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Lion

February 16, 2024

For other uses, see Lion (disambiguation).

The lion (Panthera leo) is a large cat of the genus Panthera, native to Africa and India. It has a muscular, broad-chested body; a short, rounded head; round ears; and a hairy tuft at the end of its tail. It is sexually dimorphic; adult male lions are larger than females and have a prominent mane. It is a social species, forming groups called prides. A lion's pride consists of a few adult males, related females, and cubs. Groups of female lions usually hunt together, preying mostly on large ungulates. The lion is an apex and keystone predator; although some lions scavenge when opportunities occur and have been known to hunt humans, lions typically do not actively seek out and prey on humans.

Lion
Temporal range: Pleistocene–Present
↓
Male in Okonjima, Namibia
Female (lioness) in Okonjima
CITES Appendix II (CITES)[a][2]
Scientific classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Species:
P. leo[1]
Binomial name
Panthera leo[1]
Subspecies
P. l. leo
P. l. melanochaita
†P. l. sinhaleyus
Historical and present distribution of the lion in Africa, Asia and Europe

The lion inhabits grasslands, savannahs, and shrublands. It is usually more diurnal than other wild cats, but when persecuted, it adapts to being active at night and at twilight. During the Neolithic period, the lion ranged throughout Africa and Eurasia, from Southeast Europe to India, but it has been reduced to fragmented populations in sub-Saharan Africa and one population in western India. It has been listed as Vulnerable on the IUCN Red List since 1996 because populations in African countries have 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 for concern.

One of the most widely recognised animal symbols in human culture, the lion 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 zoological gardens across the world since the late 18th century. Cultural depictions of lions were prominent in Ancient Egypt, and depictions have occurred in virtually all ancient and medieval cultures in the lion's historic and current range.

Etymology

The English word lion is derived via Anglo-Norman liun from Latin leōnem (nominative: leō), which in turn was a borrowing from Ancient Greek λέων léōn. The Hebrew word לָבִיא lavi may also be related.[4] The generic name Panthera is traceable to the classical Latin word 'panthēra' and the ancient Greek word πάνθηρ 'panther'.[5]

Taxonomy

 
The upper cladogram is based on the 2006 study,[6][7] the lower one on the 2010[8] and 2011[9] studies.

Felis leo was the scientific name used by Carl Linnaeus in 1758, who described the lion in his work Systema Naturae.[3] The genus name Panthera was coined by Lorenz Oken in 1816.[10] 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.[1] They were distinguished mostly by the size and colour of their manes and skins.[11]

Subspecies

 
Range map showing distribution of subspecies and clades

In the 19th and 20th centuries, several lion type specimens were described and proposed as subspecies, with about a dozen recognised as valid taxa until 2017.[1] Between 2008 and 2016, IUCN Red List assessors used only two subspecific names: P. l. leo for African lion populations, and P. l. persica for the Asiatic lion population.[2][12][13] In 2017, the Cat Classification Task Force of the Cat Specialist Group revised lion taxonomy, and recognises two subspecies based on results of several phylogeographic studies on lion evolution, namely:[14]

  • P. l. leo (Linnaeus, 1758) − the nominate lion subspecies includes the Asiatic lion, the regionally extinct Barbary lion, and lion populations in West and northern parts of Central Africa.[14] Synonyms include P. l. persica (Meyer, 1826), P. l. senegalensis (Meyer, 1826), P. l. kamptzi (Matschie, 1900), and P. l. azandica (Allen, 1924).[1] Multiple authors referred to it as 'northern lion' and 'northern subspecies'.[15][16]
  • P. l. melanochaita (Smith, 1842) − includes the extinct Cape lion and lion populations in East and Southern African regions.[14] Synonyms include P. l. somaliensis (Noack 1891), P. l. massaica (Neumann, 1900), P. l. sabakiensis (Lönnberg, 1910), P. l. bleyenberghi (Lönnberg, 1914), P. l. roosevelti (Heller, 1914), P. l. nyanzae (Heller, 1914), P. l. hollisteri (Allen, 1924), P. l. krugeri (Roberts, 1929), P. l. vernayi (Roberts, 1948), and P. l. webbiensis (Zukowsky, 1964).[1][11] It has been referred to as 'southern subspecies' and 'southern lion'.[16]

