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Lion

Male


Female (Lioness)

Conservation status


Vulnerable (IUCN) Nowell & Bauer (2004). Panthera leo. 2006 IUCN Red List of Threatened Species. IUCN 2006. Retrieved on 11 May 2006. Database entry includes a lengthy justification of why this species is vulnerable

Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Felidae
Genus: Panthera
Species: P. leo
Binomial name
Panthera leo
(Linnaeus, 1758)

Distribution of lions in Africa

Synonyms
Felis leo
(Linnaeus, 1758)

The lion (Panthera leo) is a member of the family Felidae and one of four big cats in the genus Panthera. With exceptionally large males exceeding 250 kg (550 lb) in weight,Nowak, Ronald M. (1999). Walker\'s Mammals of the World. Baltimore: Johns Hopkins University Press. ISBN 0-8018-5789-9.  it is the second-largest living cat after the tiger. Wild lions currently exist in sub-Saharan Africa and in Asia with a critically endangered remnant population in northwest India, having disappeared from North Africa, the Middle East, and western Asia in historic times. Until the late Pleistocene (about 10,000 years ago), the lion was the most widespread large land mammal beside humans. They were found in most of Africa, much of Eurasia from western Europe to India, and the Bering land bridge and, in the Americas, from the Yukon to Peru.

Lions live for approximately 10–14 years in the wild, while in captivity they can live over 20 years. They typically inhabit savanna and grassland, although they may take to bush and forest. Lions are unusually 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 ungulates. The lion is an apex and keystone predator, although they will resort to scavenging if the opportunity arises. While lions, in general, do not selectively hunt humans, some have been known to become man-eaters and seek human prey.

The lion is a vulnerable species, having seen a possibly irreversible population decline of 30 to 50% over the past two decades in its African range; populations are untenable outside designated reserves and national parks. Although the cause of the decline is not well-understood, habitat loss and conflicts with humans are currently the greatest causes of concern. Lions have been kept in menageries since Roman times and have been a key species sought after and exhibited in zoos the world over since the late eighteenth century. Zoos are cooperating worldwide in breeding programs for the endangered Asiatic subspecies.

Visually, the male is highly distinctive and is easily recognized by its mane. The head of the male lion is one of the most widely recognized animal symbols in human culture. It has been depicted extensively in literature, in sculptures, in paintings, on national flags, and in contemporary films and literature.

The lioness has been recognized, however, as the pinnacle of hunting prowess from the earliest of human writings and graphic representations. The lionesses are the hunters for their pride and execute their skills with precision and complex teamwork. Each lioness develops specific skills for her role in the hunting techniques used by her pride and, generally, assumes that role during most hunts. Members of human cultures living among lions in natural habitats have understood this characteristic and often have chosen the lioness to represent their most ferocious war deities and warriors, often naming their male rulers as her "son". Examples drawn from the earliest of written records include the Egyptian pantheon deities of Sekhmet, Bast, Menhit, and Tefnut and these deities may have had precursors in Nubia and Lybia. Other Egyptian deities are quite complex and assume aspects that may include one as a lioness headed human or a lioness in specific roles. Depictions of lions hunting in groups have existed from the Upper Paleolithic period, with carvings and paintings from the Lascaux and Chauvet Caves.

