View source for Lion

{{Short description|Large cat native to Africa and India}}
{{Other uses}}
{{Distinguish|Mountain lion|Lyon|Loin}}
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{{Use British English|date=December 2024}}
{{Use dmy dates|date=December 2024}}
{{Speciesbox
| 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
| image2_caption = Female in [[Okonjima]], Namibia
| status = VU
| status_system = IUCN3.1
| status_ref = <ref name=IUCN>{{cite iucn |title=''Panthera leo'' |author1=Nicholson, S. |author2=Bauer, H. |author3=Strampelli, P. |author4=Sogbohossou, E. |author5=Ikanda, D. |author6=Tumenta, P. F. |author7=Venktraman, M. |author8=Chapron, G. |author9=Loveridge, A. |year=2024 |amends=2023 |article-number=e.T15951A259030422 |access-date=30 June 2024}}</ref>
| status2 = CITES_A2
| status2_system = CITES
| status2_ref = {{efn|Populations of India are listed in Appendix I.}}<ref name=IUCN/>
| taxon = Panthera leo<ref name=MSW3>{{MSW3 Carnivora |id=14000228 |page=546 |heading=Species ''Panthera leo''}}</ref>
| authority = ([[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]])<ref name="Linn1758" />
| subdivision_ranks = Subspecies
| subdivision = :''[[Panthera leo leo|P. l. leo]]''
:''[[Panthera leo melanochaita|P. l. melanochaita]]''
| range_map = Lion (Panthera leo) IUCN range 2023.svg
| range_map_caption = Historical lion distribution
}}

The '''lion''' ('''''Panthera leo''''') is a large [[Felidae|cat]] of the genus ''[[Panthera]]'', currently ranging only in [[Sub-Saharan Africa]] and [[India]]. It has a muscular, broad-chested [[body (biology)|body]]; a short, rounded head; round ears; and a dark, hairy tuft at the tip of its tail. It is [[sexually dimorphic]]; adult male lions are larger than females and have a more prominent mane that usually obscures the ears and extends to the shoulders.

The lion inhabits [[grassland]]s, [[savannah]]s, and [[shrubland]]s. It is an [[apex predator|apex]] and [[keystone predator]], preying mostly on medium-sized and large [[ungulate]]s. It is usually more [[diurnality|diurnal]] than other wild cats, but when persecuted, it adapts to being active [[nocturnality|at night]] and [[crepuscular|at twilight]]. It is a [[social species]], forming groups called prides. A lion pride consists of related females and cubs, and a few or one adult male who is unrelated to the females. Groups of female lions usually hunt together. Adult males often compete to keep or gain that membership in the pride.

During the [[Neolithic]] period, the lion ranged throughout Africa and Eurasia, from Southeast Europe to India, but it has been reduced to fragmented populations in sub-Saharan Africa and one population in [[western India]]. It has been listed as [[Vulnerable species|Vulnerable]] on the [[IUCN Red List]] since 1996 because populations in African countries have declined by about 43% since the early 1990s. Lion populations are untenable outside designated protected areas. Although the cause of the decline is not fully understood, [[habitat loss]] and conflicts with humans are the greatest causes for concern.

The lion has been extensively depicted in sculptures and paintings, on national flags, and in literature and films. It is one of the most widely recognised animal symbols in human culture. Lions have been kept in [[menagerie]]s since the time of the [[Roman Empire]] and have been a key species sought for exhibition in [[zoological garden]]s across the world since the late 18th century. [[Cultural depictions of lions]] have occurred worldwide, particularly as a symbol of power and royalty. They figure prominently as symbols and deities in ancient religions.

==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.  |year=1940 |chapter=πάνθηρ |chapter-url=https://www.perseus.tufts.edu/cgi-bin/ptext?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3D%2377441 |title=A Greek-English Lexicon |edition=Revised and augmented |location=Oxford |publisher=Clarendon Press |access-date=20 February 2021 |archive-date=11 April 2008 |archive-url=https://web.archive.org/web/20080411203109/http://www.perseus.tufts.edu/cgi-bin/ptext?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3D%2377441 |url-status=live }}</ref>

==Taxonomy==
{{Cladogram|align=right|caption=Phylogeny of the genus ''Panthera'' based on a 2016 [[nuclear DNA]] study<ref name=Li_al2016/>
|1={{clade
|label1=''[[Panthera]]''
|1={{clade
|1={{clade
|1=[[Snow leopard]] [[File:Schneeleoparden (Panthera uncia) Zoo Salzburg 2014 f white background.jpg|70px]]
|2=[[Tiger]] [[File:Tigress in Bandhavgarh NP white background.jpg|70px]]
}}
|2={{clade sequential
|1=[[Jaguar]] [[File:Jaguar (Panthera onca palustris) male Three Brothers River white background.JPG|70px]]
|2=[[Leopard]] [[File:Flickr - Rainbirder - Sassy Lassy white background.jpg|70px]]
|3='''Lion''' [[File:Panthera leo -Ngorongoro, Tanzania-8b white background.jpg|70px]]
}}
}}
}}
}}

''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/>

===Subspecies===
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.  |date=2008 |article-number=e.T15952A5327221 |title=''Panthera leo'' ssp. ''persica'' |doi=10.2305/IUCN.UK.2008.RLTS.T15952A5327221.en}}</ref><ref name=Henschel2015>{{cite iucn |author=Henschel, P. |author2=Bauer, H. |author3=Sogbohoussou, E. |author4=Nowell, K.  |date=2015 |doi=10.2305/IUCN.UK.2015-2.RLTS.T68933833A54067639.en |title=''Panthera leo'' West Africa subpopulation}}</ref> In 2017, the Cat Classification Task Force of the Cat Specialist Group revised lion [[Taxonomy (biology)|taxonomy]], and recognises two subspecies based on results of several [[phylogeographic]] studies on lion [[evolution]], namely:<ref name=catsg>{{cite journal |author1=Kitchener, A. C. |author2=Breitenmoser-Würsten, C. |author3=Eizirik, E. |author4=Gentry, A. |author5=Werdelin, L. |author6=Wilting, A. |author7=Yamaguchi, N. |author8=Abramov, A. V. |author9=Christiansen, P. |author10=Driscoll, C. |author11=Duckworth, J. W. |author12=Johnson, W. |author13=Luo, S.-J. |author14=Meijaard, E. |author15=O'Donoghue, P. |author16=Sanderson, J. |author17=Seymour, K. |author18=Bruford, M. |author19=Groves, C. |author20=Hoffmann, M. |author21=Nowell, K. |author22=Timmons, Z. |author23=Tobe, S.  |year=2017 |title=A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group |journal=Cat News |volume=Special Issue 11 |pages=71–73 |url=https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y#page=71 |access-date=6 August 2019 |archive-date=17 January 2020 |archive-url=https://web.archive.org/web/20200117172708/https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y#page=71 |url-status=live}}</ref>

*[[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/> It has been called '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 |volume=((Mammalia, Volume 1)) |publisher=[[Routledge]] |chapter=Felidæ; or the Cat Tribe |pages=129−148 |location=London |chapter-url=https://books.google.com/books?id=v1DPAAAAMAAJ&pg=PA147 |year=1865 |access-date=23 December 2018 |archive-date=5 May 2024 |archive-url=https://web.archive.org/web/20240505150653/https://books.google.com/books?id=v1DPAAAAMAAJ&pg=PA147#v=onepage&q&f=false |url-status=live}}</ref><ref name=Hunter2018>{{cite book |last1=Hunter |first1=L. |last2=Barrett |first2=P.  |title=The Field Guide to Carnivores of the World |edition=Second |publisher=Bloomsbury |location=London, Oxford, New York, New Delhi, Sydney |isbn=978-1-4729-5080-2 |date=2018 |chapter=Lion ''Panthera leo'' |chapter-url=https://books.google.com/books?id=4HpxDwAAQBAJ&pg=PA46 |pages=46−47 |access-date=1 December 2018 |archive-date=5 March 2024 |archive-url=https://web.archive.org/web/20240305201610/https://books.google.com/books?id=4HpxDwAAQBAJ&pg=PA46 |url-status=live}}</ref>
*[[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. |last2=Barnett |first2=R.|last3=Sandoval-Velasco |first3=M. |last4=Yamaguchi |first4=N. |last5=Vieira |first5=F. G. |last6=Mendoza |first6=M. L. Z. |last7=Liu |first7=S. |last8=Martin |first8=M. D. |last9=Sinding |first9=M-S. S. |last10=Mak |first10=S. S. T. |last11=Carøe |first11=C. |last12=Liu |first12=S. |last13=Guo |first13=C. |last14=Zheng |first14=J. |last15=Zazula |first15=G. |last16=Baryshnikov |first16=G. |last17=Eizirik |first17=E. |last18=Koepfli |first18=K.-P. |last19=Johnson |first19=W. E. |last20=Antunes |first20=A. |last21=Sicheritz-Ponten |first21=T. |last22=Gopalakrishnan |first22=S. |last23=Larson |first23=G. |last24=Yang |first24=H. |last25=O'Brien |first25=S. J. |last26=Hansen |first26=A. J. |last27=Zhang |first27=G. |last28=Marques-Bonet |first28=T. |last29=Gilbert |first29=M. T. P. |date=2020 |title=The evolutionary history of extinct and living lions |journal= Proceedings of the National Academy of Sciences |volume=117 |issue=20 |pages=10927–10934 |doi=10.1073/pnas.1919423117 |pmid= 32366643 |pmc=7245068 |bibcode=2020PNAS..11710927D |doi-access=free}}</ref>

