Editing Lion (section)

==Behaviour and ecology==
Lions spend much of their time resting; they are inactive for about twenty hours per day.<ref name="Schaller122">[[#Schaller|Schaller]], p. 122.</ref> Although lions can be active at any time, their activity generally peaks after dusk with a period of socialising, grooming, and defecating. [[Cathemerality|Intermittent bursts of activity]] continue until dawn, when hunting most often takes place. They spend an average of two hours a day walking and fifty minutes eating.<ref name="Schaller120">[[#Schaller|Schaller]], pp. 120–121.</ref>

===Group organisation===
{{multiple image|align=right|direction=vertical
|image1=Lion (Panthera leo) male and cub Etosha.jpg|caption1=Lion pride in Etosha National Park
|image2=Lions Family Portrait Masai Mara.jpg|caption2=A lioness (left) and two males in Masai Mara}}
The lion is the most social of all wild felid species, living in groups of related individuals with their offspring. Such a group is called a "[[List of animal names#L|pride]]". Groups of male lions are called "coalitions".<ref name="Schaller33">[[#Schaller|Schaller]], p. 33.</ref> Females form the stable social unit in a pride and do not tolerate outside females.<ref name="Schaller37">[[#Schaller|Schaller]], p. 37.</ref> The majority of females remain in their birth prides while all males and some females [[Biological dispersal|disperse]].<ref name=Packer33>[[#Packer|Packer]], p. 33.</ref> The average pride consists of around 15 lions, including several adult females and up to four males and their cubs of both sexes. Large prides of up to 30 individuals have been observed.<ref>[[#Schaller|Schaller]], p. 34–35.</ref> The sole exception to this pattern is the [[Tsavo lion]] pride that always had only one adult male.<ref>{{Cite journal |last=Milius |first=S. |date=2002 |title= Biology: Maneless lions live one guy per pride |journal=Society for Science & the Public |volume=161 |issue=16 |page=253 |doi=10.1002/scin.5591611614}}</ref> Prides act as [[fission–fusion society|fission–fusion societies]], and members split into subgroups that keep in contact with [[roar]]s.<ref>[[#Packer|Packer]], pp. 25, 31.</ref>

Nomadic lions range widely and move around sporadically, either in pairs or alone.<ref name="Schaller33"/> Pairs are more frequent among related males. A lion may switch lifestyles; nomads can become residents and vice versa.<ref name=Estes>{{cite book |author=Estes, R. |year=1991 |title=The behavior guide to African mammals: including hoofed mammals, carnivores, primates |publisher=University of California Press |isbn=978-0-520-08085-0 |location=Berkeley |chapter=Lion |pages=[https://archive.org/details/isbn_0520080858/page/369 369–376] |chapter-url=https://archive.org/details/isbn_0520080858/page/369 |url=https://archive.org/details/isbn_0520080858/page/369 }}</ref> Interactions between prides and nomads tend to be hostile, although pride females in [[estrus]] allow nomadic males to approach them.<ref>[[#Schaller|Schaller]], pp. 52–54.</ref> Males spend years in a nomadic phase before gaining residence in a pride.<ref>{{cite book |author=Hanby, J. P.|author2=Bygott, J. D.  |year=1979 |chapter=Population changes in lions and other predators |title=Serengeti: dynamics of an ecosystem |editor1=Sinclair, A. R. E. |editor2=Norton-Griffiths, M. |publisher=The University of Chicago Press |location=Chicago |pages=249–262}}</ref> A study undertaken in the [[Serengeti National Park]] revealed that nomadic coalitions gain residency at between 3.5 and 7.3 years of age.<ref>{{cite journal |author=Borrego, N. |author2=Ozgul, A.|author3=Slotow, R.|author4=Packer, C. |year=2018 |title=Lion population dynamics: do nomadic males matter? |journal=Behavioral Ecology |volume=29 |issue=3 |doi=10.1093/beheco/ary018 |pages=660–666 |doi-access=free}}</ref> In Kruger National Park, dispersing male lions move more than {{cvt|25|km}} away from their natal pride in search of their own territory. Female lions stay closer to their natal pride. Therefore, female lions in an area are more closely related to each other than male lions in the same area.<ref>{{cite journal |author=Van Hooft, P.|author2=Keet, D. F.  |author3=Brebner, D. K.|author4=Bastos, A. D. |year=2018 |title=Genetic insights into dispersal distance and disperser fitness of African lions (''Panthera leo'') from the latitudinal extremes of the Kruger National Park, South Africa |journal=BMC Genetics |volume=19 |issue=1 |page=21 |doi=10.1186/s12863-018-0607-x |pmid=29614950 |pmc=5883395 |doi-access=free}}</ref>

The evolution of sociability in lions was likely driven both by high [[population density]] and the clumped resources of savannah habitats. The larger the pride, the more high-quality [[Territory (animal)|territory]] they can defend; "hotspots" are near river [[confluence]]s, where they have optimal access to water, prey and vegetation cover.<ref>[[#Packer|Packer]], pp. 195–196, 222.</ref><ref>{{cite journal |last1=Mosser |first1=A. A.|last2=Kosmala |first2=M. |last3=Packer|first3=C.|year=2015 |title=Landscape heterogeneity and behavioral traits drive the evolution of lion group territoriality |journal=Behavioral Ecology |volume=26 |issue=4 |pages=1051–1059 |doi=10.1093/beheco/arv046 |doi-access=free}}</ref> A study on three lion prides in a Zimbabwean wildlife reserve revealed that the dominant pride of 12 lions had the shortest average distance to water and the smallest [[home range]] of {{cvt|130.35|km2}}; the smallest pride of four lions had the longest average distance to water and the largest home range of {{cvt|174.6|km2}}.<ref>{{cite journal |author1=Tarugara, A. |name-list-style=amp |author2=Clegg, B. W. |author3=Clegg, S. B. |year=2024 |title=Factors influencing space-use and kill distribution of sympatric lion prides in a semi-arid savanna landscape |journal=PeerJ |volume=12 |page=e16749 |doi=10.7717/peerj.16749 |doi-access=free}}</ref>

