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{{Short description|Large cat native to Africa and India}}
{{other uses}}
{{Distinguish|Mountain lion}} {{Featured article}}
{{pp|vandalism|small=y}}
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| name = Lion
| fossil_range = {{fossil range|Pleistocene|Present}}
<!--Please do not replace these featured images without a consensus-->
| image = 020 The lion king Snyggve in the Serengeti National Park Photo by Giles Laurent.jpg | image_caption = Male in [[Serengeti National Park]], Tanzania
| image2 = Okonjima Lioness.jpg
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| status = VU
| status_system = IUCN3.1
| status_ref = <ref name=IUCN>{{cite iucn |title=''Panthera leo''
| status2 = CITES_A2
| status2_system = CITES
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==Etymology==
The English word ''lion'' is derived via [[Anglo-Norman language|Anglo-Norman]] {{lang|xno|liun}} from [[Latin]] {{lang|la|leōnem}} (nominative: {{lang|la|leō}}), which in turn was a borrowing from [[Ancient Greek]] {{lang|grc|λέων}} {{lang|grc-Latn|léōn}}. The [[Hebrew language|Hebrew]] word {{lang|he|לָבִיא|rtl=yes}} {{lang|he-Latn|lavi}} may also be related.<ref>{{cite OED|lion|access-date=20 March 2022}}</ref> The generic name ''Panthera'' is traceable to the [[classical Latin]] word 'panthēra' and the [[ancient Greek]] word πάνθηρ 'panther'.<ref>{{cite book |last1=Liddell, H. G. |last2=Scott, R.
==Taxonomy==
[[File:Two cladograms for Panthera.svg|thumb|right|The upper cladogram is based on the 2006 study,<ref name=Johnson2006/><ref name="werdelin2009">{{cite journal |last1=Werdelin |first1=L. |last2=Yamaguchi |first2=N. |last3=Johnson |first3=W. E. |last4=O'Brien |first4=S. J.
''Felis leo'' was the [[scientific name]] used by [[Carl Linnaeus]] in 1758, who described the lion in his work ''[[Systema Naturae]]''.<ref name="Linn1758">{{cite book |last=Linnaeus|first= C. |year=1758 |title=Caroli Linnæi Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis |volume=Tomus I |edition=decima, reformata |location=Holmiae |publisher=Laurentius Salvius |page=41 |chapter=''Felis leo'' |chapter-url=https://archive.org/stream/mobot31753000798865#page/41/mode/2up}} {{in lang|la}}</ref> The genus name ''Panthera'' was coined by [[Lorenz Oken]] in 1816.<ref>{{cite book |last1=Oken |first1=L. |year=1816 |title=Lehrbuch der Zoologie. 2. Abtheilung |location=Jena |publisher=August Schmid & Comp. |page=1052 |chapter=1. Art, ''Panthera'' |chapter-url=https://books.google.com/books?id=S5o5AAAAcAAJ&pg=PA1052}}</ref> Between the mid-18th and mid-20th centuries, 26 lion [[Zoological specimen|specimen]]s were described and proposed as subspecies, of which 11 were recognised as [[Valid name (zoology)|valid]] in 2005.<ref name=MSW3/> They were distinguished mostly by the size and colour of their manes and skins.<ref name=Hemmer/>
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[[File:Lion subspecies distribution3.png|thumb|Range map showing distribution of subspecies and clades]]
In the 19th and 20th centuries, several lion [[type specimen]]s were described and proposed as [[subspecies]], with about a dozen recognised as [[Valid name (zoology)|valid]] [[Taxon|taxa]] until 2017.<ref name=MSW3/> Between 2008 and 2016, [[IUCN Red List]] assessors used only two subspecific names: ''P. l. leo'' for African lion populations, and ''P. l. persica'' for the Asiatic lion population.<ref name=IUCN/><ref name=Breitenmoser2008>{{cite iucn |author=Breitenmoser, U. |author2=Mallon, D. P. |author3=Ahmad Khan, J. |author4=Driscoll, C.
*[[Panthera leo leo|''P. l. leo'']] {{small|(Linnaeus, 1758)}} − the [[Nominate subspecies|nominate]] lion subspecies includes the [[Asiatic lion]], the [[Locally extinct|regionally extinct]] [[Barbary lion]], and lion populations in West and northern parts of Central Africa.<ref name=catsg/> [[Synonym (taxonomy)|Synonyms]] include ''P. l. persica'' {{small|(Meyer, 1826)}}, ''P. l. senegalensis'' {{small|(Meyer, 1826)}}, ''P. l. kamptzi'' {{small|([[Paul Matschie|Matschie]], 1900)}}, and ''P. l. azandica'' {{small|([[Joel Asaph Allen|Allen]], 1924)}}.<ref name=MSW3/> Multiple authors referred to it as 'northern lion' and 'northern subspecies'.<ref name="Wood1865">{{cite book |last=Wood |first=J. G. |author-link=John George Wood |title=The Illustrated Natural History
*[[Panthera leo melanochaita|''P. l. melanochaita'']] {{small|([[Charles Hamilton Smith|Smith]], 1842)}} − includes the extinct [[Cape lion]] and lion populations in East and Southern African regions.<ref name=catsg/> Synonyms include ''P. l. somaliensis'' {{small|(Noack 1891)}}, ''P. l. massaica'' {{small|([[Oscar Neumann|Neumann]], 1900)}}, ''P. l. sabakiensis'' {{small|([[Einar Lönnberg|Lönnberg]], 1910)}}, ''P. l. bleyenberghi'' {{small|(Lönnberg, 1914)}}, ''P. l. roosevelti'' {{small|([[Edmund Heller|Heller]], 1914)}}, ''P. l. nyanzae'' {{small|(Heller, 1914)}}, ''P. l. hollisteri'' {{small|([[Joel Asaph Allen|Allen]], 1924)}}, ''P. l. krugeri'' {{small|([[Austin Roberts (zoologist)|Roberts]], 1929)}}, ''P. l. vernayi'' {{small|(Roberts, 1948)}}, and ''P. l. webbiensis'' {{small|(Zukowsky, 1964)}}.<ref name=MSW3 /><ref name=Hemmer/> It has been referred to as 'southern subspecies' and 'southern lion'.<ref name=Hunter2018/>
However, there seems to be some degree of overlap between both groups in northern Central Africa. DNA analysis from a more recent study indicates that Central African lions are derived from both northern and southern lions, as they cluster with ''P. leo leo'' in mtDNA-based phylogenies whereas their genomic DNA indicates a closer relationship with P. ''leo melanochaita''.<ref>{{cite journal |last1=de Manuel |first1=M