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 |article-number=30807 |doi=10.1038/srep30807 |pmid=27488946 |pmc=4973251  |bibcode=2016NatSR...630807B}}</ref> [[Genome]]-wide data of a wild-born historical lion sample from Sudan showed that it clustered with ''P. l. leo'' in mtDNA-based phylogenies, but with a high affinity to ''P. l. melanochaita''; this suggested that the taxonomic position of lions in Central Africa may require revision.<ref name=DeManuel_al2020>{{cite journal |author1=de Manuel, M. |author2=Ross, B. |author3=Sandoval-Velasco, M. |author4=Yamaguchi, N. |author5=Vieira, F. G. |author6=Mendoza, M. L. Z. |author7=Liu, S. |author8=Martin, M. D. |author9=Sinding, M.-H. S. |author10=Mak, S. S. T. |author11=Carøe, C. |author12=Liu, S. |author13=Guo, C. |author14=Zheng, J. |author15=Zazula, G. |author16=Baryshnikov, G. |author17=Eizirik, E. |author18=Koepfli, K.-P. |author19=Johnson, W. E. |author20=Antunes, A. |author21=Sicheritz-Ponten, T.  |author22=Gopalakrishnan, S. |author23=Larson, G. |author24=Yang, H. |author25=O'Brien, S. J. |author26=Hansen, A. J. |author27=Zhang, G. |author28=Marques-Bonet, T. |author29=Gilbert, M. T. P. |title=The evolutionary history of extinct and living lions |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=117 |issue=20 |pages=10927–10934 |doi=10.1073/pnas.1919423117 |year=2020 |pmid=32366643 |pmc=7245068 |bibcode=2020PNAS..11710927D |doi-access=free}}</ref>

===Fossil records===
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 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.  |year=2010 |title=Two forms of cave lion: Middle Pleistocene ''Panthera spelaea fossilis'' Reichenau, 1906 and Upper Pleistocene ''Panthera spelaea spelaea'' Goldfuss, 1810 from the Bisnik Cave, Poland |journal=Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen |volume=258 |issue=3 |pages=339–351 |doi=10.1127/0077-7749/2010/0117 |bibcode=2010NJGPA.258..339M |url=https://www.researchgate.net/publication/233669138 |access-date=14 March 2019 |archive-date=25 September 2018 |archive-url=https://web.archive.org/web/20180925142142/https://www.researchgate.net/publication/233669138 |url-status=live }}</ref><ref name=Sabol2014>{{cite journal |last=Sabol |first=M. |year=2014 |title=''Panthera fossilis'' (Reichenau, 1906) (Felidae, Carnivora) from Za Hájovnou Cave (Moravia, The Czech Republic): A Fossil Record from 1987–2007 |journal=Acta Musei Nationalis Pragae, Series B, Historia Naturalis |volume=70 |issue=1–2 |pages=59–70 | doi=10.14446/AMNP.2014.59 |doi-access=free }}</ref>
*''[[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.  |year=2011 |title=Extinction chronology of the cave lion ''Panthera spelaea'' |journal=Quaternary Science Reviews |volume=30 |issue=17 |pages=2329–2340 |doi=10.1016/j.quascirev.2010.04.023 |bibcode=2011QSRv...30.2329S}}</ref> Bone fragments excavated in European, North Asian, Canadian and Alaskan caves indicate that it ranged from Europe across Siberia into western Alaska.<ref name="Hemmer2011">{{cite journal |author=Hemmer, H. |year=2011 |title=The story of the cave lion – ''Panthera Leo Spelaea'' (Goldfuss, 1810) – A review |journal=Quaternaire |volume=4|pages=201–208|url=https://www.researchgate.net/publication/285886884}}</ref> It likely derived from ''P. fossilis'',<ref name=Barnett2016>{{cite journal |last1=Barnett |first1=R. |last2=Mendoza |first2=M. L. Z. |last3=Soares |first3=A. E. R. |last4=Ho |first4=S. Y. W. |last5=Zazula |first5=G. |last6=Yamaguchi |first6=N. |last7=Shapiro |first7=B. |last8=Kirillova |first8=I. V. |last9=Larson |first9=G. |last10=Gilbert |first10=M. T. P.  |year=2016 |title=Mitogenomics of the Extinct Cave Lion, ''Panthera spelaea'' (Goldfuss, 1810), resolve its position within the ''Panthera'' cats |journal=Open Quaternary |volume=2 |article-number=4 |doi=10.5334/oq.24 |url=https://ora.ox.ac.uk/objects/uuid:9d4f84e6-64c6-49fd-a1dc-a981ba7e8028/download_file?file_format=pdf&safe_filename=Larson%2Bet%2Bal%252C%2BMitogenomics%2Bof%2Bthe%2BExtinct%2BCave%2BLion%252C%2BPanthera%2Bspelaea%2B%2528Goldfuss%252C%2B1810%2529%252C%2Bresolve%2Bits%2Bposition%2Bwit.pdf&type_of_work=Journal+article |doi-access=free |hdl=10576/22920 |hdl-access=free |access-date=18 May 2019 |archive-date=3 October 2020 |archive-url=https://web.archive.org/web/20201003085724/https://ora.ox.ac.uk/objects/uuid:9d4f84e6-64c6-49fd-a1dc-a981ba7e8028/download_file?file_format=pdf&safe_filename=Larson%2Bet%2Bal%252C%2BMitogenomics%2Bof%2Bthe%2BExtinct%2BCave%2BLion%252C%2BPanthera%2Bspelaea%2B%2528Goldfuss%252C%2B1810%2529%252C%2Bresolve%2Bits%2Bposition%2Bwit.pdf&type_of_work=Journal+article |url-status=live }}</ref> and was genetically isolated and highly distinct from the modern lion in Africa and Eurasia.<ref name="BurgerJ-Molecular-phylogeny">{{Cite journal |last1=Burger |first1=J. |year=2004 |title=Molecular phylogeny of the extinct cave lion ''Panthera leo spelaea'' |journal=[[Molecular Phylogenetics and Evolution]] |pmid=15012963 |volume=30 |issue=3 |pages=841–849 |doi=10.1016/j.ympev.2003.07.020 |url=http://www.uni-mainz.de/FB/Biologie/Anthropologie/MolA/Download/Burger%202004.pdf |last3=Loreille |first3=O. |last4=Hemmer |first4=H. |last5=Eriksson |first5=T. |last6=Götherström |first6=A. |last7=Hiller |first7=J. |last8=Collins |first8=M. J. |last9=Wess |first9=T. |last2=Rosendahl |first2=W. |last10=Alt |first10=K. W. |bibcode=2004MolPE..30..841B  |archive-url=https://web.archive.org/web/20070925204424/http://www.uni-mainz.de/FB/Biologie/Anthropologie/MolA/Download/Burger%202004.pdf |archive-date=25 September 2007 }}</ref><ref name=Barnett2016/> It is depicted in [[Paleolithic]] cave paintings, ivory carvings, and clay busts.<ref name="Packer00">{{Cite journal |author=Packer, C. |author2=Clottes, J.  |title=When Lions Ruled France |journal=Natural History |volume=109 |issue=9 |pages=52–57 |date=2000 |url=http://www.cbs.umn.edu/sites/cbs.umn.edu/files/public/downloads/When_lions_ruled_France.pdf |archive-url=https://web.archive.org/web/20151229212607/http://cbs.umn.edu/sites/cbs.umn.edu/files/public/downloads/When_lions_ruled_France.pdf |archive-date=29 December 2015 |url-status=live}}</ref>
*[[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.  |year=2017 |title=The fossil American lion (''Panthera atrox'') in South America: Palaeobiogeographical implications |journal=Comptes Rendus Palevol |volume=16 |issue=8 |pages=850–864 |doi=10.1016/j.crpv.2017.06.009 |bibcode=2017CRPal..16..850C |url=https://www.researchgate.net/publication/321056731|doi-access=free |hdl=11336/65990 |hdl-access=free }}</ref> It diverged from the cave lion around 165,000 years ago.<ref>{{Cite journal |last1=Salis |first1=Alexander T. |last2=Bray |first2=Sarah C. E. |last3=Lee |first3=Michael S. Y. |last4=Heiniger |first4=Holly |last5=Barnett |first5=Ross |last6=Burns |first6=James A. |last7=Doronichev |first7=Vladimir |last8=Fedje |first8=Daryl |last9=Golovanova |first9=Liubov |last10=Harington |first10=C. Richard |last11=Hockett |first11=Bryan |last12=Kosintsev |first12=Pavel |last13=Lai |first13=Xulong |last14=Mackie |first14=Quentin |last15=Vasiliev |first15=Sergei |date=December 2022 |title=Lions and brown bears colonized North America in multiple synchronous waves of dispersal across the Bering Land Bridge |url=https://onlinelibrary.wiley.com/doi/10.1111/mec.16267 |journal=Molecular Ecology |language=en |volume=31 |issue=24 |pages=6407–6421 |bibcode=2022MolEc..31.6407S |doi=10.1111/mec.16267 |issn=0962-1083 |pmid=34748674 |hdl-access=free |hdl=11343/299180}}</ref>