The area occupied by a pride is called a "pride area" whereas that occupied by a nomad is a "range".<ref name="Schaller33"/> Males associated with a pride patrol the fringes.<ref name=Haas2005/> Both males and females defend the pride against intruders, but the male lion is better-suited for this purpose due to its stockier, more powerful build. Some individuals consistently lead the defence against intruders, while others lag behind.<ref>{{Cite journal |last=Heinsohn|first= R. |author2=Packer, C. |year=1995 |title=Complex cooperative strategies in group-territorial African lions |journal=Science |volume=269 |issue=5228 |pages=1260–1262 |doi=10.1126/science.7652573 |pmid=7652573 |bibcode=1995Sci...269.1260H |s2cid= 35849910 |url=http://www.life.umd.edu/faculty/wilkinson/BIOL608W/Heinsohn&Packer95.pdf |archive-url=https://web.archive.org/web/20170810170702/http://www.life.umd.edu/faculty/wilkinson/BIOL608W/Heinsohn&Packer95.pdf |archive-date=10 August 2017 |url-status=live}}</ref> Lions tend to assume specific roles in the pride; slower-moving individuals may provide other valuable services to the group.<ref>{{Cite journal |last=Morell |first=V. |year=1995 |title=Cowardly lions confound cooperation theory |journal=Science |volume=269 |issue=5228 |pages=1216–1217 |doi=10.1126/science.7652566 |pmid=7652566 |bibcode=1995Sci...269.1216M |s2cid=44676637}}</ref> Alternatively, there may be rewards associated with being a leader that fends off intruders; the rank of lionesses in the pride is reflected in these responses.<ref>{{Cite journal |last=Jahn |first=G. C. |year=1996 |title=Lioness Leadership |journal=Science |volume=271 |issue=5253 |page=1215 |doi=10.1126/science.271.5253.1215a |pmid=17820922 |bibcode=1996Sci...271.1215J |s2cid=5058849 }}</ref> The male or males associated with the pride must defend their relationship with the pride from outside males who may attempt to usurp them.<ref name=Estes/>

Asiatic lion prides differ in group composition. Male Asiatic lions are solitary or associate with up to three males, forming a loose pride while females associate with up to 12 other females, forming a stronger pride together with their cubs. Female and male lions associate only when mating.<ref name=Joslin1973>{{cite book |author=Joslin, P. |year=1973 |title=The Asiatic lion: a study of ecology and behaviour |location=University of Edinburgh, UK |publisher=Department of Forestry and Natural Resources}}</ref> Coalitions of males hold territory for a longer time than single lions. Males in coalitions of three or four individuals exhibit a pronounced hierarchy, in which one male dominates the others and mates more frequently.<ref name=Chakrabarti2017>{{cite journal |author=Chakrabarti, S.  |author2=Jhala, Y. V. |author2-link=Yadvendradev Vikramsinh Jhala |year=2017 |title=Selfish partners: resource partitioning in male coalitions of Asiatic lions |journal=Behavioral Ecology |volume=28 |issue=6 |pages=1532–1539 |doi=10.1093/beheco/arx118 |pmid=29622932 |pmc=5873260}}</ref>

===Hunting and diet===
{{multiple image| perrow=1 |image2=Lions taking down cape buffalo.jpg|caption2=Four lionesses catching a buffalo in the Serengeti |image1=Lion and eland.jpg |caption1=A skeletal mount of a lion attacking a [[common eland]], on display at [[The Museum of Osteology]]
|image3=Lions and a Zebra b.jpg |caption3=Lions feeding on a zebra}}

The lion is a [[Generalist and specialist species|generalist]] [[hypercarnivore]] and is considered to be both an apex and keystone predator due to its wide prey spectrum.<ref>[[#Schaller|Schaller]], p. 208.</ref><ref name=Frank1998>{{cite book |last=Frank| first=L. G. |year=1998 |title=Living with lions: carnivore conservation and livestock in Laikipia District, Kenya |location=Mpala Research Centre, Nanyuki |publisher=US Agency for International Development, Conservation of Biodiverse Resource Areas Project, 623-0247-C-00-3002-00}}</ref> Its prey consists mainly of medium-sized to large [[ungulate]]s, particularly [[blue wildebeest]], [[plains zebra]], [[African buffalo]], [[gemsbok]] and [[giraffe]]. It also frequently takes [[common warthog]] despite it being much smaller.<ref name=Hayward2005>{{Cite journal |last1=Hayward |first1=M. W. |last2=Kerley |first2=G. I. H.  |year=2005 |title=Prey preferences of the lion (''Panthera leo'') |journal=Journal of Zoology |volume=267 |issue=3 |pages=309–322 |doi=10.1017/S0952836905007508 |citeseerx=10.1.1.611.8271 |url=http://www.zbs.bialowieza.pl/g2/pdf/1595.pdf |archive-url=https://web.archive.org/web/20150627132429/http://www.zbs.bialowieza.pl/g2/pdf/1595.pdf |archive-date=27 June 2015 |url-status=live}}</ref> In India, [[chital]] and [[sambar deer]] are the most common wild prey,<ref name=Haas2005/><ref name=Hayward2005/><ref name=Mukherjee>{{cite journal |last1=Mukherjee |first1=S. |last2=Goyal |first2=S. P. |last3=Chellam| first3=R.  |year=1994 |title=Refined techniques for the analysis of Asiatic lion ''Panthera leo persica'' scats |journal=Acta Theriologica |volume=39 |issue=4 |pages=425–430 |doi=10.4098/AT.arch.94-50 |doi-access=free}}</ref> while livestock contributes significantly to lion kills outside protected areas.<ref>{{cite journal |author1=Meena, V.  |author2=Jhala, Y. V. |author3=Chellam, R. |author4=Pathak, B. |year=2011 |title=Implications of diet composition of Asiatic lions for their conservation |journal=Journal of Zoology |volume=284 |issue=1 |pages=60–67 |doi=10.1111/j.1469-7998.2010.00780.x |doi-access=free}}</ref> It usually avoids adult [[elephant]]s, [[rhinoceros]] and [[hippopotamus]] and small prey like [[dik-dik]], [[hyrax]]es, [[hare]]s and [[monkey]]s, and seldom consumes other predators; unusual prey include small reptiles and [[Old World porcupine|porcupine]]s.<ref>[[#Schaller|Schaller]], pp. 220–221.</ref>