Lion samples from some parts of the [[Ethiopian Highlands]] cluster genetically with those from Cameroon and Chad, while lions from other areas of Ethiopia cluster with samples from East Africa. Researchers, therefore, assume Ethiopia is a contact zone between the two subspecies.<ref name=Bertola2016>{{cite journal |author1=Bertola, L. D. |author2=Jongbloed, H. |author3=Van Der Gaag, K. J. |author4=De Knijff, P. |author5=Yamaguchi, N. |author6=Hooghiemstra, H. |author7=Bauer, H. |author8=Henschel, P. |author9=White, P. A. |author10=Driscoll, C. A. |author11=Tende, T. |author12=Ottosson, U. |author13=Saidu, Y. |author14=Vrieling, K. |author15=de Iongh, H. H. |year=2016 |title=Phylogeographic patterns in Africa and High Resolution Delineation of genetic clades in the Lion (''Panthera leo'') |journal=Scientific Reports |volume=6 |page=30807 |doi=10.1038/srep30807 |pmid=27488946 |pmc=4973251
===Fossil records===
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Other lion subspecies or [[sister species]] to the modern lion existed in prehistoric times:<ref name="Christiansen08CaveLions">{{cite journal |last1=Christiansen |first1=P. |year=2008 |title=Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters |journal=[[Cladistics (journal)|Cladistics]] |volume=24 |issue=6 |pages=977–992 |doi=10.1111/j.1096-0031.2008.00226.x|pmid=34892880 |s2cid=84497516 |doi-access=free }}</ref>
*''[[Panthera leo sinhaleyus|P. l. sinhaleyus]]'' was a [[fossil]] [[carnassial]] excavated in [[Sri Lanka]], which was attributed to a lion. It is thought to have become extinct around 39,000 years ago.<ref>{{Cite journal|first1=K. |last1=Manamendra-Arachchi |first2=R. |last2=Pethiyagoda |first3=R. |last3=Dissanayake |first4=M. |last4=Meegaskumbura
*[[Panthera fossilis|''P. fossilis'']] was larger than the modern lion and lived in the [[Middle Pleistocene]]. Bone fragments were excavated in caves in the United Kingdom, Germany, Italy and Czech Republic.<ref name=Marciszak2010>{{cite journal |last1=Marciszak |first1=A. |last2=Stefaniak |first2=K.
*''[[Panthera spelaea|P. spelaea]]'', or the '''cave lion''', lived in [[Eurasia]] and [[Beringia]] during the [[Late Pleistocene]]. It became extinct due to [[climate warming]] or [[Early human migrations|human expansion]] latest by 11,900 years ago.<ref name=Stuart2011>{{cite journal |last1=Stuart |first1=A. J. |last2=Lister |first2=A. M.
*[[American lion|''P. atrox'']], or the '''American lion''', ranged in the Americas from Canada to possibly [[Patagonia]] during the Late Pleistocene.<ref name="Chimento2017">{{Cite journal |last1=Chimento |first1=N. R. |last2=Agnolin |first2=F. L.
===Evolution===
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[[File:Cave lion range.png|thumb|red ''[[Panthera spelaea]]''{{break}}blue ''[[Panthera atrox]]''{{break}}green ''Panthera leo''{{break}}{{break}}Maximal range of the modern lion{{break}}and its prehistoric relatives{{break}}in the late Pleistocene]]
The ''Panthera'' [[Lineage (evolution)|lineage]] is estimated to have [[Genetic divergence|genetically diverged]] from the [[common ancestor]] of the [[Felidae]] around {{Ma|9.32|4.47|million years ago}} to {{Ma|11.75|0.97|million years ago}}.<ref name="Johnson2006">{{cite journal |last1=Johnson |first1=W. E. |last2=Eizirik |first2=E. |last3=Pecon-Slattery |first3=J. |last4=Murphy |first4=W. J. |last5=Antunes |first5=A. |last6=Teeling |first6=E. |last7=O'Brien |first7=S. J.
Estimates for the divergence time of the modern and cave lion lineages range from 529,000 to 392,000 years ago based on [[mutation rate]] per generation time of the modern lion. There is no evidence for [[gene flow]] between the two lineages, indicating that they did not share the same geographic area.<ref name="DeManuel_al2020" /> The Eurasian and American cave lions became extinct at the end of the [[last glacial period]] without [[Mitochondrion|mitochondrial]] descendants on other continents.<ref name="BurgerJ-Molecular-phylogeny"/><ref name=Barnett>{{Cite journal |last1=Barnett |first1=R. |last2=Shapiro |first2=B. |author-link2=Beth Shapiro |last3=Barnes |first3=I. |last4=Ho |first4=S. Y. W. |last5=Burger |first5=J. |author-link5=Joachim Burger |last6=Yamaguchi |first6=N. |last7=Higham |first7=T. F. G. |last8=Wheeler |first8=H. T. |title=Phylogeography of lions (''Panthera leo'' ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity |journal=[[Molecular Ecology]] |date=2009 |volume=18 |issue=8 |pages=1668–1677 |pmid=19302360 |doi=10.1111/j.1365-294X.2009.04134.x |first9=W. |last9=Rosendahl |last10=Sher |first10=A. V. |last11=Sotnikova |first11=M. |last12=Kuznetsova |first12=T. |last13=Baryshnikov |first13=G. F. |last14=Martin |first14=L. D. |last15=Harington |first15=C. R. |last16=Burns |first16=J. A. |last17=Cooper |first17=A. |bibcode=2009MolEc..18.1668B |s2cid=46716748
Extinction of lions in southern Europe, North Africa and the Middle East interrupted gene flow between lion populations in Asia and Africa. Genetic evidence revealed numerous [[mutation]]s in lion samples from East and Southern Africa, which indicates that this group has a longer evolutionary history than genetically less diverse lion samples from Asia and West and Central Africa.<ref name=Bertola2011>{{Cite journal |last1=Bertola |first1=L. D. |last2=Van Hooft |first2=W. F. |last3=Vrieling |first3=K. |last4=Uit De Weerd |first4=D. R. |last5=York |first5=D. S. |last6=Bauer |first6=H. |last7=Prins |first7=H. H. T. |last8=Funston |first8=P. J. |last9=Udo De Haes |first9=H. A. |last10=Leirs |first10=H. |last11=Van Haeringen |first11=W. A. |last12=Sogbohossou |first12=E. |last13=Tumenta |first13=P. N. |last14=De Iongh |first14=H. H.
