Array ( [0] => {{Short description|Large cat native to Africa and Asia}} [1] => {{Other uses|Lion (disambiguation)}} [2] => {{Featured article}} [3] => {{Use dmy dates|date=January 2020}} [4] => {{Speciesbox [5] => | name = Lion [6] => | fossil_range = {{fossil range|Pleistocene|Present}} [7] => | image = Lion waiting in Namibia.jpg [8] => | image_caption = Male in [[Okonjima]], Namibia [9] => | image2 = Okonjima Lioness.jpg [10] => | image2_caption = Female (lioness) in Okonjima [11] => | status = VU [12] => | status_system = IUCN3.1 [13] => | status_ref = {{cite iucn |title=''Panthera leo'' |errata=2017 |name-list-style=amp |author=Bauer, H. |author2=Packer, C. |author3=Funston, P. F. |author4=Henschel, P. |author5=Nowell, K. |year=2016 |page=e.T15951A115130419 |doi=10.2305/IUCN.UK.2016-3.RLTS.T15951A107265605.en |access-date=15 January 2022}} [14] => | status2 = CITES_A2 [15] => | status2_system = CITES [16] => | status2_ref = {{efn|Populations of India are listed in Appendix I.}} [17] => | taxon = Panthera leo{{MSW3 Carnivora |id=14000228 |page=546 |heading=Species ''Panthera leo''}} [18] => | authority = ([[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]) [19] => | subdivision_ranks = Subspecies [20] => | subdivision = :''[[Panthera leo leo|P. l. leo]]'' [21] => :''[[Panthera leo melanochaita|P. l. melanochaita]]'' [22] => :{{dagger}}''[[Sri Lanka lion|P. l. sinhaleyus]]'' [23] => | range_map = Lion distribution.png [24] => | range_map_caption = Historical and present distribution of the lion in Africa, Asia and Europe [25] => }} [26] => [27] => The '''lion''' ('''''Panthera leo''''') is a large [[Felidae|cat]] of the genus ''[[Panthera]]'', native to [[Africa]] and [[India]]. It has a muscular, broad-chested body; a short, rounded head; round ears; and a hairy tuft at the end of its [[tail]]. It is [[sexually dimorphic]]; adult male lions are larger than females and have a prominent mane. It is a [[social species]], forming groups called prides. A lion's pride consists of a few adult males, related females, and cubs. Groups of female lions usually hunt together, preying mostly on large [[ungulate]]s. The lion is an [[apex predator|apex]] and [[keystone predator]]; although some lions scavenge when opportunities occur and have been known to hunt [[human]]s, lions typically do not actively seek out and prey on humans. [28] => [29] => The lion inhabits [[grassland]]s, [[savannah]]s, and [[shrubland]]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]]. 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. [30] => [31] => One of the most widely recognised animal symbols in human culture, the lion has been extensively depicted in sculptures and paintings, on national flags, and in literature and films. 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]] were prominent in [[Ancient Egypt]], and depictions have occurred in virtually all ancient and medieval cultures in the lion's historic and current range. [32] => [33] => ==Etymology== [34] => 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.{{cite OED|lion|access-date=20 March 2022}} The generic name ''Panthera'' is traceable to the [[classical Latin]] word 'panthēra' and the [[ancient Greek]] word πάνθηρ 'panther'.{{cite book |last1=Liddell, H. G. |last2=Scott, R. |name-list-style=amp |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}} [35] => [36] => ==Taxonomy== [37] => [[File:Two cladograms for Panthera.svg|thumb|right|The upper cladogram is based on the 2006 study,{{cite journal |last1=Werdelin|first1=L. |last2=Yamaguchi|first2=N. |last3=Johnson |first3=W. E. |last4=O'Brien|first4=S. J. |name-list-style=amp |title=Phylogeny and evolution of cats (Felidae) |journal=Biology and Conservation of Wild Felids |year=2010 |pages=59–82 |url=https://www.researchgate.net/publication/266755142}} the lower one on the 2010 and 2011 studies.]] [38] => [39] => ''Felis leo'' was the [[scientific name]] used by [[Carl Linnaeus]] in 1758, who described the lion in his work ''[[Systema Naturae]]''.{{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}} The genus name ''Panthera'' was coined by [[Lorenz Oken]] in 1816.{{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}} 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. They were distinguished mostly by the size and colour of their manes and skins. [40] => [41] => ===Subspecies=== [42] => [[File:Lion subspecies distribution3.png|thumb|Range map showing distribution of subspecies and clades]] [43] => [44] => 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. 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.{{cite iucn |author=Breitenmoser, U. |author2=Mallon, D. P. |author3=Ahmad Khan, J. and |author4=Driscoll, C. |date=2008 |volume=2008 |page=e.T15952A5327221 |title=''Panthera leo'' ssp. ''persica'' |doi=10.2305/IUCN.UK.2008.RLTS.T15952A5327221.en }}{{cite iucn |author=Henschel, P. |author2=Bauer, H. |author3=Sogbohoussou, E. |author4=Nowell, K. |name-list-style=amp |date=2015 |volume=2015 |doi=10.2305/IUCN.UK.2015-2.RLTS.T68933833A54067639.en |title=''Panthera leo'' West Africa subpopulation}} 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:{{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. |name-list-style=amp |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}} [45] => [46] => *[[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. [[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)}}. Multiple authors referred to it as 'northern lion' and 'northern subspecies'.{{cite book |last=Wood |first=J. G. |author-link=John George Wood |title=The Illustrated Natural History. Mammalia, Volume 1 |publisher=[[Routledge]] |chapter=Felidæ; or the Cat Tribe |page=129−148 |location=London |chapter-url=https://books.google.com/books?id=v1DPAAAAMAAJ&pg=PA147 |year=1865}}{{cite book |last1=Hunter |first1=L. |author-link=Luke Hunter |last2=Barrett |first2=P. |title=The Field Guide to Carnivores of the World |edition=2 |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}} [47] => *[[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. 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)}}. It has been referred to as 'southern subspecies' and 'southern lion'. [48] => [49] => 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''.{{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}} [50] => [51] => 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.{{cite journal |author1=Bertola, L. D. |author2=Jongbloed, H. |author3=Van Der Gaag, K. J. |author4=De Knijff, P. |author5=Yamaguchi, N. |author6=Hooghiemstra, H. |author7=Bauer, H. |author8=Henschel, P. |author9=White, P. A. |author10=Driscoll, C. A. |author11=Tende, T. |author12=Ottosson, U. |author13=Saidu, Y. |author14=Vrieling, K. |author15=de Iongh, H. H. |year=2016 |title=Phylogeographic patterns in Africa and High Resolution Delineation of genetic clades in the Lion (''Panthera leo'') |journal=Scientific Reports |volume=6 |page=30807 |doi=10.1038/srep30807 |pmid=27488946 |pmc=4973251 |bibcode=2016NatSR...630807B}} [[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 result suggested that the taxonomic position of lions in Central Africa may require revision.{{cite journal |author1=Manuel, M. d. |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. |name-list-style=amp |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 }} [52] => [53] => ===Fossil records=== [54] => [55] => [[File:Panthera Atrox.jpg |thumb|right|Skull of an American lion on display at the [[National Museum of Natural History]]]] [56] => [57] => Other lion subspecies or [[sister species]] to the modern lion existed in prehistoric times:{{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 }} [58] => *''[[Panthera leo sinhaleyus|P. l. sinhaleyus]]'' was a [[fossil]] [[carnassial]] excavated in [[Sri Lanka]], which was attributed to a lion. It is thought to have become extinct around 39,000 years ago.{{Cite journal|first1=K. |last1=Manamendra-Arachchi |first2=R. |last2=Pethiyagoda |first3=R. |last3=Dissanayake |first4=M. |last4=Meegaskumbura |name-list-style=amp |year=2005 |title=A second extinct big cat from the late Quaternary of Sri Lanka |journal=[[The Raffles Bulletin of Zoology]] |issue=Supplement 12 |pages=423–434 |url=http://rmbr.nus.edu.sg/rbz/biblio/s12/s12rbz423-434.pdf |archive-url=https://web.archive.org/web/20070807215533/http://rmbr.nus.edu.sg/rbz/biblio/s12/s12rbz423-434.pdf |url-status=dead |archive-date=2007-08-07 }} [59] => *[[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.{{cite journal |last1=Marciszak |first1=A. |last2=Stefaniak |first2=K. |name-list-style=amp |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 |url=https://www.researchgate.net/publication/233669138}}{{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 }} [60] => *''[[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.{{cite journal |last1=Stuart |first1=A. J. |last2=Lister |first2=A. M. |name-list-style=amp |year=2011 |title=Extinction chronology of the cave lion ''Panthera spelaea'' |journal=Quaternary Science Reviews |volume=30 |issue=17 |pages=2329–40 |doi=10.1016/j.quascirev.2010.04.023 |bibcode=2011QSRv...30.2329S}} Bone fragments excavated in European, North Asian, Canadian and Alaskan caves indicate that it ranged from Europe across Siberia into western Alaska.