Contents

Naming and etymology

The lion\'s name, similar in many Romance languages, derives from the Latin leo,Simpson DP (1979). Cassell\'s Latin Dictionary, 5th edition, London: Cassell Ltd., p. 883. ISBN 0-304-52257-0.  and before that the Ancient Greek leōn/λεων.Liddell, Henry George and Robert Scott (1980). A Greek-English Lexicon (Abridged Edition). United Kingdom: Oxford University Press. ISBN 0-19-910207-4.  The Hebrew word lavi (לָבִיא) may also be related, "Lion". Oxford English Dictionary (2nd edition). (1989). Ed. Simpson, J., Weiner, E. (eds). Oxford: Clarendon Press. ISBN 0-19-861186-2.  as well as the Ancient Egyptian rw.yourdictionary.com. As in other ancient scripts, in Ancient Egyptian only the consonants are written. No distinction was made between \'l\' and \'r\'. It was one of the many species originally described, as Felis leo, by Linnaeus in his eighteenth century work, Systema Naturae.(Latin) Linnaeus, C. (1758). Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.. Holmiae. (Laurentii Salvii)., p. 824.  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 folk etymology. Although it came into English through the classical languages, panthera is probably of East Asian origin, meaning "the yellowish animal," or "whitish-yellow"."Panther". Online Etymology Dictionary. Douglas Harper. Retrieved on 2007-07-05.

Taxonomy and evolution

Skull of a modern lion at Kruger National Park

The oldest lion-like fossil is known from Laetoli in Tanzania and is perhaps 3.5 million years old; some scientists have identified 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.Werdelin, Lars; Lewis, Margaret E. (June 2005). "Plio-Pleistocene Carnivora of eastern Africa: species richness and turnover patterns". Zoological Journal of the Linnean Society 144 (2): 121–144. The Linnean Society of London. Retrieved on 2007-07-08. The closest relatives of the lion are the other Panthera species: the tiger, the jaguar and the leopard. Morphological and genetic studies reveal that the tiger was the first of these recent species to diverge. About 1.9 million years ago the Jaguar branched off 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.Yu, Li; Ya-ping Zhang (May 2003). "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 35 (2): 483–495. doi:10.1016/j.ympev.2005.01.017.

Panthera leo itself evolved in Africa between 1 million and 800,000 years ago before spreading throughout the Holarctic region;Yamaguchi, Nobuyuki; Alan Cooper, Lars Werdelin and David W. Macdonald (August 2004). "Evolution of the mane and group-living in the lion (Panthera leo): a review". Journal of Zoology 263 (4): 329–42. doi:10.1017/S0952836904005242. ISSN 1469-7998. It appeared 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), which appeared about 300,000 years ago. During the upper Pleistocene the lion spread to North and South America, and developed into Panthera leo atrox, the American lion.Turner, Allen (1997). The big cats and their fossil relatives : an illustrated guide to their evolution and natural history. New York: Columbia University Press. ISBN 0-231-10229-1.  Lions died out in northern Eurasia and America at the end of the last glaciation, about 10,000 years ago; this may have been secondary to the extinction of megafauna.Harington, CR (1996). American Lion. Yukon Beringia Interpretive Centre website. Yukon Beringia Interpretive Centre. Retrieved on 2007-09-22.

Subspecies

Southwest African lion (Panthera leo bleyenberghi)

Traditionally 12 recent subspecies of lion were recognized, the largest of which has been recognised as the Barbary lion.Barbary Lion - Panthera leo leo - Largest Lion Subspecies Retrieved on 19 September, 2007 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; additionally, they were often based upon zoo material of unknown origin that may have had "striking, but abnormal" morphological characteristics.Grisham, Jack (2001). "Lion". Encyclopedia of the World\'s Zoos Volume 2: G–P. Ed. Catherine E. Bell. Chofago: Fitzroy Dearborn. pp.733–739. ISBN1-57958-174-9.  Today only eight subspecies are usually accepted,Burger, Joachim et al. (March 2004). "Molecular phylogeny of the extinct cave lion Panthera leo spelaea". Molecular Phylogenetics and Evolution 30 (3): 841–849. doi:10.1016/j.ympev.2003.07.020. Retrieved on 2007-09-20. but one of these, the Cape lion formerly described as Panthera leo melanochaita is probably invalid. Even the remaining seven subspecies might be too much; mitochondrial variation in recent African lions is modest, which suggests that all sub-Saharan lions could be considered a single subspecies, possibly divided in two main clades: one to the west of the Great Rift Valley and the other to the east. Lions from Tsavo in Eastern Kenya are much closer genetically to lions in Transvaal (South Africa), than to those in the Aberdare Range in Western Kenya.Barnett, Ross; Nobuyuki Yamaguchi, Ian Barnes and Alan Cooper (2006). "The origin, current diversity and future conservation of the modern lion (Panthera leo)" (PDF). Proceedings of the Royal Society B: Biological Sciences 273 (1598): 2119–2125. doi:10.1098/rspb.2006.3555. Retrieved on 2007-09-04.Dubach, Jean; et al (January 2005). "Molecular genetic variation across the southern and eastern geographic ranges of the African lion, Panthera leo". Conservation Genetics 6 (1): 15–24. doi:10.1007/s10592-004-7729-6.