Additionally, in 1938 paleontologist [[Paulus Edward Pieris Deraniyagala|Paulus Deraniyagala]] named a subspecies ''P. l. sinhaleyus'' based on two fossils, a lower left carnassial and a damaged right lower canine tooth, both excavated from deposits in [[Kuruwita]], Sri Lanka. He called it "narrower and more elongate" but otherwise failed to distinguish ''P. l. sinhaleyus'' from other lion subspecies. A subsequent study of felid fossils from the Kuruwita deposits in 2005 described the fossils in further detail but only assigned to them to ''P. leo''.<ref>{{Cite journal|last1=Deraniyagala |first1=P. E. P. |date=1938 |title=Some fossil animals from Ceylon, Part II |journal=Journal of the Royal Asiatic Society (Ceylon Branch) |volume=34 |issue=91 |pages=231–239 |jstor=45385414 }}</ref><ref>{{cite journal |first1=Kelum |last1=Manamendra-Arachchi |first2=Rohan |last2=Pethiyagoda |first3=Rajith |last3=Dissanayake |first4=Madhava |last4=Meegaskumbura |year=2005 |title=A second extinct big cat from the late Quaternary of Sri Lanka |journal=The Raffles Bulletin of Zoology |volume=Supplement 12 |pages=423–434 |url=https://www.researchgate.net/publication/237009731 }}</ref>

===Evolution===
{{multiple image|align=right|direction=vertical|image1=Panthera Atrox.jpg |thumb|caption1=Skull of an American lion on display at the [[National Museum of Natural History]]|
image2=Cave lion range.png|caption2={{legend|#fe6033|''[[Panthera spelaea]]''}}{{legend|#6363ff|''[[Panthera atrox]]''}}{{legend|#6cff8a|''Panthera leo''}}{{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|10.8}}.<ref name=Johnson2006>{{cite journal |last1=Johnson |first1=W. E. |date=2006 |title=The late miocene radiation of modern Felidae: A genetic assessment |journal=[[Science (journal)|Science]]|volume=311|issue=5757|pages=73–77 |doi=10.1126/science.1122277 |pmid=16400146 |bibcode=2006Sci...311...73J }}</ref> [[Hybrid (biology)|Hybridisation]] between lion and [[snow leopard]] ancestors possibly continued until about 2.1 million years ago.<ref name=Li_al2016>{{cite journal|last1=Li |first1=G. |last2=Davis |first2=B. W. |last3=Eizirik |first3=E. |last4=Murphy |first4=W. J. |date=2016 |title=Phylogenomic evidence for ancient hybridization in the genomes of living cats (Felidae) |journal=Genome Research |volume=26 |issue=1|pages=1–11|doi=10.1101/gr.186668.114 |pmid=26518481 |pmc=4691742 }}</ref> The lion-leopard clade was distributed in the Asian and African [[Palearctic]] since at least the early [[Pliocene]].<ref name=Tseng2014>{{cite journal |author1=Tseng, Z. J. |author2=Wang, X. |author3=Slater, G. J. |author4=Takeuchi, G. T. |author5=Li, Q. |author6=Liu, J. |author7=Xie, G. |date=2014 |title=Himalayan fossils of the oldest known pantherine establish ancient origin of big cats |journal=Proceedings of the Royal Society B: Biological Sciences |volume=281 |issue=1774 |article-number=20132686 |doi=10.1098/rspb.2013.2686|pmid=24225466 |pmc=3843846 |doi-access=free}}</ref> The earliest fossils recognisable as lions were found at [[Olduvai Gorge]] in Tanzania and are estimated to be up to 2 million years old.<ref name=werdelin2009>{{cite book|last1=Werdelin|first1=L.|last2=Yamaguchi|first2=N.|last3=Johnson|first3=W. E. |last4=O'Brien |first4=S. J.|chapter=Phylogeny and evolution of cats (Felidae)|date=2010|pages=59–82|isbn=978-0-19-923445-5|chapter-url=https://www.researchgate.net/publication/266755142 |title=Biology and Conservation of Wild Felids |publisher=Oxford University Press |access-date=10 February 2019 |archive-date=25 September 2018|archive-url=https://web.archive.org/web/20180925141956/https://www.researchgate.net/publication/266755142 |url-status=live}}</ref>

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  |url=http://www.zin.ru/Labs/theriology/eng/staff/baryshnikov/references/barnett_et_al_2009.pdf |archive-url=https://web.archive.org/web/20170808225555/http://www.zin.ru/labs/theriology/eng/staff/baryshnikov/references/barnett_et_al_2009.pdf |archive-date=8 August 2017 |url-status=live}}</ref><ref>{{cite journal |author1=Argant, A. |author2=Brugal, J.-P. |year=2017 |title=The cave lion ''Panthera (Leo) spelaea'' and its evolution: ''Panthera spelaea intermedia'' nov. subspecies |journal=Acta Zoologica Cracoviensia |volume=60 |issue=2 |pages=58–103 |doi=10.3409/azc.60_2.59 |doi-access=free}}</ref> The modern lion was probably widely distributed in Africa during the [[Middle Pleistocene]] and started to diverge in sub-Saharan Africa during the Late Pleistocene. Lion populations in East and Southern Africa became separated from populations in West and North Africa when the equatorial rainforest expanded 183,500 to 81,800 years ago.<ref name=Barnett_al2014>{{cite journal |author1=Barnett, R. |author2=Yamaguchi, N. |author3=Shapiro, B. |author4=Ho, S. Y. |author5=Barnes, I. |author6=Sabin, R. |author7=Werdelin, L. |author8=Cuisin, J.  |author9=Larson, G. |year=2014 |title=Revealing the maternal demographic history of ''Panthera leo'' using ancient DNA and a spatially explicit genealogical analysis |journal=BMC Evolutionary Biology |volume=14 |issue=1 |page=70 |doi=10.1186/1471-2148-14-70|pmid=24690312 |pmc=3997813 |doi-access=free |bibcode=2014BMCEE..14...70B}}</ref> They shared a common ancestor probably between 98,000 and 52,000 years ago.<ref name="DeManuel_al2020" /> Due to the expansion of the Sahara between 83,100 and 26,600 years ago, lion populations in West and North Africa became separated. As the rainforest decreased and thus gave rise to more open habitats, lions moved from West to Central Africa. Lions from North Africa dispersed to southern Europe and Asia between 38,800 and 8,300 years ago.<ref name=Barnett_al2014/>

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. |year=2011 |title=Genetic diversity, evolutionary history and implications for conservation of the lion (''Panthera leo'') in West and Central Africa |journal=Journal of Biogeography |volume=38 |issue=7 |pages=1356–1367 |doi=10.1111/j.1365-2699.2011.02500.x |bibcode=2011JBiog..38.1356B |s2cid=82728679}}</ref> A whole genome-wide sequence of lion samples showed that samples from West Africa shared [[alleles]] with samples from Southern Africa, and samples from Central Africa shared alleles with samples from Asia. This phenomenon indicates that Central Africa was a melting pot of lion populations after they had become isolated, possibly migrating through corridors in the [[Nile Basin]] during the early [[Holocene]].<ref name=DeManuel_al2020/>

===Hybrids===

{{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=218–234 |publisher=Springer |location=Berlin, Heidelberg |year=1967 |doi=10.1007/978-3-642-48949-5_12 |isbn=978-3-642-48949-5}}</ref> The [[liger]] is bigger than a lion and a tiger, whereas most [[tigon]]s are relatively small compared to their parents because of reciprocal gene effects.<ref name="shi">{{cite thesis |last=Shi |first=W. |year=2005 |title=Growth and Behaviour: Epigenetic and Genetic Factors Involved in Hybrid Dysgenesis |type=PhD |series=Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology |publisher=Acta Universitatis Upsaliensis |location=Uppsala |chapter=Hybrid dysgenesis effects |pages=8–10 |chapter-url=http://uu.diva-portal.org/smash/get/diva2:165749/FULLTEXT01.pdf |archive-url=https://web.archive.org/web/20190518050521/http://uu.diva-portal.org/smash/get/diva2:165749/FULLTEXT01.pdf |archive-date=18 May 2019 |url-status=live|url=http://urn.kb.se/resolve?urn=urn:nbn:se:uu:diva-4784 }}</ref><ref name="Carnivores">{{Cite book |last1=Rafferty |first1=J. P. |title=Carnivores: Meat-eating Mammals |date=2011 |location=New York |publisher=The Rosen Publishing Group |isbn=978-1-61530-340-3 |chapter=The Liger |page=120 |chapter-url=https://books.google.com/books?id=EMui7zVOqeUC&pg=PA120 |access-date=4 July 2014 |archive-date=5 May 2024 |archive-url=https://web.archive.org/web/20240505150613/https://books.google.com/books?id=EMui7zVOqeUC&pg=PA120#v=onepage&q&f=false |url-status=live }}</ref> The [[leopon]] is a hybrid between a lion and leopard.<ref>{{Cite journal |author1=Zhang, Z. |author2=Chen, J. |author3=Li, L. |author4=Tao, M. |author5=Zhang, C. |author6=Qin, Q. |author7=Xiao, J. |author8=Liu, Y. |author9=Liu, S. |title=Research advances in animal distant hybridization |year=2014 |journal=Science China Life Sciences |volume=57 |issue=9 |pages=889–902 |doi=10.1007/s11427-014-4707-1 |pmid=25091377 |s2cid=18179301 |url=https://link.springer.com/content/pdf/10.1007/s11427-014-4707-1.pdf |archive-url=https://web.archive.org/web/20181030104852/https://link.springer.com/content/pdf/10.1007%2Fs11427-014-4707-1.pdf |archive-date=30 October 2018 |url-status=live|doi-access=free }}</ref>

==Description==
{{multiple image|align=right|direction=vertical
|image1=Lioness 12.jpg|caption1=A tuft at the end of the tail is a distinct characteristic of the lion.
|image2=Description iconographique comparée du squelette et du système dentaire des mammifères récents et fossiles (Panthera leo).jpg|caption2=Skeleton}}