{{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}} It likely derived from ''P. fossilis'',{{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. |name-list-style=amp |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 |page=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 }} and was genetically isolated and highly distinct from the modern lion in Africa and Eurasia.{{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. |name-list-style=amp |url-status=dead |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 }} It is depicted in [[Paleolithic]] cave paintings, ivory carvings, and clay busts.{{Cite journal |author=Packer, C. |author2=Clottes, J. |name-list-style=amp |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=2015-12-29 |url-status=live}} [61] => *[[American lion|''P. atrox'']], or the '''American lion''', ranged in the Americas from Canada to possibly [[Patagonia]].{{Cite journal |last1=Chimento |first1=N. R. |last2=Agnolin |first2=F. L. |name-list-style=amp |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 }} It arose when a cave lion population in [[Beringia]] became isolated south of the [[Cordilleran Ice Sheet]] about 370,000 years ago.{{cite journal |last1=Harington |first1=C. R.|title=Pleistocene remains of the lion-like cat (''Panthera atrox'') from the Yukon Territory and northern Alaska |journal=Canadian Journal of Earth Sciences |volume= 6|issue= 5 |year=1969 |pages=1277–1288 |doi=10.1139/e69-127|bibcode=1969CaJES...6.1277H}}{{cite journal |last1=Christiansen |first1= P. |last2=Harris |first2=J. M. |name-list-style=amp |title=Craniomandibular morphology and phylogenetic affinities of ''Panthera atrox'': implications for the evolution and paleobiology of the lion lineage |year=2009 |journal=Journal of Vertebrate Paleontology |volume=29 |issue=3 |pages= 934–945 |doi=10.1671/039.029.0314|bibcode= 2009JVPal..29..934C |s2cid= 85975640 }} A fossil from [[Edmonton]] dates to 11,355 ± 55 years ago.{{Cite journal |doi=10.1080/08912963.2013.861462 |title=Phylogenetics of ''Panthera'', including ''Panthera atrox'', based on craniodental characters |journal=[[Historical Biology]] |volume=26 |issue=6 |pages=827–833 |url=https://www.researchgate.net/publication/265790587 |date=2014 |last1=King |first1=L. M. |last2=Wallace |first2=S. C. |bibcode=2014HBio...26..827K |s2cid=84229141 |name-list-style=amp}} [62] => [63] => ===Evolution=== [64] => [65] => [[File:Cave lion range.png|thumb|red ''[[Panthera spelaea]]''{{break}}blue ''[[Panthera atrox]]''{{break}}green ''Panthera leo''{{break}}{{break}}Maximal range of the modern lion{{break}}and its prehistoric relatives{{break}}in the late Pleistocene]] [66] => [67] => The ''Panthera'' [[Lineage (evolution)|lineage]] is estimated to have [[Genetic divergence|genetically diverged]] from the [[common ancestor]] of the [[Felidae]] around {{Ma|9.32|4.47|million years ago}} to {{Ma|11.75|0.97|million years ago}}.{{cite journal |last1=Johnson |first1=W. E. |last2=Eizirik |first2=E. |last3=Pecon-Slattery |first3=J. |last4=Murphy |first4=W. J. |last5=Antunes |first5=A. |last6=Teeling |first6=E. |last7=O'Brien |first7=S. J. |name-list-style=amp |title=The late miocene radiation of modern Felidae: A genetic assessment |journal=[[Science (journal)|Science]] |volume=311 |issue=5757 |pages=73–77 |date=2006 |pmid=16400146 |doi=10.1126/science.1122277 |bibcode=2006Sci...311...73J|s2cid=41672825 |url=https://zenodo.org/record/1230866 }}{{cite book |last1=Werdelin |first1=L. |last2=Yamaguchi |first2=N. |last3=Johnson |first3=W. E. |last4=O'Brien |first4=S. J. |name-list-style=amp |chapter=Phylogeny and evolution of cats (Felidae) |date=2010 |pages=59–82 |publisher=Oxford University Press |location=Oxford, UK |isbn=978-0-19-923445-5 |chapter-url=https://www.researchgate.net/publication/266755142 |editor1-last=Macdonald |editor1-first=D. W. |editor2-last=Loveridge |editor2-first=A. J. |title=Biology and Conservation of Wild Felids}}{{cite journal |last1=Li |first1=G. |last2=Davis |first2=B. W. |last3=Eizirik |first3=E. |last4=Murphy |first4=W. J. |name-list-style=amp |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}} Results of analyses differ in the phylogenetic relationship of the lion; it was thought to form a [[sister group]] with the [[jaguar]] that diverged {{Ma|3.46|1.22|million years ago}}, but also with the [[leopard]] that diverged {{Ma|3.1|1.95|million years ago}}{{cite journal |last1=Davis|first1=B. W. |last2=Li |first2=G. |last3=Murphy |first3=W. J. |name-list-style=amp |title=Supermatrix and species tree methods resolve phylogenetic relationships within the big cats, ''Panthera'' (Carnivora: Felidae) |journal=Molecular Phylogenetics and Evolution |year=2010 |volume=56 |issue=1 |pages=64–76 |doi=10.1016/j.ympev.2010.01.036 |pmid=20138224 |url=https://www.academia.edu/12157986}}{{cite journal |last1=Mazák|first1=J. H. |last2=Christiansen |first2=P. |last3=Kitchener |first3=A. C. |last4=Goswami |first4=A. |name-list-style=amp |title=Oldest known pantherine skull and evolution of the tiger |journal=PLOS ONE |year=2011 |volume=6 |issue=10 |pages=e25483 |doi=10.1371/journal.pone.0025483 |pmid=22016768 |pmc=3189913|bibcode=2011PLoSO...625483M|doi-access=free }} to {{Ma|4.32|0.02|million years ago}}. [[Hybrid (biology)|Hybridisation]] between lion and [[snow leopard]] ancestors possibly continued until about 2.1 million years ago. The lion-leopard clade was distributed in the Asian and African [[Palearctic]] since at least the early [[Pliocene]].{{cite journal |author1=Tseng, Z. J. |author2=Wang, X. |author3=Slater, G. J. |name-list-style=amp |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 |page=20132686 |doi=10.1098/rspb.2013.2686|pmid=24225466 |pmc=3843846 |doi-access=free }} The earliest fossils recognisable as lions were found at [[Olduvai Gorge]] in Tanzania and are estimated to be up to 2 million years old. [68] => [69] => 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. The Eurasian and American cave lions became extinct at the end of the [[last glacial period]] without [[Mitochondrion|mitochondrial]] descendants on other continents.{{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 |name-list-style=amp |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=2017-08-08 |url-status=live}}{{cite journal |author1=Argant, A. |author2=Brugal, J.-P. |name-list-style=amp |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}} 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.{{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. |name-list-style=amp |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 }} They shared a common ancestor probably between 98,000 and 52,000 years ago. 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. [70] => [71] => 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.{{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. |name-list-style=amp |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 }} 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]]. [72] => [73] => ===Hybrids=== [74] => [75] => {{Further|Panthera hybrid}} [76] => [77] => In zoos, lions have been bred with [[tiger]]s to create hybrids for the curiosity of visitors or for scientific purpose.{{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 }}{{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}} 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.{{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 |page=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=2019-05-18 |url-status=live|url=http://urn.kb.se/resolve?urn=urn:nbn:se:uu:diva-4784 }}{{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}} The [[leopon]] is a hybrid between a lion and leopard.{{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=2018-10-30 |url-status=live|doi-access=free }} [78] => [79] => ==Description== [80] => {{multiple image|align=right|direction=vertical [81] => |image1=Lioness 12.jpg|caption1=A tuft at the end of the tail is a distinct characteristic of the lion. [82] => |image2=Description iconographique comparée du squelette et du système dentaire des mammifères récents et fossiles (Panthera leo).jpg|caption2=Skeleton}} [83] => [84] => 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.{{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 |url-status=dead |archive-url=https://web.archive.org/web/20170728131140/http://www.science.smith.edu/msi/pdf/762_Panthera_leo.pdf |archive-date=28 July 2017 }} The tail of all lions ends in a dark, hairy tuft that, in some lions, conceals an approximately {{cvt|5|mm|adj=on}}-long, hard "spine" or "spur" that is formed from the final, fused sections of tail bone. The functions of the spur are unknown. The tuft is absent at birth and develops at around {{frac|5|1|2}} months of age. It is readily identifiable at the age of seven months.[[#Schaller|Schaller]], pp. 28–30. [85] => [86] => 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.{{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}}{{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}} [87] => [88] => The skeletal muscles of the lion make up 58.8% of its body weight and represent the highest percentage of muscles among mammals.