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For other uses, see Lion (disambiguation).
The lion (Panthera leo) is a species in the family Felidae and a member of the genusPanthera. It is the second largest extant species after the tiger. It exhibits a pronounced sexual dimorphism; males are larger than females with a typical weight range of 150 to 250 kg (331 to 551 lb) for the former and 120 to 182 kg (265 to 401 lb) for the latter. In addition, male lions have a prominent mane, which is perhaps the most recognisable feature of the species. Both sexes have hairy tufts at the end of their tails.
In the Pleistocene, lions were the most widespread large land mammals and ranged throughout Eurasia, Africa and North America. Today, the lion occurs in fragmented populations in Sub-Saharan Africa and one in western India. It has been listed as Vulnerable on the IUCN Red List since 1996, as populations in African range countries declined by about 43% since the early 1990s. Lion populations are untenable outside designated protected areas. Although the cause of the decline is not fully understood, habitat loss and conflicts with humans are the greatest causes of concern. The Asiatic lion and the West African lion are listed as Endangered and Critically Endangered, respectively.
The lion typically inhabits grasslands and savannahs, but is absent in dense forests. It is usually more diurnal than other big cats, but when persecuted adapts to being active at night and at twilight. A lion pride consists of a few adult males, related females and cubs. Prides vary in size and composition from three to 20 adult lions, depending on habitat and prey availability. Females cooperate when hunting and prey mostly on large ungulates, including antelope, deer, buffalo, zebra and even giraffe.
The lion is one of the most widely recognised animal symbols in human culture. It has been extensively depicted in sculptures and 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 over the world since the late 18th century. Cultural depictions of lions are known from the Upper Paleolithic period, with carvings and paintings from the Lascaux and Chauvet Caves in France dated to 17,000 years ago, through virtually all ancient and medieval cultures where they once occurred.
The lion's name, similar in many Romance languages, is derived from the LatinLatin: leo, and the Ancient Greekλέων (leon). The Hebrew word לָבִיא (lavi) may also be related.
The lion's closest relatives are the other species of the genus Panthera: the tiger, snow leopard, jaguar, and leopard. Results of phylogenetic studies published in 2006 and 2009 indicated that the jaguar and the lion belong to one sister group, which diverged about 2.06 million years ago. Results of later studies published in 2010 and 2011 indicate that the leopard and the lion belong to the same sister group, which diverged 1.95–3.10 million years ago. The lion and the snow leopard diverged about 2.1 million years ago.
In 1758, Carl Linnaeus described the lion in his work Systema Naturae and gave it the scientific nameFelis leo. Between the mid 18th and mid 20th centuries, 26 lion specimens were described and proposed as subspecies, of which 11 were recognised as valid in 2005. They were distinguished on the basis of appearance, size and colour of mane. As these characteristics vary highly between individuals, most of these forms were probably not true subspecies, especially as they were often based upon museum material with "striking, but abnormal" morphological characteristics.
Based on morphology of 58 lion skulls in three European museums, the subspecies krugeri, nubica, persica, and senegalensis were assessed distinct; but bleyenberghi overlapped with senegalensis and krugeri. The Asiatic lion persica was the most distinctive, and the Cape lion had characteristics allying it more with P. l. persica than the other sub-Saharan lions. Until 2016, eight subspecies were accepted and considered valid.
Early phylogenetic research was focused on lions from eastern and southern parts of Africa, and already showed that they can possibly be divided in two main clades: one to the west of the Great Rift Valley and the other to the east. Lions in eastern Kenya are genetically much closer to lions in Southern Africa than to lions in the Aberdare National Park in western Kenya.
In a subsequent study, tissue and bone samples of 32 lion specimens in museums were used. Results indicated that lions form three phylogeographic groups, one each in North Africa and Asia, in Central Africa and in Southern Africa.
Samples of 53 lions, both wild and captive individuals, from 15 countries were used for phylogenetic analysis. Results showed little genetic diversity between lions from Asia, West and Central Africa, whereas lions from East Africa were genetically closer to lions from Southern Africa.
