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| {{Use dmy dates|date=July 2016}}
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| {{Speciesbox
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| | taxon = Panthera leo
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| | name = Lion<ref name=MSW3>{{MSW3 Carnivora | id = 14000228 | page = }}</ref>
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| | fossil_range = {{fossil range|early Pleistocene|Recent}}
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| | status = VU
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| | status_system = iucn3.1
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| | status_ref = <ref name=IUCN>{{IUCN|assessor=Bauer, H.|assessor2=Packer, C.|assessor3=Funston, P.F.|assessor4=Henschel, P.|assessor5=Nowell, K. |year=2016|id=15951|taxon=Panthera leo|version=2016.2}}</ref>
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| | image = Lion waiting in Namibia.jpg
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| | image_caption = Male
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| | image2 = Okonjima Lioness.jpg
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| | image2_caption = Female (lioness)
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| | authority=([[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]])<ref name="Linn1758" />
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| | synonyms=<center>''Felis leo''<br /><small>Linnaeus, 1758</small><ref name="Linn1758"/></center>
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| | range_map = Lion distribution.png
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| | range_map2 = Map Guj Nat Parks Sanctuary.png
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| | range_map2_caption = Distribution of lions in India: The Gir Forest, in [[Gujarat]], is the last natural range of more than 500 wild [[Asiatic lion]]s. There are plans to reintroduce some lions to [[Kuno Wildlife Sanctuary]] in neighbouring [[Madhya Pradesh]].
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| | subdivision_ranks = [[Subspecies]]
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| | subdivision =
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| {{show
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| |''Panthera leo'' subspecies
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| ''[[Congo lion|P. l. azandica]]'' <br />
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| ''[[Southwest African lion|P. l. bleyenberghi]]'' <br />
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| ''P. l. hollisteri'' <br />
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| ''P. l. kamptzi'' <br />
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| ''P. l. krugeri'' <br />
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| ''[[Barbary lion|P. l. leo]]'' <br />
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| ''[[Marozi|P. l. maculatus]]'' <br />
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| ''P. l. nubica'' <br />
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| ''[[Asiatic lion|P. l. persica]]'' <br />
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| ''P. l. senegalensis'' <br />
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| ''P. l. verneyi'' <br />
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| ''P. l. nyanzae'' <br />
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| ''P. l. roosevelti'' <br />
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| ''P. l. somaliensis'' <br />
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| {{dagger}}''[[American lion|P. l. atrox]]'' <br />
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| {{dagger}}''[[History of lions in Europe|P. l. europaea]]'' <br />
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| {{dagger}}''[[Cape lion|P. l. melanochaita]]'' <br />
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| {{dagger}}''[[Sri Lanka lion|P. l. sinhaleyus]]'' <br />
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| {{dagger}}''[[Panthera leo spelaea|P. l. spelaea]]'' <br />
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| {{dagger}}''P. l. vereshchagini''}}
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| }}
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| The '''lion''' (''Panthera leo'') is one of the big cats in the [[genus]] ''[[Panthera]]'' and a member of the family [[Felidae]]. The commonly used term '''[[Mouse|African lion]]''' collectively denotes the several subspecies in [[Nigger|Africa.]] With some males exceeding {{convert|250|kg|abbr=on}} in weight,<ref name="nowak">{{Cite book |last=Nowak|first= Ronald M. |year=1999 |title=Walker's Mammals of the World |location=Baltimore |publisher=Johns Hopkins University Press |isbn=0-8018-5789-9}}</ref> it is the second-largest living cat after the [[tiger]]. Wild lions currently exist in [[sub-Saharan Africa]] and in [[India]] (where an [[endangered]] remnant population resides in [[Gir Forest National Park]]). In [[ancient history|ancient historic]] times, their range was in most of Africa, including North Africa, and across Eurasia from Greece and southeastern Europe to India. In the late [[Pleistocene]], about 10,000 years ago, the lion was the most widespread large land mammal after humans: ''[[Panthera leo spelaea]]'' lived in northern and western Europe and ''[[Panthera leo atrox]]'' lived in the Americas from the [[Yukon]] to [[Peru]].<ref name=Harington69/> The lion is classified as a [[vulnerable species]] by the [[International Union for Conservation of Nature]] (IUCN), having seen a major population decline in its African range of 30–50% per two decades during the second half of the twentieth century.<ref name=IUCN/> Lion populations are untenable outside designated reserves and national parks. Although the cause of the decline is not fully understood, habitat loss and conflicts with humans are the greatest causes of concern. Within Africa, the [[West African lion]] population is particularly endangered.
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| In the wild, males seldom live longer than 10 to 14 years, as injuries sustained from continual fighting with rival males greatly reduce their longevity.<ref>{{Cite book |last=Smuts |first=G. L. |year=1982 |title=Lion|publisher=Macmillan South Africa |location=Johannesburg |page=231|isbn= 0-86954-122-6}}</ref> In captivity they can live more than 20 years. They typically inhabit [[savanna]] and [[grassland]], although they may take to [[Shrub|bush]] and forest. Lions are unusually [[social animal|social]] compared to other cats. A pride of lions consists of related females and offspring and a small number of adult males. Groups of female lions typically hunt together, preying mostly on large [[ungulate]]s. Lions are [[apex predator|apex]] and [[keystone species#Predators|keystone predators]], although they are also expert [[scavengers]] obtaining over 50 percent of their food by scavenging as opportunity allows. While lions do not typically hunt humans, some have. Sleeping mainly during the day, lions are active primarily at night ([[nocturnality|nocturnal]]), although sometimes at twilight ([[crepuscular]]).<ref name="earthtouch">[http://web.archive.org/web/20120420172909/http://www.earth-touch.com/result.php?i=Lions-nocturnal-chorus Lions' nocturnal chorus]. Earth-touch.com. Retrieved on 31 July 2013.</ref><ref name="phxzoo">[http://web.archive.org/web/20120415054339/http://www.phoenixzoo.org/learn/animals/animal_detail.aspx?FACT_SHEET_ID=100001 African Lion ''Panthera leo'']. Phoenix Zoo Fact Sheet.</ref>
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| Highly distinctive, the male lion is easily recognised by its mane, and its face is one of the most widely recognised animal symbols in human culture. [[Cultural depictions of lions|Depictions]] have existed from the [[Upper Paleolithic]] period, with carvings and paintings from the [[Lascaux]] and [[Chauvet Cave]]s in France dated to 17,000 years ago, through virtually all ancient and medieval cultures where they once occurred. It has been extensively depicted in sculptures, in paintings, on national flags, and in contemporary films and literature. Lions have been kept in [[menagerie]]s since the time of the [[Roman Empire]], and have been a key species sought for exhibition in [[zoo]]s over the world since the late eighteenth century. Zoos are cooperating worldwide in breeding programs for the endangered [[Asiatic lion|Asiatic subspecies]].
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| ==Etymology==
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| The lion's name, similar in many [[Romance languages]], is derived from the [[Latin]] ''{{lang|la|leo}}'',<ref>{{Cite book | last=Simpson |first=D. P. | title=Cassell's Latin Dictionary | publisher=Cassell Ltd. | year=1979 | edition=5th | location=London | page=342 | isbn=0-304-52257-0}}</ref> and the [[Ancient Greek]] {{lang|grc|λέων}} (''{{lang|grc-Latn|leon}}'').<ref name="Liddell 1980">{{Cite book | authorlink=Henry George Liddell |last=Liddell |first=Henry George |authorlink2=Robert Scott (philologist) |last2=Scott |first2=Robert | year=1980 | title=[[A Greek-English Lexicon]] (Abridged Edition)|page=411 | publisher=[[Oxford University Press]] | location=United Kingdom | isbn=0-19-910207-4}}</ref> The [[Hebrew language|Hebrew]] word {{lang|he|לָבִיא}} ({{lang|he-Latn|''lavi''}}) may also be related.<ref>{{Cite encyclopedia | title=Lion|encyclopedia=Oxford English Dictionary|editor1=Simpson, John |editor2=Weiner, Edmund | year=1989 |edition= 2nd| location=Oxford |publisher=Clarendon Press| isbn=0-19-861186-2}}</ref> It was one of the species originally described by [[Carl Linnaeus|Linnaeus]], who gave it the name ''Felis leo'', in his eighteenth-century work, ''[[Systema Naturae]]''.<ref name="Linn1758">{{Cite book |last=Linnaeus |first=Carolus |authorlink=Carl Linnaeus |title=Systema naturae per regna tria naturae :secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. |publisher=Holmiae (Laurentii Salvii) |year=1758 |page=41 |url=http://www.biodiversitylibrary.org/page/726936 |accessdate=8 September 2008 |language=Latin |volume=1 |edition=10th}}</ref>
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| ==Taxonomy and evolution==
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| [[File:Two cladograms for Panthera (including Neofelis).png|thumbnail|left|Two cladograms proposed for ''[[Panthera]]''. The upper cladogram is based on the 2006 and 2009 studies, while the other is based on the 2010 and 2011 studies.]]
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| The lion's closest relatives are the other species of the genus ''[[Panthera]]'': the tiger, the [[snow leopard]], the [[jaguar]], and the [[leopard]]. Studies from 2006 and 2009 concluded that the jaguar is a [[Sister group|sister species]] to the lion and the leopard is a sister taxon to the jaguar/lion clade<ref name="johnson2006">{{cite journal|last1=Johnson|first1=W.E.|last2=Eizirik|first2=E.|last3=Pecon-Slattery|first3=J.|last4=Murphy|first4=W.J.|last5=Antunes|first5=A.|last6=Teeling|first6=E.|last7=O'Brien|first7=S.J.|title=The late Miocene radiation of modern Felidae: a genetic assessment|journal=Science (New York)|year=2006|volume=311|issue=5757|pages=73–7|pmid=16400146|doi=10.1126/science.1122277}}</ref><ref name="werdelin2009">{{cite journal|last1=Werdelin|first1=L.|last2=Yamaguchi|first2=N.|last3=Johnson|first3=W.E.|last4=O'Brien|first4=S.J. |title=Phylogeny and evolution of cats (Felidae)|journal=Biology and Conservation of Wild Felids |year=2010|pages=59–82|url=https://www.researchgate.net/profile/Nobuyuki_Yamaguchi3/publication/266755142_Phylogeny_and_evolution_of_cats_Felidae/links/543ba0710cf2d6698be30c5e.pdf}}</ref> while 2010 and 2011 studies have swapped the positions leopard and jaguar.<ref name="davis2010">{{cite journal|last1=Davis|first1=B.W.|last2=Li|first2=G.|last3=Murphy|first3=W.J.|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|url=http://web.csulb.edu/~acarter3/course-evolution/files/cat-phylogeny.pdf}}</ref><ref name="mazak2011">{{cite journal|last1=Mazák|first1=J.H.|last2=Christiansen|first2=P.|last3=Kitchener|first3=A.C.|last4=Goswami|first4=A.|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}}</ref> ''P. leo'' evolved in Africa between 1 million and 800,000 years ago, before spreading throughout the [[Holarctic]] region.<ref>{{Cite journal |last1= Yamaguchi |first1=Nobuyuki| last2=Cooper |first2=Alan |last3=Werdelin |first3= Lars |last4= MacDonald |first4=David W.|date=August 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=329–42 |doi=10.1017/S0952836904005242 }}</ref> It appeared in the fossil record in Europe for the first time 700,000 years ago with the subspecies ''[[Panthera leo fossilis]]'' at [[Isernia]] in Italy. From this lion derived the later cave lion (''[[Panthera leo spelaea]]''),<ref>{{Cite book |last=Turner |first=Allen |year=1997 |title=The big cats and their fossil relatives: an illustrated guide to their evolution and natural history |location=New York |publisher=Columbia University Press |isbn=0-231-10229-1 |pages=}}</ref> which appeared about 300,000 years ago. Lions died out in northern [[Eurasia]] at the end of the last [[glacier|glaciation]], about 10,000 years ago; this may have been secondary to the [[Pleistocene extinctions|extinction]] of [[Pleistocene megafauna]].<ref>{{cite web | last = Harington | first = C. R. (Dick) | title = American Lion | work = Yukon Beringia Interpretive Centre website | publisher = Yukon Beringia Interpretive Centre | year = 1996 | url =http://www.beringia.com/research/lion.html | accessdate =25 January 2011}}</ref><ref name="BurgerJ-Molecular-phylogeny"/>
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| ===Subspecies===
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| [[File:Panthera leo subspecies late 20th century.png|thumb|Range map of the commonly accepted subspecies of the lion in the late twentieth century]]
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| Traditionally, 12 recent [[subspecies]] of lion were recognised, distinguished by mane appearance, size, and distribution. Because these characteristics are very insignificant and show a high individual variability, most of these forms were probably not true subspecies, especially as they were often based upon zoo material of unknown origin that may have had "striking, but abnormal" morphological characteristics.<ref name = "zoos_encyclopedia">{{cite encyclopedia | last = Grisham | first= Jack | editor-first = Catherine E | editor-last = Bell |title=Lion |encyclopedia=Encyclopedia of the World's Zoos |volume=2: G–P |year=2001 | publisher = Fitzroy Dearborn |location=Chofago |isbn=1-57958-174-9 |pages=733–39}}</ref> Today, only eight subspecies are usually accepted,<ref name="BurgerJ-Molecular-phylogeny">{{Cite journal | last1 = Burger | first1 = Joachim |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=841–49 |doi = 10.1016/j.ympev.2003.07.020 | url= http://www.uni-mainz.de/FB/Biologie/Anthropologie/MolA/Download/Burger%202004.pdf | type = fulltext | accessdate= 20 September 2007 | last3 = Loreille | first3 = Odile | last4 = Hemmer | first4 = Helmut | last5 = Eriksson | first5 = Torsten | last6 = Götherström | first6 = Anders | last7 = Hiller | first7 = Jennifer | last8 = Collins | first8 = Matthew J. | last9 = Wess | first9 = Timothy | last2 = Rosendahl | first2 = Wilfried | last10 = Alt | first10 = Kurt W.}}</ref><ref name = "CG" /> although one of these, the [[Cape lion]], formerly described as ''Panthera leo melanochaita'', is probably invalid.<ref name="CG" /> Even the remaining seven subspecies might be too many. While the status of the Asiatic lion (''P. l. persica'') as a subspecies is generally accepted, the systematic relationships among African lions are still not completely resolved. [[Mitochondria]]l variation in living African lions seemed to be modest according to some newer studies; therefore, all sub-Saharan lions have sometimes been considered a single subspecies. However, a recent study revealed lions from western and central Africa differ genetically from lions of southern or eastern Africa. According to this study, [[Western African lion]]s are more closely related to Asian lions than to South or East African lions. These findings might be explained by a late Pleistocene extinction event of lions in western and central Africa, and a subsequent recolonisation of these parts from Asia.<ref>{{Cite journal | last1 = Bertola | 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 = 1356–67 | doi = 10.1111/j.1365-2699.2011.02500.x | first1 = L. D. | 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. | last2 = Van Hooft | first2 = W. F. | 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. }}</ref>
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| Previous studies, which were focused mainly on lions from eastern and southern parts of Africa, already showed these can be possibly divided in two main [[clade]]s: one to the west of the [[East African Rift|Great Rift Valley]] and the other to the east. Lions from [[Tsavo]] in eastern Kenya are much closer genetically to lions in [[South African Republic|Transvaal]] (South Africa), than to those in the [[Aberdare Range]] in western Kenya.<ref name = "Dubach et al., 2005">{{Cite journal |last=Dubach | first =Jean|date=January 2005 |title = Molecular genetic variation across the southern and eastern geographic ranges of the African lion, ''Panthera leo'' |journal= Conservation Genetics | volume =6 |issue=1 |pages=15–24 |doi = 10.1007/s10592-004-7729-6|display-authors=etal}}</ref> Another study revealed there are three major types of lions, one North African–Asian, one southern African and one middle African.<ref>{{Cite journal|last1=Barnett |first1=Ross |last2=Yamaguchi |first2=Nobuyuki |last3=Barnes |first3=Ian |last4=Cooper |first4=Alan |year=2006 |title=The origin, current diversity and future conservation of the modern lion (''Panthera leo'') |journal=Proceedings of the Royal Society B: Biological Sciences |pmid=16901830 |volume=273 |issue=1598 |pmc=1635511 |pages=2119–25 |doi=10.1098/rspb.2006.3555 |url=http://www.adelaide.edu.au/acad/publications/papers/Barnett%20PRS%20lions.pdf |format=PDF |accessdate=4 September 2007 |deadurl=yes |archiveurl=https://web.archive.org/web/20070808182526/http://www.adelaide.edu.au/acad/publications/papers/Barnett%20PRS%20lions.pdf |archivedate=8 August 2007 }}</ref> Conversely, Per Christiansen found that using skull morphology allowed him to identify the subspecies ''krugeri'', ''nubica'', ''persica'', and ''senegalensis'', while there was overlap between ''bleyenberghi'' with ''senegalensis'' and ''krugeri''. The Asiatic lion ''persica'' was the most distinctive, and the Cape lion had characteristics allying it more with ''P. l. persica'' than the other sub-Saharan lions. He had analysed 58 lion skulls in three European museums.<ref name=christiansen08>{{Cite journal |last=Christiansen |first=Per | year = 2008 |title=On the distinctiveness of the Cape lion (''Panthera leo melanochaita'' Smith, 1842), and a possible new specimen from the Zoological Museum, Copenhagen | journal= Mammalian Biology | volume = 73 | issue = 1 | pages = 58–65 |doi = 10.1016/j.mambio.2007.06.003}}</ref>