The lion is a muscular, broad-chested cat with a short, rounded head, a reduced neck, and round ears; males have broader heads. The fur varies in colour from light [[Buff (colour)|buff]] to silvery grey, yellowish red, and dark brown. The colours of the underparts are generally lighter. A new-born lion has dark [[Rosette (zoology)|spots]], which fade as the cub reaches adulthood, although faint spots may still be seen on the legs and underparts.<ref name=Guggisberg1975/><ref name=Haas2005>{{cite journal |author1=Haas, S. K. |author2=Hayssen, V. |author3=Krausman, P. R. |title=''Panthera leo'' |year=2005 |journal=Mammalian Species |volume=762 |pages=1–11 |url=http://www.science.smith.edu/msi/pdf/762_Panthera_leo.pdf |doi=10.1644/1545-1410(2005)762[0001:PL]2.0.CO;2 |s2cid=198968757 |archive-url=https://web.archive.org/web/20170728131140/http://www.science.smith.edu/msi/pdf/762_Panthera_leo.pdf |archive-date=28 July 2017}}</ref> The tail of all lions ends in a dark, hairy tuft that, in some lions, conceals an approximately {{cvt|5|mm}}-long, hard "spine" or "spur" composed of dermal papillae.<ref>{{Cite journal |last=Turner |first=W.C. |date=July 1873 |title=On the So-Called Prickle or Claw at the End of the Tail of the Lion and Other Felines &blobtype=pdf |journal=Journal of Anatomy and Physiology |volume=7 |issue=2 |pages=271–273 |pmid=17230977|pmc=1318943 }}</ref> The functions of the spur are unknown. The tuft is absent at birth, develops at around {{frac|5|1|2}} months of age and is readily identifiable at the age of seven months.<ref name="Schaller28">[[#Schaller|Schaller]], pp. 28–30.</ref>

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 |bibcode=1962Evolu..16..505D |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> Despite this, their thigh muscles do not exceed and only approach the mass predicted for their body mass.<ref name=":0">{{Cite book |last=Alexander |first=R. M. |url=https://www.originalwisdom.com/wp-content/uploads/bsk-pdf-manager/2020/04/Alexander_2013_Principles-of-Animal-Locomotion.pdf |title=Principles of animal locomotion. |date=2003 |publisher=Princeton University Press |pages=3, 103, 105, 143}}</ref> The lion has a high concentration of [[fast twitch muscle]] fibres, giving them quick bursts of speed but less stamina.<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://www.academia.edu/18641695 |journal=Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology |volume=159 |issue=2 |pages=125–133 |doi=10.1016/j.cbpa.2011.02.006 |hdl=2263/19598 |pmid=21320626}}</ref><ref>{{Cite journal |last1=Jacobs |first1=B. |last2=Garcia |first2=M. E. |last3=Shea-Shumsky |first3=N. B. |last4=Tennison |first4=M. E. |last5=Schall |first5=M. |last6=Saviano |first6=M. S. |last7=Tummino |first7=T. A. |last8=Bull |first8=A. J. |last9=Driscoll |first9=L. L. |last10=Raghanti |first10=M. A. |last11=Lewandowski |first11=A. H. |last12=Wicinski |first12=B. |last13=Ki Chui |first13=H. |last14=Bertelsen |first14=M. F. |last15=Walsh |first15=T. |date=2018 |title=Comparative morphology of gigantopyramidal neurons in primary motor cortex across mammals |journal=Journal of Comparative Neurology |volume=526 |issue=3 |pages=496–536 |doi=10.1002/cne.24349 |pmid=29088505 |url=https://www.researchgate.net/publication/320596823}}</ref> 

===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. |author2=Johnsingh, A. J. T.  |year=1993 |chapter=Management of Asiatic lions in the Gir Forest, India |pages=409–423 |title=Mammals as predators: the proceedings of a symposium held by the Zoological Society of London and the Mammal Society, London. Volume 65 of Symposia of the Zoological Society of London |editor1-last=Dunstone |editor1-first=N. |editor2-last=Gorman |editor2-first=M. L. |publisher=Zoological Society of London |location=London}}</ref><ref name=BCKM1993>{{cite book |author=Brakefield, T. |chapter=Lion: Sociable Simba |chapter-url=https://books.google.com/books?id=szBm5kPeC-cC&pg=PA50 |pages=50–67 |title=Big Cats: Kingdom of Might |publisher=Voyageur Press |location=London |year=1993 |isbn=978-0-89658-329-0 |access-date=1 November 2020 |archive-date=14 April 2021 |archive-url=https://web.archive.org/web/20210414114707/https://books.google.com/books?id=szBm5kPeC-cC&pg=PA50 |url-status=live}}</ref><ref name=nowak>{{Cite book |last=Nowak |first=R. M. |year=1999 |title=Walker's Mammals of the World |location=Baltimore |publisher=Johns Hopkins University Press |chapter=''Panthera leo'' |pages=832–834 |chapter-url=https://books.google.com/books?id=T37sFCl43E8C&pg=PR832 |isbn=978-0-8018-5789-8}}</ref> Accounts of a few individuals that were larger than average exist from Africa and India.<ref name=Guggisberg1975/><ref name=CAP>{{Cite book |last1=Nowell |first1=K. |last2=Jackson |first2=P. |title=Wild Cats: Status Survey and Conservation Action Plan |url=http://carnivoractionplans1.free.fr/wildcats.pdf |archive-url=https://web.archive.org/web/20050529182212/http://carnivoractionplans1.free.fr/wildcats.pdf |archive-date=29 May 2005 |url-status=live |year=1996 |publisher=IUCN/SSC Cat Specialist Group |location=Gland, Switzerland |isbn=978-2-8317-0045-8 |pages=17–21; 37–41 |chapter=African lion, ''Panthera leo'' (Linnaeus, 1758); Asiatic lion, ''Panthera leo persica'' (Meyer, 1826)}}</ref><ref name="Smuts et al., 1982">{{cite book |author=Smuts, G. L. |year=1982 |title=Lion |location=Johannesburg, South Africa |publisher=MacMillan }}</ref><ref name=Sinha1987>{{cite thesis |author=Sinha, S. P. |year=1987 |title=Ecology of wildlife with special reference to the lion (''Panthera leo persica'') in Gir Wildlife Sanctuary, Saurashtra, Gujurat |type=PhD |publisher=Saurashtra University |location=Rajkot |isbn=978-3-8443-0545-6}}</ref>

{| class="wikitable"
! Average !!Female lions !!Male lions
|-
|Head-and-body length
||{{cvt|160|-|184|cm}}<ref name=West2013>{{cite book |last1=West |first1=P. M. |last2=Packer |first2=C. |chapter=''Panthera leo'' Lion |pages=150–159 |chapter-url=https://books.google.com/books?id=B_07noCPc4kC&pg=RA4-PA150 |editor1=Kingdon, J. |editor2=Happold, D. |editor3=Butynski, T. |editor4=Hoffmann, M. |editor5=Happold, M. |editor6=Kalina, J. |title=Mammals of Africa |year=2013 |publisher=Bloomsbury Publishing |location=London |isbn=978-1-4081-8996-2}}</ref>
||{{cvt|184|-|208|cm}}<ref name=West2013/>
|-
|Tail length
||{{cvt|72|-|89.5|cm}}<ref name=West2013/>
||{{cvt|82.5|-|93.5|cm}}<ref name=West2013/>
|-
|Weight
||{{cvt|118.37|-|143.52|kg}} in Southern Africa,<ref name=Smuts_etal_1980/>{{break}}{{cvt|119.5|kg}} in East Africa,<ref name=Smuts_etal_1980/>{{break}}{{cvt|110|-|120|kg}} in India<ref name=Chellam1993/>
||{{cvt|186.55|-|225|kg}} in Southern Africa,<ref name=Smuts_etal_1980/>{{break}}{{cvt|174.9|kg}} in East Africa,<ref name=Smuts_etal_1980/>{{break}}{{cvt|160|-|190|kg}} in India<ref name=Chellam1993/>
|}

===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=N. |last2=Cooper |first2=A. |last3=Werdelin |first3=L. |last4=MacDonald |first4=D. 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/> Mutations in the genes [[microphthalmia-associated transcription factor]] and [[tyrosinase]] are possibly responsible for the colour of manes.<ref>{{Cite journal |last=Tefera |first=M. |date=2003 |title=Phenotypic and reproductive characteristics of lions (''Panthera leo'') at Addis Ababa Zoo |journal=Biodiversity & Conservation |volume=12 |issue=8 |pages=1629–1639 |doi=10.1023/A:1023641629538|bibcode=2003BiCon..12.1629T }}</ref><ref>{{Cite journal |last1=Barazandeh |first1=M. |last2=Kriti |first2=D. |last3=Fickel |first3=J. |last4=Nislow |first4=C. |date=2024 |title=The Addis Ababa Lions: Whole-genome sequencing of a rare and precious population |journal=Genome Biology and Evolution |volume=16 |issue=2 |article-number=evae021 |doi=10.1093/gbe/evae021 |pmc=10871700 |pmid=38302110}}</ref> 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, India |publisher=Dorling Kindersley India |isbn=978-0-14-302998-4}}</ref>{{page needed | date=April 2025}}