{{Cite journal |last=Davis|first=D. D. |date=1962 |title=Allometric relationships in Lions vs. Domestic Cats |journal=Evolution |volume=16 |issue=4 |pages=505–514 |doi=10.1111/j.1558-5646.1962.tb03240.x |doi-access=free}}{{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}} [89] => [90] => ===Size=== [91] => Among felids, the lion is second only to the tiger in size. The size and weight of adult lions vary across its range and habitats.{{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 }}{{cite book |author=Chellam, R. and A. J. T. Johnsingh |year=1993 |chapter=Management of Asiatic lions in the Gir Forest, India |pages=409–23 |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}}{{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}}{{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}} Accounts of a few individuals that were larger than average exist from Africa and India.{{Cite book |last1=Nowell |first1=K. |last2=Jackson |first2=P. |title=Wild Cats: Status Survey and Conservation Action Plan|chapter-url=http://carnivoractionplans1.free.fr/wildcats.pdf |archive-url=https://web.archive.org/web/20050529182212/http://carnivoractionplans1.free.fr/wildcats.pdf |archive-date=2005-05-29 |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)}}{{cite book |author=Smuts, G. L. |year=1982 |title=Lion |location=Johannesburg, South Africa |publisher=MacMillan }}{{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=978-3844305456}} [92] => [93] => {| class="wikitable" [94] => ! Average !!Female lions !!Male lions [95] => |- [96] => |Head-and-body length [97] => ||{{cvt|160|-|184|cm}}{{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}} [98] => ||{{cvt|184|-|208|cm}} [99] => |- [100] => |Tail length [101] => ||{{cvt|72|-|89.5|cm}} [102] => ||{{cvt|82.5|-|93.5|cm}} [103] => |- [104] => |Weight [105] => ||{{cvt|118.37|-|143.52|kg}} in Southern Africa,{{break}}{{cvt|119.5|kg}} in East Africa,{{break}}{{cvt|110|-|120|kg}} in India [106] => ||{{cvt|186.55|-|225|kg}} in Southern Africa,{{break}}{{cvt|174.9|kg}} in East Africa,{{break}}{{cvt|160|-|190|kg}} in India [107] => |} [108] => [109] => ===Mane=== [110] => {{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=Male in Pendjari National Park}} [111] => [112] => The male lion's mane is the most recognisable feature of the species.{{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}} It may have evolved around 320,000–190,000 years ago.{{Cite journal |last1=Yamaguchi |first1=Nobuyuki |last2=Cooper |first2=A. |last3=Werdelin |first3=L. |last4=MacDonald |first4=David W. |date=2004 |title=Evolution of the mane and group-living in the lion (''Panthera leo''): a review |journal=Journal of Zoology |volume=263 |issue=4 |pages=329–342 |doi=10.1017/S0952836904005242}} 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. It starts growing when lions enter adolescence, when [[testosterone]] levels increase, and reach their full size at around four years old.[[#Packer|Packer]], p. 148. Cool ambient temperatures in European and North American zoos may result in a heavier mane. On average, Asiatic lions have sparser manes than African lions.{{Cite book |last=Menon |first=V. |year=2003 |title=A Field Guide to Indian Mammals |location=New Delhi |publisher=Dorling Kindersley India |isbn=978-0-14-302998-4}} [113] => [114] => 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.{{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}}[[#Packer|Packer]], pp. 137, 145. Unlike in other felid species, female lions consistently interact with multiple males at once.[[#Packer|Packer]], pp. 145. Another hypothesis suggests that the mane also serves to protect the neck in fights, but this is disputed.[[#Schaller|Schaller]], pp. 360.[[#Packer|Packer]], pp. 133. 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.[[#Packer|Packer]], pp. 133, 137, 145, 148. [115] => [116] => Almost all male lions in [[Pendjari National Park]] are either maneless or have very short manes.{{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}} [[Maneless lion]]s have also been reported in Senegal, in [[Sudan]]'s [[Dinder National Park]] and in [[Tsavo East National Park]], Kenya.{{cite magazine |last=Trivedi |first=Bijal 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 |url-status=dead |archive-date=5 June 2002 |access-date=7 July 2007}} [[castration|Castrated]] lions often have little to no mane because the removal of the [[gonad]]s inhibits testosterone production.{{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}} [117] => [118] => Rarely, lionesses (in the wild and in captivity) have been observed to grow manes.{{cite magazine |last=Gruber |first=Karl |date=24 October 2022 |access-date=19 November 2023 |title=Five wild lionesses grow a mane and start acting like males |url=https://www.newscientist.com/article/2106866-five-wild-lionesses-grow-a-mane-and-start-acting-like-males/ |magazine=New Scientist}}{{cite magazine |last=Young |first=Lauren J. |date=23 September 2016 |access-date=19 November 2023 |title=The rare case of a lioness with a mane |url=https://www.popsci.com/science/zoo-lioness-mane/ |magazine=Popular Science}} Increased testosterone may be the cause of maned lionesses reported in northern Botswana.{{cite magazine |last=Dell'Amore |first=C. |date=2016 |access-date=18 April 2016 |title=No, those aren't male lions mating. One is likely a female |url=https://www.nationalgeographic.com/animals/article/160418-lions-mating-africa-animals-science|archive-url=https://web.archive.org/web/20210228190519/https://www.nationalgeographic.com/animals/article/160418-lions-mating-africa-animals-science|url-status=dead|archive-date=28 February 2021|magazine=National Geographic}} [119] => [120] => ===Colour variation=== [121] => {{Further|White lion}} [122] => 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.{{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}} White lions are selected for breeding in captivity.{{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}} They have reportedly been bred in camps in South Africa for use as trophies to be killed during [[canned hunt]]s.{{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}} [123] => [124] => ==Distribution and habitat== [125] => [[File:Asiatic lion 01.jpg|thumb|Lion in [[Gir National Park]]]] [126] => 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]].{{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=}} 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.{{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 }} 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]]. [127] => [128] => ===Historical range=== [129] => In Africa, the range of the lion originally spanned most of the central [[African rainforest]] zone and the [[Sahara]] desert.[[#Schaller|Schaller]], p. 5. In the 1960s, it became extinct in North Africa, except in the southern part of Sudan.{{cite journal |last1=Black |first1=S. A. |last2=Fellous |first2=A. |last3=Yamaguchi |first3=N. | last4=Roberts |first4=D. L. |year=2013 |title=Examining the Extinction of the Barbary Lion and Its Implications for Felid Conservation |journal=PLOS ONE |volume=8 |issue=4 |page=e60174 |doi=10.1371/journal.pone.0060174 |pmid=23573239 |pmc=3616087 |bibcode=2013PLoSO...860174B |doi-access=free}} [130] => [131] => In southern Europe and Asia, the lion once ranged in regions where climatic conditions supported an abundance of prey.{{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|s2cid=202040786 }} In [[Ukraine]], the modern lion was present from about 6,400 to 2,000 [[Before Present|years BP]].{{Cite journal |last1=Marciszak |first1=Adrian |last2=Ivanoff |first2=Dmitry V. |last3=Semenov |first3=Yuriy A. |last4=Talamo |first4=Sahra |last5=Ridush |first5=Bogdan |last6=Stupak |first6=Alina |last7=Yanish |first7=Yevheniia |last8=Kovalchuk |first8=Oleksandr |date=15 November 2022 |title=The Quaternary lions of Ukraine and a trend of decreasing size in Panthera spelaea |url=https://link.springer.com/10.1007/s10914-022-09635-3 |journal=[[Journal of Mammalian Evolution]] |language=en |volume=30 |issue=1 |pages=109–135 |doi=10.1007/s10914-022-09635-3 |issn=1064-7554 |access-date=21 March 2024|hdl=11585/903022 |hdl-access=free }} 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. It was present in the [[Caucasus]] until the 10th century. It lived in [[Palestine (region)|Palestine]] 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.{{cite book |author=Üstay, A. H. |year=1990 |title=Hunting in Turkey |publisher=BBA |location=Istanbul}} The last live lion in Iran was sighted in 1942, about {{cvt|65|km}} northwest of [[Dezful]],{{cite book |last=Firouz |first=E.|year=2005 |url={{Google books |plainurl=yes |id=t2EZCScFXloC |page=66}}|title=The complete fauna of Iran |publisher=I. B. Tauris| isbn=978-1-85043-946-2 |pages=5–67}} although the corpse of a lioness was found on the banks of the [[Karun]] river in [[Khuzestan province]] in 1944. It once ranged from [[Sind]] and [[Punjab]] in Pakistan to [[Bengal]] and the [[Narmada River]] in central India.{{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}} [132] => [133] => ==Behaviour and ecology== [134] => Lions spend much of their time resting; they are inactive for about twenty hours per day.[[#Schaller|Schaller]], p. 122. 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.