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| Based on recent genetic studies, the Cat Classification Task Force of the IUCN SSC Cat Specialist Group has provisionally proposed to assign the lions occurring in Asia and West, Central and North Africa to the subspecies ''Panthera leo leo'' and the lions inhabiting South and East Africa to the subspecies ''Panthera leo melanochaita''.<ref>{{cite web|title=IUCN Cat Specialist Group|url=http://www.catsg.org/index.php?id=108}}</ref> The [[U.S. Fish and Wildlife Service]] has followed this revised taxonomic classification, as being based on "the best available scientific and commercial information", in listing these two subspecies as, respectively, endangered and threatened.<ref>{{cite journal |date=23 December 2015 |title=Endangered and Threatened Wildlife and Plants; Listing Two Lion Subspecies |url=https://www.gpo.gov/fdsys/pkg/FR-2015-12-23/pdf/2015-31958.pdf |journal=[[Federal Register]] |publisher=[[Federal government of the United States]] |id=80 FR 79999 |access-date=26 December 2015}}</ref>
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| The majority of lions kept in zoos are hybrids of different subspecies. Approximately 77% of the captive lions registered by the [[International Species Information System]] are of unknown origin. Nonetheless, they might carry genes that are extinct in the wild, and might be therefore important to maintain overall genetic variability of the lion.<ref name="CG"/> It is believed that those lions, imported to Europe before the middle of the nineteenth century, were mainly either Barbary lions from North Africa or lions from the Cape.<ref>{{cite journal | last1 = Barnett | first1 = Ross | last2 = Yamaguchi | first2 = Nobuyuki | last3 = Shapiro | first3 = Beth | last4 = Nijman | first4 = Vincent | 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}}</ref>
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| ====Recent====
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| Eight recent ([[Holocene]]) subspecies are recognised today:
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| {| class="wikitable sortable"
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| ! Subspecies !! Description !! Image
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| |'''[[Barbary lion]]''' (''P. l. leo''), also called the '''Atlas lion''' or '''North African lion''' || Formerly found in [[Algeria]], [[Morocco]], [[Tunisia]], and [[Egypt]], this is the nominate lion subspecies from North Africa. It is extinct in the wild due to excessive hunting; the last wild Barbary lion was killed in Morocco in 1920.<ref name=CAP>{{Cite book |last=Nowell |first=Kristin |last2=Jackson |first2=Peter |title= Wild Cats: Status Survey and Conservation Action Plan|url=http://carnivoractionplans1.free.fr/wildcats.pdf |format=PDF |year=1996 |publisher=IUCN/SSC Cat Specialist Group |location= Gland, Switzerland |isbn=2-8317-0045-0 |pages= 17–21, 37–41|chapter= Panthera Leo}}</ref><ref name="Black et al.2013">{{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 | url = | journal = PLoS ONE | volume = 8 | issue = 4| page = e60174 | doi = 10.1371/journal.pone.0060174 | bibcode = 2013PLoSO...860174B }}</ref> This is one of the largest of the lion subspecies,<ref name="yadav">{{Cite book |last=Yadav|first=P. R.|title=Vanishing And Endangered Species|url=https://books.google.com/?id=ecgil9AfV1QC&pg=PA178|accessdate=14 December 2010|year=2004|publisher=Discovery Publishing House|isbn=81-7141-776-0|pages=176–78}}</ref> with reported lengths of {{convert|3.0|–|3.3|m|ft|abbr=on}} and weights of more than {{convert|200|kg|lb|abbr=on}} for males. It appears to be more closely related to the Asiatic rather than sub-Saharan lions. A number of animals in captivity are likely to be Barbary lions,<ref>{{Cite journal |doi=10.1007/s10344-005-0009-z|url=http://www.uni-mainz.de/FB/Biologie/Anthropologie/MolA/Download/Burger%20Hemmer%202006.pdf |title=Urgent call for further breeding of the relic zoo population of the critically endangered Barbary lion (''Panthera leo leo'' Linnaeus 1758) |year=2006 |last=Burger |first=Joachim |journal=European Journal of Wildlife Research |volume=52 |page=54|last2=Hemmer|first2=Helmut}}</ref><!-- cites two previous sentences --> particularly the 90 animals descended from the Moroccan Royal collection at Rabat Zoo.<ref>{{Cite journal |doi=10.1007/s10344-009-0280-5 |title=Maintaining the genetic health of putative Barbary lions in captivity: an analysis of Moroccan Royal Lions |year=2010 |last=Black |first=Simon |last2=Yamaguchi |first2=Nobuyuki |last3=Harland |first3=Adrian |last4=Groombridge |first4=Jim |journal=European Journal of Wildlife Research |volume=56 |pages=21–31}}</ref>
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| North Africa: ([[Algeria]], [[Egypt]], [[Libya]], [[Morocco]] and [[Tunisia]])
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| ||[[File:Sultan the Barbary Lion.jpg|150px]]
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| |'''[[Asiatic lion]]''' (''P. l. persica''), also known as '''Indian lion''' or '''Persian lion''' || Is found in [[Gir Forest National Park]] of northwestern [[India]]. Once was widespread from [[Turkey]], across Southwest Asia, to India and [[Pakistan]],<ref name="Asiaticweb">{{cite web | title=Asiatic Lion—History | work=Asiatic Lion Information Centre | publisher=Wildlife Conservation Trust of India | year=2006 | url=http://www.asiaticlion.org/asiatic-lion-history.htm | accessdate=15 September 2007}}</ref> now 523 exist in and near the Gir Forest of [[Gujarat]].<ref name="Singh, Gibson">{{Cite journal |author1=Singh, H. S.|author2=Gibson, 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}}
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| </ref> Genetic evidence suggests its ancestors split from the ancestors of sub-Saharan African lions between 203 and 74 thousand years ago.<ref name="BurgerJ-Molecular-phylogeny"/>
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| Southern Europe: ([[Albania]], [[Bulgaria]], [[Greece]], [[Kosovo]], [[Republic of Macedonia|Macedonia]], [[Montenegro]] and [[Serbia]])
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| <br />
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| West Asia: ([[Armenia]], [[Baluchistan]], Georgia, [[Iran]], [[Iraq]], [[Israel]], [[Jordan]], [[Kuwait]], [[Lebanon]], [[Mesopotamia]], [[Oman]], Russia, [[Saudi Arabia]], [[Syria]], [[Turkey]], [[United Arab Emirates]] and [[Yemen]])
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| <br />
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| South Asia: ([[Afghanistan]], [[Bangladesh]], [[India]] and [[Pakistan]])
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| ||[[File:Asiatic lion 03.jpg|150px]]
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| |'''[[West African lion]]''' (''P. l. senegalensis''), also known as '''Senegal lion''' || Found in [[West Africa|western Africa]], from [[Senegal]] to the [[Central African Republic]].<ref name="HMW, 2009">Sunquist, M. E. and Sunquist, F. C. (2009). ''Family Felidae (Cats).'' In: Don E. Wilson, Russell A. Mittermeier (eds.): ''Handbook of the Mammals of the World. Volume 1: Carnivores.'' Lynx Edicions, 2009, ISBN 978-84-96553-49-1, pp. 137 ff.</ref><ref name="EoM">{{Cite book |last=Macdonald|first=David Wayne|title=The Encyclopedia of Mammals|year=2006|isbn=0-87196-871-1|pages=628–635}}</ref> It is currently listed as [[critically endangered]] in 2015. It is among the smallest of the Sub-Saharan African lions.
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| Western Africa: ([[Benin]], [[Burkina Faso]], [[Ghana]],<ref>{{cite journal|author=F. M. Angelici|author2=A. Mahama|author3=L. Rossi|last-author-amp=yes|title=The lion in Ghana: its historical and current status|journal=Animal Biodiversity and Conservation|date=20 July 2015|volume=38.2|pages=151–162|url=http://abc.museucienciesjournals.cat/files/ABC_38-2_pp_151-162-color.pdf|accessdate=26 February 2016}}</ref> Mali, Mauritania, Nigeria, and [[Senegal]])
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| <br />
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| Central Africa: ([[Cameroon]], [[Chad]], [[Central African Republic]], [[Nigeria]], and [[Niger]])
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| ||[[File:West African male lion.jpg|150px]]
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| |-
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| |'''[[Masai lion]]''' (''P. l. nubica''), also known as the '''East African lion'''|| Found in East Africa, from [[Ethiopia]] and [[Kenya]] to [[Tanzania]] and [[Mozambique]];<ref name="EoM"/> a local population is known as the [[Tsavo lion]].
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| Eastern Africa: ([[Djibouti]], [[Ethiopia]], [[Eritrea]], [[Kenya]], [[Somalia]], [[South Sudan]], [[Sudan]], [[Tanzania]], [[Uganda]])
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| ||[[File:African Lion 3.jpg|150px]]
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| |'''[[Congo lion]]''' (''P. l. azandica''), also known as '''Northeast Congo lion'''|| Found in the northeastern parts of the [[Democratic Republic of the Congo|Congo]].<ref name="HMW, 2009"/> It is currently extinct in [[Rwanda]].
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| Central Africa: ([[Central African Republic]], [[Democratic Republic of the Congo]], [[Rwanda]], South Sudan, Sudan, Uganda)
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| ||[[File:Murchison falls lion.jpg|150px]]
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| |'''[[Southwest African lion]]''' (''P. l. bleyenberghi''), also known as '''Katanga lion''' || Found in southwestern Africa and the Democratic Republic of the Congo. It is among the largest subspecies of African lions.
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| Southern Africa: ([[Angola]], [[Botswana]], [[Katanga Province|Katanga]] (Democratic Republic of the Congo), [[Namibia]], [[Zambia]], [[Zimbabwe]])<ref name="EoM"/>
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| ||[[File:P l Bleyenberghi 1.jpg|150px]]
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| |'''[[Transvaal lion]]''' (''P. l. krugeri''), also known as '''Southeast African lion'''|| Found in the [[South African Republic|Transvaal]] region of southeastern Africa, including [[Kruger National Park]].<ref name="EoM"/>
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| Southern Africa: (Botswana, [[Mozambique]], South Africa, [[Swaziland]], Zimbabwe)
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| ||[[File:Krugerparkléiw2.jpg|150px]]
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| |'''[[Ethiopian lion]]''' (''P. l. roosevelti''), also known as '''Addis Ababa lion'''|| A newly discerned lion subspecies could exist in captivity in Ethiopia's capital city of [[Addis Ababa]].<ref>{{cite web |title=A New, Genetically Distinct Lion Population is Found|url=http://newswatch.nationalgeographic.com/2012/11/30/a-new-genetically-distinct-lion-population-is-found/|work=News Watch|publisher=National Geographic Society|accessdate=23 January 2013|date=30 November 2012|quote=The Addis Ababa zoo lions have dark manes and small bodies, unlike other African lions. But life in captivity can sometimes influence appearance. A team of researchers, led by the Max Planck Institute for Evolutionary Anthropology in Germany and the University of York in the UK, checked to see if the lions really are different by comparing DNA samples of 15 lions from the zoo to six populations of wild lions. Their genetic analysis revealed that the gene sequence of all fifteen lions were unique and showed little sign of inbreeding. The study was recently published in the European Journal of Wildlife Research.}}</ref> Researchers compared the [[microsatellite]] variations over ten [[Locus (genetics)|loci]] of fifteen lions in captivity with those of six different wild lion populations. They determined that these lions are genetically unique and presumably that "their wild source population is similarly unique." These lions—with males that have a distinctly dark and luxuriant mane seem to define a new subspecies perhaps native only to [[Ethiopia]]. These lions were part of a collection of the late [[Haile Selassie I]] of Ethiopia.<ref>{{Cite journal |last=Bruche|first=Susann|author2=Gusset, Markus |author3=Lippold, Sebastian |author4=Barnett, Ross |author5=Eulenberger, Klaus |author6=Junhold, Jörg |author7=Driscoll, Carlos A. |author8= Hofreiter, Michael |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}}</ref>
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| Northeastern Africa: ([[Ethiopia]])
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| |}
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| ====Pleistocene====
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| Several additional subspecies of lion existed in prehistoric times:
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| * ''[[Panthera leo fossilis|P. l. fossilis]]'', known as the Middle Pleistocene European cave lion, flourished about 500,000 years ago; fossils have been recovered from Germany and Italy. It was larger than today's African lions, reaching sizes comparable to the American cave lion and slightly larger than the Upper Pleistocene European cave lion.<ref name = "BurgerJ-Molecular-phylogeny" /><ref>{{Cite book | language = German | first = Ernst | last = Probst | title = Deutschland in der Urzeit | publisher = Orbis | year = 1999 | isbn = 3-572-01057-8 }}</ref>
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| [[Image:Lions painting, Chauvet Cave (museum replica).jpg|thumb|Cave lions in the [[Chauvet Cave]], France]]
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| * ''[[Panthera leo spelaea|P. l. spelaea]]'', known as the European cave lion, Eurasian cave lion, or Upper Pleistocene European cave lion, occurred in Eurasia 300,000 to 10,000 years ago.<ref name="BurgerJ-Molecular-phylogeny" /> This species is known from [[Paleolithic]] [[cave painting]]s, [[ivory]] carvings, and clay busts,<ref name="Packer00">{{Cite journal | last=Packer | first=Craig |author2=Jean Clottes | title= When Lions Ruled France | journal=Natural History | pages=52–57 |date=November 2000}}</ref> indicating it had protruding ears, tufted tails, perhaps faint tiger-like stripes, and at least some had a ruff or primitive mane around their necks, possibly indicating males.<ref name="Koenigswald02" />
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| * ''[[American lion|P. l. atrox]]'', known as the American lion or American cave lion, was abundant in the Americas from Canada to [[Peru]] in the Pleistocene Epoch until about 10,000 years ago. This form is the sister clade of ''P. l. spelaea'', and likely arose when an early ''P. l. spelaea'' population became isolated south of the [[Cordilleran Ice Sheet|North American continental ice sheet]] about 0.34 Mya.<ref name= Barnett>{{Cite journal | last1=Barnett|first1=Ross| last2= Shapiro|first2= Beth| authorlink2= Beth Shapiro| last3 = Barnes | first3 =Ian| last4=Ho|first4=Simon Y.W.| last5=Burger|first5=Joachim| authorlink5=Joachim Burger | last6 =Yamaguchi|first6= Nobuyuki| last7= Higham|first7= Thomas FG| last8= Wheeler|first8= H Todd| title=Phylogeography of lions (''Panthera leo'' ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity| journal=[[Molecular Ecology]]|date=April 2009| volume= 18|issue= 8 | pages = 1668–77| pmid=19302360| doi=10.1111/j.1365-294X.2009.04134.x|url= http://www.uni-mainz.de/FB/Biologie/Anthropologie/MolA/Download/Barnett%20et%20al.%202009.pdf | accessdate=23 November 2011| first9=Wilfried| last9=Rosendahl| last10=Sher| first10=Andrei V.| last11=Sotnikova| first11=Marina| last12=Kuznetsova| first12=Tatiana| last13=Baryshnikov| first13=Gennady F.| last14=Martin| first14=Larry D.| last15=Harington| first15=C. Richard| last16=Burns| first16=James A.| last17=Cooper| first17=Alan}}</ref> One of the largest purported lion subspecies to have existed, its body length is estimated to have been {{convert|1.6|–|2.5|m|ft|abbr=on}}.<ref name="Quaternary extinctions">{{Cite book |last=Martin|first=Paul S. |title= Quaternary Extinctions |year=1984 |publisher= University of Arizona Press|location= Tucson, [[Arizona|AZ]] |url = https://books.google.com/?id=O8SgPwAACAAJ|isbn=0-8165-1100-4}}</ref>
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| ====Dubious====
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| * ''P. l. youngi'' or ''[[Panthera youngi]]'', flourished 350,000 years ago.<ref name= Harington69>{{Cite journal |last=Harington |first=C. R. "Dick |year= 1969 |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 |pages= 1277–88|doi= 10.1139/e69-127}}</ref> Its relationship to the extant lion subspecies is obscure, and it probably represents a distinct species.
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| * ''P. l. sinhaleyus'', known as the [[Sri Lanka lion]], appears to have become extinct around 39,000 years ago. It is only known from two teeth found in deposits at [[Kuruwita]]. Based on these teeth, [[Paul E. P. Deraniyagala|P. Deraniyagala]] erected this subspecies in 1939.<ref>{{Cite journal | first1 =Kelum | last1 = Manamendra-Arachchi | first2 = Rohan | last2 = Pethiyagoda | first3 = Rajith | last3 = Dissanayake | first4 = Madhava | last4 = Meegaskumbura |year=2005 |title= A second extinct big cat from the late Quaternary of Sri Lanka |journal= [[The Raffles Bulletin of Zoology]] |volume= Supplement 12 |pages= 423–34| format = PDF | url = http://rmbr.nus.edu.sg/rbz/biblio/s12/s12rbz423-434.pdf |publisher= National University of Singapore | accessdate = 31 July 2007}}</ref>
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| * ''[[Panthera leo vereshchagini|P. l. vereshchagini]]'', the Beringian cave lion of [[Sakha Republic|Yakutia]] (Russia), [[Alaska]] (United States), and the [[Yukon Territory]] (Canada), has been considered a subspecies separate from ''P. l. spelaea'' on morphological grounds. However, [[mitochondrial DNA]] sequences obtained from cave lion fossils from Europe and Alaska were indistinguishable.<ref name=Barnett />
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| [[File:Iranian - Cylinder Seal with a Lion Hunt - Walters 42797.jpg|thumb|A [[cylinder seal]] from [[Elam]] (now Iran) featuring an Elamite adaptation of the Babylonian theme of the lion hunt. 800–600 BC. Underbelly hair is visible. Now at the [[Walters Art Museum]].]]
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| * ''P. l. mesopotamica'' or ''[[Mesopotamian lion]]'', flourished during the Neo-Assyrian Period (approximately 1000–600 BC).<ref name= Ashrafian2011>{{Cite journal |last= Ashrafian |first= Hutan |year= 2011 |title= An Extinct Mesopotamian Lion Subspecies |journal= Veterinary Heritage | volume= 34 | issue= 2 | pages= 47–49}}</ref> It inhabited the Mesopotamian Plain where it probably represents a distinct sub-species. Nearly all ancient Mesopotamian representations of male lions demonstrate full underbelly hair in which until recently was only identified in the Barbary lion (''Panthera leo leo'') from Northern Africa and in most Asiatic lions (''P. l. persica'') from captivity in colder climates. Ancient evidence from adjacent landmasses reveal no substantiation for lions with underbelly hair in this manner so that the distinct phenotype of depicted lions in ancient Mesopotamia (including Babylon, Elam and ancient Persia) represent an extinct sub-species. Many of the images of these lions are derived from lion hunting sculptures so that the extinction of this sub-species likely resulted from overhunting in the ancient world.