Young lions first display stalking behaviour at around three months of age, although they do not participate in hunting until they are almost a year old and begin to hunt effectively when nearing the age of two.<ref name=Schaller153>[[#Schaller|Schaller]], p. 153.</ref> Single lions are capable of bringing down zebra and wildebeest, while larger prey like buffalo and giraffe are riskier.<ref name=Estes/> In [[Chobe National Park]], large prides have been observed hunting [[African bush elephant]]s up to around 15 years old in exceptional cases, with the victims being calves, juveniles, and even subadults.<ref>{{cite journal |last1=Joubert |first1=D. |year=2006 |title=Hunting behaviour of lions (''Panthera leo'') on elephants (''Loxodonta africana'') in the Chobe National Park, Botswana |journal=African Journal of Ecology |volume=44 |issue=2 |pages=279–281 |doi=10.1111/j.1365-2028.2006.00626.x|bibcode=2006AfJEc..44..279J}}</ref><ref>{{cite journal |last1=Power |first1=R. J. |last2=Compion |first2=R. X. S.  |title=Lion predation on elephants in the Savuti, Chobe National Park, Botswana |journal=African Zoology |volume=44 |issue=1 |pages=36–44 |doi=10.3377/004.044.0104 |year=2009 |s2cid=86371484 |url=https://www.researchgate.net/publication/232693088 |access-date=20 April 2018 |archive-date=31 August 2018 |archive-url=https://web.archive.org/web/20180831035431/https://www.researchgate.net/publication/232693088 |url-status=live}}</ref> In typical group hunts, each lioness has a favoured position in the group, either stalking prey on the "wing", then attacking, or moving a smaller distance in the centre of the group and capturing prey fleeing from other lionesses. Males attached to prides do not usually participate in group hunting.<ref name=hunt>{{Cite journal |last=Stander |first=P. E. |title=Cooperative hunting in lions: the role of the individual |journal=Behavioral Ecology and Sociobiology |volume=29 |issue=6 |pages=445–454 |year=1992 |doi=10.1007/BF00170175 |bibcode=1992BEcoS..29..445S |s2cid=2588727 |url=http://lchc.ucsd.edu/MCA/Mail/xmcamail.2011_09.dir/pdfeL5GzNL2FL.pdf |archive-url=https://web.archive.org/web/20150518095055/http://lchc.ucsd.edu/MCA/Mail/xmcamail.2011_09.dir/pdfeL5GzNL2FL.pdf |archive-date=18 May 2015 |url-status=live}}</ref>
Some evidence suggests, however, that males are just as successful as females; they are typically solo hunters who ambush prey in small bushland.<ref>{{cite journal |last1=Loarie, S. R. |last2=Tambling, C. J. |last3=Asner, G. P.  |year=2013 |title=Lion hunting behaviour and vegetation structure in an African savanna |journal=Animal Behaviour |volume=85 |issue=5 |pages=899–906 |doi=10.1016/j.anbehav.2013.01.018 |hdl=2263/41825 |hdl-access=free |s2cid=53185309 |url=https://repository.up.ac.za/bitstream/2263/41825/1/Loarie_Lion_2013.pdf |archive-url=https://web.archive.org/web/20170816101650/http://repository.up.ac.za/bitstream/2263/41825/1/Loarie_Lion_2013.pdf |archive-date=16 August 2017 |url-status=live}}</ref> They may join in the hunting of large, slower-moving prey like buffalo; and even hunt them on their own. Moderately-sized hunting groups generally have higher success rates than lone females and larger groups.<ref>[[#Packer|Packer]], p. 150, 153, 164–165.</ref>

