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| {{otheruses}} | |||
| {{Taxobox | |||
| | color = pink | |||
| | name = Lion<ref name=MSW3>{{MSW3 Wozencraft | pages = 546}}</ref> | |||
| | status = VU | |||
| | status_system = iucn3.1 | |||
| | trend = down | |||
| | status_ref =<ref name=IUCN>{{IUCN2006|assessors=Nowell & Bauer|year=2004|id=15951|title=Panthera leo|downloaded=11 May 2006}} Database entry includes a lengthy justification of why this species is vulnerable </ref> | |||
| | image = Lion waiting in Nambia.jpg | |||
| | image_width = 250px | |||
| | regnum = ]ia | |||
| | phylum = ] | |||
| | classis = ] | |||
| | ordo = ] | |||
| | familia = ] | |||
| | genus = '']'' | |||
| | species = '''''P. leo''''' | |||
| | binomial = ''Panthera leo'' | |||
| | binomial_authority = (], ]) | |||
| | synonyms=<center>'''''Felis leo '''''<br /><small>(], ])</small></center> | |||
| | range_map = Lion_distribution.svg | |||
| | range_map_width = 250px | |||
| | range_map_caption = Distribution of Lions in Africa | |||
| }} | |||
| The '''lion''' (''Panthera leo'') is a member of the family ] and one of four "]s" in the ] '']''. Reaching up to 225 ] (500 ]), the lion is only slightly eclipsed in size by the ] amongst felines, and both are significantly larger than any other cat. Although lions were once widespread throughout much of ], ], and ], they currently exist in the wild only in Africa, with a ] remnant population in ]. As a whole, the species is listed as "]"; it has seen a significant and possibly irreversible decline in numbers in its African range.<ref name=IUCN/> | |||
| In the wild, lions live for approximately 10–14 years, while in captivity they can live over 20 years. Today lions are found only in tropical climates, but during the ], they penetrated far north (even to ]). They typically range across ] and grassland, although they may take to bush and ]. It is an ] and ]. Unusually for a cat, lions hunt together. Groups of female lions typically bring down prey, mostly large ]s. The lion ''pride'' consists of related females and offspring and a small number of resident males. Lions are territorial and the ''pride'', though not strictly hierarchical, is dominated by an adult male or coalition of males. | |||
| The ] lion is highly distinctive and usually instantly recognized by its ]. The lion, particularly the face of the male, is one of the most widely recognized animal symbols in human ]. It has been extensively depicted on sculptures, in paintings, on national flags and in films. | |||
| ==Naming and etymology== | |||
| The Lion's name, similar in many languages, derives from the ] ''leo'',<ref>{{cite book | last = Simpson | first = D.P. | title = Cassell's Latin Dictionary | publisher = Cassell Ltd. | date = 1979 | edition = 5 | location = London | pages = 883 | id = ISBN 0-304-52257-0}}</ref> and before that the ] ''leōn''/λεων.<ref name="Liddell 1980">{{cite book | author = ] and ] | year = 1980 | title = ] (Abridged Edition) | publisher = ] | location = United Kingdom | id = ISBN 0-19-910207-4}}</ref> The ] word ''lavi'' (לָבִיא) may also be related,<ref>{{Cite encyclopedia| title=Lion|encyclopedia=Oxford English Dictionary|editor=Simpson, J., Weiner, E. (eds)| year=1989 |edition= 2nd edition| location=Oxford |publisher=Clarendon Press|id= ISBN 0-19-861186-2}}</ref> as well as the Ancient Egyptian ''rw''.<ref>. As in other ancient scripts, in Ancient Egyptian only the consonants are written. No distinction was made between 'l' and 'r'.</ref> It was one of the many species originally described, as ''Felis leo'', by ] in his 18th century work, '']''.<ref>{{la icon}} {{cite book | last=Linnaeus | first=C | authorlink=Carolus Linnaeus | title=Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. | publisher=Holmiae. (Laurentii Salvii). | date=1758| pages=824 | url=http://dz1.gdz-cms.de/index.php?id=img&no_cache=1&IDDOC=265100}}</ref> | |||
| The generic component of its scientific designation, ''Panthera leo'', is often presumed to derive from Greek ''pan-'' ("all") and ''ther'' ("beast"), but this may be a ]. Although it came into English through the classical languages, ''panthera'' is probably of East Asian origin, meaning "the yellowish animal," or "whitish-yellow".<ref>{{cite web | url = http://www.etymonline.com/index.php?term=panther | title = "Panther" | work = Online Etymology Dictionary | publisher = ] | accessdate = 2007-07-05}}</ref> | |||
| ==Physical characteristics== | |||
| ], Kenya]] | |||
| The lion is a large and muscular feline with a compact build. With short, powerful legs, a strong jaw, and long canine teeth, the lion can bring down and kill large prey.<ref name=Honolulu>{{cite web |url=http://www.honoluluzoo.org/lion.htm |title=Lion |accessdate=2007-07-12 |publisher=Honolulu Zoo}}</ref> Lion coloration varies from light buff to yellowish, reddish or dark ochraceous brown. The color of the manes varies from blond to black. The underparts are generally lighter and the tail tuft is black. | |||
| Average listed weights for the male lion are between 150–225 kg (330–500 lb), and 120–150 kg (260–330 lb) for females.<ref>{{cite web|url=http://www.bbc.co.uk/nature/wildfacts/factfiles/14.shtml|title=BBC Wildfacts – Lion}}</ref> Nowell and Jackson report average weights of 181 kg for males and 126 kg for females.<ref name=CAP> Nowell, K. and Jackson, P. (compilers and editors) 1996. IUCN/SSC Cat Specialist Group. IUCN, Gland, Switzerland. (see ''Panthera Leo'', pp. 17 – 21.)</ref> Head and body length is 170–250 cm (5 ft 7 in–8 ft 2 in) in males and 140–175 cm (4 ft 7 in–5 ft 9 in) in females; shoulder height is about 123 cm (4 ft) in males and 100 cm (3 ft 3 in) in females. The tail length is 70–100 cm (2 ft 3 in–3 ft 3 in).<ref>Ronald M. Nowak: ''Walker's Mammals of the World''. Johns Hopkins University Press, 1999 ISBN 0-8018-5789-9</ref> The tail ends in a hairy tuft. The tuft conceals a spine, approximately 5 mm long, formed of the final sections of tail bone fused together. The lion is the only felid to have tuffed tail and the function of the tuft and spine are unknown. In the wild, lions live up to 16 years of age, while in captivity they can live ten years longer.<ref name=Honolulu/> | |||
