Lion
Male Female (Lioness)
Conservation status
Vulnerable (IUCN) [1]
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. Reaching 272 kg (600 lb), it is the second-largest cat after the tiger. They currently exist in the wild 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 man. 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 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 lion pride consists of related females and offspring and a small number of dominant males. Groups of female lions typically hunt together, preying mostly on large ungulates. The lion is an apex and keystone predator, though 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;^[1] 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 18th century. Zoos are cooperating worldwide in breeding programs for the endangered Asiatic subspecies.

The male lion is highly distinctive and is easily recognized by its mane. The lion, particularly the face of the male, is one of the most widely recognized animal symbols in human culture. Depictions have existed from the Upper Paleolithic period, with carvings and paintings from the Lascaux and Chauvet Caves, through virtually all ancient and medieval cultures where they historically occurred. It has been extensively depicted in literature, in sculptures, in paintings, on national flags, and in contemporary films and literature.

Naming and etymology

The lion's name, similar in many languages, derives from the Latin leo,^[2] and before that the Ancient Greek leōn/λεων.^[3] The Hebrew word lavi (לָבִיא) may also be related,^[4] as well as the Ancient Egyptian rw.^[5] It was one of the many species originally described, as Felis leo, by Linnaeus in his 18th century work, Systema Naturae.^[6] 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".^[7]

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.^[8] 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.^[9]

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

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.^[14] 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 original who may have had "striking, but abnormal" morphological characteristics.^[15] Today only eight subspecies are usually accepted,^[16][12] but one of these, the Cape lion formerly described as Panthera leo melanochaita is probably invalid.^[16] 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.^[17][18]

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.^[19]
• 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.^[20]
• 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.^[16]

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).^[12] 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).^[21]
• 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.^[12] It is known from Paleolithic cave paintings, ivory carvings, and clay busts,^[22] 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 necks.^[23]
• 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.^[24][12]

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.^[25]
• 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.^[26] 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.^[27]

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.^[28] 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.

Average listed weights for the lions are between 150–225 kg (330–500 lb) for males, and 120–150 kg (260–330 lb) for females.^[29] 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).^[20] 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).^[30] 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.^[31]

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.^[32] 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.^[33] 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.^[34] In prides led by a coalition of two or three males, it is possible that lionesses solicit mating more actively with heavily maned lions.^[33]

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.^[34] 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.^[16][35] However the males of the Asiatic subspecies are characterized by sparser manes than average African lions.^[36]

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

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.^[37] Cave paintings of European cave lions show exclusively animals with no mane or just the hint of a mane, suggesting they were more or less maneless.^[23]

White lions

The white lion is not a distinct subspecies, but a special morph with a genetic condition, leucism,^[15] 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.^[38] They have been reportedly bred in camps in South Africa for use as trophies for canned hunts.^[39]

Confirmation of the actual existence of the White lion only came in the late 20th 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.^[40]

Hybrids

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

Further information: Panthera hybrid, liger and tigon

Lions have also been known to breed with tigers (most often the Amur and Bengal subspecies) to create hybrids called ligers and tigons.^[41] They have also been crossed with leopards to produce leopons,^[42] 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.^[43] 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 10 to 12 feet in length, and can be between 800 and 1,000 pounds or more.^[43] The less common tigon is a cross between the lioness and the male tiger.^[44]

Biology and behaviour

Lions spend much of their time resting and are inactive for about 20 hours per day.^[45] 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.^[46]

Hunting and diet

While a lion's teeth are sharp, prey is usually killed by strangulation.

Lions are powerful animals that usually hunt in groups and stalk their chosen prey. They can reach speeds of 59 km/h (40 mph),^[47] though only for short bursts,