However, there seems to be some degree of overlap between both groups in northern Central Africa. DNA analysis from a more recent study indicates, that Central African lions are derived from both northern and southern lions, as they cluster with P. leo leo in mtDNA-based phylogenies whereas their genomic DNA indicates a closer relationship with P. leo melanochaita.[17]

Lion samples from some parts of the Ethiopian Highlands cluster genetically with those from Cameroon and Chad, while lions from other areas of Ethiopia cluster with samples from East Africa. Researchers, therefore, assume Ethiopia is a contact zone between the two subspecies.[18] Genome-wide data of a wild-born historical lion sample from Sudan showed that it clustered with P. l. leo in mtDNA-based phylogenies, but with a high affinity to P. l. melanochaita. This result suggested that the taxonomic position of lions in Central Africa may require revision.[19]

Fossil records

 
Skull of an American lion on display at the National Museum of Natural History

Other lion subspecies or sister species to the modern lion existed in prehistoric times:[20]

Evolution

 
red Panthera spelaea
blue Panthera atrox
green Panthera leo

Maximal range of the modern lion
and its prehistoric relatives
in the late Pleistocene

The Panthera lineage is estimated to have genetically diverged from the common ancestor of the Felidae around 9.32 to 4.47 million years ago to 11.75 to 0.97 million years ago.[6][33][34] Results of analyses differ in the phylogenetic relationship of the lion; it was thought to form a sister group with the jaguar (P. onca) that diverged 3.46 to 1.22 million years ago,[6] but also with the leopard (P. pardus) that diverged 3.1 to 1.95 million years ago[8][9] to 4.32 to 0.02 million years ago. Hybridisation between lion and snow leopard (P. uncia) ancestors possibly continued until about 2.1 million years ago.[34] The lion-leopard clade was distributed in the Asian and African Palearctic since at least the early Pliocene.[35] The earliest fossils recognisable as lions were found at Olduvai Gorge in Tanzania and are estimated to be up to 2 million years old.[33]

Estimates for the divergence time of the modern and cave lion lineages range from 529,000 to 392,000 years ago based on mutation rate per generation time of the modern lion. There is no evidence for gene flow between the two lineages, indicating that they did not share the same geographic area.[19] The Eurasian and American cave lions became extinct at the end of the last glacial period without mitochondrial descendants on other continents.[27][36][37] The modern lion was probably widely distributed in Africa during the Middle Pleistocene and started to diverge in sub-Saharan Africa during the Late Pleistocene. Lion populations in East and Southern Africa became separated from populations in West and North Africa when the equatorial rainforest expanded 183,500 to 81,800 years ago.[38] They shared a common ancestor probably between 98,000 and 52,000 years ago.[19] Due to the expansion of the Sahara between 83,100 and 26,600 years ago, lion populations in West and North Africa became separated. As the rainforest decreased and thus gave rise to more open habitats, lions moved from West to Central Africa. Lions from North Africa dispersed to southern Europe and Asia between 38,800 and 8,300 years ago.[38]

Extinction of lions in southern Europe, North Africa and the Middle East interrupted gene flow between lion populations in Asia and Africa. Genetic evidence revealed numerous mutations in lion samples from East and Southern Africa, which indicates that this group has a longer evolutionary history than genetically less diverse lion samples from Asia and West and Central Africa.[39] A whole genome-wide sequence of lion samples showed that samples from West Africa shared alleles with samples from Southern Africa, and samples from Central Africa shared alleles with samples from Asia. This phenomenon indicates that Central Africa was a melting pot of lion populations after they had become isolated, possibly migrating through corridors in the Nile Basin during the early Holocene.[19]

Hybrids

Further information: Panthera hybrid

In zoos, lions have been bred with tigers to create hybrids for the curiosity of visitors or for scientific purpose.[40][41] The liger is bigger than a lion and a tiger, whereas most tigons are relatively small compared to their parents because of reciprocal gene effects.[42][43] The leopon is a hybrid between a lion and leopard.[44]

Description

 
A tuft at the end of the tail is a distinct characteristic of the lion.
 
