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The lion (Panthera leo) is one of the four big cats in the genus Panthera, and a member of the family Felidae. With some males exceeding 250 kg (550 lb) in weight,[4] it is the second-largest living cat after the tiger. Wild lions currently exist in Sub-Saharan Africa and in Asia with an endangered remnant population in Gir Forest National Park in India, having disappeared from North Africa and Southwest Asia in historic times. Until the late Pleistocene, about 10,000 years ago, the lion was the most widespread large land mammal after humans. They were found in most of Africa, across Eurasia from western Europe to India, and in the Americas from the Yukon to Peru.[5] The lion is a vulnerable species, having seen a possibly irreversible population decline of thirty to fifty percent over the past two decades in its African range.[2] Lion populations are untenable outside designated reserves and national parks. Although the cause of the decline is not fully understood, habitat loss and conflicts with humans are currently the greatest causes of concern. Within Africa, the West African lion population is particularly endangered.
Lions live for ten to fourteen years in the wild, while in captivity they can live longer than twenty years. In the wild, males seldom live longer than ten years, as injuries sustained from continual fighting with rival males greatly reduce their longevity.[6] 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. Lions are apex and keystone predators, although they scavenge as opportunity allows. While lions do not typically hunt humans, some have been known to do so.
Highly distinctive, the male lion is easily recognised by its mane, and its face is one of the most widely recognised animal symbols in human culture. 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 once occurred. It has been extensively depicted in sculptures, in paintings, on national flags, and in contemporary films and literature. Lions have been kept in menageries since the time of the Roman Empire and have been a key species sought for exhibition in zoos the world over since the late eighteenth century. Zoos are cooperating worldwide in breeding programs for the endangered Asiatic subspecies.
Etymology
The lion's name, similar in many Romance languages, is derived from the Latin leo;[7] and the Ancient Greek λέων (leon).[8] The Hebrew word לָבִיא (lavi) may also be related.[9] It was one of the many species originally described by Linnaeus, who gave it the name Felis leo, in his eighteenth century work, Systema Naturae.[3]
Taxonomy and evolution
The lion is a species of the genus Panthera and its closest relatives are the other species of this genus: the tiger, the jaguar, and the leopard. Panthera leo itself evolved in Africa between 1 million and 800,000 years ago, before spreading throughout the Holarctic region.[10] It appeared in the fossil record in Europe for the first time 700,000 years ago with the subspecies Panthera leo fossilis at Isernia in Italy. From this lion derived the later cave lion (Panthera leo spelaea), which appeared about 300,000 years ago.[11] Lions died out in northern Eurasia at the end of the last glaciation, about 10,000 years ago;[12] this may have been secondary to the extinction of Pleistocene megafauna.[13]
Subspecies
Traditionally, twelve recent subspecies of lion were recognised, distinguished by mane appearance, size, and distribution. Because these characteristics are very insignificant and show a high individual variability, most of these forms were probably not true subspecies, especially as they were often based upon zoo material of unknown origin that may have had "striking, but abnormal" morphological characteristics.[14] Today only eight subspecies are usually accepted,[12][15] although one of these, the Cape lion, formerly described as Panthera leo melanochaita, probably is invalid.[15] Even the remaining seven subspecies might be too many. While the status of the Asiatic lion (P. l. persica) as a subspecies is generally accepted, the systematic relationships among African lions are still not completely resolved. Mitochondrial variation in living African lions seemed to be modest according to some younger studies and therefore all sub-Saharan lions sometimes have been considered a single subspecies. However, a recent study revealed, that lions from western and central Africa differ genetically from lions of southern or eastern Africa. According to this study, Western African lions are more closely related to Asian lions, than to South or East African lions. These findings might be explained by a late Pleistocene extinction event of lions in western and central Africa and a subsequent recolonisation of these parts from Asia.[16] Previous studies, which were focusing mainly on lions from eastern and southern parts of Africa already showed that these can be 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] Another study, revealed, that there are three major types of lions, one North African–Asian, one southern African and one middle African.[18] Conversely, Per Christiansen found that using skull morphology allowed him to identify the subspecies krugeri, nubica, persica, and senegalensis, while there was overlap between bleyenberghi with senegalensis and krugeri. The Asiatic lion persica was the most distinctive, and the Cape lion had characteristics allying it more with persica than the other subsaharan lions. He had analysed 58 lion skulls in three European museums.[19]
Recent
Eight recent (Holocene) subspecies are recognised today:
- P. l. persica, known as the Asiatic lion or South Asian, Persian, or Indian Lion, once was widespread from Turkey, across Southwest Asia, to Pakistan, India, and even to Bangladesh. However, large prides and daylight activity made them easier to poach than tigers or leopards; now around 300 exist in and near the Gir Forest of India.[20] Genetic evidence suggests its ancestors split from the ancestors of subsaharan African lions between 74 and 203 thousand years ago.[12]
- P. l. leo, known as the Barbary lion, originally ranged from Morocco to Egypt. It is extinct in the wild due to excessive hunting, as the last wild Barbary lion was killed in Morocco in 1922.[21] This was one of the largest of the lion subspecies,[22] with reported lengths of 3–3.3 metres (10–10.8 ft) and weights of more than 200 kilograms (440 lb) for males. It appears to be more closely related to the Asiatic rather than subsaharan lions. There are a number of animals in captivity likely to be Barbary lions,[23] particularly 90 animals descended from the Moroccan Royal collection at Rabat Zoo.[24]
- P. l. senegalensis, known as the West African Lion, is found in western Africa, from Senegal to the Central African Republic.[25][26]
- P. l. azandica, known as the Northeast Congo Lion, is found in the northeastern parts of the Congo.[25]
- P. l. nubica, known as the East African, Massai Lion is found in east Africa, from Ethiopia and Kenya to Tanzania and Mozambique.[26], a local population is known as Tsavo Lion.