Lions are not particularly known for their stamina. For instance, a lioness's heart comprises only 0.57% of her body weight and a male's is about 0.45% of his body weight, whereas a hyena's heart comprises almost 1% of its body weight.<ref>[[#Schaller|Schaller]], p. 248.</ref> Thus, lions run quickly only in short bursts at about {{cvt|48-59|km/h}} and need to be close to their prey before starting the attack.<ref name=Schaller2478>[[#Schaller|Schaller]], pp. 233, 247–248</ref> They take advantage of factors that reduce visibility; many kills take place near some form of cover or at night.<ref name=Schaller237>[[#Schaller|Schaller]], p. 237.</ref> One study in 2018 recorded a lion running at a top speed of {{cvt|74.1|km/h}}.<ref>{{cite journal |author=Wilson |first1=A. M. |last2=Hubel |first2=T. Y. |last3=Wilshin |first3=S. D. |last4=Lowe |first4=J. C. |last5=Lorenc |first5=M. |last6=Dewhirst |first6=O. P. |last7=Bartlam-Brooks |first7=H. L. |last8=Diack |first8=R.  |last9=Bennitt |first9=E. |last10=Golabek |first10=K. A. |last11=Woledge |first11=R. C. |year=2018 |title=Biomechanics of predator–prey arms race in lion, zebra, cheetah and impala |url=https://rvc-repository.worktribe.com/preview/1388812/11143.pdf |url-status=live |journal=Nature |volume=554 |issue=7691 |pages=183–188 |bibcode=2018Natur.554..183W |doi=10.1038/nature25479 |pmid=29364874 |s2cid=4405091 |archive-url=https://web.archive.org/web/20200305065622/https://researchonline.rvc.ac.uk/id/eprint/11143/1/11143.pdf |archive-date=5 March 2020}}</ref> The lion accelerates at the start of the chase by a rate of {{cvt|9.5|m/s|kph mph}} per second, whereas zebras, wildebeest and [[Thomson's gazelle]] accelerate by a rate of {{cvt|5|m/s|kph mph}} per second, {{cvt|5.6|m/s|kph mph}} per second, and {{cvt|4.5|m/s|kph mph}} per second respectively; acceleration appears to be more important than steady displacement speed in lion hunts.<ref>{{Cite book |author=Alexander |first=R. McNeill |title=Mammals as Predators: The Proceedings of a Symposium Held by The Zoological Society of London and Mammal Society: London, 22nd and 23rd November 1991 |date=1993 |publisher=Oxford University Press |isbn=978-0-19-854067-0 |editor1=Dunstone, N. |place=Oxford |pages=1–13 |chapter=Legs and locomotion of carnivora |doi=10.1093/oso/9780198540670.003.0001 |editor2=Gorman, M. L. |chapter-url=https://archive.org/details/mammalsaspredato0000unse/mode/1up}}</ref> The lion's attack is short and powerful; it attempts to catch prey with a fast rush and final leap, usually pulls it down by the rump, and kills with a clamping bite to the [[Throat clamp|throat]] or [[Muzzle clamp|muzzle]]. It can hold the prey's throat for up to 13 minutes, until the prey stops moving.<ref>[[#Schaller|Schaller]], p. 244, 263–267.</ref> It has a [[Bite force quotient|bite force]] from 1593.8 to 1768 [[Newton (unit)|Newtons]] at the canine tip and up 4167.6 Newtons at the [[carnassial]] notch.<ref>{{Cite journal |last=Thomason |first=J. J. |date=1991 |title=Cranial strength in relation to estimated biting forces in some mammals |journal=Canadian Journal of Zoology |volume=69 |issue=9 |pages=2326–2333 |doi=10.1139/z91-327|bibcode=1991CaJZ...69.2326T}}</ref><ref>{{Cite journal |last1=Wroe |first1=S. |last2=McHenry |first2=C. |last3=Thomason |first3=J. J. |date=2005 |title=Bite club: comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa |journal=Proceedings of the Royal Society B: Biological Sciences |volume=272 |issue=1563 |pages=619–625 |doi=10.1098/rspb.2004.2986 |pmc=1564077 |pmid=15817436}}</ref><ref>{{Cite journal |last1=Van der Meijden |first1=A. |last2=González-Gómez |first2=J. C. |last3=Pulido-Osorio |first3=M. D. |last4=Herrel |first4=A. |date=2023 |title=Measurement of voluntary bite forces in large carnivores using a semi-automated reward-driven system |journal=Journal of Experimental Biology |volume=226 |issue=7 |doi=10.1242/jeb.245255 |page=jeb245255 |pmid=36939369 |bibcode=2023JExpB.226B5255V}}</ref> 
[[File:Male Lion and Cub Chitwa South Africa Luca Galuzzi 2004.JPG |thumb|Male lion and cub with mostly eaten buffalo carcass in [[Sabi Sand Game Reserve]]]]
Lions typically consume prey at the location of the hunt but sometimes drag large prey into cover.<ref name=Schaller2706/> They tend to squabble over kills, particularly the males. Cubs suffer most when food is scarce but otherwise all pride members eat their fill, including old and crippled lions, which can live on leftovers.<ref name=Estes/> Large kills are shared more widely among pride members.<ref name=Schaller133>[[#Schaller|Schaller]], p. 133.</ref> An adult lioness requires an average of about {{cvt|5|kg}} of meat per day while males require about {{cvt|7|kg}}.<ref>[[#Schaller|Schaller]], p. 276.</ref> Lions gorge themselves and eat up to {{cvt|30|kg}} in one session.<ref name=simba>{{Cite book |last=Guggisberg |first=C. A. W. |title=Simba: the life of the lion. |year=1961 |publisher=Howard Timmins |location=Cape Town}}</ref> If it is unable to consume all of the kill, it rests for a few hours before continuing to eat. On hot days, the pride retreats to shade with one or two males standing guard.<ref name=Schaller2706>[[#Schaller|Schaller]], pp. 270–76.</ref> Lions defend their kills from scavengers such as vultures and hyenas.<ref name=Estes/>

Lions scavenge on [[carrion]] when the opportunity arises, scavenging animals dead from natural causes such as disease or those that were killed by other predators. Scavenging lions keep a constant lookout for circling vultures, which indicate the death or distress of an animal.<ref name=Schaller213>[[#Schaller|Schaller]], p. 213–216.</ref> Most carrion on which both hyenas and lions feed upon are killed by hyenas rather than lions.<ref name=nowak/> Carrion is thought to provide a large part of lion diet.<ref name=AWF>{{cite web |title=Behavior and Diet |work=African Wildlife Foundation website |publisher=African Wildlife Foundation |year=1996 |url=http://www.awf.org/wildlife-conservation/lion |access-date=6 June 2014 |archive-date=19 July 2019 |archive-url=https://web.archive.org/web/20190719112213/https://www.awf.org/wildlife-conservation/lion |url-status=live}}</ref>

===Predatory competition===
{{multiple image |perrow=1 |image1=Lioness vs. Hyena in KNP 07.jpg|caption1=Lioness chasing a spotted hyena in Kruger National Park |image2=Lioness vs Leopard 9 July 2016 Latest Sightings 1.png |caption2=Lioness stealing a kill from a leopard in Kruger National Park}}