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| * ''P. l. europaea'', known as the [[European lion]], was probably identical with ''Panthera leo persica'' or ''Panthera leo spelea''. It became extinct around 100 AD due to persecution and [[over-exploitation]]. It inhabited the [[Balkans]], the [[Italian Peninsula]], southern France, and the [[Iberian Peninsula]]. It was a very popular object of hunting among ancient [[Ancient Rome|Romans]] and [[ancient Greece|Greeks]].
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| * ''P. l. maculatus'', known as the [[marozi]] or spotted lion, sometimes is believed to be a distinct subspecies, but may be an adult lion that has retained its juvenile spotted pattern. If it was a subspecies in its own right, rather than a small number of aberrantly coloured individuals, it has been extinct since 1931. A less likely identity is a natural leopard-lion [[Panthera hybrid|hybrid]] commonly known as a [[leopon]].<ref name="shuker">{{Cite book | last=Shuker | first=Karl P. N. | authorlink=Karl Shuker | title=Mystery Cats of the World | publisher=Robert Hale | year=1989 | isbn=0-7090-3706-6}}</ref>
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| ===Hybrids===
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| {{further information|Panthera hybrid|Liger|Tigon}}
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| [[File:Lion Cubs Phinda 2011.ogv|thumb|right|Video of lioness and her cubs in the wild, South Africa]]
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| Lions have been known to [[biological reproduction|breed]] with tigers (most often the [[Siberian tiger|Siberian]] and [[Bengal tiger|Bengal]] subspecies) to create hybrids called [[liger]]s and [[tiglon]]s (or tigons).<ref>{{Cite book |last=Guggisberg |first=Charles Albert Walter |title=Wild Cats of the World |year=1975 |publisher=Taplinger Publishing |location=New York |isbn=0-8008-8324-1 }}</ref> They also have been crossed with [[leopards]] to produce [[leopon]]s,<ref>{{Cite book |last=Doi |first=H |last2=Reynolds |first2=B |title=The Story of Leopons |year=1967 |publisher=Putnam |location=New York |oclc=469041}}</ref> and [[jaguar]]s to produce [[Panthera hybrid#Jaglion|jaglions]]. The [[marozi]] is reputedly a spotted lion or a naturally occurring leopon, while the [[Congolese spotted lion]] is a complex lion-jaguar-leopard hybrid called a lijagulep. Such hybrids were once commonly bred in zoos, but this is now discouraged due to the emphasis on conserving species and subspecies. Hybrids are still bred in private menageries and in zoos in China.
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| The '''liger''' is a cross between a male lion and a tigress.<ref name=Carnivores>{{Cite book |last1=Rafferty|first1=John P.|title=Carnivores|date=2011|publisher=The Rosen Publishing Group|isbn=9781615303403|page=120|url=https://books.google.com/books?id=EMui7zVOqeUC&pg=PA120&dq=liger+lion+tigress&hl=en&sa=X&ei=56G2U6vSHou3yATBr4DoCw&ved=0CCIQ6AEwAQ#v=onepage&q=liger%20lion%20tigress&f=false|accessdate=4 July 2014}}</ref> Because the growth-inhibiting gene from the female tiger mother is absent, the growth-promoting gene passed on by the male lion father is unimpeded by a regulating gene and the resulting ligers grow far larger than either parent. They share physical and behavioural qualities of both parent species (spots and stripes on a sandy background). Male ligers are sterile, but female ligers often are fertile. Males have about a 50% chance of having a mane, but if they grow them, their manes will be modest: around 50% the size of a pure lion mane.
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| Ligers are much bigger than normal lions, typically {{convert|3.65|m|ft|abbr=on}} in length, and can weigh up to {{convert|500|kg|lb|abbr=on}}.<ref name=shi/>
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| The less common '''tiglon''' or '''tigon''' is a cross between a lioness and a male tiger.<ref>{{cite web |url=http://www.britannica.com/EBchecked/topic/595552/tigon|title= Tigon—Encyclopædia Britannica Article|accessdate=25 January 2011}}</ref> In contrast to ligers, tigons are often relatively small in comparison to their parents, because of reciprocal gene effects.<ref name=shi>{{cite book |last=Shi |first=Wei |year=2005|title=Growth and Behviour: Epigenetic and Genetic Factors Involved in Hybrid Dysgenesis|work=Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology|volume=11|page=9|isbn=91-554-6147-6|url=http://urn.kb.se/resolve?urn=urn:nbn:se:uu:diva-4784|publisher=Acta Universitatis Upsaliensis|location=Uppsala}}</ref>
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| ==Characteristics==
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| [[File:Lion and eland.jpg|thumb|right|A skeletal mount of an African lion attacking a [[common eland]] on display at [[The Museum of Osteology]], [[Oklahoma City, Oklahoma]].]]
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| Of the living felids the lion is second only to the tiger in length and weight. Its skull is very similar to that of the tiger, although the frontal region is usually more depressed and flattened, with a slightly shorter postorbital region and broader nasal openings than that of a tiger. However, due to the amount of skull variation in the two species, usually only the structure of the lower jaw can be used as a reliable indicator of species.<ref name="USSR">{{Cite book | last = Heptner |first=V. G. |last2=Sludskii |first2=A. A.| title = Mammals of the Soviet Union, Volume II, Part 2 | year = 1992| isbn = 90-04-08876-8 | publisher = Brill | location = Leiden u.a.}}</ref> Lion colouration varies from light buff to yellowish, reddish, or dark ochraceous brown. The underparts are generally lighter and the tail tuft is black. Lion cubs are born with brown [[Rosette (zoology)|rosettes]] (spots) on their body, rather like those of a leopard. Although these fade as lions reach adulthood, faint spots often may still be seen on the legs and underparts, particularly on lionesses.
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| Lions are the only members of the cat family to display obvious [[sexual dimorphism]] – that is, males and females look distinctly different. They also have specialised roles that each gender plays in the pride. For instance, the lioness, the hunter, lacks the male's thick mane. The colour of the male's mane varies from blond to black, generally becoming darker as the lion grows older. The most distinctive characteristic shared by both females and males is that the tail ends in a hairy tuft. In some lions, the tuft conceals a hard "spine" or "spur", approximately 5 mm long, formed of the final sections of tail bone fused together. The lion is the only felid to have a tufted tail – the function of the tuft and spine are unknown. Absent at birth, the tuft develops around {{frac|5|1|2}} months of age and is readily identifiable at 7 months.<ref name="Schaller28">[[#Schaller|Schaller]], p. 28.</ref>
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| Nowak indicates the typical weight range of lions as {{convert|150 to 250|kg|lb|0|abbr=on}} for males and {{convert|120 to 185|kg|lb|0|abbr=on}} for females.<ref name="nowak"/> The size of adult lions varies across their range with those from the southern African populations in Zimbabwe, the Kalahari and Kruger Park averaging around {{convert|189.6|kg|lb|abbr=on}} and {{convert|126.9|kg|lb|abbr=on}} in males and females respectively compared to {{convert|174.9|kg|lb|abbr=on}} and {{convert|119.5|kg|lb|abbr=on}} of male and female lions from East Africa.<ref name=Smuts_etal_1980>{{Cite journal |last=Smuts|first=G. L.|author2=Robinson, G.A.|author3=Whyte, I.J.|title=Comparative growth of wild male and female lions (Panthera leo)|journal=Journal of Zoology|year=1980|volume=190|issue=3|pages=365–373|url=http://onlinelibrary.wiley.com/doi/10.1111/j.1469-7998.1980.tb01433.x/abstract|doi=10.1111/j.1469-7998.1980.tb01433.x}}</ref> Reported body measurements in males are head-body lengths ranging from {{convert|170|to|250|cm|ftin|abbr=on}}, tail lengths of {{convert|90|–|105|cm|ftin|abbr=on}}. In females reported head-body lengths range from {{convert|140|to|175|cm|ftin|abbr=on}}, tail lengths of {{convert|70|–|100|cm|ftin|abbr=on}},<ref name="nowak"/> however, the frequently cited maximum head and body length of {{convert|250|cm|ftin|abbr=on}} fits rather to extinct Pleistocene forms, like the [[American lion]], with even large modern lions measuring several centimetres less in length.<ref>[[Vratislav Mazák|Mazák, Vratislav]] (April 2004) ''Der Tiger'', Westarp Wissenschaften, 5th ed., pp. 178 ff., ISBN 3-89432-759-6.</ref> Record measurements from hunting records are supposedly a total length of nearly {{convert|3.6|m|ft|abbr=on}} for a male shot near Mucsso, southern [[Angola]] in October 1973 and a weight of {{convert|313|kg|lb|abbr=on}} for a male shot outside [[Hectorspruit]] in eastern [[Transvaal Province|Transvaal]], South Africa in 1936.<ref>Wood, The Guinness Book of Animal Facts and Feats. Sterling Pub Co Inc. (1983), ISBN 978-0-85112-235-9</ref> Another notably outsized male lion, which was shot near [[Mount Kenya]], weighed in at {{convert|272|kg|lb|abbr=on}}.<ref name="CAP"/>
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| ===Mane===
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| [[File:HansomeLion 002.jpg|thumb|right|During agonistic confrontations with other lions, the mane makes the lion appear larger.]]
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| The mane of the adult male lion, unique among cats, is one of the most distinctive characteristics of the species. In rare cases a female lion can have a mane.<ref>{{cite web|author=Readhead, Harry|date=23 December 2015|title=Scientists discover 'genderfluid' lioness who looks, acts and roars like a male|url=http://metro.co.uk/2015/12/23/scientists-discover-genderfluid-lioness-who-looks-acts-and-roars-like-a-male-5582846/|journal=Metro|accessdate=25 December 2015}}</ref><ref>{{cite web | last=Trivedi | first=Bijal P. | title=Are Maneless Tsavo Lions Prone to Male Pattern Baldness? | publisher=National Geographic | year=2005 | url=http://news.nationalgeographic.com/news/2002/04/0412_020412_TVtsavolions.html | accessdate=7 July 2007}}</ref> The presence, absence, colour, and size of the mane is associated with genetic precondition, sexual maturity, climate, and [[testosterone]] production; the rule of thumb is the darker and fuller the mane, the healthier the lion. Sexual selection of mates by lionesses favours males with the densest, darkest mane.<ref name="Trivedi02">{{cite web | last=Trivedi | first =Bijal P. | title=Female Lions Prefer Dark-Maned Males, Study Finds | work=National Geographic News | publisher=National Geographic | date=22 August 2002 | url=http://news.nationalgeographic.com/news/2002/08/0822_020822_TVlion.html| accessdate=1 September 2007}}</ref> Research in [[Tanzania]] also suggests mane length signals fighting success in male–male relationships. Darker-maned individuals may have longer reproductive lives and higher offspring survival, although they suffer in the hottest months of the year.<ref name=West>{{Cite journal |last=West |first=Peyton M. |author2=Packer, Craig |date=August 2002 |title=Sexual Selection, Temperature, and the Lion's Mane |journal=[[Science (journal)|Science]] |volume=297 |issue=5585 |pages= 1339–43 |doi=10.1126/science.1073257 |pmid=12193785}}</ref>
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| [[File:Maneless lion from Tsavo East National Park.png|thumb|A maneless lion.]]
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| Scientists once believed that the distinct status of some subspecies could be justified by [[morphology (biology)|morphology]], including the size of the mane. Morphology was used to identify subspecies such as the [[Barbary lion]] and [[Cape lion]]. Research has suggested, however, that environmental factors such as average [[room temperature|ambient temperature]] influence the colour and size of a lion's mane.<ref name=West/> The cooler ambient temperature in European and North American [[zoo]]s, for example, may result in a heavier mane. Thus the mane is not an appropriate marker for identifying subspecies.<ref name="CG">{{Cite journal |last=Barnett |first=Ross |author2=Yamaguchi, Nobuyuki |author3=Barnes, Ian|author4=Cooper, Alan |date=August 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=507–14 |doi=10.1007/s10592-005-9062-0}}</ref><ref name="yamaguchi-haddane">{{Cite journal |last=Yamaguchi |first=Nobuyuki |author2=Haddane, B. |year=2002 |title=The North African Barbary Lion and the Atlas Lion Project |journal=International Zoo News |volume=49 |pages=465–81}}</ref> The males of the Asiatic subspecies, however, are characterised by sparser manes than average African lions.<ref name="Menon">{{Cite book |last=Menon |first=Vivek |year=2003 |title=A Field Guide to Indian Mammals |location=Delhi |publisher=Dorling Kindersley India |isbn=0-14-302998-3| pages=}}</ref>
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| In the [[Pendjari National Park]] area almost all males are maneless or have very weak manes.<ref>Schoe, Marjolein; Sogbohossou, Etotépé A.; Kaandorp, Jacques and de Iongh, Hans (2010) ''Progress Report – collaring operation Pendjari Lion Project, Benin.'' Funded by the Dutch Zoo Conservation Fund.</ref> Maneless male lions have also been reported from [[Senegal]], from Sudan ([[Dinder National Park]]), and from [[Tsavo East National Park]] in Kenya, and the original male white lion from Timbavati also was maneless. The testosterone hormone has been linked to mane growth; therefore, [[castration|castrated]] lions often have minimal to no mane, as the removal of the gonads inhibits testosterone production.<ref>{{Cite journal | first = Linda | last = Munson| date = 3 March 2006 | title = Contraception in felids | journal = Theriogenology| pmid = 16626799 | volume = 66 | issue = 1 | pages = 126–34 | doi = 10.1016/j.theriogenology.2006.03.016 }}</ref> In addition, increased testosterone may be the cause of the maned lionesses of northern Botswana.<ref>{{Cite web|last=Dell'Amore|first=Christine|date=18 April 2016|accessdate=18 April 2016|title=No, Those Aren't Male Lions Mating. One Is Likely a Female|url=http://news.nationalgeographic.com/2016/04/160418-lions-mating-africa-animals-science/}}</ref>
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| [[Cave painting]]s of extinct European [[Panthera leo spelaea|cave lions]] almost exclusively show animals with no manes, suggesting that either they were maneless,<ref name="Koenigswald02">{{Cite book |last=Koenigswald |first=Wighart von |year=2002 |title=Lebendige Eiszeit: Klima und Tierwelt im Wandel |location=Stuttgart |publisher=Theiss |isbn=3-8062-1734-3| pages=|language=de}}</ref> or that the paintings depict lionesses as seen hunting in a group.
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| ===White lions===
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| [[File:White Lion.jpg|thumb|left|[[White lion]]s owe their colouring to a recessive allele; they are rare forms of the subspecies [[Transvaal lion]].]]
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| The [[white lion]] is not a distinct subspecies, but a special [[Polymorphism (biology)|morph]] with a genetic condition, [[leucism]],<ref name="zoos_encyclopedia"/> that causes paler colouration akin to that of the [[white tiger]]; the condition is similar to [[melanism]], which causes [[black panther]]s. They are not albinos, having normal pigmentation in the eyes and skin. White Transvaal lion (''Panthera leo krugeri'') individuals occasionally have been encountered in and around [[Kruger National Park]] and the adjacent [[Timbavati Private Game Reserve]] in eastern South Africa, but are more commonly found in [[captivity (animal)|captivity]], where breeders deliberately select them. The unusual cream colour of their coats is due to a [[recessive allele]].<ref>{{Cite book |last=McBride |first=Chris |title=The White Lions of Timbavati |year=1977 |publisher=E. Stanton |location=Johannesburg |isbn= 0-949997-32-3 }}</ref> Reportedly, they have been bred in camps in South Africa for use as trophies to be killed during [[canned hunt]]s.<ref>{{Cite book |last=Tucker|first=Linda |title=Mystery of the White Lions—Children of the Sun God|year=2003 |publisher=Npenvu Press|location=Mapumulanga |isbn= 0-620-31409-5}}</ref>
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| {{clear}}
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| ==Behaviour==
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| [[File:Panthera leo stretching (Etosha, 2012).jpg|thumbnail|left|Adult male lion stretching.]]
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| Lions spend much of their time resting and are inactive for about 20 hours per day.<ref name="Schaller122">[[#Schaller|Schaller]], p. 122.</ref> Although lions can be active at any time, their activity generally peaks after dusk with a period of socialising, grooming, and defecating. Intermittent bursts of activity follow through the night hours until dawn, when hunting most often takes place. They spend an average of two hours a day walking and 50 minutes eating.<ref name="Schaller120">[[#Schaller|Schaller]], pp. 120–21.</ref>
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| ===Group organisation===
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| [[File:Lions Family Portrait Masai Mara.jpg|thumb|right|Two males and a female lion.]]
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| Lions are the most socially inclined of all wild [[felid]]s, most of which remain quite solitary in nature. The lion is a predatory [[carnivore]] with two types of social organization. Some lions are ''residents,'' living in groups of related lionesses, their mates, and offspring. Such a group is called a ''pride''.<ref name="Schaller33">[[#Schaller|Schaller]], p. 33.</ref> Females form the stable [[social unit]] in a pride and do not tolerate outside females.<ref name="Schaller37">[[#Schaller|Schaller]], p. 37.</ref> Membership only changes with the births and deaths of lionesses,<ref name="Schaller39">[[#Schaller|Schaller]], p. 39.</ref> although some females do leave and become nomadic.<ref name="Schaller44">[[#Schaller|Schaller]], p. 44.</ref> Although extremely large prides, consisting of up to 30 individuals, have been observed, the average pride consists of five or six females, their cubs of both sexes, and one or two males (known as a ''coalition'' if more than one) who mate with the adult females. The number of adult males in a coalition is usually two but may increase to as many as four before decreasing again over time.<ref name="Schaller44"/> The sole exception to this pattern is the [[Tsavo lion]] pride which always has just one adult male.<ref>{{Cite journal |last=Milius |first=Susan |date=April 2002 |title= Biology: Maneless lions live one guy per pride|journal=Society for Science & the Public |volume=161 |issue=16 |page=253 |doi=10.1002/scin.5591611614}}</ref> Male cubs are excluded from their maternal pride when they reach maturity at around 2–3 years of age.<ref name="Schaller44"/> The second organizational behaviour is labeled ''nomads'', who range widely and move about sporadically, either singularly or in pairs.<ref name="Schaller33"/> Pairs are more frequent among related males who have been excluded from their birth pride. Note that a lion may switch lifestyles; nomads may become residents and vice versa. Males, as a rule, live at least some portion of their lives as nomads, and some are never able to join another pride. A female who becomes a nomad has much greater difficulty joining a new pride, as the females in a pride are related, and they reject most attempts by an unrelated female to join their family group.