Results of another phylogeographic study indicate that southeastern Ethiopia, western Somalia and northern Kenya are genetic admixture regions between lions from Central Africa and Southern Africa, and that lions in the northern part of Central Africa are genetically closer to lions in North and West Africa, and those in the southern part of Central Africa closer to lions in Southern Africa.
The majority of lions kept in zoos are hybrids of different subspecies. Approximately 77% of the captive lions registered by the International Species Information System are of unknown origin. Nonetheless, they might carry genes that are extinct in the wild, and might be therefore important to maintain overall genetic variability of the lion. It is thought that those lions, imported to Europe before the middle of the 19th century, were mainly either Barbary lions from North Africa, or Cape lions from Southern Africa.
Between 2008 and 2016, IUCN Red List assessors for lions used only two subspecific names, P. l. leo for African lion populations and P. l. persica for the Asiatic lion population. In 2017, the Cat Classification Task Force of the Cat Specialist Group assigned the lion populations in Asia and West, Central and North Africa to P. l. leo, and those in Southern and East Africa to P. l. melanochaita.
The following table is based on the classification of the species Panthera leo provided in Mammal Species of the World. It also reflects the classification used by IUCN Red List assessors and the revision by the Cat Classification Task Force:
|North African lion (P. l. leo)(Linnaeus, 1758), syn.P. l. nubica (de Blainville, 1843), P. l. somaliensis (Noack, 1891)||This is the nominate lion subspecies. In North Africa, lions are regionally extinct in the wild due to excessive hunting; the last known Barbary lion was killed in Morocco in 1942. Small groups of lions may have survived until the 1960s.|
A few captive lions are likely from North Africa, particularly the 90 individuals descended from the Moroccan Royal collection at Rabat Zoo. It is genetically more closely related to the Asiatic lion than to lions in East and Southern Africa.
In Algeria, Egypt, Libya, Morocco and Tunisia, lions are regionally extinct.
|Asiatic lion (P. l. leo)formerly (P. l. persica) (Meyer, 1826)||Today, the Asiatic lion population survives only in India's state of Gujarat and is listed as Endangered. Until the late 19th century, its historical range included eastern Turkey, Iran, the former Sind Province to Central India.|
The Indian population recovered from the brink of extinction to 411 individuals in 2010. It is protected in the Gir National Park and four protected areas in the region.
Results of phylogeographic studies suggest that its ancestors split from lions in Sub-Saharan Africa between 203 and 74 thousand years ago. Its closest relatives are North African and West African lions.
|West African lion (P. l. leo)formerly (P. l. senegalensis) (Meyer, 1826), syn. P. l. kamptzi (Matschie, 1900)||The type specimen originated in Senegal.|
This population has been listed as Critically Endangered in 2015 and survives in West Africa from Senegal, Burkina Faso, Benin to Niger and Nigeria. It is possibly extinct in Mauritania, Mali, Ghana, Guinea, Côte d'Ivoire, Gambia, Guinea-Bissau, Sierra Leone and Togo.
|Central African lion (P. l. leo)formerly P. l. azandica (Allen, 1924)||The type specimen was a male lion from northeastern Belgian Congo.|
The population occurs in the northeastern parts of the Democratic Republic of the Congo and Uganda. It is locally extinct in Rwanda.
|Cape lion (P. l. melanochaita)(Smith, 1842)||The type specimen originated at the Cape of Good Hope. The population lived in the Cape Province and Natal, South Africa.|
|East African lion (P. l. melanochaita), (P. l. leo)formerly P. l. massaica (Neumann, 1900), syn. P. l. sabakiensis (Lönnberg, 1910), P. l. roosevelti (Heller, 1914); P. l. nyanzae (Heller, 1914); P. l. hollisteri (Allen), 1924)P. l. webbiensis (Zukowsky, 1964)||Several type specimen were described from East African range countries.|
In East Africa, lion populations occur in Ethiopia, Kenya, Somalia, South Sudan, Tanzania and Uganda, but are regionally extinct in Djibouti, Egypt and Eritrea.
|Southern African lion (P. l. melanochaita) (P. l. leo)formerly P. l. bleyenberghi (Lönnberg, 1914), P. l. krugeri (Roberts, 1929), syn. P. l. vernayi (Roberts, 1948)||Several type specimen were described from Southern African range countries. Recorded body weights of individuals indicate that it is the heaviest of wild African lions.|
In Southern Africa lions occur in Namibia, Angola and northern Botswana. In the southwestern part of the Democratic Republic of the Congo, they are considered regionally extinct.
