magazine]]}}</ref> One female adjacent to the [[San Andres Mountains]] 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>

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 [[Plant litter|duff]] with their hind feet, and [[Territorial marking|mark]] them with [[urine]] and sometimes [[feces]].<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=[[University of Wisconsin, Stevens Point]]}}</ref>

Cougars communicate with various vocalizations. Aggressive sounds include growls, spits, snarls, and hisses. During the mating season, estrus females produce [[Cat communication#Call|caterwauls]] 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>

===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 [[Cub (organism)|cub]] in the [[Santa Monica Mountains National Recreation Area|Santa Monica Mountains]]  |image2=Mountain lion kittens.jpg |caption2=Cubs}}

Females 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.<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)032[0711:ATFNDS]2.0.CO;2 |s2cid=85819163}}</ref>

[[Gestation]] 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" />

Juveniles remain with their mothers for one to two years.<ref name="Naughton" /> When a female reaches estrous again, her offspring must [[Biological dispersal|disperse]] 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=[[University of Wisconsin, Stevens Point]]}}</ref> One study has shown a high [[mortality rate]] among cougars that travel farthest from their maternal range, often due to conflicts with other cougars.<ref name="Utah" /> 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.<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 [[hunter]]s on [[Vancouver Island]].<ref name="CAP" /> Cougars may live as long as 20 years in captivity. Causes of death in the wild include disability and disease, competition with other cougars, starvation, accidents, and, where allowed, hunting. The [[feline immunodeficiency virus]] 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>