===Hybrids===
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{{Further|Panthera hybrid}}
In zoos, lions have been bred with [[tiger]]s to create hybrids for the curiosity of visitors or for scientific purpose.<ref>{{cite journal |author=Pocock, R. I. |year=1898 |title=Lion-Tiger Hybrid |journal=Nature |volume=58 |issue=1496 |page=200 |doi=10.1038/058200b0 |url=https://zenodo.org/record/1889713 |bibcode=1898Natur..58Q.200P |s2cid=4056029 |doi-access=free |access-date=7 December 2019 |archive-date=26 March 2022 |archive-url=https://web.archive.org/web/20220326042949/https://zenodo.org/record/1889713 |url-status=live }}</ref><ref>{{cite book |author=Benirschke, K. |title=Comparative Aspects of Reproductive Failure |chapter=Sterility and Fertility of Interspecific Mammalian Hybrids |pages=
==Description==
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Its skull is very similar to that of the tiger, although the frontal region is usually more depressed and flattened and has a slightly shorter [[postorbital]] region and broader nasal openings than those of the tiger. Due to the amount of skull variation in the two species, usually only the structure of the lower jaw can be used as a reliable indicator of species.<ref name=Pocock1939>{{cite book |author=Pocock, R. I. |year=1939 |title=The Fauna of British India, including Ceylon and Burma |volume=((Mammalia. Volume 1)) |publisher=Taylor and Francis Ltd. |location=London |chapter=''Panthera leo'' |pages=212–222 |chapter-url=https://archive.org/stream/PocockMammalia1/pocock1#page/n261/mode/2up}}</ref><ref name=USSR>{{Cite book |last1=Heptner |first1=V. G. |last2=Sludskii |first2=A. A. |orig-date=1972 |year=1992 |title=Mlekopitajuščie Sovetskogo Soiuza. Moskva: Vysšaia Škola |trans-title=Mammals of the Soviet Union, Volume II, Part 2 |publisher=Smithsonian Institution and the National Science Foundation |location=Washington DC |chapter=Lion |chapter-url=https://archive.org/stream/mammalsofsov221992gept#page/82/mode/2up |pages=83–95 |isbn=978-90-04-08876-4}}</ref>
The skeletal muscles of the lion make up 58.8% of its body weight and represent the highest percentage of muscles among mammals.<ref>{{Cite journal |last=Davis |first=D. D. |date=1962 |title=Allometric relationships in Lions vs. Domestic Cats |journal=Evolution |volume=16 |issue=4 |pages=505–514 |doi=10.1111/j.1558-5646.1962.tb03240.x |doi-access=free}}</ref><ref>{{Cite book |last=Calder |first=W. A. |chapter=Skeletal muscle |pages=17–21 |chapter-url=https://books.google.com/books?id=-iBS6-2OO3wC&pg=PA19 |title=Size, Function, and Life History |date=1996 |publisher=Courier Corporation |isbn=978-0-486-69191-6 |access-date=21 July 2021 |archive-date=5 March 2024 |archive-url=https://web.archive.org/web/20240305201618/https://books.google.com/books?id=-iBS6-2OO3wC&pg=PA19#v=onepage&q&f=false |url-status=live}}</ref> The lion has a high concentration of glycolytic [[Fast twitch muscle|fast-twitch muscle]] fibers.<ref>{{Cite journal |last1=Kohn |first1=T. A. |last2=Burroughs |first2=R. |last3=Hartman |first3=M. J. |last4=Noakes |first4=T. D. |date=2011 |title=Fiber type and metabolic characteristics of lion (''Panthera leo''), Caracal (''Caracal caracal'') and human skeletal muscle. |url=https://repository.up.ac.za/bitstream/2263/19598/1/Kohn_Fiber%282011%29.pdf |journal=Comparative Biochemistry and Physiology Part A: Molecular
===Size===
Among felids, the lion is second only to the tiger in size.<ref name=Haas2005/> The size and weight of adult lions vary across its range and habitats.<ref name=Smuts_etal_1980>{{Cite journal |last=Smuts|first=G. L. |author2=Robinson, G. A. |author3=Whyte, I. J. |title=Comparative growth of wild male and female lions (''Panthera leo'') |journal=Journal of Zoology |year=1980 |volume=190 |issue=3 |pages=365–373 |doi=10.1111/j.1469-7998.1980.tb01433.x}}</ref><ref name=Chellam1993>{{cite book |author=Chellam, R.