[[#Schaller|Schaller]], pp. 120–21. [135] => [136] => ===Group organisation=== [137] => {{multiple image|align=right|direction=vertical [138] => |image1=Lion (Panthera leo) male and cub Etosha.jpg|caption1=Lion pride in Etosha National Park [139] => |image2=Lions Family Portrait Masai Mara.jpg|caption2=A lioness (left) and two males in Masai Mara}} [140] => 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".[[#Schaller|Schaller]], p. 33. Females form the stable [[social unit]] in a pride and do not tolerate outside females.[[#Schaller|Schaller]], p. 37. The majority of females remain in their birth prides while all males and some females will [[Biological dispersal|disperse]].[[#Packer|Packer]], p. 33. The average pride consists of around 15 lions, including several adult females and up to four males and their cubs of both sexes. Large prides, consisting of up to 30 individuals, have been observed.[[#Schaller|Schaller]], p. 34–35. The sole exception to this pattern is the [[Tsavo lion]] pride that always has just one adult male.{{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}} Prides act as [[fission–fusion society|fission–fusion societies]], and members will split into subgroups that keep in contact with [[roar (vocalization)|roars]].[[#Packer|Packer]], pp. 25, 31. [141] => [142] => Nomadic lions range widely and move around sporadically, either in pairs or alone. Pairs are more frequent among related males. A lion may switch lifestyles; nomads can become residents and vice versa.{{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 }} Interactions between prides and nomads tend to be hostile, although pride females in [[estrus]] allow nomadic males to approach them.[[#Schaller|Schaller]], pp. 52–54. Males spend years in a nomadic phase before gaining residence in a pride.{{cite book |author=Hanby, J. P.|author2=Bygott, J. D. |name-list-style=amp |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}} A study undertaken in the [[Serengeti National Park]] revealed that nomadic coalitions gain residency at between 3.5 and 7.3 years of age.{{cite journal |author=Borrego, N. |name-list-style=amp |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}} 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.{{cite journal |author=van Hooft, P.|author2=Keet, D.F. |name-list-style=amp |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 }} [143] => [144] => The evolution of sociability in lions was likely driven both by high population density and the clumped resources of savannah habitats. The larger the pride, the more high-quality [[Territory (animal)|territory]] they can defend; "hotspots" being near river [[confluence]]s, where the cats have better access to water, prey and shelter (via vegetation).[[#Packer|Packer]], pp. 195–196, 222.{{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}} The area occupied by a pride is called a "pride area" whereas that occupied by a nomad is a "range". Males associated with a pride patrol the fringes. Both males and females defend the pride against intruders, but the male lion is better-suited for this purpose due to its stockier, more powerful build. Some individuals consistently lead the defense against intruders, while others lag behind.{{Cite journal |last=Heinsohn|first= R. |author2=C. Packer |name-list-style=amp |year=1995 |title=Complex cooperative strategies in group-territorial African lions |journal=Science |volume=269 |issue=5228 |pages=1260–1262 |doi=10.1126/science.7652573 |pmid=7652573|bibcode=1995Sci...269.1260H |s2cid= 35849910 |url=http://www.life.umd.edu/faculty/wilkinson/BIOL608W/Heinsohn&Packer95.pdf |archive-url=https://web.archive.org/web/20170810170702/http://www.life.umd.edu/faculty/wilkinson/BIOL608W/Heinsohn&Packer95.pdf |archive-date=2017-08-10 |url-status=live}} Lions tend to assume specific roles in the pride; slower-moving individuals may provide other valuable services to the group.{{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}} 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.{{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 }} The male or males associated with the pride must defend their relationship with the pride from outside males who may attempt to usurp them. [[Dominance hierarchy|Dominance hierarchies]] do not appear to exist among individuals of either sex in a pride.[[#Packer|Packer]], pp. 42, 57. [145] => [146] => 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.{{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}} 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.{{cite journal |author=Chakrabarti, S. |name-list-style=amp |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}} [147] => [148] => ===Hunting and diet=== [149] => {{multiple image| perrow=1 |image2=Lions taking down cape buffalo.jpg|caption2=Four lionesses catching a buffalo in the Serengeti |image3=Lion and eland.jpg |caption3=A skeletal mount of a lion attacking a [[common eland]], on display at [[The Museum of Osteology]] [150] => |image1=Male Lion and Cub Chitwa South Africa Luca Galuzzi 2004.JPG |caption1=Male lion and cub feeding on a [[Cape buffalo]] in [[Sabi Sand Game Reserve]]}} [151] => [152] => 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.[[#Schaller|Schaller]], pp. 208.{{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}} Its prey consists mainly of [[ungulate]]s, particularly [[blue wildebeest]], [[plains zebra]], [[African buffalo]], [[common warthog]], [[gemsbok]] and [[giraffe]].{{Cite journal |last1=Hayward |first1=M. W. |last2=Kerley |first2=G. I. H. |name-list-style=amp |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 |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=2015-06-27 |url-status=live |citeseerx=10.1.1.611.8271}} In India, [[chital]] and [[sambar deer]] are the most common wild prey,{{cite journal |last1=Mukherjee |first1=S. |last2=Goyal |first2=S. P. |last3=Chellam| first3=R. |name-list-style=amp |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}} while livestock contributes significantly to lion kills outside protected areas.{{cite journal |author1=Meena, V. |name-list-style=amp |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}} They usually avoid fully grown adult [[elephant]]s, [[rhinoceros]]es and [[hippopotamus]] and small prey like [[dik-dik]], [[hyrax]]es, [[hare]]s and [[monkey]]s.[[#Schaller|Schaller]], p. 195. Unusual prey include [[Old World porcupine|porcupine]]s and small reptiles. Lions kill other predators but seldom consume them.[[#Schaller|Schaller]], pp. 220–221. [153] => [154] => 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.[[#Schaller|Schaller]], p. 153. Single lions are capable of bringing down zebra and wildebeest, while larger prey like buffalo and giraffe are riskier. 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.{{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 }}{{cite journal |last1=Power |first1=R. J. |last2=Compion |first2=R. X. S. |name-list-style=amp |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}} In typical 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.{{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 |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=2015-05-18 |url-status=live}} Some evidence suggests, however, that males are just as successful as females; they are typically solo hunters who ambush prey in small bushland.{{cite journal |last1=Loarie, S. R. |last2=Tambling, C. J. |last3=Asner, G. P. |name-list-style=amp |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 |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=2017-08-16 |url-status=live |hdl-access=free}} They may join in the hunting of large, slower-moving prey like buffalo; and even hunt them on their own. Moderately-sized hunting groups generally have higher success rates than lone females and larger groups.[[#Packer|Packer]], p. 150, 153, 164–165. [155] => [156] => 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.[[#Schaller|Schaller]], p. 248. 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.[[#Schaller|Schaller]], pp. 233, 247–248 One study in 2018 recorded a lion running at a top speed of {{cvt|74.1|km/h}}.{{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. |name-list-style=amp |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=2020-03-05}} They take advantage of factors that reduce visibility; many kills take place near some form of cover or at night.[[#Schaller|Schaller]], p. 237. 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.[[#Schaller|Schaller]], p. 244, 263–267. It has a [[Bite force quotient|bite force]] of 1314.7 [[Newton (unit)|Newton]] at the canine tip and 2023.7 Newton at the carnassial notch.{{Cite journal |last1=Christiansen |first1=P. |last2=Wroe |first2=S. |date=2007 |title=Bite forces and evolutionary adaptations to feeding ecology in carnivores |journal=Ecology |volume=88 |issue=2 |pages=347–358 |doi=10.1890/0012-9658(2007)88[347:bfaeat]2.0.co;2|pmid=17479753 }} [157] => [158] => Lions typically consume prey at the location of the hunt but sometimes drag large prey into cover. 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. Large kills are shared more widely among pride members.