Skeleton

The lion is a muscular, broad-chested cat with a short, rounded head, a reduced neck, and round ears; males have broader heads. The fur varies in colour from light buff to silvery grey, yellowish red, and dark brown. The colours of the underparts are generally lighter. A new-born lion has dark spots, which fade as the cub reaches adulthood, although faint spots may still be seen on the legs and underparts.[45][46] The tail of all lions ends in a dark, hairy tuft that, in some lions, conceals an approximately 5 mm (0.20 in)-long, hard "spine" or "spur" that is formed from the final, fused sections of tail bone. The functions of the spur are unknown. The tuft is absent at birth and develops at around 5+1⁄2 months of age. It is readily identifiable at the age of seven months.[47]

Its skull is very similar to that of the tiger, although the frontal region is usually more depressed and flattened and has a slightly shorter postorbital region and broader nasal openings than those of the 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.[48][49]

The skeletal muscles of the lion make up 58.8% of its body weight and represent the highest percentage of muscles among mammals.[50][51]

Size

Among felids, the lion is second only to the tiger in size.[46] The size and weight of adult lions vary across its range and habitats.[52][53][54][55] Accounts of a few individuals that were larger than average exist from Africa and India.[45][56][57][58]

Average Female lions Male lions
Head-and-body length 160–184 cm (63–72 in)[59] 184–208 cm (72–82 in)[59]
Tail length 72–89.5 cm (28.3–35.2 in)[59] 82.5–93.5 cm (32.5–36.8 in)[59]
Weight 118.37–143.52 kg (261.0–316.4 lb) in Southern Africa,[52]
119.5 kg (263 lb) in East Africa,[52]
110–120 kg (240–260 lb) in India[53]		 186.55–225 kg (411.3–496.0 lb) in Southern Africa,[52]
174.9 kg (386 lb) in East Africa,[52]
160–190 kg (350–420 lb) in India[53]

Mane

 
A six-year-old male in Phinda Private Game Reserve
 
Male in Pendjari National Park

The male lion's mane is the most recognisable feature of the species.[11] It may have evolved around 320,000–190,000 years ago.[60] It grows downwards and backwards, covering most of the head, neck, shoulders, and chest. The mane is typically brownish and tinged with yellow, rust, and black hairs.[46] It starts growing when lions enter adolescence, when testosterone levels increase, and reach their full size at around four years old.[61] Cool ambient temperatures in European and North American zoos may result in a heavier mane.[62] On average, Asiatic lions have sparser manes than African lions.[63]

This feature likely evolved to signal the fitness of males to females. The mane of the lion is thought to serve mating purposes. Males with darker manes appear to have greater reproductive success and are more likely to remain in a pride for longer. They have longer and thicker hair and higher testosterone levels, but they are also more vulnerable to heat stress.[64][65] Unlike in other felid species, female lions consistently interact with multiple males at once.[66] Another hypothesis suggests that the mane also serves to protect the neck in fights, but this is disputed.[67][68] During fights, including those involving maneless females and adolescents, the neck is not targeted as much as the face, back, and hindquarters. Injured lions also begin to lose their manes.[69]

Almost all male lions in Pendjari National Park are either maneless or have very short manes.[70] Maneless lions have also been reported in Senegal, in Sudan's Dinder National Park and in Tsavo East National Park, Kenya.[71] Castrated lions often have little to no mane because the removal of the gonads inhibits testosterone production.[72]

Rarely, lionesses (in the wild and in captivity) have been observed to grow manes.[73][74] Increased testosterone may be the cause of maned lionesses reported in northern Botswana.[75]

Colour variation

Further information: White lion

The white lion is a rare morph with a genetic condition called leucism, which is caused by a double recessive allele. It is not albino; it has normal pigmentation in the eyes and skin. White lions have occasionally been encountered in and around Kruger National Park and the adjacent Timbavati Private Game Reserve in eastern South Africa. They were removed from the wild in the 1970s, thus decreasing the white lion gene pool. Nevertheless, 17 births have been recorded in five prides between 2007 and 2015.[76] White lions are selected for breeding in captivity.[77] They have reportedly been bred in camps in South Africa for use as trophies to be killed during canned hunts.[78]