| ====Mane==== | |||
| ], ]]] | |||
| The mane of the male lion, unique amongst cats, is one of the most distinct characteristics of the species. The presence, absence, color, and size of the mane is associated with sexual maturity, climate and testosterone production.<ref> The mane aids the lion during face offs with spotted hyenas (the lion's main competitor) and with other lions and makes the lion look bigger than he really is, providing an excellent intimidation display. | |||
| {{cite web | last = Trivedi | first = BP | title = Are Maneless Tsavo Lions Prone to Male Pattern Baldness? | publisher = National Geographic | date = 2005 | url = http://news.nationalgeographic.com/news/2002/04/0412_020412_TVtsavolions.html | accessdate = 2007-07-07}}</ref> Research in Tanzania suggests mane darkness correlates to nutrition and testosterone, and that 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. |coauthors=Parker, Craig |year=2002 |month=August |title=Sexual Selection, Temperature, and the Lion's Mane |journal=] |volume=297 |issue=5585 |pages= 1339 - 1343 |url=http://www.sciencemag.org/cgi/content/abstract/297/5585/1339 |accessdate= 2007-07-12}}</ref> It is possible that lionesses more actively solicit mating with heavily maned lions in prides led by a coalition of 2 or 3 males.{{Fact|date=July 2007}} | |||
| ] | |||
| Maneless lions have been reported in ] and ] in Kenya, and the original male white lion from Timbavati was also maneless. ] lions have minimal manes. Manelessness is also found in inbred lion populations; inbreeding also results in poor fertility.<ref>Trivedi, Bijal P., "" in ''] News'', 12 June 2002. Retrieved 17 July 2007.</ref> | |||
| Scientists once believed that the distinct status of some subspecies could be justified by ], including the size of the mane. Morphology was used to identify subspecies such as the ] and ]. Research has suggested, however, that various ] factors influence the color and size of a lion's mane, such as the ].<ref name=West/> The cooler ambient temperature in European and North American ]s, for example, can result in a heavy mane. Thus the mane is an inappropriate marker for identifying subspecies.<ref name="Conservation-Genetics:Preserving-Genetic-Diversity">Barnett, R., N. Yamaguchi, I. Barnes & A. Cooper. 2006. : Implications for its ''ex situ'' conservation. Conservation Genetics.</ref><ref>Yamaguchi, N. & Haddane, B. (2002). The North African Barbary lion and the Atlas Lion Project. International Zoo News 49: 465-481.</ref> | |||
| ==Biology and behaviour== | |||
| ===Hunting and diet=== | |||
| ]]] | |||
| Female lions usually hunt at night or dawn and in packs. Their ] consists mainly of larger mammals, with a preference for ]s, ]s, ]s, ]s, ]s, and ]s. Notable exceptions to a lions usual diet include buffalo bulls in their prime and very large, fully grown male giraffes.<ref name="home">{{citeweb|url=http://home.intekom.com/ecotravel/Guides/Wildlife/Vertebrates/Mammals/Big_5/Lion/african-lion-hunting-habits.htm | |||
| |title=African Lion Hunting Habits in South Africa and Southern Africa|accessmonthday=31 July|accessyear=2007}}</ref> Many other species are hunted based on availability, mainly ungulates of a weight between 50 and 300 kg, like ], ], ] and ] in Africa or ], ]s and several deer species in India. Occasionally they take relatively small species like ]s or ]s. Very large species such as the ], the ] and the ] are generally avoided due to the danger they present to lions or the effort required to capture them.<ref>{{cite journal|author=Hayward, Matt W. and Kerley, Graham|title=Prey preferences of the lion (''Panthera leo'')|journal=Journal of Zoology|volume=267|issue=3|year=2005|pages=309-322|doi=10.1017/S0952836905007508}}</ref> In some areas, lions specialise on rather atypical prey-species; this is the case at the ] river, where they constantly prey on young ]s.<ref>Kemp, Leigh, "." Retrieved 17 July 2007.</ref> It is reported that the lions, driven by extreme hunger, started taking down baby elephants, then moved on to adolescents and occasionally fully grown adults.<ref>{{cite web |title=King of the jungle defies nature with new quarry |author=Damien Whitworth |date=October 9, 2006 |publisher=The Australian |url=http://www.news.com.au/story/0,23599,20547955-38195,00.html |accessdate=2007-07-20 }}</ref> Lions will even kill other predators such as ]s, ]s, ]s and ]s. An adult female lion requires about 5 kg (11 lb) of meat per day, a male about 7 kg (15.4 lb).<ref>{{cite web |title=Lions |publisher=Honolulu Zoo |url=http://www.honoluluzoo.org/lion.htm |accessdate=2007-07-20 }}</ref> | |||
| ] | |||
| Young lions first attempt to hunt at around three months of age, though generally not effectively until they are two years old.{{Fact|date=July 2007}} | |||
| Lions can reach speeds of 50 mph,<ref>{{cite web |title=Speed of Animals |publisher=infoplease.com |url=http://www.infoplease.com/ipa/A0004737.html |accessdate=2007-07-20 }}</ref> though only for short bursts, so they have to be quite nearby their prey before starting the attack. They sneak up to the victim until they reach a distance of about 30 ] (98 ]) or less. Usually several lions work together and encircle the herd from different points. The attack is short and powerful, and the lion tries to catch the victim with a fast rush and some final leaps. The prey is usually killed by strangulation.<ref>{{cite web |title=About lions - Ecology and behaviour |author=Dr Gus Mills |publisher=African Lion Working Group |url=http://www.african-lion.org/lions_e.htm |accessdate=2007-07-20 }}</ref> | |||