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> 

Almost all male lions in [[Pendjari National Park]] are either maneless or have very short manes.<ref name=Schoeetal2010>{{cite book |author1=Schoe, M. |author2=Sogbohossou, E. A. |author3=Kaandorp, J. |author4=De Iongh, H. |title=Progress Report—collaring operation Pendjari Lion Project, Benin |publisher=The Dutch Zoo Conservation Fund (for funding the project) |year=2010}}</ref> [[Maneless lion]]s have also been reported in Senegal, in [[Sudan]]'s [[Dinder National Park]] and in [[Tsavo East National Park]], Kenya.<ref>{{cite magazine |last=Trivedi |first=B. P. |title=Are maneless Tsavo Lions prone to male pattern baldness? |magazine=National Geographic |year=2005 |url=http://news.nationalgeographic.com/news/2002/04/0412_020412_TVtsavolions.html |archive-url=https://web.archive.org/web/20020605184139/http://news.nationalgeographic.com/news/2002/04/0412_020412_TVtsavolions.html |archive-date=5 June 2002 |access-date=7 July 2007}}</ref> [[Castration|Castrated]] lions often have little to no mane because the removal of the [[gonad]]s inhibits testosterone production.<ref>{{Cite journal |first=L. |last=Munson |date=2006 |title=Contraception in felids |journal=Theriogenology |pmid=16626799 |volume=66 |issue=1 |pages=126–134 |doi=10.1016/j.theriogenology.2006.03.016}}</ref> Rarely, both wild and captive lionesses have manes.<ref>{{cite magazine |last=Gruber |first=K. |date=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=L. J. |date=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 |magazine=National Geographic |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|archive-date=28 February 2021}}</ref>

===Colour variation===
{{Further|White lion}}
The white lion is a rare [[Polymorphism (biology)|morph]] with a genetic condition called [[leucism]], which is caused by a double [[recessive allele]]. It is not albino; it has normal pigmentation in the eyes and skin. White lions have occasionally been encountered in and around [[Kruger National Park]] and the adjacent [[Timbavati Private Game Reserve]] in eastern South Africa. They were removed from the wild in the 1970s, thus decreasing the white lion [[gene pool]]. Nevertheless, 17 births have been recorded in five prides between 2007 and 2015.<ref>{{cite journal |last1=Turner|first1=J. A.|last2= Vasicek|first2=C. A. |last3= Somers |first3=M. J. |year=2015 |title=Effects of a colour variant on hunting ability: the white lion in South Africa |journal=Open Science Repository Biology |page=e45011830}}</ref> White lions are selected for breeding in captivity.<ref>{{Cite book |last=McBride |first=C. |title=The White Lions of Timbavati |year=1977 |publisher=E. Stanton |location=Johannesburg |isbn= 978-0-949997-32-6}}</ref> They have reportedly been bred in camps in South Africa for use as trophies to be killed during [[canned hunt]]s.<ref>{{Cite book |last=Tucker|first=L. |title=Mystery of the White Lions—Children of the Sun God |year=2003 |publisher=Npenvu Press |location=Mapumulanga |isbn=978-0-620-31409-1}}</ref>

==Distribution and habitat==
[[File:Lions.Gir5.jpg|thumb|Lion in [[Gir National Park]], India]]
African lions live in scattered populations across sub-Saharan Africa. The lion prefers grassy plains and [[savannah]]s, scrub bordering rivers, and open woodlands with bushes. It rarely enters closed forests. On [[Mount Elgon]], the lion has been recorded up to an elevation of {{cvt|3600|m}} and close to the snow line on [[Mount Kenya]].<ref name=Guggisberg1975>{{Cite book |last=Guggisberg |first=C. A. W. |title=Wild Cats of the World |year=1975 |publisher=Taplinger Publishing |location=New York |isbn=978-0-8008-8324-9 |chapter=Lion ''Panthera leo'' (Linnaeus, 1758) |pages=138–179 |chapter-url=}}</ref> Savannahs with an annual rainfall of {{cvt|300|to|1500|mm}} make up the majority of lion habitat in Africa, estimated at {{cvt|3390821|km2}} at most, but remnant populations are also present in tropical moist forests in West Africa and [[Afromontane|montane]] forests in East Africa.<ref name=Riggio_al2013>{{cite journal |author=Riggio, J. |author2=Jacobson, A. |author3=Dollar, L. |author4=Bauer, H. |author5=Becker, M. |author6=Dickman, A. |author7=Funston, P. |author8=Groom, R. |author9=Henschel, P. |author10=de Iongh, H. |author11=Lichtenfeld, L. |author12=Pimm, S. |year=2013 |title=The size of savannah Africa: a lion's (''Panthera leo'') view |journal=Biodiversity Conservation |volume=22 |issue=1 |pages=17–35|doi=10.1007/s10531-012-0381-4 |doi-access=free |bibcode=2013BiCon..22...17R }}</ref> The Asiatic lion now survives only in and around [[Gir National Park]] in Gujarat, western India. Its habitat is a mixture of dry savannah forest and very dry, deciduous [[scrub forest]].<ref name=Breitenmoser2008/>

===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 |article-number=e60174 |doi=10.1371/journal.pone.0060174 |pmid=23573239 |pmc=3616087 |bibcode=2013PLoSO...860174B |doi-access=free }}</ref>

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 }}</ref> In [[Hungary]], the modern lion was present from about 4,500 to 3,200 years  [[Before Present]].<ref>{{cite journal |author1=Daróczi-Szabó, M. |author2=Kovács, Z. E. |author3=Raczky, P. |author4=Bartosiewicz, L.  |year=2020 |title=Pending danger: Recent copper age lion (''Panthera leo'' L., 1758) finds from Hungary |journal=International Journal of Osteoarchaeology |volume=30 |issue=4 |pages=469–481 |doi=10.1002/oa.2875|doi-access=free }}</ref> In [[Ukraine]], the modern lion was present from about 6,400 to 2,000 years Before Present.<ref name=Marciszak2022 /> In Greece, it was common, as reported by [[Herodotus]] in 480 BC; it was considered rare by 300 BC and [[Local extinction|extirpated]] by AD 100.<ref name=Guggisberg1975 />

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.  |title=Chronological distribution of the tiger ''Panthera tigris'' and the Asiatic lion ''Panthera leo persica'' in their common range in Asia |journal=[[Mammal Review]] |volume=49 |issue=4 |pages=340–353 |doi=10.1111/mam.12166 |date=2019 |bibcode=2019MamRv..49..340S |s2cid=202040786}}</ref> It was present in the [[Caucasus]] until the 10th century.<ref name=USSR/> It lived in the [[Levant]] until the [[Middle Ages]] and in Southwest Asia until the late 19th century. By the late 19th century, it had been extirpated in most of Turkey.<ref>{{cite book |author=Üstay, A. H. |year=1990 |title=Hunting in Turkey |publisher=BBA |location=Istanbul}}</ref> The last live lion in Iran was sighted in 1942, about {{cvt|65|km}} northwest of [[Dezful]],<ref name=Firouz05>{{cite book|last=Firouz|first=E.|year=2005|url=https://books.google.com/books?id=t2EZCScFXloC&pg=PA66|title=The complete fauna of Iran |publisher=I. B. Tauris |isbn=978-1-85043-946-2 |pages=5–67|access-date=25 September 2019|archive-date=5 May 2024 |archive-url=https://web.archive.org/web/20240505151453/https://books.google.com/books?id=t2EZCScFXloC&pg=PA66|url-status=live}}</ref> although the corpse of a lioness was found on the banks of the [[Karun]] river in [[Khuzestan province]] in 1944.<ref name=simba /> It once ranged from [[Sind]] and [[Punjab]] in Pakistan to [[Bengal]] and the [[Narmada River]] in central India.<ref name=Kinnear1920>{{cite journal |author=Kinnear, N. B. |year=1920 |title=The past and present distribution of the lion in southeastern Asia |journal=Journal of the Bombay Natural History Society |volume=27 |pages=34–39 |url=https://archive.org/stream/journalofbombayn27192022bomb#page/32/mode/2up}}</ref>

==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 |article-number=e16749 |doi=10.7717/peerj.16749 |pmid=38282863 |pmc=10821722 |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%26Packer95.pdf |archive-url=https://web.archive.org/web/20170810170702/http://www.life.umd.edu/faculty/wilkinson/BIOL608W/Heinsohn%26Packer95.pdf |archive-date=10 August 2017 |url-status=live |access-date=1 January 2019 }}</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 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 studied in [[Queen Elizabeth National Park]] increased their chances of success when hunting at night and moonless nights, when the prey was solitary and the cover of bushes dense; the chances of success increased when they attacked from a distance of {{cvt|20|m}} and with a grass cover of {{cvt|80|cm}} tall.<ref>{{Cite journal |last=Orsdol |first=K. G. V. |date=1984 |title=Foraging behaviour and hunting success of lions in Queen Elizabeth National Park, Uganda |journal=African Journal of Ecology |url=https://www.researchgate.net/publication/227938875 |volume=22 |issue=2 |pages=79–99 |doi=10.1111/j.1365-2028.1984.tb00682.x}}</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> Analysis of films of lion hunting has found that lions have an initial acceleration of 9.5 [[Metre per second squared|m/s²]], while a [[Thomson's gazelle]] has an acceleration of only 4.5 m/s², even though lions only reached 50 km/h (31 mph) compared to the gazelles' nearly 97 km/h (60 mph); lions stalk their prey in order to attack them unexpectedly, as their prey begins to run unprepared while the predator is better prepared to run.<ref name=":0" /> 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 to 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=978-1-62654-905-0 |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=978-0-7607-4767-4 |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 |article-number=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=R. D.| last2=Briggs |first2=M. B.| last3=Reeves| first3=J. J.}}</ref><ref>{{Cite book |last=Asdell |first=S. 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 also head-rub and roll around with each other before mounting each other.<ref>{{Cite book |last=Bagemihl |first=B. |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=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/>