Recent

Eight recent subspecies are recognized today:

  • P. l. persica, known as the Asiatic- or South Asian, Persian or Indian lion, was once widespread from Turkey, across the Middle East, to Pakistan, India and even Bangladesh. However, large prides and daylight activity made it easier to poach than tigers or leopards; now around 300 exist in and near the Gir Forest of India.Wildlife Conservation Trust of India (2006). Asiatic Lion - History. Asiatic Lion Information Centre. Wildlife Conservation Trust of India. Retrieved on 2007-09-15.
  • P. l. leo, known as the Barbary lion, is extinct in the wild due to excessive hunting, although captive individuals may still exist. This was the largest of the lion subspecies, at 3–3.5m approx., and weighing over 150 kilograms and more. They ranged from Morocco to Egypt. The last wild Barbary lion was killed in Morocco in 1922.
  • 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 Congo.
  • P. l. nubica, known as the East African- or Massai lion, is found in East Africa, from Ethiopia and Kenya to Tanzania and Mozambique.
  • P. l. bleyenberghi, known as the Southwest African- or Katanga lion. It is found in southwestern Africa, Zimbabwe, Angola, Katanga (Zaire).
  • P. l. krugeri, known as the Southeast African- or Transvaal lion, is found in the Transvaal region of South eastern Africa, including Kruger National Park.
  • P. l. melanochaita, known as the Cape lion, became extinct in the wild around 1860. Results of mitochondrial DNA 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.

Prehistoric

Several additional subspecies of lion existed in prehistoric times:

  • P. l. atrox, known as the American lion or American cave lion, was abundant in the Americas from Alaska to Peru in the Pleistocene Epoch 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). 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).Martin, P.S. (1984). Quaternary Extinctions. Tucson, Arizona: University of Arizona Press. ISBN 0-8165-1100-4. 
  • P. l. fossilis, known as the Early Middle Pleistocene European cave lion, flourished about 500,000 years ago; fossils have been recovered from Germany and Italy.

Cave lions, Chamber of Felines, Lascaux caves

  • 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. This species is known from Paleolithic cave paintings (such as the one displayed to the right), ivory carvings, and clay busts, Packer, Craig; Jean Clottes (November 2000). "When Lions Ruled France" (PDF). Natural History: 52–57. Retrieved on 2007-08-27. indicating it had protruding ears, tufted tails, perhaps faint tiger-like stripes, and that at least some males had a ruff or primitive mane around their necks. With this example being a hunting scene it is likely that it depicts females hunting for the pride using the same strategy as their contemporary relatives and males may not be part of the subject.
  • P. l. vereshchagini, known as the East Siberian- or Beringian cave lion, was found in Yakutia (Russia), Alaska (USA), and the Yukon Territory (Canada). Analysis of skulls and mandibles of this lion demonstrate that it is distinct—larger than the European cave lion and smaller than the American cave lion with differing skull proportions.Baryshnikov, G.F.; G. Boeskorov (2001). "The Pleistocene cave lion, Panthera spelaea (Carnivora, Felidae) from Yakutia, Russia". Cranium 18 (1): 7–24.