| Because lions hunt in open spaces, where they are easily seen by their prey, cooperating hunting increases the likelihood of a successful hunt, especially of larger species. Teamwork also enables them to defend their prey more easily against other large predators like hyenas, which can be attracted by ]s over kilometers in open savannas. Lionesses do most of the hunting; males attached to ] do not usually participate, except in the case of large animals such as buffalo. In group hunts, each lioness has a favored 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>{{cite journal|author=Stander, P. E.|title=Cooperative hunting in lions: the role of the individual|journal=Behavioral Ecology and Sociobiology|volume=29|issue=6|year=1992|doi=10.1007/BF00170175|pages=445—454}}</ref> | |||
| ===Reproduction and sexuality=== | |||
| ] | |||
| Lions do not ] at any specific time of year, and the females are ].<ref>. Retrieved 17 July 2007.</ref> Like other cats, the male lion's penis has spines which point backwards. Upon withdrawal of the penis, the spines rake the walls of the female's vagina, which may cause ovulation. Furthermore, cats are ]; that is, a female may mate with more than one male when she is in heat, meaning different cubs in a litter may have different fathers.{{Fact|date=July 2007}} During a mating bout, which could last several days, the couple frequently copulate twenty to forty times a day and are likely to forgo hunting. In captivity, lions reproduce very well. | |||
| The average gestation period is around 110 days,<ref>{{cite book |last=Schaller |first=G.B. |year=1972 |title= The Serengeti Lion|publisher= University of Chicago Press|location=Chicago |isbn= }}</ref> and the female gives birth to a litter of one to four cubs. The females in a pride will synchronize their reproductive cycles so that they cooperate in the raising and suckling of the young, who suckle indiscriminately from any or all of the nursing females in the pride. Cubs are weaned after six to seven months. In the wild, competition for food is fierce, and as many as 80% of the cubs will die before the age of two.{{Fact|date=July 2007}} | |||
| When one or more new males take over a pride and oust the previous master(s), the conquerors often kill any remaining cubs.<ref name="Packpus83">{{cite journal |author=Packer, C., Pusey, A. E. |year=1983 |month=May |title= Adaptations of female lions to infanticide by incoming males|journal= American Naturalist|volume=121 |issue=5 |pages=716-728 |doi= |url=http://www.lionresearch.org/current_docs/6.pdf |accessdate=2007-07-08}}</ref> This is explained by the fact that the females would not become fertile and receptive until the cubs grow up or die. The male lions reach maturity at about 3 years of age and are capable of taking over another pride at 4–5 years old. They begin to age, and thus weaken, at around 8. This leaves a short window for their own offspring to be born and mature—the fathers have to procreate as soon as they take over the pride. The lioness will often attempt to defend her cubs fiercely from a usurping male, but such actions are rarely successful, as he usually kills all the previous top male's cubs that are less than two years old and the female is much lighter and has less strength than the male. However, success is more likely when a group of 3 or 4 mothers within the pride join forces against one male.<ref name="Packpus83"/> | |||
| One scientific study reports that both males and females may interact ].<ref>Bruce Bagemihl, ''Biological Exuberance: Animal Homosexuality and Natural Diversity,'' St. Martin's Press, 1999; pp.302-305. In his discussion of lion same-sex relations, Bagemihl is making use of published work by: J.B. Cooper, "An Exploratory Study on African Lions" in ''Comparative Psychology Monographs'' 17:1-48; R.L. Eaton, "The Biology and Social Behavior of Reproduction in the Lion" in Eaton, ed. ''The World's Cats,'' vol. II; pp.3-58; Seattle, 1974; G.B. Schaller, ''The Serengeti Lion''; University of Chicago Press, 1972</ref><ref>Suvira Srivastav, </ref> Male lions pair-bond for a number of days and initiate homosexual activity with affectionate nuzzling and caressing, leading to mounting and thrusting. A study found that about 8% of mountings have been observed to occur with other males, while female pairings are held to be fairly common in captivity but have not been observed in the wild. | |||
| ===Social behavior=== | |||
| ] | |||
| ], ]]] | |||
| Lions are predatory ]s who manifest two types of social organization. Some are ''residents,'' living in groups, called ''prides''. The pride consists of related females, their cubs of both sexes, and a group of one to four males known as a ''coalition'' who mate with the adult females. Others are ''nomads'', ranging widely, either singularly or in pairs. | |||
| Being smaller and lighter than males, lionesses are more agile and faster and do the pride's hunting, while the stronger males patrol the territory and protect the pride, for which they take the "lion's share" of the females' prey. When resting, lions seem to enjoy good fellowship with lots of touching, head rubbing, licking and purring. But when it comes to food, each lion looks out for itself. Squabbling and fighting are common, with adult males usually eating first, followed by the females and then the cubs. | |||
| Why ]—the most pronounced in any cat species—has developed in lions 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 "cheaters" reduce per capita caloric intake. Other benefits include possible ] (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/> | |||