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| [[File:Tree-climbing lions (Panthera leo).jpg|thumb|Tree-climbing lions of [[Ishasha]], [[Uganda]]]]
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| The area a pride occupies is called a ''pride area'', whereas that by a nomad is a ''range''.<ref name="Schaller33"/> The males associated with a pride tend to stay on the fringes, patrolling their [[territory (animal)|territory]]. Why [[social behavior|sociality]] – the most pronounced in any cat species – has developed in lionesses is the subject of much debate. Increased hunting success appears an obvious reason, but this is less than sure upon examination: coordinated hunting does allow for more successful predation but also ensures that non-hunting members reduce per capita calorific intake; however, some take a role raising cubs, who may be left alone for extended periods of time. Members of the pride regularly tend to play the same role in hunts and hone their skills. The health of the hunters is the primary need for the survival of the pride, and they are the first to consume the prey at the site it is taken. Other benefits include possible [[kin selection]] (better to share food with a related lion than with a stranger), protection of the young, maintenance of territory, and individual insurance against injury and hunger.<ref name=CAP/>
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| Lionesses do most of the hunting for their pride. They are more effective hunters, as they are smaller, swifter, and more agile than the males and unencumbered by the heavy and conspicuous mane, which causes overheating during exertion. They act as a coordinated group with members who perform the same role consistently in order to stalk and bring down the prey successfully. Smaller prey is eaten at the location of the hunt, thereby being shared among the hunters; when the kill is larger it often is dragged to the pride area. There is more sharing of larger kills,<ref name="Schaller133">[[#Schaller|Schaller]], p. 133.</ref> although pride members often behave aggressively toward each other as each tries to consume as much food as possible. Near the conclusion of the hunt, males have a tendency to dominate the kill once the lionesses have succeeded. They are more likely to share this with the cubs than with the lionesses, but males rarely share food they have killed by themselves.
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| Both males and females can defend the pride against intruders, but the male lion is better-suited for this purpose due to its stockier, more powerful build.<ref>[http://www.lionconservationfund.org/lion_adopt.html Join the Pride]. Lion Conservation Fund. Retrieved on 31 July 2013.</ref> Some individuals consistently lead the defence against intruders, while others lag behind.<ref>{{Cite journal |last=Heinsohn|first= R. |author2=C. Packer |year=1995 |title=Complex cooperative strategies in group-territorial African lions |journal=Science |volume=269 |issue=5228 |pages=1260–62 |doi=10.1126/science.7652573 |pmid=7652573}}</ref> Lions tend to assume specific roles in the pride. Those lagging behind may provide other valuable services to the group.<ref>{{Cite journal |last=Morell|first= V. |year=1995 |title=Cowardly lions confound cooperation theory |journal=Science |volume=269 |issue=5228 |pages=1216–17|doi=10.1126/science.7652566 |pmid=7652566}}</ref> An alternative hypothesis is that there is some reward associated with being a leader who fends off intruders, and the rank of lionesses in the pride is reflected in these responses.<ref>{{Cite journal |last=Jahn|first= Gary C. |year=1996 |title=Lioness Leadership |journal=Science |volume=271 |issue=5253 |page=1215 |doi=10.1126/science.271.5253.1215a |pmid=17820922 }}</ref> The male or males associated with the pride must defend their relationship to the pride from outside males who attempt to take over their relationship with the pride.
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| ===Hunting and diet===
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| [[File:Female Lion.JPG|thumb|left|A lion's teeth are typical of a carnivore]]
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| [[File:Lions eat heron for lunch!.webm|thumb|Lions feeding on a heron]]
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| Lions prefer to [[Scavenger|scavenge]] when the opportunity presents itself<ref name = "Schaller213" /> with [[carrion]] providing more than 50% of their diet.<ref name=AWF>>{{cite web | title = Behavior and Diet | work = African Wildlife Foundation website | publisher = African Wildlife Foundation | year = 1996 | url =http://www.awf.org/wildlife-conservation/lion | accessdate =6 June 2014}}</ref> They scavenge animals either dead from [[natural causes]] (disease) or killed by other predators, and keep a constant lookout for circling vultures, being keenly aware that they indicate an animal dead or in distress.<ref name = "Schaller213">[[#Schaller|Schaller]], p. 213.</ref> In fact, most dead prey on which both hyenas and lions feed upon are killed by the hyenas instead of the lions.<ref name="nowak" />
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| The lionesses do most of the hunting for the pride. The male lion associated with the pride usually stays and watches over young cubs until the lionesses return from the hunt. Typically, several work together and encircle the herd from different points. Once they have closed in on the herd, they usually target the animal closest to them. The attack is short and powerful; they attempt to catch the victim with a fast rush and final leap. The prey usually is killed by [[strangulation]],<ref>{{cite web | title = About lions—Ecology and behaviour | first = Gus | last = Mills | publisher = African Lion Working Group | url= http://www.african-lion.org/lions_e.htm | accessdate = 20 July 2007}}</ref> which can cause [[cerebral ischemia]] or [[asphyxia]] (which results in [[hypoxemia|hypoxemic]], or "general", [[hypoxia (medical)|hypoxia]]). The prey also may be killed by the lion enclosing the animal's mouth and nostrils in its jaws (which would also result in asphyxia).<ref name="nowak" />
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| Lions usually hunt in coordinated groups and stalk their chosen prey. However, they are not particularly known for their stamina – for instance, a lioness' heart makes up only 0.57% of her body weight (a male's is about 0.45% of his body weight), whereas a hyena's heart is close to 1% of its body weight.<ref>[[#Schaller|Schaller]], p. 248.</ref> Thus, they only run fast in short bursts,<ref name="Schaller2478">[[#Schaller|Schaller]], pp. 247–48.</ref> and need to be close to their prey before starting the attack. They take advantage of factors that reduce visibility; many kills take place near some form of cover or at night.<ref name="Schaller237">[[#Schaller|Schaller]], p. 237.</ref> They sneak up to the victim until they reach a distance of approximately {{convert|30|m|ft|abbr=off}} or less.
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| [[File:Serengeti Lion Running saturated.jpg|thumb|left|Lioness in a burst of speed while hunting.]]
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| The [[predation|prey]] consists mainly of medium-sized mammals, with a preference for [[wildebeest]], [[zebra]]s, [[African buffalo|buffalo]], and [[warthog]]s in Africa and [[nilgai]], [[boar|wild boar]], and several [[deer]] species in India. Many other species are hunted, based on availability, mainly [[ungulate]]s weighing between {{convert|50|and|300|kg|lb|abbr=on}} such as [[kudu]], [[hartebeest]], [[gemsbok]], and [[taurotragus|eland]].<ref name="nowak" /> Occasionally, they take relatively small species such as [[Thomson's gazelle]] or [[springbok (antelope)|springbok]]. Lions hunting in groups are capable of taking down most animals, even healthy adults, but in most parts of their range they rarely attack very large prey such as fully grown male [[giraffe]]s due to the danger of injury. Giraffes and buffaloes are almost invulnerable to a solitary lion as well.<ref name="Schaller259">[[#Schaller|Schaller]], p. 259.</ref>
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| Extensive studies show that lionesses normally prey on mammals with an average weight of {{convert|126|kg|abbr=on}}, while kills made by male lions average {{convert|399|kg|abbr=on}}.<ref>{{Cite journal | author1=Radloff, Frans G. T.|author2=Du Toit|author3=Johan T. |year= 2004 |title= Large predators and their prey in a southern African savanna: a predator's size determines its prey size range |journal=Journal of Animal Ecology |volume=73 |pages=410–423 |doi=10.1111/j.0021-8790.2004.00817.x}}</ref> In Africa, wildebeest rank at the top of preferred prey (making nearly half of the lion prey in the [[Serengeti]]) followed by zebra.<ref>{{Cite book |isbn=88-8095-752-X| title = The Art of Being a Lion | page = 186 | first1 = Christine | last1 = Denis-Huot | first2 = Michel | last2 = Denis-Huot | publisher = Friedman/Fairfax | year = 2002}}</ref> Lions do not prey on fully grown adult [[elephant]]s; most adult [[hippopotamus]]es, [[rhinoceros]]es, and smaller [[gazelle]]s, [[impala]], and other agile antelopes are generally excluded. However, giraffes and buffaloes are often taken in certain regions. For instance, in Kruger National Park, giraffes are regularly hunted.<ref name=Pienaar1969>{{Cite journal |last1=Pienaar|first1=U. de V.|title=Predator–prey relationships among the larger mammals of the Kruger National Park|journal=Koedoe|date=1969|volume=12|issue=1|doi=10.4102/koedoe.v12i1.753|url=http://www.koedoe.co.za/index.php/koedoe/article/view/753}}</ref> In Manyara Park, Cape buffaloes constitute as much as 62% of the lion's diet,<ref>{{Cite journal | title = Among the Elephants | first1 = Iain | last1 = Douglas-Hamilton | first2 = Oria | last2 = Douglas-Hamilton | year = 1975 }}</ref> due to the high number density of buffaloes. Occasionally hippopotamus is also taken, but adult rhinoceroses are generally avoided. Warthogs are often taken depending on availability.<ref>{{Cite journal |last1 =Hayward|first1 = Matt W. | last2 = Kerley | first2 = Graham | year = 2005 | title= Prey preferences of the lion (''Panthera leo'') | journal= Journal of Zoology | volume = 267 | issue = 3 | pages = 309–22 |doi= 10.1017/S0952836905007508}}</ref> The lions of [[Chobe National Park#Geography and ecosystems|Savuti]], Botswana, have adapted to hunting young elephants during the dry season, and a pride of 30 lions has been recorded killing individuals between the ages of four and eleven years.<ref>{{Cite journal |last=Power |first=R. J. |last2=Shem Compion |first2=R. X.|year=2009|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}}</ref> In the [[Kalahari desert]] in South Africa, black-maned lions may chase baboons up a tree, wait patiently, then attack them when they try to escape:
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| [[File:Lion and baboon 1.jpg|thumb|250px|Baboons trapped up a tree by lions.]]
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| Lions also attack domestic livestock and in India cattle contribute significantly to their diet.<ref name = "Menon" /> Lions are capable of killing other predators such as [[leopard]]s, [[cheetah]]s, [[hyena]]s, and [[African wild dog|wild dogs]], though (unlike most felids) they seldom devour the competitors after killing them. A lion may gorge itself and eat up to {{convert|30|kg|lb|abbr=on}} in one sitting;<ref name="simba">{{Cite book | last= Guggisberg|first= Charles Albert Walter| title = Simba: the life of the lion. |year=1961 |publisher=Howard Timmins |location=Cape Town}}</ref> if it is unable to consume all the kill it will rest for a few hours before consuming more. On a hot day, the pride may retreat to shade leaving a male or two to stand guard.<ref name="Schaller2706">[[#Schaller|Schaller]], pp. 270–76.</ref> As revealed by fossil evidence at Olduvai, Tanzania, lions will occasionally drag their carcasses to a more sheltered spot to eat (as tigers and leopards are known to do), allowing them to consume more of a carcass without interference from scavengers.<ref>http://blogs.scientificamerican.com/laelaps/olduvai-s-lions-have-a-bone-to-pick/</ref> An adult lioness requires an average of about {{convert|5|kg|lb|abbr=on}} of meat per day, a male about {{convert|7|kg|lb|abbr=on}}.<ref>[[#Schaller|Schaller]], p. 276.</ref>
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| [[File:Lions taking down cape buffalo.jpg|thumb|right|250px|Four lionesses catch a cape buffalo.]]
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| Because lionesses hunt in open spaces where they are easily seen by their prey, cooperative hunting increases the likelihood of a successful hunt; this is especially true with larger species. Teamwork also enables them to defend their kills more easily against other large predators such as hyenas, which may be attracted by [[vulture]]s from kilometres away in open savannas. Lionesses do most of the hunting; males attached to prides do not usually participate in hunting, except in the case of larger quarry such as giraffe and buffalo. 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 in flight from other lionesses.<ref name=hunt>{{Cite journal | last = Stander | first = PE | title= Cooperative hunting in lions: the role of the individual |url=http://lchc.ucsd.edu/MCA/Mail/xmcamail.2011_09.dir/pdfeL5GzNL2FL.pdf |journal=Behavioral Ecology and Sociobiology|volume=29|issue=6|pages=445–54|year=1992|doi = 10.1007/BF00170175}}</ref> There is evidence that male lions are just as successful at hunting as females; they are solo hunters who ambush prey in small bush.<ref>{{cite journal |author=Scott R. Loariea |author2=Craig J. Tamblingb |author3=Gregory P. Asnera|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}}</ref> Young lions first display stalking behaviour around three months of age, although they do not participate in hunting until they are almost a year old. They begin to hunt effectively when nearing the age of two.<ref name="Schaller153">[[#Schaller|Schaller]], p. 153.</ref>
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| ====Predator competition====
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| [[File:Hyenas Fight Against Lions Over a Kill HD 10.png|thumb|left|Lion attacked by [[spotted hyena]]s]]
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| Lions and [[spotted hyena]]s occupy the same [[ecological niche]], meaning they compete for prey and carrion in the areas where they coexist. A review of data across several studies indicates a dietary overlap of 58.6%.<ref name="prey">{{Cite journal | url = http://www.zbs.bialowieza.pl/g2/pdf/1598.pdf | format = PDF | title= Prey preferences of the spotted hyaena (''Crocuta crocuta'') and degree of dietary overlap with the lion (''Panthera leo'')| last =Hayward | first = Matt W.| journal =Journal of Zoology |year= 2006|volume= 270|pages= 606–14 | doi = 10.1111/j.1469-7998.2006.00183}}</ref> Lions typically ignore spotted hyenas unless the lions are on a kill or are being harassed by the hyenas, while the latter tend to visibly react to the presence of lions whether there is food or not. Lions seize the kills of spotted hyenas: in the [[Ngorongoro crater]], it is common for lions to subsist largely on kills stolen from hyenas, causing the hyenas to increase their kill rate.<ref name="Kruuk21">{{Cite book |chapter = Interactions between Hyenas and other Carnivorous Animals | first = Hans | last = Kruuk | title = The Spotted Hyena: A Study of Predation and Social Behaviour | publisher = The University of Chicago Press | place = Chicago | year = 1972|isbn=0-226-45508-4 }}</ref> On the other hand, in Northern Botswana's Chobe National Park, the situation is reversed: hyenas frequently challenge lions and steal their kills: they obtain food from 63% of all lion kills.<ref name="Conservation">{{Cite book | title = Carnivore Conservation |edition=1st |editor1-first=John L. |editor1-last=Gittleman |editor2-first=Stephan M. |editor2-last=Funk |editor3-first=David W. |editor3-last=Macdonald |editor4-first=Robert K. |editor4-last=Wayne| publisher = Cambridge University Press | year = 2001|isbn=978-0-521-66232-1}}</ref> When confronted on a kill by lions, spotted hyenas may either leave or wait patiently at a distance of {{convert|30|–|100|m|ft|abbr=on}} until the lions have finished,<ref name="schaller272">[[#Schaller|Schaller]], p. 272.</ref> but they are also bold enough to feed alongside lions, and even force the lions off a kill. The two species may attack one another even when there is no food involved for no apparent reason.<ref name="schaller273">[[#Schaller|Schaller]], pp. 273–274.</ref><ref name="enemies">{{cite video|people=Joubert, Dereck and Joubert, Beverley|title= Eternal Enemies: Lions and Hyenas |medium=DVD | publisher = [[National Geographic Channel|National Geographic]]|year= 1992}}</ref> Lion predation can account for up to 71% of hyena deaths in [[Etosha]]. Spotted hyenas have adapted by frequently mobbing lions that enter their territories.<ref>{{Cite journal |title=Competitive interactions between spotted hyenas and lions in the Etosha National Park, Namibia| last1 =Trinkel | first1 = Martina | last2 = Kastberger | first2 = Gerald |journal=African Journal of Ecology|volume=43|issue=3|year=2005|pages=220–24|doi = 10.1111/j.1365-2028.2005.00574.x}}</ref> Experiments on captive spotted hyenas revealed that specimens with no prior experience with lions act indifferently to the sight of them, but will react fearfully to the scent.<ref name="Kruuk21" /> The size of male lions allows them occasionally to confront hyenas in otherwise evenly matched brawls and so to tip the balance in favour of the lions.