Lionesses of reproductive age within a pride have a similar number of surviving cubs, which they initially keep hidden. Lionesses without offspring do not assist in the care of the communal litter.<ref>{{Cite journal |last1=Packer |first1=C. |last2=Pusey |first2=A. E. |last3=Eberly |first3=L. E. |date=2001 |title=Egalitarianism in female African lions |journal=Science |volume=293 |issue=5530 |pages=690–693 |doi=10.1126/science.1062320 |pmid=11474110 |url=https://www.researchgate.net/publication/11868804}}</ref>
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 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 |trans-title=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 | archive-url=https://web.archive.org/web/20070929044925/http://www.arushatimes.co.tz/2006/36/features_10.htm | archive-date=29 September 2007 }}</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 |hdl=11568/174438 |hdl-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-date=1967 |editor=Morris, D. |publisher=Aldine |location=Chicago |date=2011 |isbn=978-0-202-36816-0}}</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>

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/>

{{listen|filename=Lion raring-sound1TamilNadu178.ogg|title=Lion roar|description=A captive lion roaring}}
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 roaring, [[growling]] and [[snarling]]. Other sounds produced include puffing, humming, and exhalations similar to purring, while cubs communicate by [[meow]]ing and bleating. Roaring is used to advertise the animal's presence. Lions most often roar at night, a sound that can be heard from a distance of {{cvt|8|km}}.<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>

==Conservation==
The lion is listed as [[Vulnerable species|Vulnerable]] on the [[IUCN Red List]]. The Indian population is listed on [[CITES Appendix I]] and the African population on [[CITES Appendix II]].<ref name="IUCN" />

===In Africa===
[[File:Lion.ogv|thumb|right|Video of a wild lioness]]

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 |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 |pages=882−892 |doi=10.1644/13-mamm-a-306  |doi-access=free }}{{open access}}</ref> The West African lion population is isolated from the one in Central Africa, with little or no exchange of breeding individuals. In 2015, it was estimated that this population consists of about 400 animals, including fewer than 250 mature individuals. They persist in three protected areas in the region, mostly in one population in the [[W National Park|W]] [[Arli National Park|A]] [[Pendjari National Park|P]] protected area complex, shared by [[Benin]], [[Burkina Faso]] and [[Niger]]. This population is listed as [[Critically Endangered]].<ref name=Henschel2015/> Field surveys in the [[W-Arly-Pendjari Complex|WAP ecosystem]] revealed that lion occupancy is lowest in the W National Park, and higher in areas with permanent staff and thus better protection.<ref>{{cite journal |last1=Henschel |first1=P. |last2=Petracca |first2=L. S. |last3=Hunter |first3=L. T. |last4=Kiki |first4=M. |last5=Sewadé |first5=C. |last6=Tehou |first6=A. |last7=Robinson |first7=H. S. |year=2016 |title=Determinants of distribution patterns and management needs in a critically endangered lion ''Panthera leo'' population |journal=Frontiers in Ecology and Evolution |volume=4 |issue=4 |page=110 |doi=10.3389/fevo.2016.00110 |doi-access=free|bibcode=2016FrEEv...4..110H }}</ref>

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-access=free }}</ref> In 2015, an adult male lion and a female lion were sighted in Ghana's [[Mole National Park]]. These were the first sightings of lions in the country in 39 years.<ref>{{cite journal |last1=Angelici |first1=F. M. |last2=Rossi |first2=L. |year=2017 |title=Further lion, ''Panthera leo senegalensis'' Meyer, 1826, sightings in Mole National Park, Ghana, and possible first serval ''Leptailurus serval'' Schreber, 1776 record after 39 years (Mammalia Felidae) |journal=Biodiversity Journal |volume=8 |issue=2 |pages=749–752 |url=http://www.biodiversityjournal.com/pdf/8(2)_749-752.pdf |access-date=9 March 2018 |archive-date=10 March 2018 |archive-url=https://web.archive.org/web/20180310074200/http://www.biodiversityjournal.com/pdf/8(2)_749-752.pdf }}</ref> In the same year, a population of up to 200 lions that was previously thought to have been [[Local extinction|extirpated]] was filmed in the [[Alatash National Park]], Ethiopia, close to the Sudanese border.<ref name="NewScientist2016">{{cite magazine|magazine=New Scientist|last=Wong|first=S. |title=Hidden population of up to 200 lions found in remote Ethiopia|date=2016|url=https://www.newscientist.com/article/2075740-hidden-population-of-up-to-200-lions-found-in-remote-ethiopia/|access-date=2 February 2016|archive-date=1 February 2016|archive-url=https://web.archive.org/web/20160201230053/https://www.newscientist.com/article/2075740-hidden-population-of-up-to-200-lions-found-in-remote-ethiopia/|url-status=live}}</ref><ref name="BBC Ethiopian lion, 2016">{{cite news|date=2016|url=https://www.bbc.com/news/world-africa-35460573|title=Lions rediscovered in Ethiopia's Alatash National Park|publisher=BBC News|access-date=1 February 2016|archive-date=1 February 2016|archive-url=https://web.archive.org/web/20160201144220/http://www.bbc.com/news/world-africa-35460573|url-status=live}}</ref>

In 2005, Lion Conservation Strategies were developed for West and Central Africa, and 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=19 September 2015 |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=19 June 2015 |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>

===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 |pages=32−40}}</ref><ref>{{cite journal |author=Singh, A. P. |year=2017 |title=The Asiatic Lion (''Panthera leo persica''): 50 Years Journey for Conservation of an Endangered Carnivore and its Habitat in Gir Protected Area, Gujarat, India |journal=Indian Forester |volume=143 |issue=10 |pages=993–1003}}</ref><ref>{{cite journal |author=Singh, H. S. |year=2017 |title=Dispersion of the Asiatic lion ''Panthera leo persica'' and its survival in human-dominated landscape outside the Gir forest, Gujarat, India |journal=Current Science |volume=112 |issue=5 |pages=933–940 |doi=10.18520/cs/v112/i05/933-940 |doi-access=free}}</ref> In 2017, about 650 individuals were recorded during the Asiatic Lion Census.<ref>{{cite news|title=Lion population roars to 650 in Gujarat forests|url=http://timesofindia.indiatimes.com/city/ahmedabad/Lion-population-roars-to-650-in-Gujarat-forests/articleshow/59907625.cms|author=Kaushik, H.|newspaper=The Times of India|date=2017|access-date=9 August 2017|archive-date=8 August 2017|archive-url=https://web.archive.org/web/20170808193821/http://timesofindia.indiatimes.com/city/ahmedabad/lion-population-roars-to-650-in-gujarat-forests/articleshow/59907625.cms|url-status=live}}</ref>

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===
[[File:India Animals.jpg|thumb|Captive male Asiatic lions in [[Sanjay Gandhi National Park]], India]]

Lions imported to Europe before the middle of the 19th century were possibly foremost Barbary lions from North Africa, or Cape lions from Southern Africa.<ref>{{cite journal |last1=Barnett |first1=R. |last2=Yamaguchi |first2=N. |last3=Shapiro |first3=B. |last4=Nijman |first4=V. |year=2007 |title=Using ancient DNA techniques to identify the origin of unprovenanced museum specimens, as illustrated by the identification of a 19th century lion from Amsterdam |url=http://dpc.uba.uva.nl/cgi/t/text/get-pdf?c=ctz;idno=7602a02 |journal=Contributions to Zoology |volume=76 |issue=2 |pages=87–94 |access-date=27 July 2012 |archive-url=https://web.archive.org/web/20110522041326/http://dpc.uba.uva.nl/cgi/t/text/get-pdf?c=ctz;idno=7602a02 |archive-date=22 May 2011 |doi=10.1163/18759866-07602002 |s2cid=2131247}}</ref> Another 11 animals thought to be Barbary lions kept in [[Addis Ababa Zoo]] are descendants of animals owned by [[Haile Selassie I of Ethiopia|Emperor Haile Selassie]]. WildLink International in collaboration with [[Oxford University]] launched an ambitious International [[Barbary lion|Barbary Lion Project]] with the aim of identifying and breeding Barbary lions in captivity for eventual reintroduction into a national park in the [[Atlas Mountains]] of Morocco.<ref name="yamaguchi-haddane">{{Cite journal |last=Yamaguchi |first=N. |author2=Haddane, B. |year=2002 |title=The North African Barbary Lion and the Atlas Lion Project |journal=International Zoo News |volume=49 |pages=465–481}}</ref> However, a genetic analysis showed that the captive lions at Addis Ababa Zoo were not Barbary lions, but rather closely related to wild lions in Chad and Cameroon.<ref name="Bruche_al2012">{{Cite journal |last=Bruche |first=S. |author2=Gusset, M. |author3=Lippold, S. |author4=Barnett, R. |author5=Eulenberger, K. |author6=Junhold, J. |author7=Driscoll, C. A. |author8=Hofreiter, M. |title=A genetically distinct lion (''Panthera leo'') population from Ethiopia |journal=European Journal of Wildlife Research |year=2012 |pages=215–225 |doi=10.1007/s10344-012-0668-5 |volume=59 |issue=2 |s2cid=508478}}</ref>