| Both males and females will defend the pride against intruders. Some individual lions consistently lead the defense against intruders, while others lag behind.<ref>{{cite journal |author=Heinsohn, R. and Packer, C. |year=1995 |month= |title=Complex cooperative strategies in group-territorial African lions |journal=Science |volume= |issue=269 |pages=1260-1262}}</ref> These “laggards” are not punished by leaders. Possibly laggards provide other services to the group so that leaders forgive them.<ref>{{cite journal |author=Morell, V. |year=1995 |title=Cowardly lions confound cooperation theory |journal=Science |volume= |issue=269 |pages=1216-1217}}</ref> An alternative hypothesis is that there is some reward associated with being a leader who fends off intruders.<ref>{{cite journal |author=Jahn, G. C. |year=1996 |title= Lioness Leadership|journal=Science |volume= |issue=271 |pages=1215}}</ref> | |||
| Typically, males will not tolerate outside males, and females will not tolerate outside females. Males are expelled from the pride or leave on their own when they reach maturity. | |||
| Lions spend a lot of their time resting. They are inactive for about 20 hours per day. | |||
| ==Population and distribution== | |||
| ]s. Plans are afoot to re-introduce some to ] in the neighboring State of ] in India.]] | |||
| In relatively recent times the habitat of lions spanned the southern parts of ], ranging from ] to ], and most of ] except the central ]-zone and the ] desert. The last lions in ] died out in historic times. In the ], their last European outpost, a population of the ] survived until the 10th century.<ref>* V. G. Heptner and A. A. Sludskii: ''Mammals of the Sowjetunion Vol. II, Part 2 CARNIVORA (Hyaenas and Cats)''. Leiden, New York, 1989 ISBN 900408876 8</ref> Between the late 19th and early 20th century they also became extinct in ] and the ]. Now, most lions live in eastern and southern Africa, and their numbers are rapidly decreasing. Currently, estimates of the African lion population range between 16,000 and 30,000 living in the wild, down from early 1990s estimates that ranged as high as 100,000; increased contact with human development is cited as the primary cause.<ref>{{cite web |title=NATURE. The Vanishing Lions |publisher=PBS |url=http://www.pbs.org/wnet/nature/vanishinglions/index.html |accessdate=2007-07-20 }}</ref> The remaining populations are often geographically isolated from each other, which can lead to ], and consequently, a lack of ]. | |||
| The ] (''Panthera leo persica''), which in historical times ranged from ] to India and from the Caucasus to ], was eradicated from ] by the ] and from most of the rest of Asia after the arrival of readily available firearms in the 18th century. By the late 19th century the lion had disappeared from ],<ref>{{cite book | author=Ustay, A.H.|year=1990|title= Hunting in Turkey|publisher=BBA, Istanbul}}</ref><ref>{{cite web |title=Past and present distribution of the lion in North Africa and Southwest Asia. |date=2001 |publisher=Asiatic Lion Information Centre |url=http://www.asiatic-lion.org/distrib.html |accessdate=2006-06-01 }}</ref> while the last sighting of a live Asiatic Lion in ] was in 1941 (between ] and Jahrom, ] province). In 1944, the corpse of a lioness was found on the banks of ] river, ] province, ]. There are no subsequent reliable reports from ].<ref>{{cite book | author=Guggisberg, C.A.W. |year=1961 |title= Simba: The Life of the Lion | publisher=Howard Timmins, Cape Town}}</ref> The subspecies now survives only in and around the ] of northwestern India. About 300 lions live in a 1412 km² (558 ]s) sanctuary in the state of ], which covers most of the forest. Their numbers remain stable. | |||
| Until the late ] (about 10,000 years ago), lions were also found in the ] and in northern Eurasia. The most famous of these prehistoric subspecies were the ] (''Panthera leo spelaea'') and the ] (''Panthera leo atrox'') (not to be confused with the ], ''Puma concolor'', also known as the Mountain Lion). | |||
| ==Taxonomy and evolution== | |||
| ] | |||
| The oldest ] record of a cat strongly resembling a lion is known from Laetoli in ] and is perhaps 3.5 million years old. Some scientists identify the material as ''Panthera leo''. These records are not well-substantiated, and all that can be said is that they pertain to a ''Panthera''-like felid. The oldest confirmed records of ''Panthera leo'' in Africa are about 2 million years younger.<ref>{{cite journal |last=Werdelin |first=Lars |coauthors=Lewis, Margaret E. |year=2005 |month=June |title=Plio-Pleistocene Carnivora of eastern Africa: species richness and turnover patterns |journal=Zoological Journal of the Linnean Society |volume=144|issue=2 |pages=121-144 |publisher=The Linnean Society of London |url=http://www.ingentaconnect.com/content/bsc/zoj/2005/00000144/00000002/art00001 |accessdate=2007-07-08}}</ref> | |||
| The closest relatives of the lion are the other '']'' species, by name the tiger, the jaguar and the leopard. Morphological and genetic studies reveal, that the tiger was the first of these recent species to demerge from the other. About 1.9 million years ago the Jaguar demerged from the remaining group, which contained ancestors of the leopard and lion. The Lion and leopard subsequently separated about 1 to 1.25 million years ago from each other.<ref>Li Yua and Ya-ping Zhang: ''Phylogenetic studies of pantherine cats (Felidae) based on multiple genes, with novel application of nuclear β-fibrinogen intron 7 to carnivores''. Molecular Phylogenetics and Evolution, Volume 35, Issue 2 , May 2005, Pages 483-495 . </ref> | |||