Lions and [[spotted hyena]]s occupy a similar ecological niche and compete for prey and carrion; a review of data across several studies indicates a dietary overlap of 58.6%.<ref name="prey">{{Cite journal |title=Prey preferences of the spotted hyaena (''Crocuta crocuta'') and degree of dietary overlap with the lion (''Panthera leo'') |url=http://www.zbs.bialowieza.pl/g2/pdf/1598.pdf |archive-url=https://web.archive.org/web/20110430024111/http://www.zbs.bialowieza.pl/g2/pdf/1598.pdf |archive-date=30 April 2011 |url-status=live |last=Hayward |first=M. W. |journal=Journal of Zoology |year=2006 |volume=270 |issue=4 |pages=606–614 |doi=10.1111/j.1469-7998.2006.00183.x}}</ref> Lions typically ignore hyenas unless they are on a kill or are being harassed, while the latter tend to visibly react to the presence of lions with or without the presence of food. In the [[Ngorongoro crater]], lions subsist largely on kills stolen from hyenas, causing them to increase their kill rate.<ref name="Kruuk">{{Cite book |first=H. |last=Kruuk |title=The Spotted Hyena: A Study of Predation and Social Behaviour |publisher=Echo Point Books & Media |year=2014|isbn=978-1626549050 |pages=128–138|edition=2nd}}</ref> In Botswana's Chobe National Park, the situation is reversed as hyenas there frequently challenge lions and steal their kills, obtaining food from 63% of all lion kills.<ref name="Conservation">{{Cite book |chapter=Interspecific competition and the population biology of extinction-prone carnivores |last1=Creel |first1= S.|last2= Spong |first2=G. |last3=Creel|first3= N. |pages=35–60 |chapter-url=https://books.google.com/books?id=v39RdyYUfRIC&pg=PA36 |title=Carnivore Conservation |edition=First |editor1-first=J. L. |editor1-last=Gittleman |editor2-first=S. M. |editor2-last=Funk |editor3-first=D. W. |editor3-last=Macdonald |editor4-first=R. K. |editor4-last=Wayne |publisher=Cambridge University Press |year=2001 |isbn=978-0-521-66232-1}}</ref> When confronted on a kill, hyenas may either leave or wait patiently at a distance of {{cvt|30|-|100|m}} until the lions have finished.<ref name="schaller272">[[#Schaller|Schaller]], p. 272.</ref> Hyenas may feed alongside lions and force them off a kill. The two species attack one another even when there is no food involved for no apparent reason.<ref name="schaller273">[[#Schaller|Schaller]], pp. 273–74.</ref> Lions can account for up to 71% of hyena deaths in [[Etosha National Park]]. Hyenas have adapted by frequently mobbing lions that enter their home ranges.<ref>{{Cite journal |title=Competitive interactions between spotted hyenas and lions in the Etosha National Park, Namibia |last1=Trinkel |first1=M. |last2=Kastberger |first2=G. |journal=African Journal of Ecology |volume=43 |issue=3 |year=2005 |pages=220–224 |doi=10.1111/j.1365-2028.2005.00574.x|bibcode=2005AfJEc..43..220T }}</ref> When the lion population in Kenya's [[Masai Mara National Reserve]] declined, the spotted hyena population increased rapidly.<ref>{{cite journal |last1=Green |first1= D. S.| last2= Johnson-Ulrich |first2=L.| last3=Couraud | first3=H. E. |last4=Holekamp |first4=K. E. |year=2018 |title=Anthropogenic disturbance induces opposing population trends in spotted hyenas and African lions |journal=Biodiversity and Conservation |volume=27 |issue=4 |pages=871–889 |doi=10.1007/s10531-017-1469-7 |bibcode= 2018BiCon..27..871G|s2cid= 44240882}}</ref>

Lions tend to dominate [[cheetah]]s and leopards, steal their kills and kill their cubs and even adults when given the chance.<ref>{{Cite book|last1=Denis-Hoot|first1=C. |last2= Denis-Hoot|first2= M. |year=2003 |title=The Art of Being a Lion |publisher=Barnes & Noble |location=New York |isbn=9780760747674 |page=198}}</ref> Cheetahs often lose their kills to lions or other predators.<ref>{{Cite journal |last1=O'Brien |first1=S. J. |last2=Wildt |first2 =D. E. |last3=Bush |first3=M. |year=1986 |title=The Cheetah in Genetic Peril |journal=Scientific American |volume=254 |issue=5 |pages=68–76 |doi=10.1038/scientificamerican0586-84 |url=http://www.catsg.org/cheetah/05_library/5_3_publications/N_and_O/OBrien_et_al_1986_Cheetah_in_genetic_peril.pdf |archive-url=https://web.archive.org/web/20110513071205/http://www.catsg.org/cheetah/05_library/5_3_publications/N_and_O/OBrien_et_al_1986_Cheetah_in_genetic_peril.pdf |archive-date=13 May 2011 |url-status=live|bibcode=1986SciAm.254e..84O}}</ref> A study in the Serengeti ecosystem revealed that lions killed at least 17 of 125 cheetah cubs born between 1987 and 1990.<ref name="laurenson">{{cite journal |last1=Laurenson |first1=M. K. |title=High juvenile mortality in cheetahs (''Acinonyx jubatus'') and its consequences for maternal care |journal=Journal of Zoology |year=1994 |volume=234 |issue=3 |pages=387–408 |doi=10.1111/j.1469-7998.1994.tb04855.x |url=http://www.catsg.org/cheetah/05_library/5_3_publications/L/Laurenson_1994_Cheetah_cub_mortality_-_maternal_care.pdf |archive-url=https://web.archive.org/web/20171120042410/http://www.catsg.org/cheetah/05_library/5_3_publications/L/Laurenson_1994_Cheetah_cub_mortality_-_maternal_care.pdf |archive-date=20 November 2017 |url-status=live}}</ref> Cheetahs avoid their competitors by hunting at different times and habitats.<ref>{{cite journal|last1=Rostro-García |first1=S. |last2=Kamler |first2=J. F. |last3=Hunter |first3=L. T. B.  |year=2015 |title=To kill, stay or flee: the effects of lions and landscape factors on habitat and kill site selection of cheetahs in South Africa |journal=PLOS ONE |volume=10 |issue=2 |page=e0117743 |pmid=25693067 |doi=10.1371/journal.pone.0117743 |pmc=4333767|bibcode=2015PLoSO..1017743R |doi-access=free}}</ref> Leopards, by contrast, do not appear to be motivated by an avoidance of lions, as they use heavy vegetation regardless of whether lions are present in an area and both cats are active around the same time of day. In addition, there is no evidence that lions affect leopard abundance.<ref>{{cite journal|last1=Miller|first1=J. R. B.|last2=Pitman|first2=R. T.|last3=Mann|first3=G. K. H.|last4=Fuller|first4=A. K.|last5=Balme|first5=G. A.|year=2018|title=Lions and leopards coexist without spatial, temporal or demographic effects of interspecific competition|journal=Journal of Animal Ecology|volume=87|issue=6|pages=1709–1726|doi=10.1111/1365-2656.12883|pmid=30010193 |bibcode=2018JAnEc..87.1709M }}</ref> Leopards take refuge in trees, though lionesses occasionally attempt to climb up and retrieve their kills.<ref name="Schaller293">[[#Schaller|Schaller]], p. 293.</ref>

