Naming and etymology
The lion's name, similar in many Romance languages, derives from the Latin leo,[3] and before that the Ancient Greek leōn/λεων.[4] The Hebrew word lavi (לָבִיא) may also be related,[5] as well as the Ancient Egyptian rw.[6] It was one of the many species originally described, as Felis leo, by Linnaeus in his eighteenth century work, Systema Naturae.[7] 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".[8]

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.[9] 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.[10]
Panthera leo itself evolved in Africa between 1 million and 800,000 years ago before spreading throughout the Holarctic region;[11] 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.[12] Lions died out in northern Eurasia and America at the end of the last glaciation, about 10,000 years ago;[13] this may have been secondary to the extinction of megafauna.[14]

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.[15] 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.[16] Today only eight subspecies are usually accepted,[17][13] but one of these, the Cape lion formerly described as Panthera leo melanochaita is probably invalid.[17] 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.[18][19]

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.[20]
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.[21]
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.[17]

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).[13] 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).[22]
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.[13] This species is known from Paleolithic cave paintings (such as the one displayed to the right), ivory carvings, and clay busts, [23] 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.[24] 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.[25][13]

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.[26]
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.[27] 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.[28]

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.[29] They have also been crossed with leopards to produce leopons,[30] 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.[31] 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.[31] The less common tigon is a cross between the lioness and the male tiger.[32]

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.[33] 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.[2] 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).[21] 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).[2] 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.[34]

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.[35] 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.[36] 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.[37] In prides including a coalition of two or three males, it is possible that lionesses solicit mating more actively with heavily maned lions.[36]
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.[37] 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.[17][38] However the males of the Asiatic subspecies are characterized by sparser manes than average African lions.[39]

A maneless male lion, who also has little body hair - from Tsavo East National Park, Kenya
Maneless lions have been reported in Senegal and Tsavo East National Park in Kenya, and the original male white lion from Timbavati was also maneless. Castrated lions have minimal manes. The lack of a mane is found in inbred lion populations; inbreeding also results in poor fertility.[40]
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,[24] 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,[16] 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) have been occasionally encountered in and around the Kruger National Park and the adjacent Timbavati Private Game Reserve in eastern South Africa, but are more commonly found in captivity, where breeders deliberately select them. The unusual cream color of their coats is due to a recessive gene.[41] They have been reportedly bred in camps in South Africa for use as trophies for canned hunts.[42]
Confirmation of the actual existence of the White lion only came in the late twentieth century. For hundreds of years prior, the White lion had been a figment of legend circulating through South Africa, the white pelage of the animal said to represent the goodness in all creatures. Claimed sightings were first reported in the early 1900s, and continued, infrequently, for almost 50 years until, in 1975, a litter of white lion cubs were found at Timbavati Game Reserve.[43]

Biology and behavior
Lions spend much of their time resting and are inactive for about 20 hours per day.[44] 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.[45]

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),[46] although, only for short bursts,[47] 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.[48] 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.[49]
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.[2] Occasionally, they take relatively small species such as Thomson's gazelle or springbok. 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.[50]

Lioness closing in on a zebra, a typical prey
They normally feed on mammals no larger than 550 kg, which excludes most adult hippopotamuses, rhinoceroses, elephants, giraffes, and buffalos.[51] In some areas, they specialise in hunting atypical prey-species; this is the case at the Savuti river, where they prey on young elephants.[52] 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.[53] In the Kruger National Park, giraffes are regularly hunted.[54] Lions also attack domestic livestock; in India cattle contribute significantly to their diet.[39] 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.[55] A lion may gorge itself and eat up to 30 kg (66 lb) in one sitting;[56] 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.[57] An adult lioness requires an average of about 5 kg (11 lb) of meat per day, a male about 7 kg (15.4 lb).[58]

Lions hunting down an African buffalo

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.[59]
Males attached to prides do not usually participate, except when hunting large animals such as buffalo and giraffe. 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.[60]