Additional lion subspecies, or sister species to the modern lion, existed in prehistoric times:
- Bone fragments classified under Panthera leo fossilis were excavated in Germany, United Kingdom, Italy and Czech Republic, and are estimated at between 680,000 and 600,000 years old. It was larger than today's African lions, reaching sizes comparable to the American cave lion and slightly larger than the Upper Pleistocene European cave lion.
- Bone fragments classified under Panthera leo spelaea were excavated in Spain, France, Italy, Belgium, Germany, Poland, Czech Republic, and Hungary, and are estimated at between 109,000 and 14,000 years old. This lion-like cat occurred in Eurasia about 300,000 to 10,000 years ago, and is known as the European cave lion, Eurasian cave lion, or Upper Pleistocene European cave lion. It is depicted in Paleolithiccave paintings, ivory carvings, and clay busts, which show it with protruding ears, tufted tails, and faint tiger-like stripes. A few had a ruff or primitive mane around their necks, possibly indicating males, but many scenes show hunting behavior.
- P. l. atrox or P. atrox, known as the American lion or American cave lion, existed in the Americas from Canada to Peru in the Pleistocene Epoch until about 10,000 years ago. This form is a sister clade of P. l. spelaea, and likely arose when an early P. l. spelaea population became isolated south of the North American continental ice sheet about 0.34 Mya. It is among the largest purported lion subspecies to have existed, its body length is estimated to have been 1.6–2.5 m (5.2–8.2 ft).
- P. l. youngi or Panthera youngi, flourished 350,000 years ago. Its relationship to the extant lion subspecies is obscure, and it probably represents a distinct species.
- P. l. vereshchagini was proposed as subspecies of the spelaea cave lion on the basis of skulls and teeth found in Yakutia, Russia, which were smaller in size than average spelaea bone fragments. Analysis of mitochondrial DNA sequences obtained from cave lion fossils provided no evidence for a distinct subspecific status of these samples.
- P. l. mesopotamica was described on the basis of a relief from the Neo-Assyrian Period, about 1000–600 BC, in ancient Mesopotamia.
- P. l. europaea was proposed for subfossil remains of lions excavated in Southern Europe that date to the Late Neolith to the Early Iron Age.
- P. l. maculatus, known as the Marozi or spotted lion, sometimes is thought 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.
Further information: Panthera hybrid, Liger, and Tigon
Lions have been known to breed with tigers (most often the Siberian and Bengal tigers) to create hybrids called 'ligers' and 'tiglons' (or 'tigons'). They also have been crossed with leopards to produce leopons. Such hybrid breeding 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. Because the growth-inhibiting gene from the female tiger mother is absent, the growth-promoting gene passed on by the male lion father is unimpeded by a regulating gene and 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 often are fertile. Males have about a 50% chance of having a mane, but if they grow them, their manes will be modest: around 50% the size of a pure lion mane. Ligers are much bigger than normal lions and tigers, typically 3.65 m (12.0 ft) in length, and can weigh up to 500 kg (1,100 lb).
The less common tiglon or tigon is a cross between a lioness and a male tiger. In contrast to ligers, tigons are often relatively small in comparison to their parents, because of reciprocal gene effects.