Distribution and habitat

 
Lion in Gir National Park

African lions live in scattered populations across sub-Saharan Africa. The lion prefers grassy plains and savannahs, scrub bordering rivers, and open woodlands with bushes. It rarely enters closed forests. On Mount Elgon, the lion has been recorded up to an elevation of 3,600 m (11,800 ft) and close to the snow line on Mount Kenya.[45] Savannahs with an annual rainfall of 300 to 1,500 mm (12 to 59 in) make up the majority of lion habitat in Africa, estimated at 3,390,821 km2 (1,309,203 sq mi) at most, but remnant populations are also present in tropical moist forests in West Africa and montane forests in East Africa.[79] The Asiatic lion now survives only in and around Gir National Park in Gujarat, western India. Its habitat is a mixture of dry savannah forest and very dry, deciduous scrub forest.[12]

Historical range

In Africa, the range of the lion originally spanned most of the central African rainforest zone and the Sahara desert.[80] In the 1960s, it became extinct in North Africa, except in the southern part of Sudan.[81][79][82]

In southern Europe and Asia, the lion once ranged in regions where climatic conditions supported an abundance of prey.[83] In Greece, it was common, as reported by Herodotus in 480 BC; it was considered rare by 300 BC and extirpated by AD 100.[45] It was present in the Caucasus until the 10th century.[49] It lived in Palestine until the Middle Ages and in Southwest Asia until the late 19th century. By the late 19th century, it had been extirpated in most of Turkey.[84] The last live lion in Iran was sighted in 1942, about 65 km (40 mi) northwest of Dezful,[85] although the corpse of a lioness was found on the banks of the Karun river in Khuzestan province in 1944.[86] It once ranged from Sind and Punjab in Pakistan to Bengal and the Narmada River in central India.[87]

Behaviour and ecology

Lions spend much of their time resting; they are inactive for about twenty hours per day.[88] 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 continue until dawn, when hunting most often takes place. They spend an average of two hours a day walking and fifty minutes eating.[89]

Group organisation

 
Lion pride in Etosha National Park
 
A lioness (left) and two males in Masai Mara

The lion is the most social of all wild felid species, living in groups of related individuals with their offspring. Such a group is called a "pride". Groups of male lions are called "coalitions".[90] Females form the stable social unit in a pride and do not tolerate outside females.[91] The majority of females remain in their birth prides while all males and some females will disperse.[92] 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 been observed.[93] The sole exception to this pattern is the Tsavo lion pride that always has just one adult male.[94] Prides act as fission–fusion societies, and members will split into subgroups that keep in contact with roars.[95]

Nomadic lions range widely and move around sporadically, either in pairs or alone.[90] Pairs are more frequent among related males. A lion may switch lifestyles; nomads can become residents and vice versa.[96] Interactions between prides and nomads tend to be hostile, although pride females in estrus allow nomadic males to approach them.[97] Males spend years in a nomadic phase before gaining residence in a pride.[98] A study undertaken in the Serengeti National Park revealed that nomadic coalitions gain residency at between 3.5 and 7.3 years of age.[99] In Kruger National Park, dispersing male lions move more than 25 km (16 mi) away from their natal pride in search of their own territory. Female lions stay closer to their natal pride. Therefore, female lions in an area are more closely related to each other than male lions in the same area.[100]

The evolution of sociability in lions was likely driven both by high population density and the clumped resources of savannah habitats. The larger the pride, the more high-quality territory they can defend; "hotspots" being near river confluences, where the cats have better access to water, prey and shelter (via vegetation).[101][102] The area occupied by a pride is called a "pride area" whereas that occupied by a nomad is a "range".[90] Males associated with a pride patrol the fringes.[46] 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 defense against intruders, while others lag behind.[103] Lions tend to assume specific roles in the pride; slower-moving individuals may provide other valuable services to the group.[104] Alternatively, there may be rewards associated with being a leader that fends off intruders; the rank of lionesses in the pride is reflected in these responses.[105] The male or males associated with the pride must defend their relationship with the pride from outside males who may attempt to usurp them.[96] Dominance hierarchies do not appear to exist among individuals of either sex in a pride.[106]