Dubious

  • 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, P. Deraniyagala erected this subspecies in 1939.Kelum Manamendra-Arachchi, Rohan Pethiyagoda, Rajith Dissanayake, Madhava Meegaskumbura (2005). "A second extinct big cat from the late Quaternary of Sri Lanka.". The Raffles Bulletin of Zoology Supplement 12: 423–434. National University of Singapore. Retrieved on 2007-07-31.
  • P. l. europaea, known as the European lion, was probably identical with Panthera leo persica or Panthera leo spelea; its status as subspecies is unconfirmed. It became extinct around 100 AD due to persecution and over-exploitation. Inhabited the Balkans, the Italian Peninsula, southern France and the Iberian Peninsula. It was a very popular object of hunting among Romans, Greeks and Macedonians.
  • P. l. youngi or Panthera youngi , known as the North-Eastern Pleistocene China cave lion, flourished 350,000 years ago.Harington, CR (1969). "Pleistocene remains of the lion-like cat (Panthera atrox) from the Yukon Territory and northern Alaska". Canadian Journal Earth Sciences 6 (5): 1277–88. Its relationship to the extant lion subspecies is obscure, and probably represents a distinct species.
  • P. l. maculatus, known as the Marozi 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 hybrid commonly known as a leopon.Shuker, Karl P.N. (1989). Mystery Cats of the World. Robert Hale. ISBN 0-7090-3706-6. 

Hybrids

A liger is the offspring of a male lion and female tiger.

Further information: Panthera hybrid, liger and tigon

Lions also have been known to breed with tigers (most often the Amur and Bengal subspecies) to create hybrids called ligers and tigons.Guggisberg, C. A. W. (1975). Wild Cats of the World. New York: Taplinger Publishing. ISBN 0-8008-8324-1.  They have also been crossed with leopards to produce leopons,Doi H, Reynolds B (1967). The Story of Leopons. New York: Putnam. OCLC 469041.  and jaguars to produce 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.

The liger is a cross between a male lion and a tigress.Markel, Scott; Darryl León (2003). Sequence Analysis in a Nutshell: a guide to common tools and databases. Sebastopol, California: O\'Reily. ISBN 0-596-00494-X.  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 their manes will be modest: around 50% of a pure lion mane. Ligers are typically between 3.0 and 3.7 m (10 to 12 feet) in length, and can be between 360 and 450 kg (800 to 1,000 pounds) or more. The less common tigon is a cross between the lioness and the male tiger.tigon - Encyclopædia Britannica Article. Retrieved on 12 September, 2007.

Physical characteristics

During confrontations with others, the mane makes the lion look bigger than he really is.

The lion is the second largest feline after the tiger. With powerful legs, a strong jaw, and long canine teeth, the lion can bring down and kill large prey.Lion. Honolulu Zoo. Retrieved on 2007-07-12. Lion coloration varies from light buff to yellowish, reddish or dark ochraceous brown. The underparts are generally lighter and the tail tuft is black. The color of the mane varies from blond to black.

Weights for adult lions generally lie between 150–250 kg (330–550 lb) for males, and 120–180 kg (260–400 lb) for females.Nowak, Ronald M. (1999). Walker\'s Mammals of the World. Baltimore: Johns Hopkins University Press. ISBN 0-8018-5789-9.  Nowell and Jackson report average weights of 181 kg for males and 126 kg for females; one male shot near Mount Kenya was weighed at 272 kg (600 lb).Nowell K, Jackson P (1996). "Panthera Leo", [http://carnivoractionplans1.free.fr/wildcats.pdf Wild Cats: Status Survey and Conservation Action Plan] (PDF), Gland, Switzerland: IUCN/SSC Cat Specialist Group, 17–21. ISBN 2-8317-0045-0.  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).Nowak, Ronald M. (1999). Walker\'s Mammals of the World. Baltimore: Johns Hopkins University Press. ISBN 0-8018-5789-9.  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 a tufted tail and the function of the tuft and spine are unknown. Absent at birth, the tuft develops around 5½ months of age and readily identifiable at 7 months.Schaller, p. 28