| 700,000 years ago, ''Panthera leo'' appeared in Europe for the first time with the subspecies '']'' at ] in ]. From this lion derived the later ] (''Panthera leo spelaea''), which appeared about 300,000 years ago. During the upper ] the lion spread to North and South America, and developed into ''Panthera leo atrox'', the ].<ref>A. Turner: ''The big cats and their fossil relatives''. Columbia University Press, 1997. ISBN 0-231-10229-1</ref> | |||
| Lions were common in northern ] and America during the upper Pleistocene, but died out there at the end of the last ], about 10,000 years ago. | |||
| ===Subspecies=== | |||
| ] in ] is the natural home of the Asiatic lion but this animal was born in captivity.]] | |||
| ] | |||
| Traditionally 12 recent subspecies of lion were recognized. The major differences between these subspecies are location, mane appearance, size and distribution. Because these characteristics are very insignificant and show a high individual variability, most of these forms were debatable and probably invalid.<ref>* V. G. Heptner and A. A. Sludskii: ''Mammals of the Sowjetunion Vol. II, Part 2 CARNIVORA (Hyaenas and Cats)''. Leiden, New York, 1989 ISBN 900408876 8</ref> Today usually only eight subspecies are accepted.<ref name="Conservation-Genetics:Preserving-Genetic-Diversity" /><ref name="BurgerJ-Molecular-phylogeny">Burger, Joachim ''et al.'' (2004): (PDF) ] Vol.30, p.841-849.</ref> But one of them, the Cape lion formerly described as ''Panthera leo melanochaita'' is probably invalid.<ref name="Conservation-Genetics:Preserving-Genetic-Diversity" /> | |||
| Even the remaining seven subspecies might be too much, mitochondrial variation in recent African lions is modest, what suggests that all sub-Saharan lions could be considered a single subspecies, possibly divided in two main clades: one to the west of the ] and the other to the east. Lions from ] in Eastern Kenya genetically differ minimally from lions in ] (South Africa), but markedly from those in the ] in Western Kenya.<ref>Ross Barnett ''et al.'' (2006): (PDF) Proceedings of the Royal Society B, doi: 10.1098/rspb.</ref><ref>Jean Dubach ''et al.'' Molecular genetic variation across the southern and eastern geographic ranges of the African lion, Panthera leo. Conservation Genetics 6: 15–24, 2005. .</ref> | |||
| ====Recent subspecies==== | |||
| Seven recent subspecies are recognized today: | |||
| * '''''P. l. persica''''', known as the ''']''' or '''South Asian lion''' was once widespread from Turkey, across the Middle East, to ], India and even ]. However large prides and daylight activity made it easier to poach than tigers or leopards; now around 350 currently exist in and near the Gir Forest of India. | |||
| * '''''P. l. leo''''', known as the ''']''', is extinct at least in the wild and was believed to be extinct in captivity. This was the largest of the Lion subspecies, which ranged from Morocco to Egypt. The last wild Barbary lion was killed in Morocco in 1922 due to excessive hunting. Barbary Lions were kept by Roman emperors to take part in the gladiator arenas. Roman notables, including ], ], and ], often ordered the mass slaughter of Barbary lions—up to 400 at a time.<ref>{{cite web | last = Welman | first = N | title = Lions | work = Alexander the Great's Home on the Web | publisher = Thomas William-Powlett | date = 1994 | url = http://www.pothos.org/alexander.asp?paraID=103&keyword_id=6&title=Lions | accessdate = 2007-07-08}}</ref> | |||
| * '''''P. l. senegalensis''''', known as the '''West African lion''', is found in Western Africa, from Senegal to Nigeria. | |||
| * '''''P. l. azandica''''', known as the '''North East Congo lion''', is found in the Northeastern parts of the ]. | |||
| * '''''P. l. nubica''''', known as the '''East African-''' or '''Massai lion''', occurs in East Africa, from Ethiopia and Kenia to Tansania and Mozambique. | |||
| * '''''P. l. bleyenberghi''''', known as the '''Southwest African-''' or '''Katanga lion'''. South western Africa, ], ], ] (]). | |||
| * '''''P. l. krugeri''''', known as the '''Southeast African-''' or '''Transvaal lion''', is found in South eastern Africa, ]. | |||
| ====Prehistoric subspecies==== | |||
| Besides these subspecies there are also some prehistoric ones. | |||
| <!-- prose form please --> | |||
| * '''''P. l. atrox''''', known as the ''']''' or '''North American cave lion''', was abundant in the Americas from ] to ] in the Pleistocene Period until about 10,000 years ago. This form as well as the cave lion are sometimes considered to represent separate species, but recent phylogenetic studies lead to suggest, that they are in fact subspecies of the lion (''Panthera leo''). <ref name="BurgerJ-Molecular-phylogeny" /> One of the largest lion subspecies to have existed, its body length is estimated to have been 1.6–2.5 m (5–8 ft).<ref name="Quaternary extinctions">{{cite book |author=Martin, P.S. |title=Quaternary Extinctions |year=1984 |publisher= University of Arizona Press|location= Tucson, Arizona|isbn=0-8165-1100-4 }}</ref> | |||
| * '''''P. l. fossilis''''', known as the ''']''', flourished about 500,000 years ago; fossils have been recovered from ] and ]. It was larger than today's African lions, with a maximum head and body length of 2.40 m (8 ft).{{Fact|date=July 2007}} | |||
| * '''''P. l. spelaea''''', known as the ''']''', Eurasian cave lion or Upper Pleistocene European cave lion occurred in Eurasia 300,000 to 10,000 years ago.<ref name="BurgerJ-Molecular-phylogeny" /> It is known from ] ], ] carvings, and clay busts, indicating it had protruding ears, tufted tails, faint tiger-like stripes, and that at least some males had a "ruff" or primitive mane around their neck.{{Fact|date=July 2007}} | |||