Asiatic lion prides differ in group composition. Male Asiatic lions are solitary or associate with up to three males, forming a loose pride while females associate with up to 12 other females, forming a stronger pride together with their cubs. Female and male lions associate only when mating.[107] Coalitions of males hold territory for a longer time than single lions. Males in coalitions of three or four individuals exhibit a pronounced hierarchy, in which one male dominates the others and mates more frequently.[108]

Hunting and diet

 
Male lion and cub feeding on a Cape buffalo in Sabi Sand Game Reserve
 
Four lionesses catching a buffalo in the Serengeti
 
A skeletal mount of a lion attacking a common eland, on display at The Museum of Osteology

The lion is a generalist hypercarnivore and is considered to be both an apex and keystone predator due to its wide prey spectrum.[109][110] Its prey consists mainly of ungulates particularly blue wildebeest, plains zebra, African buffalo, common warthog, gemsbok and giraffe.[111] In India, chital and sambar deer are the most common wild prey,[46][111][112] while livestock contributes significantly to lion kills outside protected areas.[113] They usually avoid fully grown adult elephants, rhinoceroses and hippopotamus and small prey like dik-dik, hyraxes, hares and monkeys.[111][114] Unusual prey include porcupines and small reptiles. Lions kill other predators but seldom consume them.[115]

Young lions first display stalking behaviour at around three months of age, although they do not participate in hunting until they are almost a year old and begin to hunt effectively when nearing the age of two.[116] Single lions are capable of bringing down zebra and wildebeest, while larger prey like buffalo and giraffe are riskier.[96] In Chobe National Park, large prides have been observed hunting African bush elephants up to around 15 years old in exceptional cases, with the victims being calves, juveniles, and even subadults.[117][118] In typical hunts, each lioness has a favoured 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 fleeing from other lionesses. Males attached to prides do not usually participate in group hunting.[119] Some evidence suggests, however, that males are just as successful as females; they are typically solo hunters who ambush prey in small bushland.[120] They may join in the hunting of large, slower-moving prey like buffalo; and even hunt them on their own. Moderately-sized hunting groups generally have higher success rates than lone females and larger groups.[121]

Lions are not particularly known for their stamina. For instance, a lioness's heart comprises only 0.57% of her body weight and a male's is about 0.45% of his body weight, whereas a hyena's heart comprises almost 1% of its body weight.[122] Thus, lions run quickly only in short bursts at about 48–59 km/h (30–37 mph) and need to be close to their prey before starting the attack.[123] One study in 2018 recorded a lion running at a top speed of 74.1 km/h (46.0 mph).[124] They take advantage of factors that reduce visibility; many kills take place near some form of cover or at night.[125] The lion's attack is short and powerful; it attempts to catch prey with a fast rush and final leap, usually pulls it down by the rump, and kills with a clamping bite to the throat or muzzle. It can hold the prey's throat for up to 13 minutes, until the prey stops moving.[126]

Lions typically consume prey at the location of the hunt but sometimes drag large prey into cover.[127] They tend to squabble over kills, particularly the males. Cubs suffer most when food is scarce but otherwise all pride members eat their fill, including old and crippled lions, which can live on leftovers.[96] Large kills are shared more widely among pride members.[128] An adult lioness requires an average of about 5 kg (11 lb) of meat per day while males require about 7 kg (15 lb).[129] Lions gorge themselves and eat up to 30 kg (66 lb) in one session.[86] If it is unable to consume all of the kill, it rests for a few hours before continuing to eat. On hot days, the pride retreats to shade with one or two males standing guard.[127] Lions defend their kills from scavengers such as vultures and hyenas.[96]

Lions scavenge on carrion when the opportunity arises, scavenging animals dead from natural causes such as disease or those that were killed by other predators. Scavenging lions keep a constant lookout for circling vultures, which indicate the death or distress of an animal.[130] Most carrion on which both hyenas and lions feed upon are killed by hyenas rather than lions.[55] Carrion is thought to provide a large part of lion diet.[131]