Mane

Thermal image of a lion, showing the insulating manes

The mane of the male lion, unique amongst cats, is one of the most distinctive characteristics of the species. It makes the lion appear larger, providing an excellent intimidation display; this aids the lion during confrontations with other lions and with the species\' chief competitor in Africa, the spotted hyena. Trivedi, Bijal P. (2005). Are Maneless Tsavo Lions Prone to Male Pattern Baldness?. National Geographic. Retrieved on 2007-07-07. The presence, absence, color, 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.Trivedi, Bijal P. (August 22, 2002). Female Lions Prefer Dark-Maned Males, Study Finds. National Geographic News. National Geographic. Retrieved on 2007-09-01. 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.West, Peyton M.; Craig Parker (August 2002). "Sexual Selection, Temperature, and the Lion\'s Mane". Science 297 (5585): 1339–1343. doi:10.1126/science.1073257. In prides including a coalition of two or three males, it is possible that lionesses solicit mating more actively with heavily maned lions.

Scientists once believed that the distinct status of some subspecies could be justified by 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 influence the color and size of a lion\'s mane, such as the ambient temperature. The cooler ambient temperature in European and North American zoos, for example, can result in a heavy mane. Thus the mane is an inappropriate marker for identifying subspecies.Barnett, Ross; Nobuyuki Yamaguchi, Ian Barnes and Alan Cooper (August 2006). "Lost populations and preserving genetic diversity in the lion Panthera leo: Implications for its ex situ conservation". Conservation Genetics 7 (4): 507–514. doi:10.1007/s10592-005-9062-0. Yamaguchi, Nobuyuki; B. Haddane (2002). "The North African Barbary lion and the Atlas Lion Project". International Zoo News 49: 465–481. However the males of the Asiatic subspecies are characterized by sparser manes than average African lions.Menon, Vivek (2003). A Field Guide to Indian Mammals. Delhi: Dorling Kindersley India. ISBN 0-14-302998-3. 

A maneless male lion, who also has little body hair - from Tsavo East National Park, Kenya

Maneless male lions have been reported in Senegal and Tsavo East National Park in Kenya, and the original male white lion from Timbavati also was maneless. Castrated lions have minimal manes. The lack of a mane is found in inbred lion populations; inbreeding also results in poor fertility.Trivedi, Bijal P. (12 June 2002). To Boost Gene Pool, Lions Artificially Inseminated. National Geographic News. National Geographic. Retrieved on 2007-09-20.

Lioness showing the ruff that sometimes leads to misidentification as a male

Many lionesses have a ruff that may be apparent in certain poses. Sometimes it is indicated in sculptures and drawings, especially ancient artwork, and is misinterpreted as a mane, hence interpreted as a male. It differs, however, from the mane of the male in that the ruff of a lioness is at the jaw line below the ears, of much less hair length, and frequently, not noticeable; whereas the mane of the male extends above the ears, often obscuring the outline of the ears entirely.

Cave paintings of extinct European cave lions exclusively show animals with no mane or, just the hint of a mane, suggesting to some that they were more or less maneless,(German) Koenigswald, Wighart von (2002). Lebendige Eiszeit: Klima und Tierwelt im Wandel. Stuttgart: Theiss. ISBN 3-8062-1734-3.  however, the females hunting for a pride are more likely the subjects of the drawings—since they are shown in a group related to hunting—so these images are not reliable to make a judgment about whether the males had manes. The drawings do suggest that the extinct species used the same social orginazation and hunting strategies as contemporary prides.