Lions similarly dominate [[African wild dog]]s, taking their kills and dispatching pups or adult dogs. Population densities of wild dogs are low in areas where lions are more abundant.<ref>{{cite journal |title=Conserving the African wild dog ''Lycaon pictus''. I. Diagnosing and treating causes of decline |last1=Woodroffe |first1=R. |last2=Ginsberg |first2=J. R.  |year=1999 |journal=Oryx |volume=33 |pages=132–142 |doi=10.1046/j.1365-3008.1999.00052.x |issue=2 |doi-access=free}}</ref><!--cites two previous refs--> However, there are a few reported cases of old and wounded lions falling prey to wild dogs.<ref name=Pienaar1969>{{Cite journal |last1=Pienaar |first1=U. de V. |title=Predator–prey relationships among the larger mammals of the Kruger National Park |journal=Koedoe |date=1969 |volume=12 |issue=1 |pages=108–176 |doi=10.4102/koedoe.v12i1.753 |doi-access=free}}</ref><ref name="Schaller, p. 188">[[#Schaller|Schaller]], p. 188.</ref>

===Reproduction and life cycle===

{{multiple image|align=right|direction=vertical
|image1=Panthera leo massaica mating.jpg|caption1=Lions mating at Masai Mara
|image2=Lion_cub,_Masai_Mara,_Kenya.jpg|caption2=A lion cub in Masai Mara}}

Most lionesses reproduce by the time they are four years of age.<ref name="Schaller29">[[#Schaller|Schaller]], p. 29.</ref> Lions do not mate at a specific time of year and the females are [[polyestrous]].<ref name="Schaller174">[[#Schaller|Schaller]], p. 174.</ref> Like those of other cats, the male lion's penis has [[Penile spines|spines]] that point backward. During withdrawal of the penis, the spines rake the walls of the female's vagina, which may cause [[ovulation]].<ref>{{Cite journal | doi=10.1002/zoo.1430130403|title = Spontaneous and induced ovulation in the lion (Panthera leo)| journal=Zoo Biology| volume=13| issue=4| pages=301–307|year = 1994|last1 = Schramm|first1 = Ralph Dee| last2=Briggs| first2=Michael B.| last3=Reeves| first3=Jerry J.}}</ref><ref>{{Cite book |last=Asdell |first=Sydney A. |title=Patterns of mammalian reproduction |year=1993 |orig-date=1964 |publisher=Cornell University Press |location=Ithaca |isbn=978-0-8014-1753-5 |url=https://archive.org/details/asdellspatternso00hays }}</ref> A lioness may mate with more than one male when she is [[Estrous cycle|in heat]].<ref name="Schaller142">[[#Schaller|Schaller]], p. 142.</ref> Lions of both sexes may be involved in group [[Homosexuality in animals|homosexual]] and courtship activities. Males will also head-rub and roll around with each other before mounting each other.<ref>{{Cite book |last=Bagemihl |first=Bruce |year=1999 |title=Biological Exuberance: Animal Homosexuality and Natural Diversity |location=New York |publisher=St. Martin's Press |isbn=978-0-312-19239-6 |pages=[https://archive.org/details/biologicalexuber00bage/page/302 302–305] |url=https://archive.org/details/biologicalexuber00bage/page/302 }}</ref><ref>[[#Schaller|Schaller]], p. 137.</ref> [[Generation time|Generation length]] of the lion is about seven years.<ref>{{cite journal |title=Generation length for mammals |author=Pacifici, M.|author2= Santini, L.|author3= Di Marco, M.|author4= Baisero, D.|author5= Francucci, L.|author6= Grottolo Marasini, G.|author7= Visconti, P. |author8= Rondinini, C. |journal=Nature Conservation |year=2013 |issue=5 |pages=87–94}}</ref> The average gestation period is around 110{{spaces}}days;<ref name="Schaller174"/> the female gives birth to a litter of between one and four cubs in a secluded den, which may be a thicket, a reed-bed, a cave, or some other sheltered area, usually away from the pride. She will often hunt alone while the cubs are still helpless, staying relatively close to the den.<ref name="Scott45">[[#Scott|Scott]], p. 45.</ref> Lion cubs are born blind, their eyes opening around seven days after birth. They weigh {{convert|1.2|–|2.1|kg|lb|abbr=on}} at birth and are almost helpless, beginning to crawl a day or two after birth and walking around three weeks of age.<ref name="Schaller143">[[#Schaller|Schaller]], p. 143.</ref> To avoid a buildup of scent attracting the attention of predators, the lioness moves her cubs to a new den site several times a month, carrying them one-by-one by the nape of the neck.<ref name="Scott45"/>

Usually, the mother does not integrate herself and her cubs back into the pride until the cubs are six to eight weeks old.<ref name="Scott45"/> Sometimes the introduction to pride life occurs earlier, particularly if other lionesses have given birth at about the same time.<ref name=Estes/><ref name=Schaller148/> When first introduced to the rest of the pride, lion cubs lack confidence when confronted with adults other than their mother. They soon begin to immerse themselves in the pride life, however, playing among themselves or attempting to initiate play with the adults.<ref name=Schaller148/> Lionesses with cubs of their own are more likely to be tolerant of another lioness's cubs than lionesses without cubs. Male tolerance of the cubs varies—one male could patiently let the cubs play with his tail or his mane, while another may snarl and bat the cubs away.<ref>[[#Scott|Scott]], p. 46.</ref>
[[File:Lion Cubs Phinda 2011.ogv|thumb|right|Video of a lioness and her cubs in Phinda Reserve]]

Pride lionesses often synchronise their reproductive cycles and communal rearing and suckling of the young, which suckle indiscriminately from any or all of the nursing females in the pride. The synchronisation of births is advantageous because the cubs grow to being roughly the same size and have an equal chance of survival, and sucklings are not dominated by older cubs.<ref name=Estes/><ref name=Schaller148>[[#Schaller|Schaller]], p. 147-49.</ref> Weaning occurs after six or seven months. Male lions reach maturity at about three years of age and at four to five years are capable of challenging and displacing adult males associated with another pride. They begin to age and weaken at between 10 and 15 years of age at the latest.<ref>{{Cite book |last=Crandall |first=L. S. |title=The management of wild animals in captivity |year=1964 |publisher=University of Chicago Press |location=Chicago |oclc=557916}}</ref>