Predatory competition

 
Lioness chasing a spotted hyena in Kruger National Park
 
Lioness stealing a kill from a leopard in Kruger National Park

Lions and spotted hyenas occupy a similar ecological niche and compete for prey and carrion; a review of data across several studies indicates a dietary overlap of 58.6%.[132] Lions typically ignore hyenas unless they are on a kill or are being harassed, while the latter tend to visibly react to the presence of lions with or without the presence of food. In the Ngorongoro crater, lions subsist largely on kills stolen from hyenas, causing them to increase their kill rate.[133] In Botswana's Chobe National Park, the situation is reversed as hyenas there frequently challenge lions and steal their kills, obtaining food from 63% of all lion kills.[134] When confronted on a kill, hyenas may either leave or wait patiently at a distance of 30–100 m (98–328 ft) until the lions have finished.[135] Hyenas may feed alongside lions and force them off a kill. The two species attack one another even when there is no food involved for no apparent reason.[136] Lions can account for up to 71% of hyena deaths in Etosha National Park. Hyenas have adapted by frequently mobbing lions that enter their home ranges.[137] When the lion population in Kenya's Masai Mara National Reserve declined, the spotted hyena population increased rapidly.[138]

Lions tend to dominate cheetahs and leopards, steal their kills and kill their cubs and even adults when given the chance.[139] Cheetahs often lose their kills to lions or other predators.[140] A study in the Serengeti ecosystem revealed that lions killed at least 17 of 125 cheetah cubs born between 1987 and 1990.[141] Cheetahs avoid their competitors by hunting at different times and habitats.[142] Leopards take refuge in trees, but lionesses occasionally attempt to climb up and retrieve their kills.[143]

Lions similarly dominate African wild dogs, taking their kills and slaying pups or adult dogs. Population densities of wild dogs are low in areas where lions are more abundant.[144] However, there are a few reported cases of old and wounded lions falling prey to wild dogs.[145][146]

Reproduction and life cycle

 
Lions mating at Masai Mara
 
A lion cub in Masai Mara

Most lionesses reproduce by the time they are four years of age.[147] Lions do not mate at a specific time of year and the females are polyestrous.[148] Like those of other cats, the male lion's penis has spines that point backward. During withdrawal of the penis, the spines rake the walls of the female's vagina, which may cause ovulation.[149][150] A lioness may mate with more than one male when she is in heat.[151] Lions of both sexes may be involved in group homosexual and courtship activities. Males will also head-rub and roll around with each other before mounting each other.[152][153] Generation length of the lion is about seven years.[154] The average gestation period is around 110 days;[148] the female gives birth to a litter of between one and four cubs in a secluded den, which may be a thicket, a reed-bed, a cave, or some other sheltered area, usually away from the pride. She will often hunt alone while the cubs are still helpless, staying relatively close to the den.[155] Lion cubs are born blind, their eyes opening around seven days after birth. They weigh 1.2–2.1 kg (2.6–4.6 lb) at birth and are almost helpless, beginning to crawl a day or two after birth and walking around three weeks of age.[156] To avoid a buildup of scent attracting the attention of predators, the lioness moves her cubs to a new den site several times a month, carrying them one-by-one by the nape of the neck.[155]

Usually, the mother does not integrate herself and her cubs back into the pride until the cubs are six to eight weeks old.[155] Sometimes the introduction to pride life occurs earlier, particularly if other lionesses have given birth at about the same time.[96][157] When first introduced to the rest of the pride, lion cubs lack confidence when confronted with adults other than their mother. They soon begin to immerse themselves in the pride life, however, playing among themselves or attempting to initiate play with the adults.[157] Lionesses with cubs of their own are more likely to be tolerant of another lioness's cubs than lionesses without cubs. Male tolerance of the cubs varies—one male could patiently let the cubs play with his tail or his mane, while another may snarl and bat the cubs away.[158]

Video of a lioness and her cubs in Phinda Reserve

Pride lionesses often synchronise their reproductive cycles and communal rearing and suckling of the young, which suckle indiscriminately from any or all of the nursing females in the pride. The synchronisation of births is advantageous because the cubs grow to being roughly the same size and have an equal chance of survival, and sucklings are not dominated by older cubs.[96][157] Weaning occurs after six or seven months. Male lions reach maturity at about three years of age and at four to five years are capable of challenging and displacing adult males associated with another pride. They begin to age and weaken at between 10 and 15 years of age at the latest.[159]