White lions

White lions owe their coloring to a recessive gene; they are rare forms of the subspecies Panthera leo krugeri

The white lion is not a distinct subspecies, but a special morph with a genetic condition, leucism, that causes paler colouration akin to that of the white tiger; the condition is similar to melanism, which causes black panthers. White animals of the Transvaal lion (Panthera leo krugeri) occasionally have been encountered in and around the Kruger National Park and the adjacent Timbavati Private Game Reserve in eastern South Africa, but more commonly, are found in captivity, where breeders deliberately select them. The unusual cream color of their coats is due to a recessive gene.McBride, Chris (1977). The White Lions of Timbavati. Johannesburg: E. Stanton. ISBN 0-949997-32-3.  Reportedly, they have been bred in camps in South Africa for use as trophies for canned hunts.Tucker, Linda (2003). Mystery of the White Lions - Children of the Sun God. Mapumulanga: Npenvu Press. ISBN 0-620-31409-5. 

Confirmation of the existence of white lions only came in the late twentieth century. For hundreds of years prior, the white lion had been thought to be a figment of legend circulating in South Africa, the white pelage of the animal said to represent the goodness in all creatures. Sightings were first reported in the early 1900s, and continued, infrequently, for almost fifty years until, in 1975, a litter of white lion cubs was found at Timbavati Game Reserve.The rare white lions Retrieved on 20 September, 2007.

Biology and behavior

Lions spend much of their time resting and are inactive for about 20 hours per day.Schaller, p. 122 Although lions can be active at any time, their activity generally peaks after dusk with a period of socializing, grooming and defecating. Intermittent bursts of activity follow through the night hours to dawn, when hunting most often takes place. They spend an average two hours a day walking and 50 minutes eating.Schaller, p. 120–1

Hunting and diet

While a lioness such as this has very sharp teeth, prey is usually killed by strangulation

Lionesses are powerful animals who usually hunt in groups and stalk their chosen prey. They can reach speeds of 59 km/h (40 mph),Schaller, p. 233 although, only for short bursts,Schaller, p. 247–8 so they have 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.Schaller, p. 237 They sneak up to the victim until they reach a distance of approximately 30 m (98 feet) or less. Typically, several female lions work together and encircle the herd from different points. Once they have closed with a herd, they usually target the closest prey. 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.Dr Gus Mills. About lions - Ecology and behaviour. African Lion Working Group. Retrieved on 2007-07-20.

The prey consists mainly of large mammals, with a preference for wildebeest, impalas, zebras, buffalo, and warthogs in Africa and nilgai; wild boar and several deer species in India. Many other species are hunted, based on availability. Mainly this will include ungulates weighing between 50 and 300 kg such as kudu, hartebeest, gemsbok, and eland.Ronald M. Nowak: Walker\'s Mammals of the World. Occasionally, they take relatively small species such as Thomson\'s gazelle or springbok. Lions living near the Namib coast feed extensively on seals.http://www.5050.co.za/inserts.asp?ID=8258 While hunting in groups, they are capable of taking down most animals, even healthy adults, but they rarely attack very large prey such as buffalo bulls or fully grown male giraffes, due to the danger of injury.African Lion Hunting Habits in South Africa and Southern Africa. Retrieved on 31 July, 2007.

They normally feed on mammals no larger than 550 kg, which excludes most adult hippopotamuses, rhinoceroses, elephants, giraffes, and buffalos.Hayward, Matt W.; Graham Kerley (2005). "Prey preferences of the lion (Panthera leo)". Journal of Zoology 267 (3): 309–322. doi:10.1017/S0952836905007508. In some areas, they specialise in hunting atypical prey-species; this is the case at the Savuti river, where they prey on young elephants.Kemp, Leigh. The Elephant Eaters of the Savuti. go2africa. Park guides in the area reported that the lions, driven by extreme hunger, started taking down baby elephants, and then moved on to adolescents and, occasionally, fully grown adults.Whitworth, Damien. "King of the jungle defies nature with new quarry", The Australian, October 9, 2006. Retrieved on 2007-07-20.  In the Kruger National Park, giraffes are regularly hunted.Pienaar U de V (1969). "Predator-prey relationships amongst the larger mammals of the Kruger National Park". Koedoe 12: 108-76. Lions also attack domestic livestock; in India cattle contribute significantly to their diet.Vivek Menon: A Field Guide to Indian Mammals. They are capable of killing other predators such as leopards, cheetahs, hyenas, and wild dogs, as well as scavenging animals either dead from natural causes or killed by other predators.Schaller, p. 213 A lion may gorge itself and eat up to 30 kg (66 lb) in one sitting;Guggisberg, C. A. W. (1961). Simba: the life of the lion.. Cape Town: Howard Timmins.  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.Schaller, p. 270–76 An adult lioness requires an average of about 5 kg (11 lb) of meat per day, a male about 7 kg (15.4 lb).Lions. Honolulu Zoo. Retrieved on 2007-07-20.