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| Lions tend to dominate smaller felines such as [[cheetah]]s and [[leopard]]s where they co-occur, stealing their kills and killing their cubs and even adults when given the chance. The cheetah has a 50% chance of losing its kill to lions or other predators.<ref>{{Cite journal | last1 =O'Brien | first1 = Stephen J | last2 = Wildt | first2 = David E | last3 = Bush | first3 = Mitchell|year= 1986| title= The Cheetah in Genetic Peril| format = PDF | journal = Scientific American|issue= 254|pages= 68–76| url= http://www.catsg.org/cheetah/05_library/5_3_publications/N_and_O/OBrien_et_al_1986_Cheetah_in_genetic_peril.pdf}}</ref> Lions are major killers of cheetah cubs, up to 90% of which are lost in their first weeks of life due to attacks by other predators. Cheetahs avoid competition by hunting at different times of the day and hide their cubs in thick brush. Leopards also use such tactics, but have the advantage of being able to subsist much better on small prey than either lions or cheetahs. Also, unlike cheetahs, leopards can climb trees and use them to keep their cubs and kills away from lions; however, lionesses will occasionally be successful in climbing to retrieve leopard kills.<ref name = "Schaller293">[[#Schaller|Schaller]], p. 293.</ref> Similarly, lions dominate [[African wild dog]]s, not only taking their kills but also preying on young and (rarely) adult dogs. Population densities of wild dogs are low in areas where lions are more abundant.<ref>{{cite journal | title = Conserving the African wild dog ''Lycaon pictus''. I. Diagnosing and treating causes of decline | last1 =Woodroffe | first1 = Rosie | last2 = Ginsberg | first2 = Joshua R |year=1999| journal =Oryx | volume = 33| pages =132–42|doi= 10.1046/j.1365-3008.1999.00052.x |issue=2}}</ref><!-- cites two previous refs --> However, there are a few reported cases of old and wounded lions falling prey to wild dogs.<ref name=Pienaar1969/><ref>[[#Schaller|Schaller]], p. 188.</ref>
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| The [[Nile crocodile]] is the only [[Sympatry|sympatric]] predator (besides humans) that can singly threaten the lion. Depending on the size of the crocodile and the lion, either can lose kills or carrion to the other. Lions have been known to kill crocodiles venturing onto land,<ref>{{cite web |url= http://www.pbs.org/wgbh/nova/transcripts/2509crocs.html |title=Crocodiles! | publisher = PBS | work = Nova | type = transcript |date= 28 April 1998 |accessdate=21 November 2010}}</ref> while the reverse is true for lions entering waterways, as evidenced by the occasional lion claw found in crocodile stomachs.<ref name="Guggisberg">{{Cite book | last = Guggisberg | first = Charles Albert Walter |url =https://books.google.com/?id=92NuQgAACAAJ| title = Crocodiles: Their Natural History, Folklore, and Conservation| year = 1972 | page = 195 | isbn = 0-7153-5272-5 | publisher = David & Charles | location = Newton Abbot}}</ref>
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| ====Man-eating====
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| {{Main article|Man-eater}}
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| While lions do not usually hunt people, some (usually males) seem to seek out human prey; one well-publicised case includes the [[Tsavo maneaters]], where 28 officially recorded railway workers building the [[Uganda Railway|Kenya-Uganda Railway]] were taken by lions over nine months during the construction of a bridge over the [[Tsavo River]] in Kenya in 1898.<ref name=Patterson>{{cite book | last = Patterson |first=Bruce D. | year = 2004 | title = The Lions of Tsavo: Exploring the Legacy of Africa's Notorious Man-Eaters | publisher = McGraw-Hill | isbn = 0-07-136333-5}}</ref> The hunter who killed the lions wrote a book detailing the animals' predatory behaviour. The lions were larger than normal, lacked manes, and one seemed to suffer from tooth decay. The infirmity theory, including tooth decay, is not favoured by all researchers; an analysis of teeth and jaws of man-eating lions in museum collections suggests that while tooth decay may explain some incidents, prey depletion in human-dominated areas is a more likely cause of lion predation on humans.<ref>{{Cite journal |last=Patterson |first=Bruce D. |author2=Neiburger, Ellis J.|author3=Kasiki, Samuel M. |date=February 2003 |title=Tooth Breakage and Dental Disease as Causes of Carnivore–Human Conflicts |journal=Journal of Mammalogy |volume=84 |issue=1 |pages=190–96|doi=10.1644/1545-1542(2003)084<0190:TBADDA>2.0.CO;2}}</ref>
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| In their analysis of Tsavo and general man-eating, Kerbis Peterhans and Gnoske acknowledge that sick or injured animals may be more prone to man-eating, but that the behaviour is "not unusual, nor necessarily 'aberrant'" where the opportunity exists; if inducements such as access to livestock or human corpses are present, lions will regularly prey upon human beings. The authors note that the relationship is well-attested among other pantherines and primates in the paleontological record.<ref>{{Cite journal |url=http://www.man-eater.info/gpage6.html |archiveurl=https://web.archive.org/web/20071030031426/http://www.man-eater.info/gpage6.html |archivedate=30 October 2007 |accessdate=7 July 2007 |last=Peterhans |first=Julian C. Kerbis |author2=Thomas Patrick 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
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| }}</ref>
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| The lion's 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 – a number far exceeding the more famed "Tsavo" incidents of a century earlier. The incidents occurred near [[Selous Game Reserve|Selous National Park]] in [[Rufiji River|Rufiji District]] and in [[Lindi Region|Lindi Province]] near the Mozambican border. While the expansion of villagers into bush country is one concern, the authors argue that conservation policy must mitigate the danger because, in this case, conservation contributes directly to human deaths. Cases in Lindi have been documented where lions seize humans from the center of substantial villages.<ref name="Packer05">{{Cite journal |last=Packer |first=C. |author2=Ikanda, D. |author3=Kissui, B. |author4= Kushnir, H. |date=August 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}}</ref> Another study of 1,000 people attacked by lions in southern Tanzania between 1988 and 2009 found that the weeks following the [[full moon]] (when there was less moonlight) were a strong indicator of increased night attacks on people.<ref name="fullmoon">{{cite journal | title=Fear of Darkness, the Full Moon and the Nocturnal Ecology of African Lions | last=Packer |first=C | journal=[[PLoS ONE]] |date=July 2011 | volume=6 | issue=7 | doi=10.1371/journal.pone.0022285 |author2=Swanson A |author3=Ikanda D |author4=Kushnir H | editor1-last=Rands | editor1-first=Sean A. | pages=e22285}}</ref>
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| [[File:Lionsoftsavo2008.jpg|thumb|The Tsavo Man-Eaters on display in the [[Field Museum of Natural History]] in [[Chicago, Illinois]] ]]
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| Author Robert R. Frump wrote in ''The Man-eaters of Eden'' that Mozambican refugees regularly crossing Kruger National Park at night in South Africa are attacked and eaten by the lions; park officials have conceded that man-eating is a problem there. Frump believes thousands may have been killed in the decades after [[South Africa under apartheid|apartheid]] sealed the park and forced the refugees to cross the park at night. For nearly a century before the border was sealed, Mozambicans had regularly walked across the park in daytime with little harm.<ref>{{Cite book |last=Frump |first=RR |title=The Man-Eaters of Eden: Life and Death in Kruger National Park |year=2006 |publisher=The Lyons Press |isbn=1-59228-892-8}}</ref>
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| Packer estimates more than 200 Tanzanians are killed each year by lions, [[crocodile]]s, elephants, hippos, and snakes, and that the numbers could be double that amount, with lions thought to kill at least 70 of those. Packer has documented that between 1990 and 2004, lions attacked 815 people in Tanzania, killing 563. Packer and Ikanda are among the few [[conservation movement|conservationists]] who believe western conservation efforts must take account of these matters not just because of ethical concerns about human life, but also for the long term success of conservation efforts and lion preservation.<ref name="Packer05"/>
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| A man-eating lion was killed by game scouts in Southern Tanzania in April 2004. It is believed to have killed and eaten at least 35 people in a series of incidents covering several villages in the Rufiji Delta coastal region.<ref>{{Cite news|title=Toothache 'made lion eat humans' |last=Dickinson |first=Daniel |date=19 October 2004 |publisher=BBC News |url=http://news.bbc.co.uk/2/hi/africa/3756180.stm |accessdate=25 January 2011}}</ref> Dr Rolf D. Baldus, the [[GTZ]] wildlife programme coordinator, commented that it was likely that the lion preyed on humans because it had a large [[abscess]] underneath a [[molar (tooth)|molar]] that was cracked in several places. He further commented that "This lion probably experienced a lot of pain, particularly when it was chewing."<ref>{{Cite journal |last=Baldus |first=R |date=March 2006 |title=A man-eating lion (''Panthera leo'') from Tanzania with a toothache |journal=European Journal of Wildlife Research |volume=52 |issue= 1 |pages= 59–62|doi=10.1007/s10344-005-0008-0}}</ref> GTZ is the German development cooperation agency and has been working with the Tanzanian government on wildlife conservation for nearly two decades. As in other cases this lion was large, lacked a mane, and had a tooth problem.
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| The "All-Africa" record of man-eating generally is considered to be not Tsavo, but incidents in the early 1930s through the late 1940s in what was then [[Tanganyika]] (now Tanzania). [[George Gilman Rushby|George Rushby]], game warden and professional hunter, eventually dispatched the pride, which over three generations is thought to have killed and eaten 1,500 to 2,000 people in what is now [[Njombe]] district.<ref>{{Cite book |last=Rushby |first=George G. |title= No More the Tusker |year= 1965 |publisher=W. H. Allen |location=London}}</ref>
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| ===Reproduction and life cycle===
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| {{commons category|Mating lions}}
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| Most lionesses will have reproduced by the time they are four years of age.<ref name="Schaller29">[[#Schaller|Schaller]], p. 29.</ref> Lions do not [[mating|mate]] at any specific time of year, and the females are [[polyestrous]].<ref name="Schaller174">[[#Schaller|Schaller]], p. 174.</ref> As with other [[Cat's penis|cats' penises]], the male lion's penis has [[penile spines|spines that point backward]]. During withdrawal of the penis, the spines rake the walls of the female's [[vagina]], which may cause ovulation.<ref>{{Cite book|last=Asdell|first=Sydney A.|title=Patterns of mammalian reproduction |year=1993|origyear= 1964|publisher=Cornell University Press|location=Ithaca|isbn=978-0-8014-1753-5}}</ref> A lioness may mate with more than one male when she is [[Estrous cycle|in heat]].<ref name="Schaller142">[[#Schaller|Schaller]], p. 142.</ref>
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| The average gestation period is around 110 days,<ref name="Schaller174"/> the female giving birth to a litter of one to 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 rest of the pride. She will often hunt by herself while the cubs are still helpless, staying relatively close to the thicket or den where the cubs are kept.<ref name="Scott45">[[#Scott|Scott]], p. 45.</ref> The cubs themselves are born blind – their eyes do not open until roughly a week after birth. They weigh {{convert|1.2|–|2.1|kg|lb|abbr=on}} at birth and are almost helpless, beginning to crawl a day or two after birth and walking around three weeks of age.<ref name="Schaller143">[[#Schaller|Schaller]], p. 143.</ref> 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, to prevent scent from building up at a single den site and thus avoiding the attention of predators that may harm the cubs.<ref name="Scott45"/>
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| Usually, the mother does not integrate herself and her cubs back into the pride until the cubs are six to eight weeks old.<ref name="Scott45"/> Sometimes this introduction to pride life occurs earlier, however, particularly if other lionesses have given birth at about the same time. For instance, lionesses in a pride often synchronise their reproductive cycles so that they cooperate in the raising and suckling of the young (once the cubs are past the initial stage of isolation with their mother), who suckle indiscriminately from any or all of the nursing females in the pride. In addition to greater protection, the synchronization of births also has an advantage in that the cubs end up being roughly the same size, and thus have an equal chance of survival. If one lioness gives birth to a litter of cubs a couple of months after another lioness, for instance, then the younger cubs, being much smaller than their older brethren, usually are dominated by larger cubs at mealtimes – consequently, death by starvation is more common among the younger cubs.
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| In addition to starvation, cubs also face many other dangers, such as predation by jackals, hyenas, leopards, martial eagles, and snakes. Even buffaloes, should they catch the scent of lion cubs, often stampede toward the thicket or den where they are being kept, doing their best to trample the cubs to death while warding off the lioness. Furthermore, when one or more new males oust the previous male(s) associated with a pride, the conqueror(s) often kill any existing young cubs,<ref name="Packpus83">{{Cite journal |last=Packer |first=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}}</ref> perhaps because females do not become fertile and receptive until their cubs mature or die. All in all, as many as 80% of the cubs will die before the age of two.<ref>{{Cite book |last=Macdonald |first=David |year=1984 |title=The Encyclopedia of Mammals|publisher=Facts on File|location=New York |page=31|isbn= 0-87196-871-1}}</ref>
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| <gallery mode = packed heights = 110px>
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| File:Panthera leo -Ngorongoro Conservation Area, Tanzania -mating-8.jpg|Mating
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| File:PregnantLioness.jpg|A pregnant lioness (right)
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| File:Black-maned lion and cub 1.jpg|A tolerant black-maned male
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| File:Black-maned lion and cub 2.jpg|
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| File:Black-maned lion and cub 3.jpg|
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| File:Black-maned lion and cub 4.jpg|
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| </gallery>
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| When first introduced to the rest of the pride, the cubs initially lack confidence when confronted with adult lions 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. The tolerance of the male lions toward the cubs varies – sometimes, a male will patiently let the cubs play with his tail or his mane, whereas another may snarl and bat the cubs away.<ref>[[#Scott|Scott]], p. 46.</ref>
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| [[File:Male Lion and Cub Chitwa South Africa Luca Galuzzi 2004.JPG|thumb|Male lions are generally more likely to share food caught by the pride with cubs than with lionesses, but rarely share their own catches with others.]]
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| Weaning occurs after six to seven months. Male lions reach maturity at about 3 years of age and, at 4–5 years of age, are capable of challenging and displacing the adult male(s) associated with another pride. They begin to age and weaken between 10 and 15 years of age at the latest,<ref>{{Cite book |last=Crandall |first=Lee S. |title=The management of wild animals in captivity |year=1964 |publisher=University of Chicago Press |location=Chicago |oclc=557916}}</ref> if they have not already been critically injured while defending the pride (once ousted from a pride by rival males, male lions rarely manage a second take-over). This leaves a short window for their own offspring to be born and mature. If they are able to procreate as soon as they take over a pride, potentially, they may have more offspring reaching maturity before they also are displaced. A lioness often will attempt to defend her cubs fiercely from a usurping male, but such actions are rarely successful. He usually kills all of the existing cubs who are less than two years old. A lioness is weaker and much lighter than a male; success is more likely when a group of three or four mothers within a pride join forces against one male.<ref name="Packpus83"/>
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| Contrary to popular belief, it is not only males that are ousted from their pride to become nomads, although most females certainly do remain with their birth pride. However, when the pride becomes too large, the next generation of female cubs may be forced to leave to eke out their own territory. Furthermore, when a new male lion takes over the pride, subadult lions, both male and female, may be evicted.<ref>[[#Scott|Scott]], p. 68.</ref> Life is harsh for a female nomad. Nomadic lionesses rarely manage to raise their cubs to maturity, without the protection of other pride members.
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| Canadian researcher [[Bruce Bagemihl]] reports that both males and females may interact [[homosexuality in animals|homosexually]]. Lions are shown to be involved in group homosexual and courtship activities. Male lions will also head rub and roll around with each other before simulating sex together.<ref>{{Cite book |last=Bagemihl |first=Bruce |year=1999 |title=Biological Exuberance: Animal Homosexuality and Natural Diversity |location=New York |publisher=St. Martin's Press |isbn=0-312-19239-8| pages=302–05}}</ref>
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| ===Health===
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| [[File:TB-Infected Lion.png|thumb|left|Lion showing advanced signs of [[bovine tuberculosis]]]]
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| Although adult lions have no natural predators, evidence suggests that the majority die violently from humans or other lions.<ref name="Schaller183">[[#Schaller|Schaller]], p. 183.</ref> Lions often inflict serious injuries on each other, either members of different prides encountering each other in territorial disputes, or members of the same pride fighting at a kill.<ref name="Schaller1889">[[#Schaller|Schaller]], pp. 188–89.</ref> Crippled lions and lion cubs may fall victim to hyenas, leopards, or be trampled by buffalo or elephants, and careless lions may be maimed when hunting prey.<ref name="Schaller1890">[[#Schaller|Schaller]], pp. 189–90.</ref>
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| [[File:Lake-Nakuru-Lions-in-Tree.JPG|thumb|Lions seeking refuge from flies by climbing a tree.]]