[[#Schaller|Schaller]], p. 133. An adult lioness requires an average of about {{cvt|5|kg}} of meat per day while males require about {{cvt|7|kg}}.[[#Schaller|Schaller]], p. 276. Lions gorge themselves and eat up to {{cvt|30|kg}} in one session.{{Cite book |last=Guggisberg |first=C. A. W. |title=Simba: the life of the lion. |year=1961 |publisher=Howard Timmins |location=Cape Town}} 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.[[#Schaller|Schaller]], pp. 270–76. Lions defend their kills from scavengers such as vultures and hyenas. [159] => [160] => 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.[[#Schaller|Schaller]], p. 213–216. Most carrion on which both hyenas and lions feed upon are killed by hyenas rather than lions. Carrion is thought to provide a large part of lion diet.{{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}} [161] => [162] => ===Predatory competition=== [163] => {{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}} [164] => [165] => 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%.{{Cite journal |title=Prey preferences of the spotted hyaena (''Crocuta crocuta'') and degree of dietary overlap with the lion (''Panthera leo'') |url=http://www.zbs.bialowieza.pl/g2/pdf/1598.pdf |archive-url=https://web.archive.org/web/20110430024111/http://www.zbs.bialowieza.pl/g2/pdf/1598.pdf |archive-date=2011-04-30 |url-status=live |last=Hayward |first=M. W. |journal=Journal of Zoology |year=2006 |volume=270 |issue=4 |pages=606–614 |doi=10.1111/j.1469-7998.2006.00183.x}} 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.{{Cite book |first=H. |last=Kruuk |title=The Spotted Hyena: A Study of Predation and Social Behaviour |publisher=Echo Point Books & Media |year=2014|isbn=978-1626549050 |pages=128–138|edition=2nd}} 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.{{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}} 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.[[#Schaller|Schaller]], p. 272. 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.[[#Schaller|Schaller]], pp. 273–74. 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.{{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 }} When the lion population in Kenya's [[Masai Mara National Reserve]] declined, the spotted hyena population increased rapidly.{{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}} [166] => [167] => Lions tend to dominate [[cheetah]]s and leopards, steal their kills and kill their cubs and even adults when given the chance.{{Cite book|last1=Denis-Hoot|first1=C. |last2= Denis-Hoot|first2= M. |year=2003 |title=The Art of Being a Lion |publisher=Barnes & Noble |location=New York |isbn=9780760747674 |page=198}} Cheetahs often lose their kills to lions or other predators.{{Cite journal |last1=O'Brien |first1=S. J. |last2=Wildt |first2 =D. E. |last3=Bush |first3=M. |year=1986 |title=The Cheetah in Genetic Peril |journal=Scientific American |volume=254 |issue=5 |pages=68–76 |doi=10.1038/scientificamerican0586-84 |url=http://www.catsg.org/cheetah/05_library/5_3_publications/N_and_O/OBrien_et_al_1986_Cheetah_in_genetic_peril.pdf |archive-url=https://web.archive.org/web/20110513071205/http://www.catsg.org/cheetah/05_library/5_3_publications/N_and_O/OBrien_et_al_1986_Cheetah_in_genetic_peril.pdf |archive-date=2011-05-13 |url-status=live|bibcode=1986SciAm.254e..84O}} A study in the Serengeti ecosystem revealed that lions killed at least 17 of 125 cheetah cubs born between 1987 and 1990.{{cite journal |last1=Laurenson |first1=M. K. |title=High juvenile mortality in cheetahs (''Acinonyx jubatus'') and its consequences for maternal care |journal=Journal of Zoology |year=1994 |volume=234 |issue=3 |pages=387–408 |doi=10.1111/j.1469-7998.1994.tb04855.x |url=http://www.catsg.org/cheetah/05_library/5_3_publications/L/Laurenson_1994_Cheetah_cub_mortality_-_maternal_care.pdf |archive-url=https://web.archive.org/web/20171120042410/http://www.catsg.org/cheetah/05_library/5_3_publications/L/Laurenson_1994_Cheetah_cub_mortality_-_maternal_care.pdf |archive-date=2017-11-20 |url-status=live}} Cheetahs avoid their competitors by hunting at different times and habitats.{{cite journal|last1=Rostro-García |first1=S. |last2=Kamler |first2=J. F. |last3=Hunter |first3=L. T. B. |name-list-style=amp |year=2015 |title=To kill, stay or flee: the effects of lions and landscape factors on habitat and kill site selection of cheetahs in South Africa |journal=PLOS ONE |volume=10 |issue=2 |page=e0117743 |pmid=25693067 |doi=10.1371/journal.pone.0117743 |pmc=4333767|bibcode=2015PLoSO..1017743R |doi-access=free}} 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 effect leopard abundance.{{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 }} Leopards take refuge in trees, though lionesses occasionally attempt to climb up and retrieve their kills.[[#Schaller|Schaller]], p. 293. [168] => [169] => 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.{{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. |name-list-style=amp |year=1999 |journal=Oryx |volume=33 |pages=132–142 |doi=10.1046/j.1365-3008.1999.00052.x |issue=2 |doi-access=free}} However, there are a few reported cases of old and wounded lions falling prey to wild dogs.{{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}}[[#Schaller|Schaller]], p. 188. [170] => [171] => ===Reproduction and life cycle=== [172] => [173] => {{multiple image|align=right|direction=vertical [174] => |image1=Panthera leo massaica mating.jpg|caption1=Lions mating at Masai Mara [175] => |image2=Lion_cub,_Masai_Mara,_Kenya.jpg|caption2=A lion cub in Masai Mara}} [176] => [177] => Most lionesses reproduce by the time they are four years of age.[[#Schaller|Schaller]], p. 29. Lions do not mate at a specific time of year and the females are [[polyestrous]].[[#Schaller|Schaller]], p. 174. 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]].{{Cite journal | doi=10.1002/zoo.1430130403|title = Spontaneous and induced ovulation in the lion (Panthera leo)| journal=Zoo Biology| volume=13| issue=4| pages=301–307|year = 1994|last1 = Schramm|first1 = Ralph Dee| last2=Briggs| first2=Michael B.| last3=Reeves| first3=Jerry J.}}{{Cite book |last=Asdell |first=Sydney A. |title=Patterns of mammalian reproduction |year=1993 |orig-date=1964 |publisher=Cornell University Press |location=Ithaca |isbn=978-0-8014-1753-5 |url=https://archive.org/details/asdellspatternso00hays }} A lioness may mate with more than one male when she is [[Estrous cycle|in heat]].[[#Schaller|Schaller]], p. 142. Lions of both sexes may be involved in group [[Homosexuality in animals|homosexual]] and courtship activities. Males will also head-rub and roll around with each other before mounting each other.{{Cite book |last=Bagemihl |first=Bruce |year=1999 |title=Biological Exuberance: Animal Homosexuality and Natural Diversity |location=New York |publisher=St. Martin's Press |isbn=978-0-312-19239-6 |pages=[https://archive.org/details/biologicalexuber00bage/page/302 302–05] |url=https://archive.org/details/biologicalexuber00bage/page/302 }}[[#Schaller|Schaller]], p. 137. [[Generation time|Generation length]] of the lion is about seven years.{{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}} The average gestation period is around 110{{spaces}}days; 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.[[#Scott|Scott]], p. 45. 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.[[#Schaller|Schaller]], p. 143. 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. [178] => [179] => Usually, the mother does not integrate herself and her cubs back into the pride until the cubs are six to eight weeks old. Sometimes the introduction to pride life occurs earlier, particularly if other lionesses have given birth at about the same time. 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. 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.[[#Scott|Scott]], p. 46. [180] => [[File:Lion Cubs Phinda 2011.ogv|thumb|right|Video of a lioness and her cubs in Phinda Reserve]] [181] => [182] => 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.[[#Schaller|Schaller]], p. 147-49. 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.{{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}} [183] => [184] => 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.{{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=716–28 |doi= 10.1086/284097|s2cid=84927815 }} 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,[[#Schaller|Schaller]], p. 44. while some females may leave when they reach the age of two. When a new male lion takes over a pride, adolescents both male and female may be evicted.[[#Scott|Scott]], p. 68. [185] => [186] => ===Health and mortality=== [187] => [188] => [[File:Lake-Nakuru-Lions-in-Tree.JPG|thumb|Lions in a tree near [[Lake Nakuru]]]] [189] => [190] => Lions may live 12–17 years in the wild. Although adult lions have no natural predators, evidence suggests most die violently from attacks by humans or other lions.[[#Schaller|Schaller]], p. 183. 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.[[#Schaller|Schaller]], pp. 188–89. 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.[[#Schaller|Schaller]], pp. 189–90. [[Nile crocodile]]s may also kill and eat lions, evidenced by the occasional lion claw found in crocodile stomachs.