The hunters of a pride sharing a zebra where the kill occurred

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 prey more easily against other large predators such as hyenas, which may be attracted by vultures over kilometers in open savannas. Lionesses do most of the hunting. In typical 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.Stander, P. E. (1992). "Cooperative hunting in lions: the role of the individual". Behavioral Ecology and Sociobiology 29 (6): 445—454. doi:10.1007/BF00170175.

Males attached to prides do not usually participate in the hunting by the lionesses, except when the pride is hunting large animals such as giraffe and buffalo, males lions also hunt when they are bachelors which have no pride of their own. Male lions have also been observed and recorded hunting in groups.

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.Schaller, p. 153 Depending upon how quickly they mature, two or three is the age at which males are excluded from the pride.

Reproduction

Most lionesses will have reproduced by the time they are four years of age.Schaller, p. 29 Lions do not mate at any specific time of year, and the females are polyestrous.Schaller, p. 174 As with 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.Asdell, Sydney A. [1964] (1993). Patterns of mammalian reproduction. Ithaca: Cornell University Press. ISBN 978-0-8014-1753-5.  A lioness may mate with more than one male when she is in heat;Schaller, p. 142 during a mating bout, which could last several days, the couple copulates twenty to forty times a day and are likely to forgo eating. Lions reproduce very well in captivity.

During a mating bout, a couple may copulate twenty to forty times a day for several days; the ruff of the female is clearly visible

The average gestation period is around 110 days, the female giving birth to a litter of one to four cubs. Lionesses 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 usually are born and initially kept hidden from view in thickets or sheltered areas. They weigh 1.2–2.1 kg at birth and are almost helpless, beginning to crawl a day or two after birth and walking around three weeks of age.Schaller, p. 143 Weaning occurs 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.Macdonald, David (1984). The Encyclopedia of Mammals. New York: Facts on File, 31. ISBN 0-87196-871-1. 

When one or more new males oust the previous male(s) associated with a pride, the conqueror(s) often kill any existing young cubs, Packer, C., Pusey, A. E. (May 1983). "Adaptations of female lions to infanticide by incoming males". American Naturalist 121 (5): 716–728. Retrieved on 2007-07-08. perhaps, because females do not become fertile and receptive until their cubs mature or die. 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.Crandall, Lee S. (1964). The management of wild animals in captivity. Chicago: University of Chicago Press. OCLC 557916.  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 rarely are successful. He usually kills all of the existing cubs who are less than two years old. The lioness is much lighter and has less strength than the male, however, success is more likely when a group of three or four mothers within the pride join forces against one male.

One scientific study reports that both males and females may interact homosexually.Bagemihl, Bruce (1999). Biological Exuberance: Animal Homosexuality and Natural Diversity. New York: St. Martin\'s Press, pp. 302–305. ISBN 0-312-19239-8.  Srivastav, Suvira (15–31 December 2001). Lion, Without Lioness. TerraGreen: News to Save the Earth. Terragreen. Retrieved on 2007-09-02. 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. Female pairings are held to be fairly common in captivity, but have not been observed in the wild.