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| Various species of [[tick]] commonly infest the ears, neck and groin regions of most lions.<ref name="Schaller184">[[#Schaller|Schaller]], p. 184.</ref><ref>{{Cite book |last=Yeoman |first=Guy Henry|author2=Walker, Jane Brotherton |title=The ixodid ticks of Tanzania|year=1967 |publisher=Commonwealth Institute of Entomology |location=London |oclc=955970}}</ref> Adult forms of several species of the tapeworm genus ''[[Taenia (tapeworm)|Taenia]]'' have been isolated from intestines, the lions having ingested larval forms from antelope meat.<ref>{{Cite journal |last=Sachs |first=R |year=1969|title=Untersuchungen zur Artbestimmung und Differenzierung der Muskelfinnen ostafrikanischer Wildtiere [Differentiation and species determination of muscle-cysticerci in East African game animals] |journal=Zeitschrift für tropenmedizin und Parasitologie |volume=20 |issue=1 |pages=39–50 |pmid=5393325|language=de}}</ref> Lions in the [[Ngorongoro Conservation Area|Ngorongoro Crater]] were afflicted by an outbreak of stable fly (''[[stable fly|Stomoxys calcitrans]]'') in 1962; this resulted in lions becoming covered in bloody bare patches and emaciated. Lions sought unsuccessfully to evade the biting flies by climbing trees or crawling into hyena burrows; many perished or emigrated as the population dropped from 70 to 15 individuals.<ref>{{Cite journal |last=Fosbrooke |first=Henry |year=1963 |title=The stomoxys plague in Ngorongoro |journal=East African Wildlife Journal |volume=1 |issue=6|pages=124–26 |doi=10.1111/j.1365-2028.1963.tb00190.x}}</ref> A more recent outbreak in 2001 killed six lions.<ref name="Nkwame06">{{Cite news| last=Nkwame|first= Valentine M | title=King of the jungle in jeopardy | publisher=The Arusha Times | date=9 September 2006 | url=http://www.arushatimes.co.tz/2006/36/features_10.htm | accessdate=4 September 2007}}</ref> Lions, especially in captivity, are vulnerable to the [[canine distemper]] virus (CDV), [[feline immunodeficiency virus]] (FIV), and [[feline infectious peritonitis]] (FIP).<ref name="zoos_encyclopedia"/> CDV is spread through domestic dogs and other [[carnivora|carnivores]]; a 1994 outbreak in [[Serengeti National Park]] resulted in many lions developing neurological symptoms such as seizures. During the outbreak, several lions died from pneumonia and [[encephalitis]].<ref>{{Cite journal |last=Roelke-Parker |first=M. E. |date=February 1996 |title=A canine distemper epidemic in Serengeti lions (''Panthera leo'') |journal= Nature|volume=379 |issue= 6564|pages=441–45 |doi=10.1038/379441a0|pmid=8559247|last2=Munson|first2=Linda|last3=Packer|first3=Craig|last4=Kock|first4=Richard|last5=Cleaveland|first5=Sarah|last6=Carpenter|first6=Margaret|last7=O'Brien|first7=Stephen J.|last8=Pospischil|first8=Andreas|last9=Hofmann-Lehmann|first9=Regina |displayauthors=9|first10=Hans|last10=Lutz|first11=George L. M.|last11=Mwanengele|first12=M. N.|last12=Mgasa|first13=G. A.|last13=Machange|first14=Brian A.|last14=Summers|first15=Max J. G.|last15=Appel}}</ref> FIV, which is similar to HIV while not known to adversely affect lions, is worrisome enough in its effect in domestic cats that the Species Survival Plan recommends systematic testing in captive lions. It occurs with high to endemic frequency in several wild lion populations, but is mostly absent from Asiatic and Namibian lions.<ref name="zoos_encyclopedia"/>''
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| ===Communication===
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| [[File:Lion cub with mother.jpg|thumb|right|Head rubbing and licking are common social behaviours within a pride]]
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| When resting, lion socialisation occurs through a number of behaviours, and the animal's expressive movements are highly developed. The most common peaceful tactile gestures are head rubbing and [[social licking]],<ref name="Schaller85">[[#Schaller|Schaller]], p. 85.</ref> which have been compared with grooming in primates.<ref>{{Cite book |editor=Desmond Morris|last=Sparks |first=J |title=Primate Ethology |chapter=Allogrooming in primates:a review |year=1967 |publisher=Aldine |location=Chicago |isbn=0-297-74828-9}} (2007 edition: 0-202-30826-X)</ref> Head rubbing – nuzzling one's forehead, face and neck against another lion – appears to be a form of greeting,<ref>{{Cite book |last=Leyhausen |first=Paul |title=Verhaltensstudien an Katzen|edition= 2nd |year=1960 |publisher=Paul Parey |location=Berlin |isbn=3-489-71836-4 |language=de}}</ref> as it is seen often after an animal has been apart from others, or after a fight or confrontation. Males tend to rub other males, while cubs and females rub females.<ref name="Schaller858">[[#Schaller|Schaller]], pp. 85–88.</ref> Social licking often occurs in tandem with head rubbing; it is generally mutual and the recipient appears to express pleasure. The head and neck are the most common parts of the body licked, which may have arisen out of utility, as a lion cannot lick these areas individually.<ref name="Schaller8891">[[#Schaller|Schaller]], pp. 88–91.</ref>
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| {{sound|filename=Lion raring-sound1TamilNadu178.ogg|title=Lion roar|description=A lion in captivity roaring}}
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| Lions have an array of facial expressions and body postures that serve as visual gestures.<ref name="Schaller92102">[[#Schaller|Schaller]], pp. 103–115.</ref> Their repertoire of vocalisations is also large; variations in intensity and pitch, rather than discrete signals, appear central to communication. Most lion vocals are variations of growling/snarling, miaowing and roaring. Other sounds produced include purring, puffing, bleating and humming.<ref name="Schaller10313"/> Lions tend to [[roar (animal)|roar]] in a very characteristic manner, starting with a few deep, long roars that trail off into a series of shorter ones.<ref>{{Cite journal |url=http://roberteklund.info/pdf/Ananthakrishnan_et_al_2011_LionRoars.pdf |title=An acoustic analysis of lion roars. II: Vocal tract characteristics |last=Ananthakrishnan |first=G. |last2=Eklund |first2=Robert |last3=Peters |first3=Gustav |last4=Mabiza |first4=Evans |journal=Speech, Music and Hearing Quarterly Progress and Status Report TMH-QPSR |volume=51 |page=5 |year=2011}}</ref><ref>{{Cite journal |url=http://roberteklund.info/pdf/Eklund_et_al_2011_LionRoars.pdf |title=An acoustic analysis of lion roars. I: Data collection and spectrogram and waveform analyses |last=Eklund |first=Robert |last2=Peters |first2=Gustav |last3=Ananthakrishnan |first3=G |last4=Mabiza |first4=Evans |journal=Speech, Music and Hearing Quarterly Progress and Status Report TMH-QPSR |volume=51 |page=1 |year=2011}}</ref>
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| They most often roar at night; the sound, which can be heard from a distance of {{convert|8|km}}, is used to advertise the animal's presence.<ref name="Schaller10313">[[#Schaller|Schaller]], pp. 103–113.</ref> Lions have the loudest roar of any big cat.
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| ==Distribution and habitat==
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| [[File:India Animals.jpg|thumb|Two male [[Asiatic lions]]. The wild population of the endangered Asiatic lions is restricted to the [[Gir Forest National Park]] in western India.<ref>{{Cite book
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| |last = Miller
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| |first = Brian
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| |title = Endangered animals: a reference guide to conflicting issues
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| |publisher = Greenwood Publishing Group|year= 2000
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| |isbn = 0-313-30816-0}}</ref>]]
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| In Africa, lions can be found in savanna grasslands with scattered ''[[Acacia]]'' trees, which serve as shade;<ref>{{Cite book |last=Rudnai|first= Judith A. |title=The social life of the lion |year=1973 |publisher=s.n. |location=Wallingford |isbn=0-85200-053-7 }}</ref> their habitat in India is a mixture of dry savanna forest and very dry deciduous scrub forest.<ref>{{cite web | title=The Gir—Floristic | work=Asiatic Lion Information Centre | publisher=Wildlife Conservation Trust of India | year=2006 | url=http://www.asiaticlion.org/gir-floristic.htm | accessdate=14 September 2007}}</ref> The habitat of lions originally spanned the southern parts of [[Eurasia]], ranging from [[Greece]] to India, and most of Africa except the central [[rainforest]]-zone and the [[Sahara]] desert. [[Herodotus]] reported that lions had been common in Greece in 480 BC; they attacked the baggage camels of the Persian king [[Xerxes I of Persia|Xerxes]] on his march through the country. [[Aristotle]] considered them rare by 300 BC. By 100 AD they were extirpated.<ref name="Schaller5">[[#Schaller|Schaller]], p. 5.</ref> A population of [[Asiatic lion]]s survived until the tenth century in the [[Caucasus]], their last European outpost.<ref>{{Cite book | last=Heptner| first= V. G.|author2=A. A. Sludskii | year=1989| title=Mammals of the Soviet Union: Volume 1, Part 2: Carnivora (Hyaenas and Cats)|publisher=Amerind| location=New York| isbn=90-04-08876-8}}</ref>
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| The species was eradicated from [[Palestine (region)|Palestine]] by the [[Middle Ages]] and from most of the rest of Asia after the arrival of readily available firearms in the eighteenth century. Between the late nineteenth and early twentieth centuries, they became extinct in North Africa and Southwest Asia. By the late nineteenth century, the lion had disappeared from [[Turkey]] and most of northern India,<ref name=zoos_encyclopedia /><ref>{{cite web |title=Past and present distribution of the lion in North Africa and Southwest Asia. |year=2001 |publisher=Asiatic Lion Information Centre |url=http://www.asiatic-lion.org/distrib.html |accessdate=1 June 2006|archiveurl=https://web.archive.org/web/20080510060942/http://www.asiatic-lion.org/distrib.html|archivedate=10 May 2008}}</ref> while the last sighting of a live Asiatic lion in [[Iran]] was in 1941 (between [[Shiraz]] and Jahrom, [[Fars Province]]), although the corpse of a lioness was found on the banks of the [[Karun]] river, [[Khūzestān Province]] in 1944. There are no subsequent reliable reports from [[Iran]].<ref name="simba"/> The subspecies now survives only in and around the [[Gir Forest National Park|Gir Forest]] of northwestern India.<ref name="Asiaticweb"/> Approximately 500 lions live in the area of the {{convert|1412|km2|sqmi|abbr=on}} sanctuary in the state of [[Gujarat]], which covers most of the forest. Their numbers have increased from 180 to 523 animals mainly because the natural prey species have recovered.<ref name="Singh, Gibson"/><ref>{{cite web | title=Asiatic Lion—Population | work=Asiatic Lion Information Centre | publisher=Wildlife Conservation Trust of India | year=2006 | url=http://www.asiaticlion.org/population-gir-forests.htm | accessdate=15 September 2007}}</ref>
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| ==Population and conservation status==
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| {{Main article|Lion hunting}}
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| [[File:Lion Gir.jpg|thumb|The Asiatic lion, whose habitat once ranged from the Mediterranean to north-west Indian subcontinent, is today found only in the [[Gir Forest]] of [[Gujarat]], India. It was estimated that only 523 Asiatic lions survive in the wild.<ref>{{cite web |url=http://www.outlookindia.com/article.aspx?220509 |title=The Mane Don't Fit |last=Desai |first=Darshan |work=Outlook India Magazine |publisher=Outlookindia.com |date=23 June 2003 |accessdate=21 November 2010}}</ref>]]
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| Most lions now live in eastern and southern Africa, and their numbers there are rapidly decreasing, with an estimated 30–50% decline per 20 years in the late half of the twentieth century.<ref name=IUCN/> Estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004,<ref>{{Cite journal |last=Bauer |first=H. |last2=Van Der Merwe |first2=S. |year= 2002|title=The African lion database |journal=Cat news |volume=36 |pages=41–53}}</ref><ref>{{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}}</ref> down from early 1990s estimates that ranged as high as 100,000 and perhaps 400,000 in 1950. Primary causes of the decline include disease and human interference.<ref name=IUCN/> Habitat loss and conflicts with humans are considered the most significant threats to the species.<ref name="awf">{{cite web |title=AWF Wildlife: Lion|url=http://www.awf.org/content/wildlife/detail/lion|publisher=[[African Wildlife Foundation]]|accessdate=29 August 2007}}</ref><ref>{{cite web |title=NATURE. The Vanishing Lions |publisher=PBS |url=http://www.pbs.org/wnet/nature/episodes/the-vanishing-lions/introduction/545/ |accessdate=25 January 2011}}</ref> The remaining populations are often geographically isolated from one another, which can lead to [[inbreeding]], and consequently, reduced [[genetic diversity]]. Therefore, the lion is considered a [[vulnerable species]] by the [[International Union for Conservation of Nature]], while the Asiatic subspecies is [[endangered]].<ref>{{IUCN2010.4|assessor=Breitenmoser, U.|assessor2=Mallon, D.P.|assessor3=Ahmad Khan, J.|assessor4=Driscoll, C.|last-assessor-amp=yes|year=2008|id=15952|title=Panthera leo ssp. persica|downloaded=13 December 2010}}</ref> The lion population in the region of West Africa is isolated from lion populations of Central Africa, with little or no exchange of breeding individuals. The number of mature individuals in West Africa is estimated by two separate recent surveys at 850–1,160 (2002/2004). There is disagreement over the size of the largest individual population in West Africa: the estimates range from 100 to 400 lions in [[Burkina Faso]]'s [[Arly-Singou]] ecosystem.<ref name=IUCN/> Another population in northwestern Africa is found in [[Waza National Park]], where approximately 14–21 animals persist.<ref>{{Cite journal |last=Tumenta |first=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}}</ref>
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| [[File:The Big Game of Africa (1910) - Male lion Sotik Plains May 1909.png|left|thumb|Black maned male lion, shot in the Sotik Plains, Kenya (May 1909)]]
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| Conservation of both African and Asian lions has required the setup and maintenance of national parks and game reserves; among the best known are [[Etosha National Park]] in Namibia, [[Serengeti National Park]] in Tanzania, and [[Kruger National Park]] in eastern South Africa. The [[Ewaso Lions]] Project protects lions in the [[Samburu National Reserve]], [[Buffalo Springs National Reserve]] and [[Shaba National Reserve]] of the [[Ewaso Ng'iro]] ecosystem in Northern [[Kenya]].<ref>{{cite web
| |
| |url = http://www.theeastafrican.co.ke/magazine/-/434746/657312/-/15kjo9jz/-/index.html
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| |title = Winning with lions
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| |publisher = The East African Magazine
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| }}</ref> Outside these areas, the issues arising from lions' interaction with livestock and people usually results in the elimination of the lions.<ref>{{cite web | last=Roach | first=John | title=Lions Vs. Farmers: Peace Possible? | work=National Geographic News | publisher=National Geographic | date=16 July 2003 | url=http://news.nationalgeographic.com/news/2003/07/0716_030716_lions.html | accessdate=1 September 2007}}</ref> In India, the last refuge of the Asiatic lion is the {{convert|1412|km2|sqmi|abbr=on}} [[Gir Forest National Park]] in western India, which had approximately 180 lions in 1974 and about 400 in 2010.<ref name="Singh, Gibson"/> As in Africa, numerous human habitations are close by with the resultant problems between lions, livestock, locals and wildlife officials.<ref name="C4WDefault-10.1046/j.1523-1739.1994.08020501.x">{{cite journal |url=http://onlinelibrary.wiley.com/doi/10.1046/j.1523-1739.1994.08020501.x/abstract;jsessionid=FF3227E78A0339404CEBBA045818D361.f04t04 |title=Lion‐Human Conflict in the Gir Forest, India |date=1 June 1994 |publisher=Blackwell Science Inc. |issn=1523-1739 |accessdate=1 May 2014 |doi=10.1046/j.1523-1739.1994.08020501.x |issue=2 |volume=8 |pages=501–7 |journal=Conservation Biology |first1=Vasant K. |last1=Saberwal |first2=James P. |last2=Gibbs |first3=Ravi |last3=Chellam |first4=A. J. T. |last4=Johnsingh}}</ref> The establishment of a second independent population of [[Asiatic lion]]s is planned at the [[Kuno Wildlife Sanctuary]] in the Indian state of [[Madhya Pradesh]].<ref>{{Cite news |last=Santoshi |first=Neeraj |date=25 February 2016 |title=Lions not in, Madhya Pradesh moves to populate Kuno with tigers |journal=Hindustan Times |url=http://indiatoday.intoday.in/story/girs-lions-may-soon-be-shifted-to-palpur-kuno-wildlife-sanctuary-in-mp/1/711359.html |accessdate=1 September 2016}}</ref>
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| The former popularity of the Barbary lion as a zoo animal has meant that scattered lions in captivity are likely to be descended from Barbary lion stock. This includes lions at [[Port Lympne Zoo|Port Lympne Wild Animal Park]] in Kent, England that are descended from animals owned by the [[King of Morocco]].<ref>{{cite web |url=http://www.bigcatrescue.org/barbary_lion_news.htm |archiveurl=https://web.archive.org/web/20051217091555/http://www.bigcatrescue.org/barbary_lion_news.htm |archivedate=17 December 2005 |title=Barbary Lion News |accessdate=24 September 2007}}</ref> Another eleven animals believed to be Barbary lions were found in [[Addis Ababa]] zoo, descendants of animals owned by [[Haile Selassie I of Ethiopia|Emperor Haile Selassie]]. WildLink International, in collaboration with [[Oxford University]], launched their ambitious International [[Barbary lion#The Barbary Lion Project|Barbary Lion Project]] with the aim of identifying and breeding Barbary lions in captivity for eventual reintroduction into a national park in the [[Atlas Mountains]] of [[Morocco]].<ref name="yamaguchi-haddane"/>
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| Following the discovery of the decline of lion population in Africa, several coordinated efforts involving lion [[Wildlife conservation|conservation]] have been organised in an attempt to stem this decline.
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| Lions are one species included in the [[Species Survival Plan]], a coordinated attempt by the [[Association of Zoos and Aquariums]] to increase its chances of survival. The plan was originally started in 1982 for the Asiatic lion, but was suspended when it was found that most Asiatic lions in North American zoos were not [[genetic pollution|genetically pure]], having been hybridised with African lions. The African lion plan started in 1993, focusing especially on the South African subspecies, although there are difficulties in assessing the genetic diversity of captive lions, since most individuals are of unknown origin, making maintenance of genetic diversity a problem.<ref name="zoos_encyclopedia"/>
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| In 2015, a population of lions that was previously believed extirpated was filmed in the Alatash National Park, Ethiopia, close to the Sudanese border. This population may possibly number up to 200 animals.<ref>{{cite news|publisher=New Scientist|title=Hidden population of up to 200 lions found in remote Ethiopia|date=1 February 2016|url=https://www.newscientist.com/article/2075740-hidden-population-of-up-to-200-lions-found-in-remote-ethiopia/|accessdate=2 February 2016}}</ref><ref>{{cite news|date=1 February 2016|url=http://www.bbc.com/news/world-africa-35460573|title=Lions rediscovered in Ethiopia's Alatash National Park|publisher=BBC News|accessdate=1 February 2016}}</ref>
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| ===In captivity===
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| [[File:Lion cubs (4031355883).jpg|thumb|Lion cubs at Clifton Zoological Gardens, England, 1854]]
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| [[File:MP-panthera leo krugeri 8.jpg|thumb|Male African lion of the Transvaal subspecies (''P. l. krugeri'')]]
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| Lions are part of a group of exotic animals that are the core of [[zoo]] exhibits since the late eighteenth century; members of this group are invariably large vertebrates and include elephants, rhinoceroses, hippopotamuses, large primates, and other big cats; zoos sought to gather as many of these species as possible.<ref name="dc81">[[#Courcy|de Courcy]], p. 81</ref> Although many modern zoos are more selective about their exhibits,<ref>[[#Courcy|de Courcy]], p. 82</ref> there are more than 1,000 African and 100 Asiatic lions in zoos and wildlife parks around the world. They are considered an ambassador species and are kept for tourism, education and conservation purposes.<ref name="WAZA">{{cite web |last1= Dollinger | first1= P |last2= Geser | first2= S| title= Lion: In the Zoo (subpage) |work=Visit the Zoo|publisher=WAZA (World Association of Zoos and Aquariums)|url= http://www.waza.org/en/zoo/visit-the-zoo/cats-1254385523/panthera-leo| accessdate=5 April 2011}}</ref> Lions can reach an age of over 20 years in captivity; Apollo, a resident lion of [[Honolulu Zoo]] in [[Honolulu]], Hawaii, died at age 22 in August 2007. His two sisters, born in 1986, were still alive in August 2007.<ref>{{Cite news | last= Aguiar |first= Eloise|date=August 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| accessdate=4 September 2007}}</ref> Breeding programs need to note origins to avoid breeding different subspecies and thus reducing conservation value.<ref>{{Cite book | url= https://books.google.com/?id=vLZKnsCk89wC&pg=PA437|accessdate=12 September 2010| last1= Frankham |first1= Richard |last2= Ballou |first2= Jonathan |last3= Briscoe |first3= David |title= Introduction to Conservation Genetics |page=437|publisher=Cambridge University Press| year= 2009| isbn= 0-521-70271-2| location= Cambridge, England}}</ref> However, several Asiatic-African lion crosses have been bred.<ref name="AviseHamrick1996">{{Cite book |last1=Avise|first1=J. C.|last2=Hamrick|first2=J. L.|title=Conservation Genetics|date=31 January 1996|publisher=Springer Science & Business Media|isbn=9780412055812|page=67|quote=Furthermore, when Asiatic lions were inadvertently bred to African lion subspecies in North America, the fecundity, reproductive success, and spermatozoal development improved dramatically (Box 3.3; O'Brien et al., 1987b).}}</ref>
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| At the ancient Egyptian cities of [[Taremu]] and [[Per-Bast]] were temples to the lioness goddesses of Egypt, [[Sekhmet]] and [[Bast (goddess)|Bast]] and at Taremu there was a temple to the son of the deity, [[Maahes]] the lion prince, where live lions were kept and allowed to roam within his temple. The Greeks called the city Leontopolis, the "City of Lions" and documented that practice.