{{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=}} [191] => [192] => [[Tick]]s commonly infest the ears, neck and groin regions of the lions.[[#Schaller|Schaller]], p. 184.{{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}} Adult forms of several [[tapeworm]] species of the genus ''[[Taenia (tapeworm)|Taenia]]'' have been isolated from lion intestines, having been ingested as larvae in [[antelope]] meat.{{Cite journal |last=Sachs |first=R. |year=1969|title=Untersuchungen zur Artbestimmung und Differenzierung der Muskelfinnen ostafrikanischer Wildtiere [Differentiation and species determination of muscle-cysticerci in East African game animals] |journal=Zeitschrift für Tropenmedizin und Parasitologie |volume=20 |issue=1 |pages=39–50 |pmid=5393325|language=de}} 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.{{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}} A more recent outbreak in 2001 killed six lions.{{Cite news | last=Nkwame | first=V. M. |title=King of the jungle in jeopardy | work=The Arusha Times| date=9 September 2006 | url=http://www.arushatimes.co.tz/2006/36/features_10.htm | access-date=4 September 2007 | url-status=dead | archive-url=https://web.archive.org/web/20070929044925/http://www.arushatimes.co.tz/2006/36/features_10.htm | archive-date=29 September 2007 | df=dmy-all}} [193] => [194] => Captive lions have been infected with [[canine distemper]] virus (CDV) since at least the mid-1970s.{{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=}} 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]].{{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 }} [[Feline immunodeficiency virus]] and [[lentivirus]] also affect captive lions.{{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 }}{{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 }} [195] => [196] => ===Communication=== [197] => [[File:Lion cub with mother.jpg|thumb|Head rubbing among pride members is a common social behaviour.]] [198] => [199] => 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]],[[#Schaller|Schaller]], p. 85. which have been compared with the role of [[allogrooming]] among primates.{{Cite book |last=Sparks |first=J. |title=Primate Ethology |chapter=Allogrooming in primates: a review |pages=148–175 |year=1967 |editor=Morris, D. |publisher=Aldine |location=Chicago |edition=2011 |isbn=9780202368160}} Head rubbing, nuzzling the forehead, face and neck against another lion appears to be a form of greeting{{Cite book |last=Leyhausen |first=P. |title=Verhaltensstudien an Katzen |edition=Second |year=1960 |publisher=Paul Parey |location=Berlin |isbn=978-3-489-71836-9 |language=de}} 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.[[#Schaller|Schaller]], pp. 85–88. 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.[[#Schaller|Schaller]], pp. 88–91. [200] => {{listen|filename=Lion raring-sound1TamilNadu178.ogg|title=Lion roar|description=A captive lion roaring}} [201] => [202] => Lions have an array of facial expressions and body postures that serve as visual gestures.[[#Schaller|Schaller]], pp. 103–117. 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.[[#Schaller|Schaller]], p. 95. Lions also use chemical and visual marking; males [[Territory (animal)#Scent marking|spray urine]]{{cite journal |author1=Andersen, K. F. |name-list-style=amp |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}} and scrape plots of ground and objects within the territory. [203] => [204] => The lion's repertoire of vocalisations is large; variations in intensity and pitch appear to be central to communication. Most lion vocalisations are variations of [[growling]], [[snarling]], meowing and roaring. Other sounds produced include purring, puffing, bleating and humming. Roaring is used to advertise its presence. Lions most often roar at night, a sound that can be heard from a distance of {{convert|8|km|0}}.[[#Schaller|Schaller]], pp. 103–113. They tend to roar in a very characteristic manner starting with a few deep, long roars that subside into grunts.{{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=2011-10-01 |url-status=live}}{{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=2011-10-01 |url-status=live}} [205] => [206] => ==Conservation== [207] => 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]]. [208] => [209] => ===In Africa=== [210] => [[File:Lion.ogv|thumb|right|Video of a wild lioness]] [211] => [212] => 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. 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. In 1975, it was estimated that since the 1950s, lion numbers had decreased by half to 200,000 or fewer.{{cite journal |last1=Myers |first1=N. |year=1975 |title=The silent savannahs |journal=International Wildlife |volume=5 |issue=5 |pages=5–10 }} Estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004.{{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}}{{Cite book |last=Chardonnet|first=P. |year=2002|title=Conservation of African lion |url=http://conservationforce.org/pdf/conservationoftheafricanlion.pdf |publisher=International Foundation for the Conservation of Wildlife|location=Paris, France |url-status=dead |archive-date=10 November 2013 |archive-url=https://web.archive.org/web/20131110184540/http://conservationforce.org/pdf/conservationoftheafricanlion.pdf}} [213] => [214] => 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.{{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 |page=882−892 |doi=10.1644/13-mamm-a-306 |doi-access=free }}{{open access}} 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]]. 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.{{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 |pages=110 |doi=10.3389/fevo.2016.00110 |doi-access=free}} [215] => [216] => A population occurs in Cameroon's [[Waza National Park]], where between approximately 14 and 21 animals persisted as of 2009.{{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 }} In addition, 50 to 150 lions are estimated to be present in Burkina Faso's [[Arly-Singou]] ecosystem.{{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 }} 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.{{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 |url-status=dead }} 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.{{cite magazine|magazine=New Scientist|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}}{{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}} [217] => [218] => In 2005, Lion Conservation Strategies were developed for West and Central Africa, and or East and Southern Africa. The strategies seek to maintain suitable habitat, ensure a sufficient wild prey base for lions, reduce factors that lead to further fragmentation of populations, and make lion–human coexistence sustainable.{{cite book |title=Conservation Strategy for the Lion West and Central Africa |publisher=IUCN |author=IUCN Cat Specialist Group |year=2006 |location=Yaounde, Cameroon |url=http://www.catsg.org/fileadmin/filesharing/3.Conservation_Center/3.4._Strategies___Action_Plans/African_lion/IUCN_CatSG_2006_West_and_Central_Africa_Lion_Conservation_Strategy.pdf |archive-url=https://web.archive.org/web/20150919114530/http://www.catsg.org/fileadmin/filesharing/3.Conservation_Center/3.4._Strategies___Action_Plans/African_lion/IUCN_CatSG_2006_West_and_Central_Africa_Lion_Conservation_Strategy.pdf |archive-date=2015-09-19 |url-status=live}}{{cite book |author=IUCN Cat Specialist Group |year=2006 |title=Conservation Strategy for the Lion ''Panthera leo'' in Eastern and Southern Africa |publisher=IUCN |location=Pretoria, South Africa |url=http://www.catsg.org/fileadmin/filesharing/3.Conservation_Center/3.4._Strategies___Action_Plans/African_lion/IUCN_CatSG_2006_East_and_South_Africa_Lion_Conservation_Strategy.pdf |archive-url=https://web.archive.org/web/20150619001314/http://www.catsg.org/fileadmin/filesharing/3.Conservation_Center/3.4._Strategies___Action_Plans/African_lion/IUCN_CatSG_2006_East_and_South_Africa_Lion_Conservation_Strategy.pdf |archive-date=2015-06-19 |url-status=live}} Lion depredation on livestock is significantly reduced in areas where herders keep livestock in improved enclosures. Such measures contribute to mitigating [[human–lion conflict]].{{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 }} [219] => [220] => ===In Asia=== [221] => [[File:Lion Gir.jpg|thumb|A lioness in Gir National Park]] [222] => 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.{{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}} 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]]. By 2015, the population had grown to 523 individuals inhabiting an area of {{cvt|7000|km2}} in Saurashtra.{{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 |page=32−40}}{{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}}{{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}} In 2017, about 650 individuals were recorded during the Asiatic Lion Census.{{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}} [223] => [224] => The presence of numerous human settlements close to Gir National Park resulted in conflict between lions, local people and their livestock.{{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}} Some consider the presence of lions a benefit, as they keep populations of crop damaging herbivores in check.