{| class="wikitable"
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===Mane===
{{multiple image|align=right|direction=vertical |image1=Lion (Panthera leo) male 6y.jpg|caption1=A six-year-old male in [[Phinda Private Game Reserve]] |image2=Lion au repos parc pendjari.jpg|caption2=Young male in Pendjari National Park}}
The male lion's mane is the most recognisable feature of the species.<ref name=Hemmer>{{cite journal |author=Hemmer, H. |year=1974 |title=Untersuchungen zur Stammesgeschichte der Pantherkatzen (''Pantherinae'') Teil 3. Zur Artgeschichte des Löwen ''Panthera (Panthera) leo'' (Linnaeus, 1758) |journal=Veröffentlichungen der Zoologischen Staatssammlung |volume=17| pages=167–280 |url=https://archive.org/stream/verfentlichungen171974zool#page/178/mode/2up}}</ref> It may have evolved around 320,000–190,000 years ago.<ref name=Yamaguchietal2004Mane>{{Cite journal |last1=Yamaguchi |first1=
This feature likely evolved to signal the [[Fitness (biology)|fitness]] of males to females. Males with darker manes appear to have greater reproductive success and are more likely to remain in a pride for longer. They have longer and thicker hair and higher testosterone levels, but they are also more vulnerable to heat stress.<ref name=PeytonPacker1992>{{cite journal |last1=Peyton |first1=P. M. |last2=Packer |first2=C. |year=2002 |title=Sexual selection, temperature, and the lion's mane |journal=Science |volume=297 |issue=5585 |pages=1339–1343 |doi=10.1126/science.1073257 |pmid=12193785 |bibcode=2002Sci...297.1339W |s2cid=15893512}}</ref><ref>[[#Packer|Packer]], pp. 137, 145.</ref> The core body temperature does apparently not increase regardless of sex, season, feeding time, length and colour of mane, but only surface temperature is affected.<ref>{{Cite journal |last1=Trethowan |first1=P. |last2=Fuller |first2=A. |last3=Haw |first3=A. |last4=Hart |first4=T. |last5=Markham |first5=A. |last6=Loveridge |first6=A. |last7=Hetem |first7=R. |last8=du Preez |first8=B. |last9=Macdonald |first9=D. W. |date=2017 |title=Getting to the core: Internal body temperatures help reveal the ecological function and thermal implications of the lions' mane |journal=Ecology and Evolution |volume=7 |issue=1 |pages=253–262 |doi=10.1002/ece3.2556 |pmc=5214092 |pmid=28070288 |bibcode=2017EcoEv...7..253T}}</ref> Unlike in other felid species, female lions consistently interact with multiple males at once.<ref>[[#Packer|Packer]], pp. 145.</ref> Another hypothesis suggests that the mane also serves to protect the neck in fights, but this is disputed.<ref>[[#Schaller|Schaller]], pp. 360.</ref><ref>[[#Packer|Packer]], pp. 133.</ref> During fights, including those involving maneless females and adolescents, the neck is not targeted as much as the face, back, and hindquarters. Injured lions also begin to lose their manes.<ref>[[#Packer|Packer]], pp. 133, 137, 145, 148.</ref> ▼
▲The male lion's mane is the most recognisable feature of the species.<ref name=Hemmer>{{cite journal |author=Hemmer, H. |year=1974 |title=Untersuchungen zur Stammesgeschichte der Pantherkatzen (''Pantherinae'') Teil 3. Zur Artgeschichte des Löwen ''Panthera (Panthera) leo'' (Linnaeus, 1758) |journal=Veröffentlichungen der Zoologischen Staatssammlung |volume=17| pages=167–280 |url=https://archive.org/stream/verfentlichungen171974zool#page/178/mode/2up}}</ref> It may have evolved around 320,000–190,000 years ago.<ref name=Yamaguchietal2004Mane>{{Cite journal |last1=Yamaguchi |first1=Nobuyuki |last2=Cooper |first2=A. |last3=Werdelin |first3=L. |last4=MacDonald |first4=David W. |date=2004 |title=Evolution of the mane and group-living in the lion (''Panthera leo''): a review |journal=Journal of Zoology |volume=263 |issue=4 |pages=329–342 |doi=10.1017/S0952836904005242}}</ref> It grows downwards and backwards, covering most of the head, neck, shoulders, and chest. The mane is typically brownish and tinged with yellow, rust, and black hairs.<ref name=Haas2005/> It starts growing when lions enter adolescence, when [[testosterone]] levels increase, and reach their full size at around four years old.<ref>[[#Packer|Packer]], p. 148.</ref> Cool ambient temperatures in European and North American zoos may result in a heavier mane.<ref name=BarnettYamaguchi2006/> On average, Asiatic lions have sparser manes than African lions.<ref name=Menon>{{Cite book |last=Menon |first=V. |year=2003 |title=A Field Guide to Indian Mammals |location=New Delhi |publisher=Dorling Kindersley India |isbn=978-0-14-302998-4}}</ref>
▲This feature likely evolved to signal the [[Fitness (biology)|fitness]] of males to females. Males with darker manes appear to have greater reproductive success and are more likely to remain in a pride for longer. They have longer and thicker hair and higher testosterone levels, but they are also more vulnerable to heat stress.<ref name=PeytonPacker1992>{{cite journal |last1=Peyton |first1=P. M. |last2=Packer |first2=C. |year=2002 |title=Sexual selection, temperature, and the lion's mane |journal=Science |volume=297 |issue=5585 |pages=1339–1343 |doi=10.1126/science.1073257 |pmid=12193785 |bibcode=2002Sci...297.1339W |s2cid=15893512}}</ref><ref>[[#Packer|Packer]], pp. 137, 145.</ref> Unlike in other felid species, female lions consistently interact with multiple males at once.<ref>[[#Packer|Packer]], pp. 145.</ref> Another hypothesis suggests that the mane also serves to protect the neck in fights, but this is disputed.<ref>[[#Schaller|Schaller]], pp. 360.</ref><ref>[[#Packer|Packer]], pp. 133.</ref> During fights, including those involving maneless females and adolescents, the neck is not targeted as much as the face, back, and hindquarters. Injured lions also begin to lose their manes.<ref>[[#Packer|Packer]], pp. 133, 137, 145, 148.</ref>