Health

Male lion in Ngorongoro Crater

Though adult lions have no natural predators, evidence suggests that the majority die violently from humans or other lions.Schaller, p. 183 Various species of tick commonly infest the ears, neck and groin regions of most lions.Schaller, p. 184Yeoman, G.; Jane B. Walker (1967). The ixodid ticks of Tanzania. London: Commonwealth Institute of Entomology. OCLC 955970.  Adult forms of several species of the tapeworm genus Taenia have been isolated from intestines, the lions having ingested larval forms from antelope meat.(German)Sachs, R (1969). "Untersuchungen zur Artbestimmung und Differenzierung der Muskelfinnen ostafrikanischer Wildtiere [Differentiation and species determination of muscle-cysticerci in East African game animals]". Zeitschrift für tropenmedizin und Parasitologie 20 (1): 39–50. PMID 5393325. Lions in the Ngorongoro Crater were afflicted by an outbreak of 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.Fosbrooke, Henry (1963). "The stomoxys plague in Ngorongoro". East African Wildlife Journal 1: 124–6. A more recent outbreak in 2001 killed six lions.Nkwame, Valentine M. "King of the jungle in jeopardy", The Arusha Times, September 9, 2006. Retrieved on 2007-09-04.  Lions, especially in captivity, are vulnerable to the Canine distemper virus (CDV), feline immunodeficiency virus (FIV) and feline infectious peritonitis (FIP). CDV is spread through domestic dogs and other carnivores; a 1994 outbreak in Serengeti National Park resulted in many lions developing neurological symptoms such as seizures. During the outbreak, several lions died from pneumonia and encephalitis.M.E. Roelke-Parker et al. (February 1996). "A canine distemper epidemic in Serengeti lions (Panthera leo)" (PDF). Nature 379: 441–445. doi:10.1038/379441a0. PMID 8559247. Retrieved on 2007-07-08. 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.

Group organization

A pride on the move near Governors Camp, in the Massai Mara, Kenya

Lions are predatory carnivores who manifest two types of social organization. Some are residents, living in groups, called prides.Schaller, p. 33 The pride usually consists of approximately five or six related females, their cubs of both sexes, and one or two males known as a coalition who mate with the adult females. The coalition of males associated with a pride are usually two, but may increase to four and decrease again over time. Male cubs are excluded from their maternal pride when they are reaching maturity.

The second organizational behaviour is labeled, nomads, who range widely and move about sporadically, either singularly or in pairs. 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 have to go through this lifestyle and some never are 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 reject most attempts by an unrelated female to join their family group.

The area a pride occupies is called a pride area, whereas that by a nomad is a range. The males associated with a pride tend to stay on the finges, patrolling their territory. Why 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 "cheaters" reduce per capita caloric intake. 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.

Lioness in a burst of speed while hunting in the Serengeti

Lionesses do the hunting for their pride. Lionesses hunt as a co-ordinated group and are smaller, swifter, and more agile than males. They are not encumbered by the heavy and conspicuous mane which causes overheating during exertion. There is no clear hierarchy with food except the females are likely to share with each other first and allow cubs and males to eat if the kill is large enough; males will take some of this food from cubs, but are more likely to share alongside the cubs than the lionesses. Male lions will never share food they have killed by themselves. 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 with larger kills.Schaller, p. 133

Both males and females defend the pride against intruders. Some individual lions consistently lead the defense against intruders, while others lag behind.Heinsohn, R.; C. Packer (1995). "Complex cooperative strategies in group-territorial African lions". Science 269 (5228): 1260–1262. doi:10.1126/science.7652573. PMID 7652573. These "laggards" are not punished by leaders. Possibly laggards provide other services to the group so that leaders forgive them.Morell, V. (1995). "Cowardly lions confound cooperation theory". Science 269 (5228): 1216–1217. doi:10.1126/science.7652566. An alterna