| * '''''P. l. vereshchagini''''', known as the '''East Siberian-''' or ''']''', was found in ] (]), ] (]), and the ] (]). Analysis of skulls and mandibles of this ] demonstrate that it is distinct—larger than the European cave lion and smaller than the American cave lion with differing ] proportions.<ref>Baryshnikov, G.F., Boeskorov, G., 2001. The Pleistocene cave lion, ''Panthera spelaea'' (Carnivora, Felidae) from Yakutia, Russia. Cranium 18, 7–24.</ref><ref>Burger, Joachim ''et al.'' (2004): ''Molecular phylogeny of the extinct cave lion'' Panthera leo spelaea. Molecular Phylogenetics and Evolution Vol.30, p.841-849. </ref> | |||
| ====Dubious subspecies==== | |||
| * '''''P. l. melanochaita''''', known as the ''']''', became extinct in the wild around 1860. Results of ] research do not support the status as a distinct subspecies. It seems probable that the Cape lion was only the southernmost population of the extant southern African lion.<ref name="Conservation-Genetics:Preserving-Genetic-Diversity" /> | |||
| * '''''P. l. sinhaleyus''''', known as the ''']''', appears to have become extinct ] 39,000 years ago. It is only known from two teeth found in deposits at ]. Based on these teeth, P. Deraniyagala erected in 1939 this subspecies.<ref>{{cite journal |author=Kelum Manamendra-Arachchi, Rohan Pethiyagoda, Rajith Dissanayake, Madhava Meegaskumbura |year=2005 |month= |title=A second extinct big cat from the late Quaternary of Sri Lanka. |journal=The Raffles Bulletin of Zoology. |volume=|issue=Supplement No. 12 |pages=423–434 |id= |url=|publisher=National University of Singapore |accessdate= 2007-07-31}}</ref> | |||
| * '''''P. l. europaea''''', known as the ''']'''; Status as subspecies is unconfirmed. (Probably identical with ''Panthera leo persica'' or ''Panthera leo spelea''.) Extinct around 100 due to persecution and over-exploitation. Inhabited the ], the ], southern ] and the ]. It was a very popular object of hunting among ], ] and ]. | |||
| * '''''P. l. youngi''''' or ''Panthera youngi'' , known as the ''']''', 350,000 years ago, relationship to the extant lion subspecies is obscure. Represents probably a distinct species. | |||
| *'''''P. l. maculatus''''', known as the ''']''' or '''Spotted lion''', is sometimes 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 colored individuals, it has been extinct since 1931. A less likely identity is a natural leopard/lion ] commonly known as a ].<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> | |||
| ==Variations== | |||
| ].]] | |||
| A number of natural variations have been observed in lion populations. Some of these have been encouraged by captive breeding. | |||
| ===White lions=== | |||
| {{main|White lion}} | |||
| Although rare, white lions are occasionally encountered in and around the Kruger National Park, ]. Their unusual color is due to a ]. A white lion has a disadvantage in regards to hunting: it can be detected by its color, unlike the regular lion which camouflages with its surroundings. White lions are born almost pure white without the normal ] spots seen in lion cubs. Their color gradually darkens to cream or ivory color, known as blonde. | |||
| ===Hybridization=== | |||
| {{further|], ] and ]}} | |||
| ] is the offspring of a male lion and female ].|200px]] | |||
| Lions have also been known to ] with ]s (most often Amur and Bengal) to create hybrids called ]s and ]s. They have also been crossed with ] to produce ]s and ]s to produce ]s. The ] is reputedly a spotted lion or a naturally occurring leopon, while the ] is a complex lion/jaguar/leopard hybrid called a ]. 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 ]. | |||
| The liger is a cross between a male lion and a tigress. Because the lion sire passes on a growth-promoting gene, but the corresponding growth-inhibiting gene from the female lion is absent, 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 are often fertile. Males have about a 50% chance of having a mane, but if they grow one the mane will be modest, around 50% again of a pure lion mane. | |||
| The less common tigon is a cross between the lioness and the male tiger. Because the male tiger does not pass on a growth-promoting gene and the lioness passes on a growth inhibiting gene, tigons are often relatively small, only weighing up to 150 kilograms (350 lb), which is about 20% smaller than lions. Like ligers, they have physical and behavioural traits from both parental species and males are sterile. | |||
| ==Conservation status== | |||
| ], ]]] | |||
| The Lion is considered vulnerable by the ], while the Asiatic subspecies is Critically Endangered. The total population of wild lions dragged from perhaps 400,000 in 1950 to an estimated size of 16,500–47,000 in 2002-2004.<ref name=IUCN/> | |||
| 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 ]'s ] ecosystem.<ref name=IUCN/> | |||
| Conservation of both African and Asian Lions has required the setup and maintenance of National Parks and Game Reserves; among the best known are ] in ], ] in Tansania and ] in eastern ]. Outside these areas, the issues arising from lions interaction with cattle and people usually results in the elimination of the former. | |||
| ]]] | |||
| Lions are vulnerable to the ] ] (CDV), which is spread though domestic dogs and other ]; an outbreak in ], Tanzania, in 1994 resulted in many lions exhibiting neurological features such as seizures, and succumbing.<ref>{{cite journal |author=Roelke-Parker, M.E., Munson, L., Packer, C., Kock, R., Cleaveland, S., Carpenter, M., O'Brien, S.J., Pospischii, A., Hoffmann-Lehmann, R., Lutz, H., Mwamengele, G.L.M., Mgasa, M.N., Machange, G.A., Summers, B., Appel, M.J.G. |year=1996 |month=February |title=A canine distemper epidemic in Serengeti lions ''(Panthera leo)'' |journal= Nature|volume=379 |issue= |pages=441-445 |id= |url=http://www.lionresearch.org/current_docs/17.pdf |format=PDF|accessdate=2007-07-08}}</ref> | |||