▲Rarely, lionesses (in the wild and in captivity) have been observed to grow manes.<ref>{{cite magazine |last=Gruber |first=Karl |date=24 October 2022 |access-date=19 November 2023 |title=Five wild lionesses grow a mane and start acting like males |url=https://www.newscientist.com/article/2106866-five-wild-lionesses-grow-a-mane-and-start-acting-like-males/ |magazine=New Scientist |archive-date=11 April 2024 |archive-url=https://web.archive.org/web/20240411033205/https://www.newscientist.com/article/2106866-five-wild-lionesses-grow-a-mane-and-start-acting-like-males/ |url-status=live}}</ref><ref>{{cite magazine |last=Young |first=Lauren J. |date=23 September 2016 |access-date=19 November 2023 |title=The rare case of a lioness with a mane |url=https://www.popsci.com/science/zoo-lioness-mane/ |magazine=Popular Science |archive-date=19 November 2023 |archive-url=https://web.archive.org/web/20231119185541/https://www.popsci.com/science/zoo-lioness-mane/ |url-status=live}}</ref> Increased testosterone may be the cause of maned lionesses reported in northern Botswana.<ref>{{cite magazine |last=Dell'Amore |first=C. |date=2016 |access-date=18 April 2016 |title=No, those aren't male lions mating. One is likely a female |url=https://www.nationalgeographic.com/animals/article/160418-lions-mating-africa-animals-science|archive-url=https://web.archive.org/web/20210228190519/https://www.nationalgeographic.com/animals/article/160418-lions-mating-africa-animals-science|url-status=dead|archive-date=28 February 2021|magazine=National Geographic}}</ref>
===Colour variation===
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===Historical range===
In Africa, the range of the lion originally spanned most of the central [[African rainforest]] zone and the [[Sahara]] desert.<ref name=Schaller5>[[#Schaller|Schaller]], p. 5.</ref> In the 1960s, it became extinct in North Africa, except in the southern part of Sudan.<ref name=Chardonnet2002/><ref name=Riggio_al2013/><ref name=Black_al2013>{{cite journal |last1=Black |first1=S. A. |last2=Fellous |first2=A. |last3=Yamaguchi |first3=N. | last4=Roberts |first4=D. L. |year=2013 |title=Examining the Extinction of the Barbary Lion and Its Implications for Felid Conservation |journal=PLOS ONE |volume=8 |issue=4 |page=e60174 |doi=10.1371/journal.pone.0060174 |pmid=23573239 |pmc=3616087 |bibcode=2013PLoSO...860174B |doi-access=free
During the mid-[[Holocene]], around 8,000-6,000 years ago, the range of lions expanded into Southeastern and Eastern Europe, partially re-occupying the range of the now extinct [[Panthera spelaea|cave lion]].<ref name=Marciszak2022>{{Cite journal |last1=Marciszak |first1=A. |last2=Ivanoff |first2=D. V. |last3=Semenov |first3=Y. A. |last4=Talamo |first4=S. |last5=Ridush |first5=B. |last6=Stupak |first6=A. |last7=Yanish |first7=Y. |last8=Kovalchuk |first8=O. |date=2022 |title=The Quaternary lions of Ukraine and a trend of decreasing size in ''Panthera spelaea'' |journal=[[Journal of Mammalian Evolution]] |volume=30 |issue=1 |pages=109–135 |doi=10.1007/s10914-022-09635-3 |hdl-access=free |hdl=11585/903022
In Asia the lion once ranged in regions where climatic conditions supported an abundance of prey.<ref>{{cite journal |last1=Schnitzler |first1=A. |last2=Hermann |first2=L.
==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.
===Group organisation===
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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 will [[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, consisting 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 has just 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 will 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.
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" being near river [[confluence]]s, where the cats have better access to water, prey and shelter (via vegetation).<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> 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 defense against intruders, while others lag behind.<ref>{{Cite journal |last=Heinsohn|first= R. |author2=Packer, C.
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.
===Hunting and diet===
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|image1=Male Lion and Cub Chitwa South Africa Luca Galuzzi 2004.JPG |caption1=Male lion and cub feeding on an African buffalo in [[Sabi Sand Game Reserve]]}}
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]],
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.
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> 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.
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/>
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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=2011-04-30 |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=2011-05-13 |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=2017-11-20 |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.
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.
===Reproduction and life cycle===
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|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
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>
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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=
===Health and mortality===
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[[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.
[[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>
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{{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.
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
==Conservation==
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Several large and well-managed protected areas in Africa host large lion populations. Where an infrastructure for wildlife tourism has been developed, cash revenue for park management and local communities is a strong incentive for lion conservation.<ref name="IUCN" /> Most lions now live in East and Southern Africa; their numbers are rapidly decreasing, and fell by an estimated 30–50% in the late half of the 20th century. Primary causes of the decline include disease and human interference.<ref name=IUCN/> In 1975, it was estimated that since the 1950s, lion numbers had decreased by half to 200,000 or fewer.<ref>{{cite journal |last1=Myers |first1=N. |year=1975 |title=The silent savannahs |journal=International Wildlife |volume=5 |issue=5 |pages=5–10 }}</ref> Estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004.<ref>{{Cite journal |last1=Bauer |first1=H. |last2=Van Der Merwe |first2=S. |year= 2002 |title=The African lion database |journal=Cat News |volume=36 |pages=41–53}}</ref><ref name=Chardonnet2002>{{Cite book |last=Chardonnet|first=P. |year=2002|title=Conservation of African lion |url=http://conservationforce.org/pdf/conservationoftheafricanlion.pdf |publisher=International Foundation for the Conservation of Wildlife|location=Paris, France |url-status=dead |archive-date=10 November 2013 |archive-url=https://web.archive.org/web/20131110184540/http://conservationforce.org/pdf/conservationoftheafricanlion.pdf}}</ref>