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| Lions were kept and bred by [[Assyria]]n kings as early as 850 BC,<ref name="Schaller5"/> and [[Alexander the Great]] was said to have been presented with tame lions by the [[Malhi]] of northern India.<ref>{{Cite book |last= Smith |first=Vincent Arthur|title= The Early History of India|year=1924|page=97|publisher=Clarendon Press|location=Oxford|isbn=}}</ref> Later in [[Roman Empire|Roman]] times, lions were kept by emperors to take part in the gladiator arenas or for executions (see [[bestiarii]], [[damnatio ad bestias]], and [[venatio]]). Roman notables, including [[Lucius Cornelius Sulla|Sulla]], [[Pompey]], and [[Julius Caesar]], often ordered the mass slaughter of hundreds of lions at a time.<ref>{{Cite book | first= Thomas |last= Wiedemann | title= Emperors and Gladiators |publisher= Routledge |year= 1995 |page= 60 | isbn= 0-415-12164-7 }}</ref> In the East, lions were tamed by Indian princes, and [[Marco Polo]] reported that [[Kublai Khan]] kept lions inside.<ref>[[#Baratay|Baratay & Hardouin-Fugier]], p. 17.</ref> The first European "zoos" spread among noble and royal families in the thirteenth century, and until the seventeenth century were called [[seraglio]]s; at that time, they came to be called [[menagerie]]s, an extension of the [[cabinet of curiosities]]. They spread from France and Italy during the [[Renaissance]] to the rest of Europe.<ref>[[#Baratay|Baratay & Hardouin-Fugier]], pp. 19–21, 42.</ref> In England, although the seraglio tradition was less developed, lions were [[Tower of London#Menagerie|kept at the Tower of London]] in a seraglio established by [[John of England|King John]] in the thirteenth century,<ref>[[#Baratay|Baratay & Hardouin-Fugier]], p. 20.</ref><ref>{{cite web |last=Owen|first=James|title=Medieval Lion Skulls Reveal Secrets of Tower of London "Zoo"|work=National Geographic Magazine|publisher=National Geographic|date=3 November 2005|url=http://news.nationalgeographic.com/news/2005/11/1103_051103_tower_lions.html| accessdate=5 September 2007}}</ref> 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|Woodstock]], near [[Oxford]]; where lions had been stocked according to [[William of Malmesbury]].<ref name="Blunt15">[[#Blunt|Blunt]], p. 15.</ref>
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| Seraglios served as expressions of the nobility's power and wealth. Animals such as big cats and elephants, in particular, symbolised power, and would be pitted in fights against each other or domesticated animals. By extension, menageries and seraglios served as demonstrations of the dominance of humanity over nature. Consequently, the defeat of such natural "lords" by a cow in 1682 astonished the spectators, and the flight of an elephant before a rhinoceros drew jeers. Such fights would slowly fade out in the seventeenth century with the spread of the menagerie and their appropriation by the commoners. The tradition of keeping big cats as pets would last into the nineteenth century, at which time it was seen as highly eccentric.<ref>[[#Baratay|Baratay & Hardouin-Fugier]], pp. 24–28.</ref>
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| [[File:Albrecht Dürer - Two seated lions - Google Art Project.jpg|thumb|[[Albrecht Dürer]], lions sketch. (c. 1520)]]
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| The presence of lions at the Tower of London was intermittent, being restocked when a monarch or his consort, such as [[Margaret of Anjou]] the wife of [[Henry VI of England|Henry VI]], either sought or were given animals. Records indicate they were kept in poor conditions there in the seventeenth century, in contrast to more open conditions in [[Florence]] at the time.<ref name="Blunt16">[[#Blunt|Blunt]], p. 16.</ref> The menagerie was open to the public by the eighteenth century; admission was a sum of three half-pence or the supply of a cat or dog for feeding to the lions.<ref name="Blunt17">[[#Blunt|Blunt]], p. 17.</ref> A rival menagerie at the [[Exeter Exchange]] also exhibited lions until the early nineteenth century.<ref>[[#Courcy|de Courcy]], pp. 8–9.</ref> The Tower menagerie was closed down by [[William IV of the United Kingdom|William IV]],<ref name="Blunt17"/> and animals transferred to the [[London Zoo]], which opened its gates to the public on 27 April 1828.<ref name="Blunt32">[[#Blunt|Blunt]], p. 32.</ref>
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| The wild animals trade flourished alongside improved colonial trade of the nineteenth century. Lions were considered fairly common and inexpensive. Although they would barter higher than tigers, they were less costly than larger, or more difficult to transport animals such as the giraffe and hippopotamus, and much less than [[giant panda]]s.<ref>[[#Baratay|Baratay & Hardouin-Fugier]], p. 122.</ref> Like other animals, lions were seen as little more than a natural, boundless commodity that was mercilessly exploited with terrible losses in capture and transportation.<ref>[[#Baratay|Baratay & Hardouin-Fugier]], pp. 114, 117.</ref> The widely reproduced imagery of the heroic hunter chasing lions would dominate a large part of the century.<ref>[[#Baratay|Baratay & Hardouin-Fugier]], p. 113.</ref> Explorers and hunters exploited a popular [[Manichaeism|Manichean]] division of animals into "good" and "evil" to add thrilling value to their adventures, casting themselves as heroic figures. This resulted in big cats, always suspected of being man-eaters, representing "both the fear of nature and the satisfaction of having overcome it."<ref>[[#Baratay|Baratay & Hardouin-Fugier]], pp. 173, 180–83.</ref>
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| [[File:Lion - melbourne zoo.jpg|thumb|Lion at [[Melbourne Zoo]] enjoying an elevated grassy area with some tree shelter]]
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| Lions were kept in cramped and squalid conditions at London Zoo until a larger lion house with roomier cages was built in the 1870s.<ref name="Blunt208">[[#Blunt|Blunt]], p. 208.</ref> Further changes took place in the early twentieth century, when [[Carl Hagenbeck]] designed enclosures more closely resembling a natural habitat, with concrete 'rocks', more open space and a moat instead of bars. He designed lion enclosures for both [[Melbourne Zoo]] and Sydney's [[Taronga Zoo]], among others, in the early twentieth century. Though his designs were popular, the old bars and cage enclosures prevailed until the 1960s in many zoos.<ref name="dc69">[[#Courcy|de Courcy]], p. 69.</ref> In the later decades of the twentieth century, larger, more natural enclosures and the use of [[mesh|wire mesh]] or [[laminated glass]] instead of lowered dens allowed visitors to come closer than ever to the animals, with some attractions even placing the den on ground higher than visitors, such as the Cat Forest/Lion Overlook of [[Oklahoma City Zoo and Botanical Garden|Oklahoma City Zoological Park]].<ref name="zoos_encyclopedia"/> Lions are now housed in much larger naturalistic areas; modern recommended guidelines more closely approximate conditions in the wild with closer attention to the lions' needs, highlighting the need for dens in separate areas, elevated positions in both sun and shade where lions can sit and adequate ground cover and drainage as well as sufficient space to roam. There have also been instances where a lion was kept by a private individual, such as the lioness [[Elsa the Lioness|Elsa]], who was raised by [[George Adamson]] and his wife [[Joy Adamson]] and came to develop a strong bond with them, particularly the latter. The lioness later achieved fame, her life being documented in a series of books and films.
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| ===Baiting and taming===
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| {{Main article|Lion-baiting|Lion taming}}
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| [[File:Lion tamer (LOC pga.03749).jpg|left|thumb|Nineteenth-century [[etching]] of a lion tamer in a cage of lions]]
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| Lion-baiting is a [[blood sport]] involving the [[animal-baiting|baiting]] of lions in combat with other animals, usually dogs. Records of it exist in ancient times through until the seventeenth century. It was finally banned in [[Vienna]] by 1800 and England in 1835.<ref>{{Cite book |last=Hone |first= William |editor= Kyle Grimes|title= The Every-Day Book |origyear=1825–1826 |url=http://www.uab.edu/english/hone/etexts/edb/day-pages/207-july26.html |accessdate=25 January 2011|year=2004 |publisher=University of Alabama at Birmingham |page=26 |chapter=July |archiveurl= https://web.archive.org/web/20080624080642/http://www.uab.edu/english/hone/etexts/edb/day-pages/207-july26.html |archivedate= 24 June 2008}}</ref><ref>{{Cite journal |last=Blaisdell |first=Warren H. |date=November 1997 |title= How A Lion Fight Caused England To Stop The Breeding of Both Ring And Pit Bulldogs|journal=American Bulldog Review |volume=3 |issue=4 |url=http://www.american-bulldog.com/how_a_lion.htm |archiveurl=https://web.archive.org/web/20080424001716/http://www.american-bulldog.com/how_a_lion.htm |archivedate=24 April 2008 |accessdate=5 September 2007}}</ref>
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| Lion taming refers to the practice of taming lions for entertainment, either as part of an established [[circus]] or as an individual act, such as [[Siegfried & Roy]]. The term is also often used for the taming and display of other big cats such as tigers, leopards, and [[cougar]]s. The practice was pioneered in the first half of the nineteenth century by Frenchman Henri Martin and American [[Isaac A. Van Amburgh|Isaac Van Amburgh]] who both toured widely, and whose techniques were copied by a number of followers.<ref name = "baratay187"/> 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 that Amburgh quickly borrowed. These acts eclipsed [[equestrianism]] acts as the central display of circus shows, but truly entered public consciousness in the early twentieth century with cinema. In demonstrating the superiority of human over animal, lion taming served a purpose similar to animal fights of previous centuries.<ref name="baratay187">[[#Baratay|Baratay & Hardouin-Fugier]], p. 187.</ref> The ultimate proof of a tamer's dominance and control over a lion is demonstrated by placing his head in the lion's mouth. The now iconic lion tamer's chair was possibly first used by American [[Clyde Beatty]] (1903–1965).<ref>{{Cite book | first=David | last=Feldman | authorlink=David Feldman (author) | year=1993 | title=How Does Aspirin Find a Headache? | publisher=HarperCollins | isbn= 0-06-016923-0 }}</ref>
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| ==Cultural significance==
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| {{Details |Cultural depictions of lions}}
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| [[File:Georgia (15).jpg|thumb|left|Georgian lion from [[Colchis]]]]
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| The lion has been an icon for humanity for thousands of years, appearing in cultures across Europe, Asia, and Africa. Despite incidents of attacks on humans, lions have enjoyed a positive depiction in culture as strong and noble. A common depiction is their representation as "king of the jungle" or "king of beasts"; hence, the lion has been a popular symbol of royalty and stateliness,<ref name="Garai73">{{Cite book |last = Garai |first=Jana |title=The Book of Symbols |year=1973 |publisher=Simon & Schuster |location=New York |isbn = 0-671-21773-9}}</ref> as well as a symbol of bravery; it is featured in several [[Aesop's Fables|fables]] of the sixth century BC [[Ancient Greece|Greek]] storyteller [[Aesop]].<ref>{{Cite book | author = Aesop | editor-last = Gibbs | editor-first = L |title= Fables |series=Oxford World's Classics |year=2002 |publisher = Oxford University Press |location=Oxford |isbn= 0-19-284050-9}}</ref>
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| Representations of lions date back to the early [[Upper Paleolithic]]. The [[löwenmensch figurine|lioness-headed]] ivory carving from Vogelherd cave in the [[Swabian Alb]] in southwestern Germany, dubbed ''Löwenmensch'' (lion-human) in German. The sculpture has been determined to be at least 32,000 years old from the [[Aurignacian]] culture,<ref name="BurgerJ-Molecular-phylogeny"/> but it may date to as early as 40,000 years ago.<ref>Martin Bailey [http://www.theartnewspaper.com/articles/Ice-Age-iLion-Mani-is-worlds-earliest-figurative-sculpture/28595 Ice Age Lion Man is world’s earliest figurative sculpture] The Art Newspaper, 31 January 2013, accessed 1 February 2013. {{webarchive |url=https://web.archive.org/web/20130508104713/http://www.theartnewspaper.com/articles/Ice-Age-iLion-Mani-is-worlds-earliest-figurative-sculpture/28595 |date=8 May 2013 }}</ref> The sculpture has been interpreted as [[anthropomorphic]], giving human characteristics to an animal, however, it also may represent a [[deity]]. Two lions were depicted mating in the Chamber of Felines in 15,000-year-old [[Paleolithic]] [[cave painting]]s in the [[Lascaux]] caves. Cave lions also are depicted in the [[Chauvet Cave]], discovered in 1994; this has been dated at 32,000 years of age,<ref name="Packer00" /> though it may be of similar or younger age to Lascaux.<ref>{{cite conference | first = Christian | last= Züchner | title= Grotte Chauvet Archaeologically Dated |date=September 1998 | location = International Rock Art Congress IRAC '98, Vila Real, Portugal | url= http://www.rupestre.net/tracce/12/chauv.html | accessdate =6 November 2010}}</ref>
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| [[File:Lion Darius Palace Louvre Sb3298.jpg|thumb|A lion depicted on a decorative panel from [[Darius I the Great]]'s palace of the [[Achaemenid Empire|Persian Empire]] (550–330 BC).]]
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| In Africa, cultural views of the lion have varied by region. In some cultures, the lion symbolises power and royalty and some powerful rulers had the word "lion" in their nickname. For example, Marijata of the [[Mali Empire]] (c. 1235–c. 1600) was given the name "Lion of Mali".<ref name="Lynch"/> Njaay, the legendary founder of the [[Waalo]] kingdom (1287–1855), is said to have been raised by lions and returned to his people part-lion to unite them using the knowledge he learned from the beasts. In much of West Africa, to be compared to a lion was considered to be one of the greatest compliments. The social hierarchies of their societies where connected to the animal kingdom and the lion represented the top class. In parts of West and East Africa, the lion is associated with healing and is seen as the link between the [[Clairvoyance|seers]] and the supernatural. In other East African traditions, the lion is the symbol of laziness.<ref>{{cite book|author1=Hogarth, C. |author2=Butler, N. |year=2004|contribution=Animal Symbolism (Africa)|title=Shamanism: An Encyclopedia of World Beliefs, Practices, and Culture, Volume 1|editor=Walter, M. N.|pages=3–6|isbn=1-57607-645-8}}</ref> In many [[Traditional African religion|folktales]], lions are portrayed as having low intelligence and are easily tricked by other animals.<ref name="Lynch">{{cite book|author=Lynch, P. A.|year=2004|title=African Mythology A to Z|publisher=Infobase Publishing|page=63|isbn=0-8160-4892-4}}</ref>
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| [[Ancient Egypt]] venerated the lioness (the fierce hunter) as their [[List of war deities|war deities]] and among those in the [[Egyptian pantheon]] are, [[Bastet|Bast]], [[Mafdet]], [[Menhit]], [[Pakhet]], [[Sekhmet]], [[Tefnut]], and the [[Sphinx]];<ref name="Garai73"/> The [[Nemean lion]] was symbolic in Ancient Greece and Rome, represented as the constellation and zodiac sign [[Leo (astrology)|Leo]], and described in mythology, where its skin was borne by the hero [[Heracles]].<ref>{{Cite book |last=Graves |first=R |authorlink=Robert Graves |title=Greek Myths |year=1955|publisher=Penguin |location=London |isbn=0-14-001026-2|pages=465–69 | chapter =The First Labour: The Nemean Lion}}</ref>
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| [[File:6 . simha vahana in front of the sanctum ; sahadeva ratha also seen behind.JPG|thumbnail|right|A lion carving in [[Mahabalipuram]], [[Tamil Nadu]], [[India]] ([[UNESCO]] [[World Heritage Site]])]]
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| The lion was a prominent symbol in ancient [[Mesopotamia]] (from [[Sumer]] up to [[Assyria]]n and [[Babylonia]]n times), where it was strongly associated with kingship.<ref>{{Cite journal |last=Cassin |first=Elena |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 | url = http://www.persee.fr/web/revues/home/prescript/article/rhr_0035-1423_1981_num_198_4_4828 |format=PDF | accessdate = 3 December 2009|language=French|doi=10.3406/rhr.1981.4828}}</ref> The classic Babylonian lion motif, found as a statue, carved or painted on walls, is often referred to as the ''striding lion of Babylon''. It is in Babylon that the biblical [[Daniel (biblical figure)|Daniel]] is said to have been delivered from the lion's den.<ref>[[Book of Daniel|Daniel]] 6</ref> The ''[[Lion Hunt of Ashurbanipal]]'' is a famous sequence of [[Assyrian palace reliefs]] from c. 640 BC, now in the [[British Museum]].