{{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}} [225] => [226] => ===Captive breeding=== [227] => [[File:India Animals.jpg|thumb|Two captive male Asiatic lions in [[Sanjay Gandhi National Park]], India]] [228] => [229] => 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.{{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 |url-status=dead |doi=10.1163/18759866-07602002 |s2cid=2131247 }} 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.{{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}} 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.{{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 }} [230] => [231] => 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.{{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 |url-status=dead }} Breeding programs need to note origins of the participating animals to avoid cross-breeding different subspecies and thus reducing their conservation value.{{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}} 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.{{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}} [232] => [233] => 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.{{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 |url-status=dead |archive-date=2006-08-24 }} [234] => [235] => ==Interactions with humans== [236] => ===In zoos and circuses=== [237] => {{multiple image|align=right|direction=vertical [238] => |image1=Lion - melbourne zoo.jpg|caption1=Lion at [[Melbourne Zoo]] [239] => |image2=Lion tamer (LOC pga.03749).jpg|caption2=19th-century [[etching]] of a lion tamer in a cage with lions and tigers}} [240] => 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,[[#Courcy|de Courcy]], p. 81-82. 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.{{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 |url-status=dead }} 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.{{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|url-status=dead}}{{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|url-status=dead}} [241] => [242] => 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.[[#Baratay|Baratay & Hardouin-Fugier]], pp. 19–21, 42. 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;[[#Baratay|Baratay & Hardouin-Fugier]], p. 20.{{cite magazine |last=Owen |first=J. |title=Medieval Lion Skulls Reveal Secrets of Tower of London "Zoo"|magazine=National Geographic Magazine |date=2005 |url=http://news.nationalgeographic.com/news/2005/11/1103_051103_tower_lions.html|archive-url=https://web.archive.org/web/20051105025912/http://news.nationalgeographic.com/news/2005/11/1103_051103_tower_lions.html|url-status=dead|archive-date=5 November 2005| access-date=5 September 2007}} 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.[[#Blunt|Blunt]], p. 15. [243] => [244] => Lions were kept in cramped and squalid conditions at [[London Zoo]] until a larger lion house with roomier cages was built in the 1870s.[[#Blunt|Blunt]], p. 208. 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.[[#Courcy|de Courcy]], p. 69. 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.{{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}} [245] => [246] => [[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. Van Amburgh performed before [[Queen Victoria]] in 1838 when he toured Great Britain. 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.[[#Baratay|Baratay & Hardouin-Fugier]], p. 187. 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).{{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}} [247] => [248] => ===Hunting and games=== [249] => {{Main|Lion hunting}} [250] => {{See also|Lion baiting}} [251] => [[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}}]] [252] => Lion hunting has occurred since ancient times and was often a royal tradition, intended to demonstrate the power of the king over nature. Such hunts took place in a reserved area in front of an audience. The monarch was accompanied by his men and controls were put in place to increase their safety and ease of killing. The earliest surviving record of lion hunting is an [[ancient Egypt]]ian inscription dated circa 1380 BC that mentions Pharaoh [[Amenhotep III]] killing 102 lions in ten years "with his own arrows". The [[Assyria]]n emperor [[Ashurbanipal]] had one of his lion hunts depicted on a sequence of [[Assyrian palace reliefs]] {{circa|640 BC}}, known as the [[Lion Hunt of Ashurbanipal]]. Lions were also hunted during the [[Mughal Empire]], where Emperor [[Jahangir]] is said to have excelled at it.[[#Jackson|Jackson]], p. 156–159. In [[Ancient Rome]], lions were kept by emperors for [[Venatio|hunts]], [[Bestiarii|gladiator fights]] and [[Damnatio ad bestias|executions]].[[#Jackson|Jackson]], p. 142. [253] => [254] => 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.{{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 }} 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.[[#Jackson|Jackson]], p. 166. The widely reproduced imagery of the heroic hunter chasing lions would dominate a large part of the century.[[#Baratay|Baratay & Hardouin-Fugier]], p. 113. [[Trophy hunting]] of lions in recent years has been met with controversy, notably with the [[killing of Cecil the lion]] in mid-2015.{{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}} [255] => [256] => ===Man-eating=== [257] => [[File:Lionsoftsavo2008.jpg|thumb|The Tsavo maneaters of East Africa on display in the [[Field Museum of Natural History]] in Chicago]] [258] => {{Further information|Man-eater#Lions}} [259] => Lions do not usually hunt humans but some (usually males) seem to seek them out. One well-publicised case is the [[Tsavo maneaters]]; in 1898, 28 officially recorded railway workers building the [[Uganda Railway]] were taken by lions over nine months during the construction of a bridge in Kenya.{{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}} 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.{{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}} Sick or injured animals may be more prone to man-eating but the behaviour is not unusual, nor necessarily aberrant.{{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}} [260] => [261] => Lions' proclivity for [[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.{{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=927–28 |doi=10.1038/436927a| pmid=16107828 |bibcode=2005Natur.436..927P |s2cid=3190757}} 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.{{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. |pages=e22285 |pmid=21799812 |pmc=3140494|bibcode=2011PLoSO...622285P|doi-access=free}} [262] => [263] => According to Robert R. Frump, Mozambican refugees regularly crossing Kruger National Park, South Africa, at night are attacked and eaten by lions; park officials have said man-eating is a problem there. Frump said thousands may have been killed in the decades after [[apartheid]] sealed the park and forced refugees to cross the park at night. For nearly a century before the border was sealed, Mozambicans had regularly crossed the park in daytime with little harm.{{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}} [264] => [265] => ===Cultural significance=== [266] => {{main|Cultural depictions of lions}} [267] => [[File:Luxor Sekhmet New Kingdom.JPG|thumb|right|upright|Granite statue of the Egyptian goddess [[Sekhmet]] from the [[Luxor Temple]], dated 1403–1365 BC, exhibited in the [[National Museum of Denmark]]]] [268] => 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. It is considered to be the monarch of the beasts; symbolizing power, royalty and protection.{{cite book | first=Hope B. | last=Werness |year=2007 |title=The Continuum Encyclopedia of Animal Symbolism in World Art |publisher=Continuum International Publishing Group |pages=254–560|isbn=978-0826419132}} Several leaders have had "lion" in their name including [[Sundiata Keita]] of the [[Mali Empire]], who was called "Lion of Mali", and [[Richard the Lionheart]] of England.[[#Jackson|Jackson]], p. 133. The male's mane makes it a particularly recognisable feature and thus has been represented more than the female.[[#Jackson|Jackson]], p. 100. Nevertheless, the lioness has also had importance as a guardian. [269] => [270] => In sub-Saharan Africa, the lion has been a common character in stories, proverbs and dances, but rarely featured in visual arts.[[#Jackson|Jackson]], p. 119. In the [[Swahili language]], the lion is known as ''simba'' which also means "aggressive", "king" and "strong". 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.{{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}} In much of [[African folklore]], the lion is portrayed as having low intelligence and is easily tricked by other animals.{{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}} In [[Ancient Egypt]], lions were linked both with the sun and the waters of the Nile. Several gods were conceived as being partially lion including the war deities [[Sekhmet]] and [[Maahes]], and [[Tefnut]] the goddess of moisture. [[Shu (Egyptian god)|Shu]], god of the air, was depicted kneeing between two lions while holding a headrest. The lions mark where the sun rises and sets and symbolise yesterday and tomorrow. In [[Nubia]], The lion-god [[Apedemak]] was associated with the flooding of the Nile.