In the [[Republic of the Congo]], [[Odzala-Kokoua National Park]] was considered a lion stronghold in the 1990s. By 2014, no lions were recorded in the protected area so the population is considered locally extinct.<ref name=carn>{{cite journal |last1=Henschel |first1=P. |last2=Malanda |first2=G.-A. |last3=Hunter |first3=L. |title=The status of savanna carnivores in the Odzala-Kokoua National Park, northern Republic of Congo |journal=Journal of Mammalogy |date=2014 |volume=95 |issue=4 |
A population occurs in Cameroon's [[Waza National Park]], where between approximately 14 and 21 animals persisted as of 2009.<ref>{{Cite journal |last1=Tumenta |first1=P. N. |last2=Kok |first2=J. S. |last3=van Rijssel |first3=J. C. |last4=Buij |first4=R. |last5=Croes |first5=B. M. |last6=Funston |first6=P. J. |last7=de Iongh |first7=H. H. |last8=de Haes |first8=H. A. Udo |year=2009 |title=Threat of rapid extermination of the lion (''Panthera leo leo'') in Waza National Park, Northern Cameroon |journal=African Journal of Ecology |doi=10.1111/j.1365-2028.2009.01181.x |pages=1–7 |volume=48 |issue=4|hdl=1887/14372 |s2cid=56451273 |hdl-access=free }}</ref> In addition, 50 to 150 lions are estimated to be present in Burkina Faso's [[Arly-Singou]] ecosystem.<ref name="Bauer & van der Merwe">{{Cite journal |last1=Bauer |first1=H. |last2=Van Der Merwe |first2=S. |doi=10.1017/S0030605304000055 |title=Inventory of free-ranging lions ''Panthera leo'' in Africa |journal=Oryx |volume=38 |year=2004 |issue=1 |pages=26–31|doi-broken-date=
In 2005, Lion Conservation Strategies were developed for West and Central Africa, and or East and Southern Africa. The strategies seek to maintain suitable habitat, ensure a sufficient wild prey base for lions, reduce factors that lead to further fragmentation of populations, and make lion–human coexistence sustainable.<ref>{{cite book |title=Conservation Strategy for the Lion West and Central Africa |publisher=IUCN |author=IUCN Cat Specialist Group |year=2006 |location=Yaounde, Cameroon |url=http://www.catsg.org/fileadmin/filesharing/3.Conservation_Center/3.4._Strategies___Action_Plans/African_lion/IUCN_CatSG_2006_West_and_Central_Africa_Lion_Conservation_Strategy.pdf |archive-url=https://web.archive.org/web/20150919114530/http://www.catsg.org/fileadmin/filesharing/3.Conservation_Center/3.4._Strategies___Action_Plans/African_lion/IUCN_CatSG_2006_West_and_Central_Africa_Lion_Conservation_Strategy.pdf |archive-date=2015-09-19 |url-status=live}}</ref><ref>{{cite book |author=IUCN Cat Specialist Group |year=2006 |title=Conservation Strategy for the Lion ''Panthera leo'' in Eastern and Southern Africa |publisher=IUCN |location=Pretoria, South Africa |url=http://www.catsg.org/fileadmin/filesharing/3.Conservation_Center/3.4._Strategies___Action_Plans/African_lion/IUCN_CatSG_2006_East_and_South_Africa_Lion_Conservation_Strategy.pdf |archive-url=https://web.archive.org/web/20150619001314/http://www.catsg.org/fileadmin/filesharing/3.Conservation_Center/3.4._Strategies___Action_Plans/African_lion/IUCN_CatSG_2006_East_and_South_Africa_Lion_Conservation_Strategy.pdf |archive-date=2015-06-19 |url-status=live}}</ref> Lion depredation on livestock is significantly reduced in areas where herders keep livestock in improved enclosures. Such measures contribute to mitigating [[human–lion conflict]].<ref>{{cite journal |title=Assessment and mitigation of human-lion conflict in West and Central Africa |author1=Bauer, H. |author2=de Iongh, H. |author3=Sogbohossou, E. |journal=Mammalia |year=2010 |volume=74 |issue=4 |pages=363–367 |doi=10.1515/MAMM.2010.048|s2cid=86228533 }}</ref>
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===In Asia===
[[File:Lion Gir.jpg|thumb|A lioness in Gir National Park]]
The last refuge of the Asiatic lion population is the {{cvt|1412|km2}} Gir National Park and surrounding areas in the [[Saurashtra (region)|region of Saurashtra]] or [[Kathiawar Peninsula]] in Gujarat State, India. The population has risen from approximately 180 lions in 1974 to about 400 in 2010.<ref name="Singh, Gibson">{{Cite journal |last1=Singh | first1=H. S. |last2=Gibson |first2=L. |title=A conservation success story in the otherwise dire megafauna extinction crisis: The Asiatic lion (''Panthera leo persica'') of Gir forest |journal=Biological Conservation |volume=144 |issue=5 |pages=1753–1757 |year=2011 |doi=10.1016/j.biocon.2011.02.009| bibcode=2011BCons.144.1753S}}</ref> It is geographically isolated, which can lead to [[inbreeding]] and reduced [[genetic diversity]]. Since 2008, the Asiatic lion has been listed as [[Endangered]] on the [[IUCN Red List]].<ref name=Breitenmoser2008/> By 2015, the population had grown to 523 individuals inhabiting an area of {{cvt|7000|km2}} in Saurashtra.<ref name=Venkataraman2016>{{cite book |author=Venkataraman, M. |year=2016 |chapter=Wildlife and human impacts in the Gir landscape |title=Human Animal Conflict in Agro-Pastoral Context: Issues & Policies |editor1=Agrawal, P. K. |editor2=Verghese, A. |editor3=Krishna, S. R. |editor4=Subaharan, K. |publisher=Indian Council of Agricultural Research |location=New Delhi |
The presence of numerous human settlements close to Gir National Park resulted in conflict between lions, local people and their livestock.<ref>{{cite journal |first1=V. K. |last1=Saberwal |first2=J. P. |last2=Gibbs |first3=R. |last3=Chellam |first4=A. J. T. |last4=Johnsingh |title=Lion-Human Conflict in the Gir Forest, India |date=1994 |doi=10.1046/j.1523-1739.1994.08020501.x |journal=Conservation Biology |issue=2 |volume=8 |pages=501–507|bibcode=1994ConBi...8..501S}}</ref><ref name=Venkataraman2016 /> Some consider the presence of lions a benefit, as they keep populations of crop damaging herbivores in check.<ref name=Meena2016>{{cite book |author=Meena, V. |year=2016 |chapter=Wildlife and human impacts in the Gir landscape |editor=Agrawal, P. K. |editor2=Verghese, A. |editor3=Radhakrishna, S. |editor4=Subaharan, K. |title=Human Animal Conflict in Agro-Pastoral Context: Issues & Policies |publisher=Indian Council of Agricultural Research |location=New Delhi}}</ref>
===Captive breeding===
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|image1=Lion - melbourne zoo.jpg|caption1=Lion at [[Melbourne Zoo]]