In 1982, the [[Association of Zoos and Aquariums]] started a [[Species Survival Plan]] for the Asiatic lion to increase its chances of survival. In 1987, it was found that most lions in North American zoos were hybrids between African and Asiatic lions.<ref name=OBrien1987>{{cite journal |author1=O'Brien, S. J. |author2=Joslin, P. |author3=Smith, G. L. III |author4=Wolfe, R. |author5=Schaffer, N. |author6=Heath, E. |author7=Ott-Joslin, J. |author8=Rawal, P. P. |author9=Bhattacharjee, K. K. |author10=Martenson, J. S. |year=1987 |title=Evidence for African origins of founders of the Asiatic lion Species Survival Plan |journal=Zoo Biology |volume=6 |issue=2 |pages=99–116 |doi=10.1002/zoo.1430060202 |url=http://dobzhanskycenter.bio.spbu.ru/pdf/sjop/MS129_O%27Brien_ZooBiol.pdf |access-date=25 May 2019 |archive-url=https://web.archive.org/web/20140225092713/http://dobzhanskycenter.bio.spbu.ru/pdf/sjop/MS129_O%27Brien_ZooBiol.pdf |archive-date=25 February 2014 }}</ref> Breeding programs need to note origins of the participating animals to avoid cross-breeding different subspecies and thus reducing their conservation value.<ref>{{Cite book |last1=Frankham |first1=R. |last2=Ballou |first2=J. |last3=Briscoe |first3=D. |title=Introduction to Conservation Genetics |publisher=Cambridge University Press |year=2009| isbn=978-0-521-70271-3 |location=Cambridge, New York, Melbourne, Madrid |edition=Second |chapter=Genetic management of Captive Populations |pages=430–452 |chapter-url=https://books.google.com/books?id=vLZKnsCk89wC&pg=PA437 |access-date=25 May 2019}}</ref> Captive breeding of lions was halted to eliminate individuals of unknown origin and [[Pedigree (animal)|pedigree]]. Wild-born lions were imported to American zoos from Africa between 1989 and 1995. Breeding was continued in 1998 in the frame of an African lion Species Survival Plan.<ref>{{cite journal |author1=Daigle, C. L. |author2=Brown, J. L. |author3=Carlstead, K. |author4=Pukazhenthi, B. |author5=Freeman, E. W. |author6=Snider, R. J. |year=2015 |title=Multi-institutional survey of social, management, husbandry and environmental factors for the SSP African lion Panthera leo population: examining the effects of a breeding moratorium in relation to reproductive success |journal=International Zoo Yearbook |volume=49 |issue=1 |pages=198–213 |doi=10.1111/izy.12073 |url=https://www.researchgate.net/publication/270901189}}</ref>

About 77% of the captive lions registered in the [[International Species Information System]] in 2006 were of unknown origin; these animals might have carried genes that are extinct in the wild and may therefore be important to the maintenance of the overall [[genetic variability]] of the lion.<ref name=BarnettYamaguchi2006>{{Cite journal |last1=Barnett |first1=R. |last2=Yamaguchi |first2 =N. |last3=Barnes | first3=I. |last4=Cooper |first4=A. |date=2006 |title=Lost populations and preserving genetic diversity in the lion ''Panthera leo'': Implications for its ''ex situ'' conservation |journal=Conservation Genetics |volume=7 |issue=4 |pages=507–514 |doi=10.1007/s10592-005-9062-0 |bibcode=2006ConG....7..507B |s2cid=24190889 |url=http://abc.zoo.ox.ac.uk/Papers/consgen06_lion.pdf |archive-url=https://web.archive.org/web/20060824064412/http://abc.zoo.ox.ac.uk/Papers/consgen06_lion.pdf |archive-date=24 August 2006}}</ref>

==Interactions with humans==
===In zoos and circuses===
{{multiple image|align=right|direction=vertical
|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]], pp. 81–82.</ref> there are more than 1,000 African and 100 Asiatic lions in zoos and wildlife parks around the world. They are considered an ambassador species and are kept for tourism, education and conservation purposes.<ref name="WAZA">{{cite web |last1= Dollinger |first1= P. |last2= Geser |first2= S. |title=Lion: In the Zoo (subpage) |work= Visit the Zoo |publisher= WAZA (World Association of Zoos and Aquariums) |url= http://www.waza.org/en/zoo/visit-the-zoo/cats-1254385523/panthera-leo |access-date= 5 April 2011 |archive-url= https://web.archive.org/web/20110929135611/http://www.waza.org/en/zoo/visit-the-zoo/cats-1254385523/panthera-leo |archive-date= 29 September 2011 }}</ref> Lions can live over twenty years in captivity; for example, three sibling lions at the [[Honolulu Zoo]] lived to the age of 22 in 2007.<ref>{{Cite news|last=Aguiar|first=E.|date=2007|title=Honolulu zoo's old lion roars no more|newspaper=Honolulu Advertiser|url=http://the.honoluluadvertiser.com/article/2007/Aug/08/ln/hawaii708080394.html|access-date=4 September 2007|archive-date=25 December 2018|archive-url=https://web.archive.org/web/20181225033236/http://the.honoluluadvertiser.com/article/2007/Aug/08/ln/hawaii708080394.html}}</ref><ref>{{Cite news|last=Lum|first=C.|date=2007|title=Zoo puts end to 2 lions' suffering|newspaper=Honolulu Advertiser|url=http://the.honoluluadvertiser.com/article/2009/Feb/25/ln/hawaii902250384.html|access-date=29 September 2020|archive-date=6 November 2020|archive-url=https://web.archive.org/web/20201106104728/http://the.honoluluadvertiser.com/article/2009/Feb/25/ln/hawaii902250384.html}}</ref>

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|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>

Lions were kept in cramped and squalid conditions at [[London Zoo]] until a larger lion house with roomier cages was built in the 1870s.<ref name=Blunt208>[[#Blunt|Blunt]], p. 208.</ref> Further changes took place in the early 20th century when [[Carl Hagenbeck]] designed enclosures with concrete "rocks", more open space and a moat instead of bars, more closely resembling a natural habitat. Hagenbeck designed lion enclosures for both [[Melbourne Zoo]] and Sydney's [[Taronga Zoo]]; although his designs were popular, the use of bars and caged enclosures prevailed in many zoos until the 1960s.<ref name=dc69>[[#Courcy|de Courcy]], p. 69.</ref> In the late 20th century, larger, more natural enclosures and the use of wire mesh or laminated glass instead of lowered dens allowed visitors to come closer than ever to the animals; some attractions such as the Cat Forest/Lion Overlook of [[Oklahoma City Zoo and Botanical Garden|Oklahoma City Zoological Park]] placed the den on ground level, higher than visitors.<ref>{{cite encyclopedia |last=Grisham |first=J. |editor-first=C. E. |editor-last=Bell |title=Lion |encyclopedia=Encyclopedia of the World's Zoos |volume=2: G–P |year=2001 |publisher=Fitzroy Dearborn |location=Chicago |isbn=978-1-57958-174-9 |pages=733–739}}</ref>

[[Lion taming]] has been part of both established [[circus]]es and individual acts such as [[Siegfried & Roy]]. The practice began in the early 19th century by Frenchman Henri Martin and American [[Isaac A. Van Amburgh|Isaac Van Amburgh]], who both toured widely and whose techniques were copied by a number of followers.<ref name=baratay187/> Martin composed a [[pantomime]] titled ''Les Lions de Mysore'' ("the lions of Mysore"), an idea Amburgh quickly borrowed. These acts eclipsed [[equestrianism]] acts as the central display of circus shows and entered public consciousness in the early 20th century with cinema. In demonstrating the superiority of human over animal, lion taming served a purpose similar to animal fights of previous centuries.<ref name=baratay187>[[#Baratay|Baratay & Hardouin-Fugier]], p. 187.</ref> The ultimate proof of a tamer's dominance and control over a lion is demonstrated by the placing of the tamer's head in the lion's mouth. The now-iconic lion tamer's chair was possibly first used by American [[Clyde Beatty]] (1903–1965).<ref>{{Cite book |first=D. |last=Feldman |author-link=David Feldman (author) |year=1993 |title=How Does Aspirin Find a Headache? |publisher=HarperCollins |location=New York |isbn=978-0-06-016923-7 |url=https://archive.org/details/howdoesaspirinfi00davi}}</ref>

===Hunting and games===
{{Main|Lion hunting}}
{{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&nbsp;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]], pp. 156–159.</ref> In [[Ancient Rome]], lions were kept by emperors for [[Venatio|hunts]], [[Bestiarii|gladiator fights]] and [[Damnatio ad bestias|executions]].<ref>[[#Jackson|Jackson]], p. 142.</ref>

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>

===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-eating animal#Lions{{!}}Man-eater lions}}
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 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=927–928 |doi=10.1038/436927a| pmid=16107828 |bibcode=2005Natur.436..927P |s2cid=3190757}}</ref> Another study of 1,000 people attacked by lions in southern Tanzania between 1988 and 2009 found that the weeks following the [[full moon]], when there was less moonlight, were a strong indicator of increased night-time attacks on people.<ref name=fullmoon>{{cite journal |title=Fear of Darkness, the Full Moon and the Nocturnal Ecology of African Lions |last=Packer |first=C. |journal=[[PLOS One]] |date=2011 |volume=6 |issue=7 |doi=10.1371/journal.pone.0022285 |author2=Swanson, A. |author3=Ikanda, D. |author4=Kushnir, H. |article-number=e22285 |pmid=21799812 |pmc=3140494|bibcode=2011PLoSO...622285P|doi-access=free}}</ref>

According to Robert R. Frump, Mozambican refugees regularly crossing Kruger National Park, South Africa, at night are attacked and eaten by lions. Frump said thousands may have been killed in the decades after [[apartheid]] sealed the park and forced refugees to cross the park at night.<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>