The lion evolved in Africa between 1 million and 800,000 years ago, from where it spread throughout the Holarctic region. The earliest fossil record in Europe was found near Pakefield in the United Kingdom and is about 680,000 years old. From this lion the late Pleistocene Eurasian cave lion probably derived about 300,000 years ago. Fossil remains found in the Cromer Forest Bed suggest that it was of a gigantic size and represented a lineage that was genetically isolated and highly distinct from lions in Africa and Asia. It was distributed throughout Europe, across Siberia and into western Alaska, via the Beringian landmass. The gradual formation of dense forest likely caused the decline of its geographic range near the end of the Late Pleistocene. Frequently encountered lion bones in cave deposits from Eemian times suggest that the cave lion survived in the Balkans and Asia Minor. There was probably a continuous population extending into India. Fossil lion remains were found in Pleistocene deposits in West Bengal. It became extinct about 10,000 years ago at the end of the last glacial period without mitochondrial descendants on other continents.
A fossil carnassial found in the Batadomba Cave indicates that Panthera leo sinhaleyus inhabited Sri Lanka during the late Pleistocene, and is thought to have become extinct around 39,000 years ago. This subspecies was described by Deraniyagala in 1939. It is distinct from the contemporary lion.
The modern lion probably originated in East and Southern Africa about 100,000 years ago. During the last glacial maximum until about 20,000 years ago, it was likely distributed throughout most of Southern and Central Africa, and expanded its range northwards during the early Holocene about 10,000 to 4,000 years ago.
|Male and female lion|
The lion is a muscular, deep-chested cat with a short, rounded head, a reduced neck and round ears. The colour of its fur varies from light buff to silverly gray, to yellowish red and dark brown. The underparts are generally lighter, and cubs are born with dark spots on their bodies. The spots fade as lions reach adulthood, although faint spots often may still be seen on the legs and underparts. The lion is the only member of the cat family that displays obvious sexual dimorphism. Males are more robust than females, have broader heads and a prominent mane, which grows downward and backward and covers most of the head, neck, shoulders, and chest. The mane is typically brownish and tinged with yellow, rust, and black hairs. The most distinctive characteristic shared by both females and males is that the tail ends in a dark, 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 cat with a tufted tail, but the function of the tuft and spine are unknown. Absent at birth, the tuft develops around 5 1⁄2 months of age and is readily identifiable at the age of seven months.
Of the living, non-hybrid felids, the lion is second only to the tiger 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 and broader nasal openings than that of a tiger. 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.
The lion and tiger are the tallest cat species in shoulder height. The typical weight range of lions is indicated as 150 to 250 kg (331 to 551 lb) for males and 120 to 182 kg (265 to 401 lb) for females. The size of adult lions varies across the global range, with those from the Southern African populations in Zimbabwe, the Kalahari and Kruger Park averaging from 187.5 to 193.3 kg (413 to 426 lb) for males and from 124.2 to 139.8 kg (274 to 308 lb) for females, compared to 174.9 kg (386 lb) and 119.5 kg (263 lb) for male and female lions respectively in East Africa. Measurements of head-body lengths in males range from 170 to 298 cm (5 ft 7 in to 9 ft 9 in), with tail lengths of 90–105 cm (2 ft 11 in–3 ft 5 in). Head-body lengths of females range from 140 to 175 cm (4 ft 7 in to 5 ft 9 in), with tail lengths of 70–100 cm (2 ft 4 in–3 ft 3 in). The frequently cited maximum head and body length of 250 cm (8 ft 2 in) fits rather to extinct Pleistocene forms, like the American lion, with even large modern lions measuring several centimeters less in length.
Record measurements from hunting records are supposedly a total length of nearly 3.6 m (12 ft) for a male lion shot near Mucusso National Park in southern Angola in October 1973. A male shot in 1936 outside Hectorspruit in eastern Transvaal Province, South Africa, weighed 313 kg (690 lb). Another notably large male lion was shot near Mount Kenya and weighed 272 kg (600 lb). There are also reports of large Asiatic lions.