| In India, the last refuge of the Asiatic Lion is the 1,412 km² (558 ]s) ] in western ] which had about 359 lions (as of April 2006). As in Africa, numerous human habitations are close by with the resultant problems between lions, livestock, locals and wildlife officials.<ref> | |||
| {{cite journal |author= Saberwal, V. K., Gibbs, J. P., Chellam, R., Johnsingh, A. J. T. |year=1994 |month=June |title= Lion-Human Conflict in the Gir Forest, India|journal=Conservation Biology |volume= |issue= |pages=501-507 |id= |url= |accessdate= |quote= }}</ref> The ] plans to establish a second independent population of ] at the ] in the Indian state of ].<ref>A.J.T. Johnsingh, (2004), ''Wildlife Institute of India Newsletter'', 11 (4)</ref> It is important to start a second population to serve as a ] for the last surviving Asiatic Lions and to help develop and maintain ] enabling the species to survive. | |||
| 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 twelve lions at ] in ], ] that are descended from animals owned by the ]. Another eleven animals believed to be Barbary lions were found in ] zoo, descendants of animals owned by ]. WildLink International, in collaboration with ], launched their ambitious International ] with the aim of identifying and breeding Barbary lions in captivity for eventual reintroduction into a National Park in the ] of ]. | |||
| ==Lions and humans== | |||
| ===Attacks on humans=== | |||
| While lions usually do not attack humans, some (usually male) lions seem to seek out human prey. Well-publicized cases include the ] and the ] man-eater. In both cases the hunters who killed the lions wrote books detailing the animals' predatory behavior. The Mfuwe and Tsavo incidents bear similarities: the lions in both incidents were larger than normal, lacked manes, and seemed to suffer from ]. Some have speculated that they might belong to an unclassified species of lion, or that they may have been sick and could not have easily caught prey.{{Fact|date=July 2007}} | |||
| The infirmity theory, including tooth decay, is not favored 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. |coauthors=Ellis J. Neiburger, Ellis J.; Kasiki, Samuel M. |year=2003 |month=February |title=Tooth Breakage and Dental Disease as Causes of Carnivore-Human Conflicts |journal=Journal of Mammalogy |volume=84 |issue=1 |pages=190–196 |id= |url=http://www.bioone.org/perlserv/?request=get-abstract&doi=10.1644%2F1545-1542(2003)084%3C0190%3ATBADDA%3E2.0.CO%3B2 |accessdate= 2007-07-06 |quote= }}</ref> In their analysis of Tsavo and man-eating generally, Peterhans and Gnoske acknowledge that sick or injured animals may be more prone to man-eating, but that the behavior is "not unusual, nor necessarily 'aberrant'" where the opportunity exists, and 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 amongst other pantherines and primates in the paleontological record.<ref>{{cite web |url=http://www.man-eater.info/gpage6.html |title= |accessdate=2007-07-07 |last=Peterhans |first=Julian C. Kerbis |coauthors=Thomas Patrick Gnoske |work=Man-eater.info |title=The Science of Man-eating }}</ref> | |||
| ] in the ]]] | |||
| The lion's proclivity for man-eating has been systematically examined. American and Tanzanian scientists report that man-eating behavior 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 ] in ] and in ] near the ] 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>{{cite journal |last=Packer |first=C. |coauthors=Ikanda, D.; Kissui, B.; Kushnir, H. |year=2005 |month=August |title=Conservation biology: lion attacks on humans in Tanzania |journal=] |volume=436 |issue=7053 |pages=927-928 |url=http://www.ncbi.nlm.nih.gov/sites/entrez?cmd=Retrieve&db=PubMed&list_uids=16107828&dopt=Citation |accessdate= 2007-07-06 |quote= }}</ref> | |||
| Author Robert R. Frump wrote in ''The Man-eaters of Eden'', published in 2006, that Mozambican refugees crossing Kruger National Park at night in ] regularly are attacked and eaten by the lions there. Park officials conceded that man-eating is a problem there. Frump believes thousands may have been killed in the decades after ] sealed the park and forced the refugees to cross the park at night. Mozambicans had for nearly a century before the border was sealed regularly walked across the park in daytime with little harm.<ref>Frump, R. (2006) ''The Man-eaters of Eden'', The Lyons Press. ISBN 1-592288-92-9</ref> | |||
| Packer estimates more than 200 Tanzanians are killed each year by lions, ]s, elephants, hippos and ]s, and that the numbers could be double that amount. Lions are thought to kill about 70 humans per year at least in Tanzania, Packer and Ikanda note. Packer and Ikanda are among the few ] 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. | |||
| 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 web |title=Toothache 'made lion eat humans' |author=Daniel Dickinson |date=October 19, 2004 |publisher=BBC News |url=http://news.bbc.co.uk/1/hi/world/africa/3756180.stm |accessdate=2007-07-20 }}</ref><ref>{{cite web |title=Terror from man-eating lions increasing in Tanzania |author=Ludger Kasumuni |date=August 28, 2006 |publisher=IPPmedia.com |url=http://www.ippmedia.com/ipp/guardian/2006/08/28/73305.html |accessdate=2007-07-20 }}</ref> Dr Rolf D. Baldus, the GTZ wildlife programme coordinator, is reported to have commented that it was likely that the lion preyed on humans because it had a large ] underneath a ] which was cracked in several places. He further commented that "This lion probably experienced a lot of pain, particularly when it was chewing." GTZ is the German development cooperation agency and has been working with the Tanzanian government on wildlife conservation for nearly two decades. Like in other cases this lion was large, lacked a mane, and had a tooth problem. | |||