[[#Jackson|Jackson]], pp. 107–108, 111. [271] => [[File:Berlín Ishtar león. 02.JPG|thumb|left|Roaring and striding lion from the Throne Room of [[Nebuchadnezzar II]], 6th century BC, from [[Babylon]], Iraq]] [272] => 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.{{Cite journal |last=Cassin |first=Elena |author-link=Elena Cassin |year=1981 |title= Le roi et le lion |journal=Revue de l'Histoire des Religions | trans-title = The king & the lion | volume=298 |issue=198–4 |pages=355–401 |language=fr|doi=10.3406/rhr.1981.4828}} The big cat was a symbol and steed of fertility goddess [[Inanna]]. Lions decorate the [[Ishtar_Gate#Ishtar_Gate_and_Processional_Way|Processional Way leading to the Ishtar Gate]] in Babylon which was built by the [[Nebuchadnezzar II]] in the 6th century BCE. The [[Lion of Babylon]] symbolized the power of the king and protector of the land against enemies, but was also invoked for good luck.{{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}} The constellation [[Leo (constellation)|Leo the lion]] was first recognised by the Sumerians around 4,000 years ago and is considered to be the fifth sign of the [[zodiac]].[[#Jackson|Jackson]], p. 109. Lions are frequently mentioned in the [[Bible]], notably in the [[Book of Daniel]], in which the [[Daniel (biblical figure)|eponymous hero]] refuses to worship [[Darius the Mede|King Darius]] and is forced to sleep in [[Daniel in the lions' den|the lions' den]] where he is miraculously unharmed ({{bibleref2|Dan|6}}). In the [[Book of Judges]], [[Samson]] kills a lion as he travels to visit a Philistine woman.({{bibleref2|Judg|14}}). The power and ferocity of the lion is invoked when describing the anger of God ({{bibleref2|Amos|3:4–8}}, {{bibleref2|Lam|3:10}}) and the menace of Israel's enemies ({{bibleref2|Psalm|17:12|22:21}}, {{bibleref2|Jer|2:30|5:6}}) and Satan ({{bibleref2|1 Pet|5:8}}). The book of Isaiah uses the imagery of a lion laying with a calf and child, and eating straw to portray the harmony of creation ({{bibleref2|Isa|11:6–7}}). In the [[Book of Revelation]], a lion, an ox, a man and an eagle are seen on a heavenly throne in [[John of Patmos|John]]'s vision;({{bibleref2|Rev|4:7}}).{{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=978-0687333653}} [273] => [[File:Cowardly lion2.jpg|thumb|upright|[[Dorothy Gale]] meets the Cowardly Lion in ''[[The Wonderful Wizard of Oz]]''. Art by [[W. W. Denslow]], 1900.]] [274] => 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 in general or deer in particular.{{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 }} In India, the [[Lion Capital of Ashoka]], erected by Emperor Ashoka in the 3rd century CE, 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]]. 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 in China and beyond for over a thousand years. In Buddhist art, lions are associated with both ''[[arhat]]s'' and [[bodhisattva]]s and may be ridden by the [[Manjushri]].[[#Jackson|Jackson]], pp. 113, 119–122, 124. [275] => [276] => 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 wears its skin. [[Lancelot]] and [[Gawain]] were also heroes slaying lions in [[Middle Ages|medieval Europe]]. In some medieval stories, lions were portrayed as allies and companions. 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]]''. The lion's role as king of the beasts has been used in the 1994 [[Disney animated feature]] film ''[[The Lion King]]''.[[#Jackson|Jackson]], pp. 7, 96, 99, 103–105, 128, 135, 150, 197. [277] => [278] => ==See also== [279] => {{div col|colwidth=15em}} [280] => * [[List of largest cats]] [281] => * [[Mapogo lion coalition]] [282] => * [[Roar (film)]] from 1981 [283] => {{div col end}} [284] => [285] => == Notes == [286] => {{Notelist}} [287] => [288] => ==References== [289] => [290] => [291] => === Citations === [292] => {{Reflist|23em}} [293] => [294] => === Books === [295] => {{Refbegin}} [296] => * {{Cite book |last1=Baratay |first1=E. |first2=E. |last2=Hardouin-Fugier |name-list-style=amp |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}} [297] => * {{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}} [298] => * {{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}} [299] => * {{cite book|last=Jackson| first= D. |year=2010|title=Lion |url=https://books.google.com/books?id=65lK7UIVRfIC |location=London |publisher=Reaktion Books |isbn=978-1861896551 |ref=Jackson}} [300] => * {{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=9780691215297|url=https://books.google.com/books?id=TYOREAAAQBAJ&q=The+Lion:+Behavior,+Ecology,+and+Conservation+of+an+Iconic+Species|ref=Packer}} [301] => * {{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}} [302] => * {{Cite book |last1=Scott |first1=J. |last2=Scott |first2=A. |year=2002 |title=Big Cat Diary: Lion |location=New York |publisher=Harper Collins |isbn=9780007146666 |ref=Scott}} [303] => {{Refend}} [304] => [305] => ==External links== [306] => {{Wiktionary}} [307] => {{Wikispecies |Panthera leo|Lion}} [308] => {{EB1911 poster|Lion}} [309] => *{{Commons-inline|Panthera leo|Lion}} [310] => * {{cite web |url=http://www.catsg.org/index.php?id=108 | title=Lion ''Panthera leo'' |author=IUCN/SSC Cat Specialist Group}} [311] => * {{cite web |url=http://www.lionconservationfund.org/ | title=Lion Conservation Fund}} [312] => * {{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 |author=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 |url-status=dead }} [313] => * {{Cite Collier's|wstitle=Lion|short=x}} [314] => [315] => {{Carnivora |Fe.}} [316] => {{Portal bar|Cats|Mammals|Animals|Biology|Africa|India}} [317] => {{Taxonbar|from1=Q140|from2=Q15294488}} [318] => {{Authority control}} [319] => [320] => [[Category:Lions| ]] [321] => [[Category:Apex predators]] [322] => [[Category:Articles containing video clips]] [323] => [[Category:Big cats]] [324] => [[Category:Extant Pliocene first appearances]] [325] => [[Category:Felids of Africa]] [326] => [[Category:Mammals described in 1758]] [327] => [[Category:Mammals of Sub-Saharan Africa]] [328] => [[Category:Mammals of South Asia]] [329] => [[Category:National symbols of Burundi]] [330] => [[Category:National symbols of Chad]] [331] => [[Category:National symbols of Eswatini]] [332] => [[Category:National symbols of Kenya]] [333] => [[Category:National symbols of Malawi]] [334] => [[Category:National symbols of Morocco]] [335] => [[Category:National symbols of Sierra Leone]] [336] => [[Category:National symbols of Singapore]] [337] => [[Category:National symbols of South Africa]] [338] => [[Category:National symbols of Sri Lanka]] [339] => [[Category:National symbols of the Republic of the Congo]] [340] => [[Category:National symbols of Togo]] [341] => [[Category:National symbols of Tunisia]] [342] => [[Category:Panthera]] [343] => [[Category:Taxa named by Carl Linnaeus]] [344] => [[Category:Vulnerable animals]] [345] => [[Category:Vulnerable biota of Africa]] [] => )
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Lion

The Wikipedia page for "Lion" provides a comprehensive overview of this large felid species known for its majestic appearance and fierce behavior. Lions are native to parts of Africa and Asia, with the majority of the population found in sub-Saharan Africa.

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Lions are native to parts of Africa and Asia, with the majority of the population found in sub-Saharan Africa. The article delves into various aspects of lion biology, including their physical characteristics, habitat, and social structure. The page highlights the distinguishing features of lions, such as their muscular build, tawny-colored fur, and iconic manes in males. It also covers aspects of their lifestyle, including their diet, hunting strategies, and their ability to roar – a unique vocalization only present in this species. One significant aspect of lion behavior is their social organization. Lions live in prides, which are family groups consisting of multiple related adult females, their offspring, and a few adult males. The article explains the roles and dynamics within a pride, including the territorial behavior, mating system, and cooperative hunting. The conservation status of lions is another important topic covered in the article, emphasizing the increasing threats faced by the species due to habitat loss, human-wildlife conflict, and poaching. The page discusses various conservation efforts and initiatives, as well as the role of captive breeding programs and protected areas in preserving lion populations. The page also provides a historical perspective on the interaction between humans and lions, covering ancient symbolism and cultural significance of lions in different civilizations. Additionally, it mentions notable lion conservationists and famous lions in popular culture. Overall, the Wikipedia page on lions offers a comprehensive understanding of these iconic creatures, encompassing their biology, behavior, conservation, and cultural significance. It serves as a valuable resource for individuals seeking knowledge on one of the most recognizable and admired animals on the planet.

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