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| [[File:Daniellion.jpg|thumb|left|[[Daniel in the lions' den]] is an account in [[Book of Daniel|Daniel]] 6 in the Bible]]
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| In the [[Talmud]], ''Chullin'' 59b, Rabbi [[Joshua ben Hananiah]] tells Emperor [[Hadrian]] about the giant lion of the forest of Bei Ilai called the ''[[Jewish mythology#The Keresh and the Tigris of the Bei Ilai|Tigris]]'', a lion so huge that the space between its ears measures 9 [[cubit]]s. The emperor asks the rabbi to call forth this lion. He reluctantly agrees. At a distance of 400 [[Cubit (Biblical)|parasangs]] from Rome it roars, and all pregnant women [[Miscarriage|miscarry]] and all the walls of Rome fall down. Then it comes to 300 parasangs and roars, and all the front teeth and molars of Roman men fall out, and even the emperor himself falls from his throne. He begs the rabbi to send it back. The rabbi prays and it returns to its place.
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| In the [[Puranic]] texts of [[Hinduism]], [[Narasimha]] ("man-lion") a half-lion, half-man incarnation or ([[avatar]]) of [[Vishnu]], is worshipped by his devotees and saved the child devotee [[Prahlada]] from his father, the evil demon king [[Hiranyakashipu]];<ref>{{Cite journal | url = http://srimadbhagavatam.com/1/3/18/en1 | title = Bhag-P | publisher = Srimadbhagavatam | at = 1.3.18 | quote = In the fourteenth incarnation, the Lord appeared as Nrisimha and bifurcated the strong body of the atheist Hiranyakasipu with His nails, just as a carpenter pierces cane. }}</ref> Vishnu takes the form of half-man/half-[[Asiatic lions|lion]], in Narasimha, having a human torso and lower body, but with a lion-like face and claws.<ref>{{cite web |url = http://srimadbhagavatam.com/7/8/19-22/en1 | title= Bhag-P | at = 7.8.19–22 | work = Srimadbhagavatam | accessdate = 21 November 2010}}</ref> [[Singh]] is an [[ancient]] Indian [[Vedic Sanskrit|vedic]] name meaning "lion" ([[Asiatic lion]]), dating back over 2000 years to [[History of India|ancient India]]. It was originally only used by [[Rajputs]] a [[Hindu]] [[Kshatriya]] or military [[caste]] in India. After the birth of the [[Khalsa]] brotherhood in 1699, the [[Sikh]]s also adopted the name "Singh" due to the wishes of [[Guru Gobind Singh]]. Along with millions of Hindu Rajputs today, it is also used by over 20 million [[Sikhs]] worldwide.<ref>{{Cite book | first = Khushwant | last = Singh | authorlink = Khushwant Singh | title = A History of the Sikhs: 1469–1838|publisher=Oxford University Press|year= 2004 |isbn=0-19-567308-5| volume = I }}</ref> Found famously on numerous [[flags]] and [[coat of arms|coats of arms]] all across Asia and Europe, the Asiatic lions also stand firm on the [[National Emblem of India]].<ref>{{cite web | title =Know India: State Emblem | work= National Portal of India | publisher = National Informatics Centre | year = 2005 | url= http://india.gov.in/knowindia/state_emblem.php | accessdate = 27 August 2007 | archiveurl = https://web.archive.org/web/20070822111444/http://india.gov.in/knowindia/state_emblem.php | archivedate = 22 August 2007}}</ref> Farther south on the [[Indian subcontinent]], the Asiatic lion is symbolic for the [[Sinhalese people|Sinhalese]],<ref>{{cite web | title = National Flag | publisher= Government of Sri Lanka | url= http://www.gov.lk/info/index.asp?mi=19&xp=52&xi=54&xl=3&o=0&t= | archiveurl = https://web.archive.org/web/20080327112704/http://www.gov.lk/info/index.asp?mi=19&xp=52&xi=54&xl=3&o=0&t= | archivedate = 27 March 2008 | accessdate= 6 August 2007}}</ref> [[Sri Lanka]]'s ethnic majority; the term derived from the Indo-Aryan ''Sinhala'', meaning the "lion people" or "people with lion blood", while a sword-wielding lion is the central figure on the national [[flag of Sri Lanka]].<ref>{{cite web | work = Constitution | year = 1978 | title=Article 6: The National Flag | publisher=Government of Sri Lanka | url = http://www.priu.gov.lk/Cons/1978Constitution/Schedle_2_Amd.html | accessdate=6 August 2007}}</ref>
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| [[File:China - Beijing 12 - lion outside the Tibetan Monastery (134036069).jpg|left|upright|thumb|A [[Chinese guardian lion]] outside [[Yonghe Temple]], Beijing]]
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| [[File:Nishi Honganji Karamon (detail).jpg|thumb|Detail of the [[Karamon]] of [[Nishi Hongan-ji]] in [[Kyoto]]; [[Momoyama period]]; [[National Treasures of Japan|National Treasure]]]]
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| The Asiatic lion is a common motif in [[Chinese art]]. They were first used in art during the late [[Spring and Autumn period]] (fifth or sixth century BC), and became much more popular during the [[Han Dynasty]] (206 BC – AD 220), when [[imperial guardian lions]] started to be placed in front of imperial palaces for protection. Because lions have never been native to China, early depictions were somewhat unrealistic; after the introduction of [[Buddhist art]] to China in the [[Tang Dynasty]] (after the sixth century AD), lions usually were wingless, with shorter, thicker bodies, and curly manes.<ref>{{Cite news | first = Li | last = Ling |date=May 2002 | url= http://www.cityu.edu.hk/ccs/Newsletter/newsletter4/Lion/Lion.htm | title = The Two-Way Process in the Age of Globalization | archiveurl = https://web.archive.org/web/20050406143133/http://www.cityu.edu.hk/ccs/Newsletter/newsletter4/Lion/Lion.htm | archivedate =6 April 2005 | others = Egan, Ronald transl. | newspaper = Ex/Change | type = newsletter | publisher = [[City University of Hong Kong]] | issue = 4 | accessdate = 26 September 2007 }}</ref> The [[lion dance]] is a form of traditional dance in [[Chinese culture]] in which performers mimic a lion's movements in a lion costume, often with musical accompaniment from cymbals, drums, and gongs. They are performed at [[Chinese New Year]], the [[Mid-Autumn Festival|August Moon Festival]] and other celebratory occasions for good luck.<ref>{{Cite journal | url = http://scripts.mit.edu/~lion-dance/about.php | publisher = MIT | title = Lion Dance Club }}</ref>
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| The [[island nation]] of Singapore derives its name from the [[Malay language|Malay]] words {{lang |ms|''singa''}} (lion) and {{lang|ms|''pora''}} (city/fortress), which in turn is from the [[Tamil language|Tamil]]-[[Sanskrit]] சிங்க ''singa'' {{lang|sa|सिंह}} {{IAST|''siṃha''}} and {{lang|sa|पुर}} புர {{IAST|''pura''}}, which is cognate to the [[Greek language|Greek]] {{lang|el|''πόλις''}}, ''pólis''.<ref>{{cite web | url = http://www.bartleby.com/61/46/S0424600.html | archiveurl= https://web.archive.org/web/20080629044446/http://www.bartleby.com/61/46/S0424600.html | archivedate=29 June 2008|title=Singapore| work=The American Heritage Dictionary of the English Language | edition = 4th |year=2000|publisher= Bartleby |accessdate=14 April 2006}}</ref> According to the [[Malay Annals]], this name was given by a fourteenth-century [[Sumatra]]n Malay prince [[Sang Nila Utama]], who, on alighting the island after a thunderstorm, spotted an auspicious beast on shore that appeared to be a lion.<ref>{{cite web | title=Early History | url=http://app.www.sg/who/30/Early-Names.aspx | archiveurl=https://web.archive.org/web/20100412092432/http://app.www.sg/who/30/Early-Names.aspx | archivedate=12 April 2010 | publisher= Ministry of Information, Communications and the Arts | location = Singapore | accessdate = 6 November 2010}}</ref>
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| The name of the nomadic [[Hadendoa]] people, inhabiting parts of [[Sudan]], [[Egypt]], and [[Eritrea]], is made up of ''haɖa'' 'lion' and (''n)ɖiwa'' 'clan'. Other variants are ''Haɖai ɖiwa'', ''Hanɖiwa'', and ''Haɖaatʼar'' (children of lioness).
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| [[File:Durga Mahishasura-mardini, the slayer of the buffalo demon.jpg|thumb|left|upright|Depiction of Goddess [[Durga]], her [[Vahana|mount]] is a lion]]
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| [[File:Lion in the garden of Palmyra Archeological Museum, 2010-04-21.jpg|thumb|Lion in Museum]]
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| "Lion" was the nickname of several medieval warrior rulers with a reputation for bravery, such as the English King [[Richard the Lionheart]],<ref name="Garai73"/> [[Henry the Lion]], ({{lang-de|link=no|Heinrich der Löwe}}), [[Duke of Saxony]], [[William the Lion]], King of Scotland, and [[Robert III of Flanders]] nicknamed "The Lion of Flanders"—a major [[Flanders|Flemish]] national icon up to the present. Lions are frequently depicted on [[coat of arms|coats of arms]], either as a device on shields themselves, or as [[supporters]], but the lioness is much more infrequent.<ref>{{cite web |title=Arms of Margaret Norrie McCain | work = The Public Register of Arms, Flags and Badges | location = [[Canada|CA]] |url = http://archive.gg.ca/heraldry/pub-reg/project-pic.asp?lang=e&ProjectID=538&ProjectElementID=1883 |accessdate = 30 June 2010}}</ref> The formal language of [[heraldry]], called [[blazon]], employs French terms to describe the images precisely. Such descriptions specified whether lions or other creatures were "rampant" or "passant", that is whether they were rearing or crouching.<ref name="Notre Dame">{{cite web | title =Heraldic Dictionary: Beasts | publisher=University of Notre Dame | url =http://www.rarebooks.nd.edu/digital/heraldry/charges/lions.html | accessdate = 20 July 2007}}</ref> The lion is used as a symbol of sporting teams, from national association football teams such as [[England national football team|England]], [[Scotland national football team|Scotland]] and [[Singapore national football team|Singapore]] to famous clubs such as the [[Detroit Lions]]<ref>{{cite web | type = official Website | title = Detroit Lions | year = 2001 | url = http://www.detroitlions.com/index.cfm?homelink=y | accessdate=8 July 2007}}</ref> of the NFL, [[Chelsea F.C.|Chelsea]]<ref name="new crest">{{Cite news| date=12 November 2004 | url=http://news.bbc.co.uk/sport1/hi/football/teams/c/chelsea/4008257.stm | title= Chelsea centenary crest unveiled | publisher = BBC | accessdate = 2 January 2007}}</ref> and [[Aston Villa F.C.|Aston Villa]] of the English [[Premier League]],<ref>{{cite web | title=The Aston Villa Crest: 2007 Onwards... | publisher=Aston Villa F.C. | year=2007 | url = http://www.avfc.premiumtv.co.uk/page/CrestTest/0,,10265,00.html | archiveurl= https://web.archive.org/web/20071011024349/http://www.avfc.premiumtv.co.uk/page/CrestTest/0,,10265,00.html | archivedate=11 October 2007 | accessdate= 6 August 2007}}</ref> (and the Premiership itself), [[Eintracht Braunschweig]] of the [[Bundesliga]], and to a host of smaller clubs around the world.
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| [[File:Christian VIII og Caroline Amalie i salvingsdragt.jpg|thumb|upright|Three silver lions both guard and are a part of the [[Throne Chair of Denmark]].<br />By [[Joseph-Désiré Court]] (1841)]]
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| Lions continue to be featured in modern literature, from the messianic [[Aslan]] in ''[[The Lion, the Witch and the Wardrobe]]'' and following books from [[The Chronicles of Narnia]] series written by [[C. S. Lewis]],<ref>{{Cite book |last=Lewis |first=Clive Staples |authorlink=C. S. Lewis |title= The Lion, the Witch and the Wardrobe | year = 1950 | publisher = HarperCollins | isbn= 0-06-023481-4}}</ref> to the comedic [[Cowardly Lion]] in ''[[The Wonderful Wizard of Oz]]''.<ref>{{cite book | last1 = Baum | first1 = L. Frank | last2 = Hearn | first2 = Michael Patrick |title=The Annotated Wizard of Oz|page=148|publisher= Clarkson N. Potter |location=New York, NY |year=1973|isbn=0-517-50086-8}}</ref> The advent of moving pictures saw the continued presence of lion symbolism; one of the most iconic and widely recognised lions is [[Leo the Lion (MGM)|Leo the Lion]], which has been the mascot for [[Metro-Goldwyn-Mayer]] (MGM) studios since the 1920s.<ref name="tvacres">{{cite web | url= http://www.tvacres.com/adanimals_leolion.htm | title = Advertising Mascots—Animals—Leo the MGM Lion (MGM Studios) | work=TV Acres}}</ref> The 1960s saw the appearance of what is possibly the most famous lioness, the Kenyan animal [[Elsa the lioness|Elsa]] in the movie ''[[Born Free]]'',<ref>{{Cite book |last=Adamson |first=George | authorlink = George Adamson |title= Bwana Game: the life story of George Adamson|year= 1969 | publisher = Fontana | isbn=0-00-612145-4 }}</ref> based on the true-life book of the same title.<ref>{{Cite book |last=Adamson |first=Joy |authorlink=Joy Adamson |title=Born Free: A Lioness of Two Worlds | year =2000|origyear=1960 |publisher=Pantheon |isbn=0-375-71438-3}}</ref> The lion's role as king of the beasts has been used in cartoons, such as the 1994 [[Walt Disney Pictures|Disney]] animated feature film ''[[The Lion King]]''.<ref>{{Cite book |last=Schweizer |first=Peter |title= Disney: The Mouse Betrayed |year= 1998 | pages = 164–69 | publisher =Regnery |location=Washington D.C. |isbn= 0-89526-387-4}}</ref>
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| ==Heraldic depictions==
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| {{Main article|Lion (heraldry)}}
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| ==See also==
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| * [[Lion lights]] (lights used to repel lions)
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| * [[Tiger versus lion]]
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| ==References==<!-- ContribZool76:87. -->
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| {{Reflist|2}}
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| ===Cited texts===
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| {{Refbegin|30em}}
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| * {{Cite book | last1 = Baratay |first1 = Eric | first2 = Elisabeth | last2 = Hardouin-Fugier |title= Zoo: a history of zoological gardens in the West |year=2002 |publisher=Reaktion Books |location=London |isbn = 1-86189-111-3|ref=Baratay}}
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| * {{Cite book |last=Blunt |first= Wilfred |title= The Ark in the Park: The Zoo in the Nineteenth Century |year = 1975 | publisher =Hamish Hamilton |location=London |isbn=0-241-89331-3|ref=Blunt}}
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| * {{Cite book |last=de Courcy |first=Catherine |title= The Zoo Story |year= 1995 |publisher= Penguin Books | location = Ringwood, Victoria |isbn=0-14-023919-7|ref=Courcy}}
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| * {{Cite book | last= Schaller |first= George B. |authorlink= George Schaller|year= 1972 |title= The Serengeti lion: A study of predator–prey relations|publisher= University of Chicago Press|location=Chicago |isbn = 0-226-73639-3 | ref =Schaller}}
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| * {{Cite book | last= Scott |first=Jonathon |last2=Scott |first2=Angela |year=2002 |title=Big Cat Diary: Lion |publisher=Harper Collins |isbn=0-00-714666-3 |ref=Scott}}
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| {{Refend}}
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| ==External links==
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| {{Wiktionary}}
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| {{wikispecies |Panthera leo|Lion}}
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| {{Collier's Poster}}
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| * {{Commons-inline|Panthera leo|Lion}}
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| * [http://www.catsg.org/index.php?id=108 Species portrait Lion; IUCN/SSC Cat Specialist Group]
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| * {{Cite journal | url = http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_leo.html | title = Animal Diversity Web – ''Panthera leo'' (lion) | publisher = U Mich }}
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| * {{Cite journal | url = http://www.awf.org/content/wildlife/detail/lion | publisher = African Wildlife Foundation | title = Lion }}
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| * [https://www.youtube.com/watch?v=LU8DDYz68kM Battle at Kruger]: Video of a pack of lions fighting against a crocodile and buffaloes over a kill.
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| * {{cite web | url = http://www.biodiversitylibrary.org/name/Felis_leo | work = Biodiversity Heritage Library | type = bibliography | title = ''Felis leo''}}
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| * {{cite web | url = http://www.biodiversitylibrary.org/name/Panthera_leo | work = Biodiversity Heritage Library | type = bibliography | title = Panthera leo}}
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| * {{cite web | url = http://ngm.nationalgeographic.com/2013/08/lion-conservation/lion-strongholds | work = National Geographic | type = bibliography | title =The state of Lions }}
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| * {{Cite journal | url = http://www.bbc.co.uk/nature/species/Lion | publisher = The BBC | title = Lion | type = news, and video clips from BBC programmes past and present }}
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| * {{cite web | url = http://www.lionconservationfund.org/ | title = Lion Conservation Fund }} Example of a fund and its projects about the research and conservation of the lion.
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| * {{cite web | url = https://edge.org/conversation/lions-africa-39s-magnificent-predators | title = Lions: africa's magnificent predators }} Description article
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| [[Category:Lions| ]]
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| [[Category:Apex predators]]
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| [[Category:Felids of Africa]]
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