When one or more new males oust the previous males associated with a pride, the victors often [[infanticide (zoology)|kill any existing young cubs]], perhaps because females do not become fertile and receptive until their cubs mature or die. Females often fiercely defend their cubs from a usurping male but are rarely successful unless a group of three or four mothers within a pride join forces against the male.<ref name="Packpus83">{{Cite journal |last1=Packer |first1=C. |last2=Pusey |first2=A. E. |date=May 1983 |title= Adaptations of female lions to infanticide by incoming males|journal= American Naturalist |volume=121 |issue=5 |pages=716–728 |doi= 10.1086/284097|bibcode=1983ANat..121..716P |s2cid=84927815 }}</ref> Cubs also die from starvation and abandonment, and predation by leopards, hyenas and wild dogs. Male cubs are excluded from their maternal pride when they reach maturity at around two or three years of age,<ref name="Schaller44">[[#Schaller|Schaller]], p. 44.</ref> while some females may leave when they reach the age of two.<ref name=Packer33/> When a new male lion takes over a pride, adolescents both male and female may be evicted.<ref>[[#Scott|Scott]], p. 68.</ref>

===Health and mortality===

[[File:Lake-Nakuru-Lions-in-Tree.JPG|thumb|Lions in a tree near [[Lake Nakuru]]]]

Lions may live 12–17 years in the wild.<ref name=Haas2005/> Although adult lions have no natural predators, evidence suggests most die violently from attacks by humans or other lions.<ref name="Schaller183">[[#Schaller|Schaller]], p. 183.</ref> Lions often inflict serious injuries on members of other prides they encounter in territorial disputes or members of the home pride when fighting at a kill.<ref name="Schaller1889">[[#Schaller|Schaller]], pp. 188–189.</ref> Crippled lions and cubs may fall victim to hyenas and leopards or be trampled by buffalo or elephants. Careless lions may be maimed when hunting prey.<ref name="Schaller1890">[[#Schaller|Schaller]], pp. 189–190.</ref>  [[Nile crocodile]]s may also kill and eat lions, evidenced by the occasional lion claw found in crocodile stomachs.<ref name="Guggisberg1972">{{Cite book |last=Guggisberg |first=C. A. W. |title=Crocodiles: Their Natural History, Folklore, and Conservation |year=1972 |page=195 |isbn=978-0-7153-5272-4 |publisher=David & Charles |location=Newton Abbot |url=<!--rem url : no preview provided-->}}</ref>

[[Tick]]s commonly infest the ears, neck and groin regions of the lions.<ref name="Schaller184">[[#Schaller|Schaller]], p. 184.</ref><ref>{{Cite book |last=Yeoman |first=Guy Henry|author2=Walker, Jane Brotherton |title=The ixodid ticks of Tanzania |year=1967 |publisher=Commonwealth Institute of Entomology |location=London |oclc=955970}}</ref> Adult forms of several [[tapeworm]] species of the genus ''[[Taenia (flatworm)|Taenia]]'' have been isolated from lion intestines, having been ingested as larvae in [[antelope]] meat.<ref>{{Cite journal |last=Sachs |first=R. |year=1969|title=Untersuchungen zur Artbestimmung und Differenzierung der Muskelfinnen ostafrikanischer Wildtiere [Differentiation and species determination of muscle-cysticerci in East African game animals] |journal=Zeitschrift für Tropenmedizin und Parasitologie |volume=20 |issue=1 |pages=39–50 |pmid=5393325|language=de}}</ref> Lions in the Ngorongoro Crater were afflicted by an outbreak of stable fly (''[[Stable fly|Stomoxys calcitrans]]'') in 1962, resulting in lions becoming emaciated and covered in bloody, bare patches. Lions sought unsuccessfully to evade the biting flies by climbing trees or crawling into hyena burrows; many died or migrated and the local population dropped from 70 to 15 individuals.<ref>{{Cite journal |last=Fosbrooke |first=H. |year=1963 |title=The stomoxys plague in Ngorongoro |journal=East African Wildlife Journal |volume=1 |issue=6 |pages=124–126 |doi=10.1111/j.1365-2028.1963.tb00190.x}}</ref> A more recent outbreak in 2001 killed six lions.<ref name="Nkwame06">{{Cite news | last=Nkwame | first=V. M. |title=King of the jungle in jeopardy | work=The Arusha Times| date=9 September 2006 | url=http://www.arushatimes.co.tz/2006/36/features_10.htm | access-date=4 September 2007 | url-status=dead | archive-url=https://web.archive.org/web/20070929044925/http://www.arushatimes.co.tz/2006/36/features_10.htm | archive-date=29 September 2007 | df=dmy-all}}</ref>

Captive lions have been infected with [[canine distemper]] virus (CDV) since at least the mid-1970s.<ref>{{Cite journal |author1=Myers, D. L. |author2=Zurbriggen, A. |author3=Lutz, H. |author4=Pospischil, A. |date=1997 |title=Distemper: not a new disease in lions and tigers |journal=Clinical Diagnostic Laboratory Immunology |volume=4 |issue=2 |pages=180–184 |doi=10.1128/CDLI.4.2.180-184.1997 |pmid=9067652 |pmc=170498 |url=}}</ref> CDV is spread by domestic dogs and other carnivores; a 1994 outbreak in Serengeti National Park resulted in many lions developing neurological symptoms such as seizures. During the outbreak, several lions died from [[pneumonia]] and [[encephalitis]].<ref>{{Cite journal|last1=Roelke-Parker|first1=M. E.|date=1996 |title=A canine distemper epidemic in Serengeti lions (''Panthera leo'') |journal=Nature |volume=379 |issue=6564 |pages=441–445 |doi=10.1038/379441a0|pmid=8559247 |last2=Munson |first2=L. |last3=Packer |first3=C. |last4=Kock |first4=R. |last5=Cleaveland |first5=S. |last6=Carpenter |first6=M. |last7=O'Brien |first7=S. J. |last8=Pospischil |first8=A. |last9=Hofmann-Lehmann |first9=R. |first10=Hans |last10=L. |first11=G. L. M. |last11=Mwanengele |first12=M. N. |last12=Mgasa |first13=G. A. |last13=Machange |first14=B. A. |last14=Summers |first15=M. J. G. |last15=Appel |bibcode=1996Natur.379..441R|pmc=7095363 }}</ref> [[Feline immunodeficiency virus]] and [[lentivirus]] also affect captive lions.<ref>{{Cite journal |author1=Bull, M. E. |author2=Kennedy-Stoskopf, S. |author3=Levine, J. F. |author4=Loomis, M. |author5=Gebhard, D. G. |author6=Tompkins, W. A. |date=2003 |title=Evaluation of T lymphocytes in captive African lions (''Panthera leo'') infected with feline immunodeficiency virus |journal=American Journal of Veterinary Research |volume=64 |issue=10 |pages=1293–1300 |doi=10.2460/ajvr.2003.64.1293|pmid=14596469 |doi-access=free }}</ref><ref>{{Cite journal |author1=Poli, A. |author2=Abramo, F. |author3=Cavicchio, P. |author4=Bandecchi, P. |author5=Ghelardi, E. |author6=Pistello, M. |date=1995 |title=Lentivirus infection in an African lion: a clinical, pathologic and virologic study |journal=Journal of Wildlife Diseases |volume=31 |issue=1 |pages=70–74 |doi=10.7589/0090-3558-31.1.70|pmid=7563428 |doi-access=free }}</ref>