| The "All-Africa" record of man-eating generally is considered to be not Tsavo, but the lesser known incidents in the late 1930s through the late 1940s in what was then Tanganyika (now Tanzania). 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 in what is now Njombe district. | |||
| Tsavo and Patterson body counts vary from a confirmed 28 victims up to a possible 140. While some authors disparage the actual number, it should be kept in mind that Patterson kept firm records of skilled-labor killed by the lions, not indigenous Africans. The toll could easily have been much higher. | |||
| ===Lions in captivity=== | |||
| ====Baiting and taming==== | |||
| {{main|Lion-baiting|Lion taming}} | |||
| Lion-baiting is a ] involving the ] of lions in combat with other animals, usually dogs. Records of it exist in ancient times through until the 17th century. It was finally banned in England in 1825. | |||
| Lion taming refers to the practice of taming lions generally for entertainment, either as part of an established ] or as an individual act, such as ]. The term is also often used for the taming and display of other big cats such as ], ] and ]. The now iconic lion tamer's chair was possibly first used by American ] (1903–1965).<ref>{{cite book | first=David | last=Feldman | authorlink=David Feldman (author) | coauthors= | year=1993 | title=How Does Aspirin Find a Headache? | edition= | publisher=HarperCollins | location= | id=ISBN 0-06-016923-0 }}</ref> | |||
| ===Cultural depictions=== | |||
| {{details|Cultural depictions of lions}} | |||
| ] | |||
| The Lion has been an icon of 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 but noble. A consistent depiction is their representation as "] of the jungle" or "king of the beasts"; hence, the lion has been a popular symbol of royalty and stateliness, as well as a symbol of bravery. ], for instance, was known as Richard the Lionheart. (] would later be adopted as an album title by ], ], and ], as well as in films and computer games.) | |||
| Representations of lions date back some 15,000 years to the earliest ] ]s: two lions were depicted mating in the Chamber of Felines in the ] caves. ] venerated the lion, with the ] and the lion-headed deity ]; ] and ] were Egyptian deities in full lion form. The ] was symbolic in Ancient Greece and Rome, represented as the constellation and zodiac sign ], and described in mythology, where its skin was borne by the hero ]. | |||
| ] | |||
| The lion was a prominent symbol in both the ] and ] periods. 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 ] is said to have been delivered from the lion's den. Such symbolism was appropriated by Saddam Hussein's regime in Iraq for their ], with the technology adapted from a Russian model. | |||
| The lion is symbolic for the ], ]'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 ]. | |||
| In the medieval period in Europe, lions were frequently depicted on ], either as a device on shields themselves, or as ]. The formal language of ], called a ], employs French terms to describe the images precisely. Such descriptions were required to specify whether lions (or other creatures) were "rampant" or "passant", that is whether they are rearing or crouching. | |||
| Lions continue to feature in modern tales, from the messianic ] in the '']'' and following books from the ] series written by ], to the comedic ] in the '']''.<ref>L. Frank Baum, Michael Patrick Hearn, ''The Annotated Wizard of Oz'', p 148, ISBN 0-517-500868 </ref> The word '']'' is Turkish for ''lion''. | |||
| The advent of moving pictures saw the continued presence of leonine symbols; one of the most iconic and widely seen and recognised lions is ], the mascot for ] (MGM) studios. The 1960s saw what is possbily the most famous lioness, Kenyan lioness ] in the movie ] appeared, based on the true-life international bestselling book of the same title. The same decade saw the first Japanese colour TV animation series ]. More recently, animated lions proved popular in ], which featured the popular song ] in its soundtrack. The lion is a common symbol or epithet for sporting teams; from national Soccer teams such as ], ] and ] to famous clubs such as the ]<ref>{{cite web | author = | title = Official Website of the Detroit Lions | publisher = Detroit Lions | date = 2001 | url = http://www.detroitlions.com/index.cfm?homelink=y | accessdate = 2007-07-08}}</ref> of the NFL and ]<ref name="new crest">{{cite news | date=] | url=http://news.bbc.co.uk/sport1/hi/football/teams/c/chelsea/4008257.stm | title=Chelsea centenary crest unveiled | publisher=BBC | accessdate=2007-01-02 | author=}}</ref> and ] of the ] to a ] around the world. Villa sport a Scottish ] on their crest, as do ] and ] of the ]. | |||
| <gallery> | |||
| Image:Male lion face.jpg|A male lion in Ngorongoro Crater, Tanzania. | |||
| Image:Lions at night.jpg|Lions hunting at night in the Sabi Sands, South Africa. | |||
| Image:Lioness small.jpg|A lioness in Ngorongoro Crater, Tanzania. | |||
| Image:Lion (Panthera leo) eye close-up.jpg|Close-up of a lion's eye | |||
| </gallery> | |||
| ==References== | |||
| {{Reflist|2}} | |||
| ==External links== | |||
| {{commons|Panthera leo|Lion}} | |||
| {{wikispecies|Panthera leo|Lion}} | |||
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| * from “ARKive images of life on Earth” website | |||
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| * – The official website of a research group at the U. of Minnesota that has conducted extensive field research on lions and has published over 50 peer-reviewed scientific articles on lions. | |||
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| * – A fund dedicated to the research and conservation of the lion | |||
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| * Photogalleries: , | |||
| * ] video: | |||
| * Short videos from Botswana: , , | |||
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