- P. l. bleyenberghi, known as the Southwest African or Katanga Lion, is found in southwestern Africa, Namibia, Botswana, Angola, Katanga (Zaire), Zambia, and Zimbabwe.[26]
- P. l. krugeri, known as the Southeast African Lion or Transvaal Lion, is found in the Transvaal region of southeastern Africa, including Kruger National Park.[26]
- 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 P. l. krugeri.[15]
Pleistocene
Several additional subspecies of lion existed in prehistoric times:
- 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. It was larger than today's African lions, reaching the American cave lion in size[12][27]
- 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] This species is known from Paleolithic cave paintings (such as the one displayed to the right), ivory carvings, and clay busts,[28] 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.[29]
Dubious
- P. l. youngi or Panthera youngi, flourished 350,000 years ago.[5] Its relationship to the extant lion subspecies is obscure, and it probably represents a distinct species.
- P. l. maculatus, known as the Marozi or Spotted lion, sometimes is believed to be a distinct subspecies, but may be an adult lion that has retained its juvenile spotted pattern. If it was a subspecies in its own right, rather than a small number of aberrantly coloured individuals, it has been extinct since 1931. A less likely identity is a natural leopard-lion hybrid commonly known as a leopon.[31]
- P. l. atrox, known as the American Lion or American cave lion, was abundant in the Americas from Canada to Peru in the Pleistocene Epoch until about 10,000 years ago. This form likely represents a separate species, possibly more closely related to the jaguar, and can not be considered a true lion.[32] One of the largest purported lion subspecies to have existed, its body length is estimated to have been 1.6–2.5 m (5–8 ft).[33]
Hybrids
Video of lion cubs in the wild, South Africa
Lions have been known to breed with tigers (most often the Siberian and Bengal subspecies) to create hybrids called ligers and tiglons.[34] They also have been crossed with leopards to produce leopons,[35] 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.[36] Because the growth-inhibiting gene from the female tiger is absent, a growth-promoting gene is passed on by the male lion, the resulting ligers grow far larger than either parent. They share physical and behavioural qualities of both parent species (spots and stripes on a sandy background). Male ligers are sterile, but female ligers are often fertile. Males have about a 50 percent chance of having a mane, but if they grow one, their manes will be modest: around 50 percent 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.[36] The less common tigon is a cross between the lioness and the male tiger.[37]
Characteristics
The lion is the tallest (at the shoulder) of all living cats, averaging about 14 cm (5.5 in) taller than the tiger. Behind only the tiger, the lion is the second largest living felid in length and weight. Its skull is very similar to that of the tiger, although the frontal region is usually more depressed and flattened, with a slightly shorter postorbital region. The lion's skull has broader nasal openings than the tiger. However, due to the amount of skull variation in the two species, usually, only the structure of the lower jaw can be used as a reliable indicator of species.[38] Lion coloration varies from light buff to yellowish, reddish, or dark ochraceous brown. The underparts are generally lighter and the tail tuft is black. Lion cubs are born with brown rosettes (spots) on their body, rather like those of a leopard. Although these fade as lions reach adulthood, faint spots often may still be seen on the legs and underparts, particularly on lionesses.
Lions are the only members of the cat family to display obvious sexual dimorphism—that is, males and females look distinctly different. They also have specialised roles that each gender plays in the pride. For instance, the lioness, the hunter, lacks the male's thick cumbersome mane. It seems to impede the male's ability to be camouflaged when stalking the prey and create overheating in chases. The colour of the male's mane varies from blond to black, generally becoming darker as the lion grows older.
During confrontations with others, the mane makes the lion look larger.