===Communication===
{{multiple image|perrow=1|image1=Lion cub with mother.jpg|caption1=Head rubbing among pride members is a common social behaviour.|image2=Lion (Panthera leo) marking its territory ... (52806595064).jpg
|caption2=A male lion raises his tail while marking his territory.}}

When resting, lion socialisation occurs through a number of behaviours; the animal's expressive movements are highly developed. The most common peaceful, tactile gestures are [[Bunting (animal behavior)|head rubbing]] and [[social licking]],<ref name=Schaller85>[[#Schaller|Schaller]], p. 85.</ref> which have been compared with the role of [[allogrooming]] among primates.<ref>{{Cite book |last=Sparks |first=J. |title=Primate Ethology |chapter=Allogrooming in primates: a review |pages=148–175 |orig-year=1967 |editor=Morris, D. |publisher=Aldine |location=Chicago |date=2011 |isbn=9780202368160}}</ref> Head rubbing, nuzzling the forehead, face and neck against another lion appears to be a form of greeting<ref>{{Cite book |last=Leyhausen |first=P. |title=Verhaltensstudien an Katzen |edition=Second |year=1960 |publisher=Paul Parey |location=Berlin |oclc=636909227 |language=de}}</ref> and is seen often after an animal has been apart from others or after a fight or confrontation. Males tend to rub other males, while cubs and females rub females.<ref name=Schaller858>[[#Schaller|Schaller]], pp. 85–88.</ref> Social licking often occurs in tandem with head rubbing; it is generally mutual and the recipient appears to express pleasure. The head and neck are the most common parts of the body licked; this behaviour may have arisen out of utility because lions cannot lick these areas themselves.<ref name=Schaller8891>[[#Schaller|Schaller]], pp. 88–91.</ref>
{{listen|filename=Lion raring-sound1TamilNadu178.ogg|title=Lion roar|description=A captive lion roaring}}

Lions have an array of facial expressions and body postures that serve as visual gestures.<ref name=Schaller92102>[[#Schaller|Schaller]], pp. 103–117.</ref> A common facial expression is the "grimace face" or [[flehmen response]], which a lion makes when sniffing chemical signals and involves an open mouth with bared teeth, raised muzzle, wrinkled nose, closed eyes and relaxed ears.<ref>[[#Schaller|Schaller]], p. 95.</ref> Lions also use chemical and visual marking;<ref name=Schaller92102/> males [[Territory (animal)#Scent marking|spray urine]]<ref name="Schaller116">[[#Schaller|Schaller]], [https://books.google.com/books?id=7ann2dYn9iYC&pg=PA116 p. 116.]</ref><ref>{{cite journal |author1=Andersen, K. F.  |author2=Vulpius, T. |year=1999 |title=Urinary volatile constituents of the lion, ''Panthera leo'' |journal=Chemical Senses |volume=24 |issue=2 |pages=179–189 |doi=10.1093/chemse/24.2.179 |pmid=10321819 |doi-access=free}}</ref> and scrape plots of ground and objects within the territory.<ref name=Schaller92102/>

The lion's repertoire of vocalisations is large; variations in intensity and pitch appear to be central to communication. Most lion vocalisations are variations of [[growling]], [[snarling]], meowing and roaring. Other sounds produced include puffing, bleating and humming. Roaring is used to advertise its presence. Lions most often roar at night, a sound that can be heard from a distance of {{convert|8|km|0}}.<ref name=Schaller10313>[[#Schaller|Schaller]], pp. 103–113.</ref> They tend to roar in a very characteristic manner starting with a few deep, long roars that subside into grunts.<ref>{{Cite journal |last1=Eklund |first1=R. |last2=Peters |first2=G. |last3=Ananthakrishnan |first3=G. |last4=Mabiza |first4=E. |title=An acoustic analysis of lion roars. I: Data collection and spectrogram and waveform analyses |journal=Speech, Music and Hearing Quarterly Progress and Status Report |volume=51 |page=1 |year=2011 |url=http://roberteklund.info/pdf/Eklund_et_al_2011_LionRoars.pdf |archive-url=https://web.archive.org/web/20111001083629/http://roberteklund.info/pdf/Eklund_et_al_2011_LionRoars.pdf |archive-date=1 October 2011 |url-status=live}}</ref><ref>{{Cite journal |last1=Ananthakrishnan |first1=G. |last2=Eklund |first2=R. |last3=Peters |first3=G. |last4=Mabiza |first4=E. |title=An acoustic analysis of lion roars. II: Vocal tract characteristics |journal=Speech, Music and Hearing Quarterly Progress and Status Report |volume=51 |page=5 |year=2011 |url=http://roberteklund.info/pdf/Ananthakrishnan_et_al_2011_LionRoars.pdf |archive-url=https://web.archive.org/web/20111001083621/http://roberteklund.info/pdf/Ananthakrishnan_et_al_2011_LionRoars.pdf |archive-date=1 October 2011 |url-status=live}}</ref>