|image2=Lion tamer (LOC pga.03749).jpg|caption2=19th-century [[etching]] of a lion tamer in a cage with lions and tigers}}
Lions are part of a group of exotic animals that have been central to zoo exhibits since the late 18th century. Although many modern zoos are more selective about their exhibits,<ref name="dc81">[[#Courcy|de Courcy]],
The first European "zoos" spread among noble and royal families in the 13th century, and until the 17th century were called [[seraglio]]s. At that time, they came to be called [[menagerie]]s, an extension of the [[cabinet of curiosities]]. They spread from France and Italy during the [[Renaissance]] to the rest of Europe.<ref>[[#Baratay|Baratay & Hardouin-Fugier]], pp. 19–21, 42.</ref> In England, although the seraglio tradition was less developed, lions were [[Tower of London#Menagerie|kept at the Tower of London]] in a seraglio established by [[John of England|King John]] in the 13th century;<ref>[[#Baratay|Baratay & Hardouin-Fugier]], p. 20.</ref><ref>{{cite magazine |last=Owen |first=J. |title=Medieval Lion Skulls Reveal Secrets of Tower of London "Zoo"|magazine=National Geographic Magazine |date=2005 |url=http://news.nationalgeographic.com/news/2005/11/1103_051103_tower_lions.html|archive-url=https://web.archive.org/web/20051105025912/http://news.nationalgeographic.com/news/2005/11/1103_051103_tower_lions.html|url-status=dead|archive-date=5 November 2005| access-date=5 September 2007}}</ref> this was probably stocked with animals from an earlier menagerie started in 1125 by [[Henry I of England|Henry I]] at his hunting lodge in [[Woodstock, Oxfordshire]], where according to [[William of Malmesbury]] lions had been stocked.<ref name=Blunt15>[[#Blunt|Blunt]], p. 15.</ref>
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{{See also|Lion baiting}}
[[File:Sculpted reliefs depicting Ashurbanipal, the last great Assyrian king, hunting lions, gypsum hall relief from the North Palace of Nineveh (Irak), c. 645-635 BC, British Museum (16722183731).jpg|thumb|[[Bas-relief]] of a wounded lioness from [[Nineveh]], {{circa|645–635 BC}}]]
Lion hunting has occurred since ancient times and was often a royal tradition, intended to demonstrate the power of the king over nature. Such hunts took place in a reserved area in front of an audience. The monarch was accompanied by his men and controls were put in place to increase their safety and ease of killing. The earliest surviving record of lion hunting is an [[ancient Egypt]]ian inscription dated circa 1380 BC that mentions Pharaoh [[Amenhotep III]] killing 102 lions in ten years "with his own arrows". The [[Assyria]]n emperor [[Ashurbanipal]] had one of his lion hunts depicted on a sequence of [[Assyrian palace reliefs]] {{circa|640 BC}}, known as the [[Lion Hunt of Ashurbanipal]]. Lions were also hunted during the [[Mughal Empire]], where Emperor [[Jahangir]] is said to have excelled at it.<ref name=Jackson156>[[#Jackson|Jackson]],
The [[Maasai people]] have traditionally viewed the killing of lions as a rite of passage. Historically, lions were hunted by individuals, however, due to reduced lion populations, elders discourage solo lion hunts.<ref>{{cite journal |last1=Hazzah |first1=L. |last2=Borgerhoof Mulder |first2=M. |last3=Frank |first3=L. |year=2009 |title=Lions and warriors: Social factors underlying declining African lion populations and the effect of incentive-based management in Kenya |journal=Biological Conservation |volume=142 |issue= 11|pages=2428–2437 |doi=10.1016/j.biocon.2009.06.006|bibcode=2009BCons.142.2428H }}</ref> During the [[Scramble for Africa|European colonisation of Africa]] in the 19th century, the hunting of lions was encouraged because they were considered pests and lion skins were sold for [[Pound sterling|£1]] each.<ref>[[#Jackson|Jackson]], p. 166.</ref> The widely reproduced imagery of the heroic hunter chasing lions would dominate a large part of the century.<ref>[[#Baratay|Baratay & Hardouin-Fugier]], p. 113.</ref> [[Trophy hunting]] of lions in recent years has been met with controversy, notably with the [[killing of Cecil the lion]] in mid-2015.<ref>{{cite news |last1=Capecchi |first1=Christina |last2=Rogers |first2=Katie |title=Killer of Cecil the lion finds out that he is a target now, of internet vigilantism |work=[[The New York Times]] |date=30 July 2015 |url=https://www.nytimes.com/2015/07/30/us/cecil-the-lion-walter-palmer.html |access-date=30 July 2015 |archive-date=29 July 2015 |archive-url=https://web.archive.org/web/20150729215743/http://www.nytimes.com/2015/07/30/us/cecil-the-lion-walter-palmer.html |url-status=live }}</ref>
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===Man-eating===
[[File:Lionsoftsavo2008.jpg|thumb|The Tsavo maneaters of East Africa on display in the [[Field Museum of Natural History]] in Chicago]]
{{Further information|Man-
Lions do not usually hunt humans but some (usually males) seem to seek them out. One well-publicised case is the [[Tsavo maneaters]]; in 1898, 28 officially recorded railway workers building the [[Uganda Railway]] were taken by lions over nine months during the construction of a bridge in Kenya.<ref name=Patterson>{{cite book |last=Patterson |first=B. D. |year=2004 |title=The Lions of Tsavo: Exploring the Legacy of Africa's Notorious Man-Eaters |publisher=McGraw Hill Professional |location=New York |isbn=978-0-07-136333-4}}</ref> The hunter who killed the lions wrote a book detailing the animals' predatory behaviour; they were larger than normal and lacked manes, and one seemed to suffer from tooth decay. The infirmity theory, including tooth decay, is not favoured by all researchers; an analysis of teeth and jaws of man-eating lions in museum collections suggests that while tooth decay may explain some incidents, prey depletion in human-dominated areas is a more likely cause of lion predation on humans.<ref>{{Cite journal |last=Patterson |first=B. D. |author2=Neiburger, E. J.|author3=Kasiki, S. M. |date=2003 |title=Tooth Breakage and Dental Disease as Causes of Carnivore–Human Conflicts |journal=Journal of Mammalogy |volume=84 |issue=1 |pages=190–196 |doi=10.1644/1545-1542(2003)084<0190:TBADDA>2.0.CO;2 |doi-access=free}}</ref> Sick or injured animals may be more prone to man-eating but the behaviour is not unusual, nor necessarily aberrant.<ref>{{Cite journal |last1=Peterhans |first1=J. C. K. |first2=T. P. | last2=Gnoske |title=The Science of Man-eating |journal=Journal of East African Natural History |volume=90 |issue=1&2 |year=2001 |pages=1–40 |doi=10.2982/0012-8317(2001)90[1:TSOMAL]2.0.CO;2 |doi-access=free}}</ref>