Weights for adult lions range between 150–250 kg (330–550 lb) for males and 120–182 kg (264–400 lb) for females.[4] 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] Lions tend to vary in size depending on their environment and area, resulting in a wide spread in recorded weights. For instance, lions in southern Africa tend to be about 5 percent heavier than those in East Africa, in general.[39]
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 up to 123 cm (4 ft) in males and as low as 91 cm (3 ft) in females.[40] The tail length is 90–105 cm (2 ft 11 in - 3 ft 5 in) in males and 70–100 cm in females (2 ft 4 in – 3 ft 3 in).[4] The longest known lion, at nearly 3.6 m (12 ft) in total length, was a black-maned male shot near Mucsso, southern Angola in October 1973; the heaviest lion known in the wild was a man-eater shot in 1936 just outside Hectorspruit in eastern Transvaal, South Africa and weighed 313 kg (690 lb).[41] Lions in captivity tend to be larger than lions in the wild—the heaviest lion on record is a male at Colchester Zoo in England named Simba in 1970, which weighed 375 kg (826 lb).[42]
The most distinctive characteristic shared by both females and males is that the tail ends in a hairy tuft. In some lions, the tuft conceals a hard "spine" or "spur", approximately 5 mm long, formed of the final sections of tail bone fused together. The lion is the only felid to have a tufted tail—the function of the tuft and spine are unknown. Absent at birth, the tuft develops around 5½ months of age and is readily identifiable at 7 months.[43]
Mane
The mane of the adult male lion, unique among 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.[44] The presence, absence, colour, and size of the mane is associated with genetic precondition, sexual maturity, climate, and testosterone production; the rule of thumb is the darker and fuller the mane, the healthier the lion. Sexual selection of mates by lionesses favors males with the densest, darkest mane.[45] 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.[46] In prides including a coalition of two or three males, it is possible that lionesses solicit mating more actively with the males who are more heavily maned.[45]
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 colour and size of a lion's mane, such as the ambient temperature.[46] The cooler ambient temperature in European and North American zoos, for example, may result in a heavier mane. Thus the mane is not an appropriate marker for identifying subspecies.[15][47] The males of the Asiatic subspecies, however, are characterised by sparser manes than average African lions.[48]
In the Pendjari National Park area amost all males are maneless or have very weak manes.[49] Maneless male lions have also been reported from Senegal and from Tsavo East National Park in Kenya, and the original male white lion from Timbavati also was maneless. The testosterone hormone has been linked to mane growth, therefore castrated lions often have minimal to no mane, as the removal of the gonads inhibits testosterone production.[50]
Cave paintings of extinct European cave lions exclusively show animals with no mane, or just the hint of a mane, suggesting that they were maneless.[29]
White lions
White lions owe their colouring 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,[14] that causes paler colouration akin to that of the white tiger; the condition is similar to melanism, which causes black panthers. They are not albinos, having normal pigmentation in the eyes and skin. White Transvaal lion (Panthera leo krugeri) individuals occasionally have been encountered in and around Kruger National Park and the adjacent Timbavati Private Game Reserve in eastern South Africa, but are more commonly found in captivity, where breeders deliberately select them. The unusual cream colour of their coats is due to a recessive gene.[51] Reportedly, they have been bred in camps in South Africa for use as trophies to be killed during canned hunts.[52]
Kevin Richardson is an animal behaviourist who works with the native big cats of Africa. He currently works in a special facility called the Kingdom of the White Lion in Broederstroom[53] which is 50 miles form Johannesburg.[54] The site was built with the help of Rodney Fuhr[55] and was made for the movie set of White Lion: Home is a Journey.[54] He has 39 white lions on-site[53] and works diligently to protect and preserve the white lion type. While the park is currently a private property, there are plans to open it to the public soon.[56]
Behaviour
Lions spend much of their time resting and are inactive for about 20 hours per day.[57] 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 until dawn, when hunting most often takes place. They spend an average of two hours a day walking and 50 minutes eating.[58]
Group organization
Lions are predatory carnivores who manifest two types of social organization. Some are residents, living in groups, called prides.[59] The pride usually consists of five or six related females, their cubs of both sexes, and one or two males (known as a coalition if more than one) who mate with the adult females (although extremely large prides, consisting of up to 30 individuals, have been observed). The number of adult males in a coalition is usually two, but may increase to four and decrease again over time. Male cubs are excluded from their maternal pride when they reach maturity.
Two lionesses and a mature male of a pride, northern Serengeti
The second organizational behaviour is labeled nomads, who range widely and move about sporadically, either singularly or in pairs.[59] 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 they 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.[59] The males associated with a pride tend to stay on the fringes, 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 members reduce per capita caloric intake, however, some take a role raising cubs, who may be left alone for extended periods of time. Members of the pride regularly tend to play the same role in hunts. The health of the hunters is the primary need for the survival of the pride and they are the first to consume the prey at the site it is taken. Other benefits include possible kin selection (better to share food with a related lion than with a stranger), protection of the young, maintenance of territory, and individual insurance against injury and hunger.[21]