The lion's mane is the most recognisable feature of the species. The mane starts growing when lions are about one year old. Mane colour varies, and darkens with age. Research results indicate that environmental factors such as average ambient temperature influence the mane's colour and size. Mane length apparently 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. 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. In the Serengeti National Park, female lions favour males as mates with dense, dark manes. The main purpose of the mane is thought to protect the lion's neck and throat in territorial fights with rivals.
Scientists once thought that distinct 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, which had the thickest, most extensive manes amongst wild lions. The cooler ambient temperature in European and North American zoos may result in a heavier mane. Thus the mane is not an appropriate marker for identifying subspecies. The males of the Asiatic subspecies, however, are characterised by sparser manes than average African lions.
In the area of Pendjari National Park, almost all West African males are maneless or have very weak manes. Maneless male African lions have also been reported from Senegal, from Sudan's Dinder National Park, and from Tsavo East National Park in Kenya. The original male white lion from Timbavati was also 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. Increased testosterone may be the cause of maned lionesses reported from northern Botswana.
Cave paintings of extinct European cave lions almost exclusively show hunting animals with no manes. Some suggest this as evidence that the males of this species were maneless, however, since the hunting usually involved groups of lionesses, this presumption remains unproven. In the Chauvet cave, there is a sketchy drawing of two maneless lions, appearing to be walking side by side. One is mostly obscured behind the other, with the former being larger than the latter, and shown with a scrotum.
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 dark skin. They are not albinos, but have normal pigmentation in the eyes and skin. White lion 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 caused by a recessive allele. Reportedly, they have been bred in camps in South Africa for use as trophies to be killed during canned hunts.
A melanistic Asiatic lion has been described from Khuzestan, Iran, by Austen Henry Layard. According to Layard it was dark brown in color, with nearly black patches.
Behaviour and ecology
Lions spend much of their time resting, and are inactive for about 20 hours per day. Although lions can be active at any time, their activity generally peaks after dusk with a period of socialising, 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.
Of all wild cat species, the lion is the most social cat, living in groups of related individuals with their offspring. Such a group is called a pride. Male lion groups are called a coalition. Females form the stable social unit in a pride and do not tolerate outside females. Membership only changes with the births and deaths of lionesses, although some females leave and become nomadic. The average pride consists of around 15 lions, including several adult females and up to four males and their cubs of both sexes. Large prides, consisting of up to 30 individuals, have also been observed. The sole exception to this pattern is the Tsavo lion pride which always has just one adult male. Male cubs are excluded from their maternal pride when they reach maturity at around 2–3 years of age.
Another lion behaviour is labeled nomads: lions 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. A lion may switch lifestyles; nomads can become residents and vice versa. Interactions between prides and nomads tend to be hostile, although pride females in estrous allow nomad males to approach them. Males spend years in a nomadic phase before gaining residence in a pride. A study in the Serengeti National Park revealed that nomadic coalitions gain residency at between 3.5 and 7.3 years of age.
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 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 calorific 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 and hone their skills. 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.
Both males and females defend the pride against intruders, but the male lion is better-suited for this purpose due to its stockier, more powerful build. Some individuals consistently lead the defence against intruders, while others lag behind. Lions tend to assume specific roles in the pride. Those lagging behind may provide other valuable services to the group. An alternative hypothesis is that there is some reward associated with being a leader who fends off intruders, and the rank of lionesses in the pride is reflected in these responses. The male or males associated with the pride must defend their relationship to the pride from outside males who attempt to take over their relationship with the pride.
Asiatic lion prides differ from African prides in group composition. Male Asiatic lions are solitary or associate with up to three males forming a loose pride. Pairs of males rest and feed together, and display marking behaviour at the same sites. Females associate with up to 12 females forming a stronger pride together with their cubs. They share large carcasses among each other, but seldom with males. Female and male lions associate only when mating. Coalitions of males hold a territory for a longer time than single lions. Males in coalitions of three to four individuals exhibit a pronounced hierarchy with one male dominating the others. Dominant males mate more frequently than their coalition partners. During a study carried out between December 2012 and December 2016, three females were observed switching mating partners in favour of the dominant male.
Hunting and diet
A lion's teeth are typical of a carnivore