Lions' proclivity for [[Man-eating animal#Lions|man-eating]] has been systematically examined. American and Tanzanian scientists report that man-eating behaviour in rural areas of Tanzania increased greatly from 1990 to 2005. At least 563 villagers were attacked and many eaten over this period. The incidents occurred near [[Selous Game Reserve]] in [[Rufiji River]] and in [[Lindi Region]] near the Mozambican border. While the expansion of villages into bush country is one concern, the authors argue conservation policy must mitigate the danger because in this case, conservation contributes directly to human deaths. Cases in Lindi in which lions seize humans from the centres of substantial villages have been documented.<ref name=Packer05>{{Cite journal |last1=Packer |first1=C. |last2=Ikanda | first2= D. |last3=Kissui| first3= B. |last4=Kushnir| first4= H. |date=2005 |title=Conservation biology: lion attacks on humans in Tanzania |journal=[[Nature (journal)|Nature]] |volume=436 |issue=7053 |pages=
According to Robert R. Frump, Mozambican refugees regularly crossing Kruger National Park, South Africa, at night are attacked and eaten by lions; park officials have said man-eating is a problem there. Frump said thousands may have been killed in the decades after [[apartheid]] sealed the park and forced refugees to cross the park at night. For nearly a century before the border was sealed, Mozambicans had regularly crossed the park in daytime with little harm.<ref>{{Cite book |last=Frump |first=R. R. |title=The Man-Eaters of Eden: Life and Death in Kruger National Park |year=2006 |publisher=The Lyons Press |isbn=978-1-59228-892-2}}</ref>
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{{main|Cultural depictions of lions}}
[[File:Luxor Sekhmet New Kingdom.JPG|thumb|right|upright|Granite statue of the Egyptian goddess [[Sekhmet]] from the [[Luxor Temple]], dated 1403–1365 BC, exhibited in the [[National Museum of Denmark]]]]
The lion is one of the most widely recognised animal symbols in human culture. It has been extensively depicted in sculptures and paintings, on national flags, and in contemporary films and literature.<ref name=Guggisberg1975/> It is considered to be the monarch of the beasts; symbolizing power, royalty and protection.<ref name=Symbolism>{{cite book | first=Hope B. | last=Werness |year=2007 |title=The Continuum Encyclopedia of Animal Symbolism in World Art |publisher=Continuum International Publishing Group |pages=
In sub-Saharan Africa, the lion has been a common character in stories, proverbs and dances, but rarely featured in visual arts.<ref>[[#Jackson|Jackson]], p. 119.</ref> In the [[Swahili language]], the lion is known as ''simba'' which also means "aggressive", "king" and "strong".<ref name=BCKM1993/> In parts of West and East Africa, the lion is associated with healing and provides the connection between [[Clairvoyance|seers]] and the supernatural. In other East African traditions, the lion represents laziness.<ref name=Hogarth>{{cite book |last1=Hogarth |first1=C. |last2=Butler |first2=N. |year=2004 |title=Shamanism: An Encyclopedia of World Beliefs, Practices, and Culture |volume=1 |editor=Walter, M. N. |isbn=978-1-57607-645-3 |chapter=Animal Symbolism (Africa) |pages=3–6 |publisher=Bloomsbury Academic |chapter-url=https://books.google.com/books?id=X8waCmzjiD4C&pg=PA3 |url=https://archive.org/details/shamanism00mari/page/3}}</ref> In much of [[African folklore]], the lion is portrayed as having low intelligence and is easily tricked by other animals.<ref name=Lynch>{{cite book |last=Lynch |first=P. A. |year=2004 |title=African Mythology A to Z |publisher=Infobase Publishing |page=63 |isbn=978-0-8160-4892-2 |chapter=Lion |chapter-url=https://archive.org/details/africanmythology00lync_0/page/63}}</ref> In [[Nubia]], The lion-god [[Apedemak]] was associated with the flooding of the Nile. In [[Ancient Egypt]], lions were linked both with the sun and the waters of the Nile. Several gods were conceived as being partially lion including the war deities [[Sekhmet]] and [[Maahes]], and [[Tefnut]] the goddess of moisture. [[Shu (Egyptian god)|Shu]], god of the air, was depicted kneeing between two lions while holding a headrest. The lions mark where the sun rises and sets and symbolise yesterday and tomorrow.<ref>[[#Jackson|Jackson]], pp. 107–108, 111.</ref>
[[File:Berlín Ishtar león. 02.JPG|thumb|left|Roaring and striding lion from the Throne Room of [[Nebuchadnezzar II]], 6th century BC, from [[Babylon]], Iraq]]
The lion was a prominent symbol in ancient [[Mesopotamia]] from [[Sumer]] up to [[Assyria]]n and [[Babylonia]]n times, where it was strongly associated with kingship.<ref>{{Cite journal |last=Cassin |first=Elena |author-link=Elena Cassin |year=1981 |title= Le roi et le lion |journal=Revue de l'Histoire des Religions | trans-title = The
[[File:Cowardly lion2.jpg|thumb|upright|[[Dorothy Gale]] meets the Cowardly Lion in ''[[The Wonderful Wizard of Oz]]''. Art by [[W. W. Denslow]], 1900.]]
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=== Books ===
{{Refbegin}}
* {{Cite book |last1=Baratay |first1=E. |first2=E. |last2=Hardouin-Fugier
* {{Cite book |last=Blunt |first=W. |year=1975 |title=The Ark in the Park: The Zoo in the Nineteenth Century |url=https://archive.org/details/isbn_241893313 |url-access=registration |location=London |publisher=Hamish Hamilton |isbn=978-0-241-89331-9 |ref=Blunt }}
* {{Cite book |last=de Courcy |first=C. |year=1995 |title=The Zoo Story |location=Ringwood, Victoria |publisher=Penguin Books |isbn=978-0-14-023919-5|ref=Courcy}}
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* {{cite web |url=http://www.catsg.org/index.php?id=108 | title=Lion ''Panthera leo'' |author=IUCN/SSC Cat Specialist Group}}
* {{cite web |url=http://www.lionconservationfund.org/ | title=Lion Conservation Fund}}
* {{cite web |url=http://www.theportugalnews.com/news/rare-desert-lion-killed-in-angola-after-supplying-unprecedented-data/32633 |title=Rare desert lion killed in Angola after supplying unprecedented data |
* {{Cite Collier's|wstitle=Lion|short=x}}
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