===Cultural significance===
{{main|Cultural depictions of lions}}
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 'King of the Beasts'<ref>[[#Jackson|Jackson]], p. 7.</ref> and has symbolised power, royalty and protection.<ref name=Symbolism>{{cite book | first=H. B. | last=Werness |year=2007 |title=The Continuum Encyclopedia of Animal Symbolism in World Art |publisher=Continuum International Publishing Group |pages=254–260|isbn=978-0-8264-1913-2}}</ref> Several leaders have had "lion" in their name including [[Sundiata Keita]] of the [[Mali Empire]], who was called "Lion of Mali",<ref name=Lynch/> and [[Richard the Lionheart]] of England.<ref>[[#Jackson|Jackson]], p. 133.</ref> The male's mane makes it a particularly recognisable feature and thus has been represented more than the female.<ref>[[#Jackson|Jackson]], p. 100.</ref> Nevertheless, the lioness has also had importance as a guardian.<ref name=Symbolism/>
[[File:Berlín Ishtar león. 02.JPG|thumb|right|Roaring and striding lion from the Throne Room of [[Nebuchadnezzar&nbsp;II]], 6th century BC, from [[Babylon]], Iraq]]
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.<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 part lion, including the war deities [[Sekhmet]] and [[Maahes]], and [[Tefnut]], the goddess of moisture.<ref>[[#Jackson|Jackson]], pp. 107–108, 111.</ref>

[[File:Cowardly lion2.jpg|thumb|upright|[[Dorothy Gale]] meets the Cowardly Lion in ''[[The Wonderful Wizard of Oz]]''. Art by [[W. W. Denslow]], 1900.]]
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=E. |author-link=Elena Cassin |year=1981 |title= Le roi et le lion |journal=Revue de l'Histoire des Religions |trans-title=The King and the Lion | volume=298 |issue=198–4 |pages=355–401 |language=fr |doi=10.3406/rhr.1981.4828}}</ref> The big cat was a symbol and steed of fertility goddess [[Inanna]].<ref name=Symbolism/> Lions decorate the [[Ishtar_Gate#Ishtar_Gate_and_Processional_Way|Processional Way leading to the Ishtar Gate]] in Babylon which was built by [[Nebuchadnezzar II]] in the 6th century BCE. The [[Lion of Babylon]] symbolised the power of the king and protection of the land against enemies, but was also invoked for good luck.<ref>{{cite journal|last=Watanabe|first=C. E.|year=2015|title=The symbolic role of animals in Babylon: a contextual approach to the lion, the bull and the mušḫuššu |journal=Iraq |volume=77 |pages=215–224 |doi=10.1017/irq.2015.17}}</ref> The constellation [[Leo (constellation)|Leo the lion]] was first recognised by the Sumerians around 4,000 years ago and is the fifth sign of the [[zodiac]]. In ancient Israel, [[Lion of Judah|a lion]] represented the [[tribe of Judah]].<ref>[[#Jackson|Jackson]], pp. 109, 115.</ref> Lions are frequently mentioned in the [[Bible]], notably in the [[Book of Daniel]], in which the [[Daniel (biblical figure)|eponymous hero]] is forced to sleep in [[Daniel in the lions' den|the lions' den]].<ref>{{cite book |editor=Sakenfeld, K. D. |year=2008 |title=New Interpreter's Dictionary of the Bible |volume=3 |chapter=Lion |author=Borowski, O. |publisher=Abingdon Press |pages=669–670 |isbn=978-0-687-33365-3}}</ref>

Indo-Persian chroniclers regarded the lion as keeper of order in the realm of animals. The [[Sanskrit]] word ''mrigendra'' signifies a lion as king of animals.<ref>{{cite journal |author=Rangarajan, M. |year=2013 |title=Animals with rich histories: the case of the lions of Gir Forest, Gujarat, India |journal=History and Theory |volume=52 |issue=4 |pages=109–127 |doi=10.1111/hith.10690 |doi-access=free}}</ref> In India, the [[Lion Capital of Ashoka]], erected by Emperor Ashoka in the 3rd century BCE, depicts four lions standing back to back. In [[Hindu mythology]], the half-lion [[Narasimha]], an avatar of the deity [[Vishnu]], battles and slays the evil ruler [[Hiranyakashipu]]. In Buddhist art, lions are associated with both ''[[arhat]]s'' and [[bodhisattva]]s and may be ridden by the [[Manjushri]]. Though they were never native to the country, lions have played important roles in [[Chinese culture]]. Statues of the beast have guarded the entrances to the imperial palace and many religious shrines. The [[lion dance]] has been performed for over a thousand years.<ref>[[#Jackson|Jackson]], pp. 113, 119–122, 124.</ref>

In [[ancient Greece]], the lion is featured in several of [[Aesop's fables]], notably [[The Lion and the Mouse]]. In [[Greek mythology]], the [[Nemean lion]] is slain by the hero [[Heracles]] who then [[Skinning|wears its skin]]. [[Lancelot]] and [[Gawain]] were also heroes slaying lions in [[Middle Ages|medieval Europe]]. Lions continue to appear in modern literature such as the [[Cowardly Lion]] in [[L. Frank Baum]]'s 1900 ''[[The Wonderful Wizard of Oz]]'', and in [[C. S. Lewis]]'s ''[[The Lion, the Witch and the Wardrobe]]''. For cinema, the lion was portrayed as the ruler of animals in the 1994 [[Disney animated feature]] film ''[[The Lion King]]''.<ref>[[#Jackson|Jackson]], pp. 7, 96, 99, 103–105, 128, 135, 150, 197.</ref>

==See also==
{{div col|colwidth=15em}}
* [[List of largest cats]]
* [[Mapogo lion coalition]] 
{{div col end}}

== Notes ==
{{Notelist}}

==References==
<!--ContribZool76:87.-->

=== Citations ===
{{Reflist|23em}}

=== Books ===
{{Refbegin}}
* {{Cite book |last1=Baratay |first1=E. |first2=E. |last2=Hardouin-Fugier  |year=2002 |title=Zoo: A History of Zoological Gardens in the West |url=https://books.google.com/books?id=V0JSVvpZvYYC&pg=PA3 |location=London |publisher=Reaktion Books |isbn=978-1-86189-111-2 |ref=Baratay |access-date=1 November 2020 |archive-date=5 May 2024 |archive-url=https://web.archive.org/web/20240505151453/https://books.google.com/books?id=V0JSVvpZvYYC&pg=PA3#v=onepage&q&f=false |url-status=live }}
* {{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}}
* {{cite book |last=Jackson |first=D. |year=2010 |title=Lion |url=https://books.google.com/books?id=65lK7UIVRfIC |location=London |publisher=Reaktion Books |isbn=978-1-86189-655-1 |ref=Jackson |access-date=18 May 2019 |archive-date=5 March 2024 |archive-url=https://web.archive.org/web/20240305201628/https://books.google.com/books?id=65lK7UIVRfIC |url-status=live }}
* {{cite book|last=Packer|first=C.|author-link=Craig Packer|year=2023|title=The Lion: Behavior, Ecology, and Conservation of an Iconic Species|publisher=Princeton University Press|isbn=978-0-691-21529-7|url=https://books.google.com/books?id=TYOREAAAQBAJ&q=The+Lion:+Behavior,+Ecology,+and+Conservation+of+an+Iconic+Species|ref=Packer|access-date=10 July 2023|archive-date=28 August 2023|archive-url=https://web.archive.org/web/20230828195723/https://books.google.com/books?id=TYOREAAAQBAJ&q=The+Lion:+Behavior,+Ecology,+and+Conservation+of+an+Iconic+Species|url-status=live}}
* {{Cite book |last=Schaller |first=G. B. |author-link=George Schaller |year=1972 |title=The Serengeti Lion: A Study of Predator–Prey Relations |location=Chicago |publisher=University of Chicago Press |isbn=978-0-226-73639-6 |url=https://books.google.com/books?id=7ann2dYn9iYC&pg=PP1 |ref=Schaller |access-date=1 November 2020 |archive-date=5 May 2024 |archive-url=https://web.archive.org/web/20240505151524/https://books.google.com/books?id=7ann2dYn9iYC&pg=PP1#v=onepage&q&f=false |url-status=live }}
* {{Cite book |last1=Scott |first1=J. |last2=Scott |first2=A. |year=2002 |title=Big Cat Diary: Lion |location=New York |publisher=Harper Collins |isbn=978-0-00-714666-6 |ref=Scott}}
{{Refend}}

==External links==
{{Wiktionary}}
{{EB1911 poster|Lion}}
* {{cite web |url=http://www.catsg.org/index.php?id=108 |title=Lion ''Panthera leo'' |author=IUCN/SSC Cat Specialist Group |access-date=16 December 2014 |archive-date=27 March 2019 |archive-url=https://web.archive.org/web/20190327090743/http://www.catsg.org/index.php?id=108 }}
* {{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 |website=The Portugal News |date=2014 |access-date=24 May 2018 |archive-url=https://web.archive.org/web/20180802223523/http://www.theportugalnews.com/news/rare-desert-lion-killed-in-angola-after-supplying-unprecedented-data/32633 |archive-date=2 August 2018 }}
* {{Cite Collier's|wstitle=Lion|short=x}}

{{Carnivora |Fe.}}
{{Taxonbar|from1=Q140|from2=Q15294488}}
{{Authority control}}
{{Portal bar|Cats|Mammals|Animals|Biology|Africa|India}}

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