When one or more new males oust the previous males associated with a pride, the victors often kill any existing young cubs, perhaps because females do not become fertile and receptive until their cubs mature or die. Females often fiercely defend their cubs from a usurping male but are rarely successful unless a group of three or four mothers within a pride join forces against the male.[160] Cubs also die from starvation and abandonment, and predation by leopards, hyenas and wild dogs. Male cubs are excluded from their maternal pride when they reach maturity at around two or three years of age,[161] while some females may leave when they reach the age of two.[92] When a new male lion takes over a pride, adolescents both male and female may be evicted.[162]

Health and mortality

 
Lions in a tree near Lake Nakuru

Lions may live 12–17 years.[46] Although adult lions have no natural predators, evidence suggests most die violently from attacks by humans or other lions.[163] Lions often inflict serious injuries on members of other prides they encounter in territorial disputes or members of the home pride when fighting at a kill.[164] Crippled lions and cubs may fall victim to hyenas and leopards or be trampled by buffalo or elephants. Careless lions may be maimed when hunting prey.[165] Nile crocodiles may also kill and eat lions, evidenced by the occasional lion claw found in crocodile stomachs.[166]

Ticks commonly infest the ears, neck and groin regions of the lions.[167][168] Adult forms of several tapeworm species of the genus Taenia have been isolated from lion intestines, having been ingested as larvae in antelope meat.[169] Lions in the Ngorongoro Crater were afflicted by an outbreak of stable fly (Stomoxys calcitrans) in 1962, resulting in lions becoming emaciated and covered in bloody, bare patches. Lions sought unsuccessfully to evade the biting flies by climbing trees or crawling into hyena burrows; many died or migrated and the local population dropped from 70 to 15 individuals.[170] A more recent outbreak in 2001 killed six lions.[171]

Captive lions have been infected with canine distemper virus (CDV) since at least the mid-1970s.[172] CDV is spread by 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.[173] Feline immunodeficiency virus and lentivirus also affect captive lions.[174][175]

Communication

 
Head rubbing among pride members is a common social behaviour.

When resting, lion socialisation occurs through a number of behaviours; the animal's expressive movements are highly developed. The most common peaceful, tactile gestures are head rubbing and social licking,[176] which have been compared with the role of allogrooming among primates.[177] Head rubbing, nuzzling the forehead, face and neck against another lion appears to be a form of greeting[178] and is seen often after an animal has been apart from others or after a fight or confrontation. Males tend to rub other males, while cubs and females rub females.[179] Social licking often occurs in tandem with head rubbing; it is generally mutual and the recipient appears to express pleasure. The head and neck are the most common parts of the body licked; this behaviour may have arisen out of utility because lions cannot lick these areas themselves.[180]

 
Lion roar
A captive lion roaring
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Lions have an array of facial expressions and body postures that serve as visual gestures.[181] A common facial expression is the "grimace face" or flehmen response, which a lion makes when sniffing chemical signals and involves an open mouth with bared teeth, raised muzzle, wrinkled nose, closed eyes and relaxed ears.[182] Lions also use chemical and visual marking;[181] males spray urine[183] and scrape plots of ground and objects within the territory.[181]

The lion's repertoire of vocalisations is large; variations in intensity and pitch appear to be central to communication. Most lion vocalisations are variations of growling, snarling, meowing and roaring. Other sounds produced include purring, puffing, bleating and humming. Roaring is used to advertise its presence. Lions most often roar at night, a sound that can be heard from a distance of 8 kilometres (5 mi).[184] They tend to roar in a very characteristic manner starting with a few deep, long roars that subside into grunts.[185][186]

Conservation

The lion is listed as Vulnerable on the IUCN Red List. The Indian population is listed on CITES Appendix I and the African population on CITES Appendix II.[2]

In Africa

Video of a wild lioness

Several large and well-managed protected areas in Africa host large lion populations. Where an infrastructure for wildlife tourism has been developed, cash revenue for park management and local communities is a strong incentive for lion conservation.[2] Most lions now live in East and Southern Africa; their numbers are rapidly decreasing, and fell by an estimated 30–50% in the late half of the 20th century. Primary causes of the decline include disease and human interference.[2] In 1975, it was estimated that since the 1950s, lion numbers had decreased by half to 200,000 or fewer.[187] Estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004.[188][81]

In the Republic of the Congo, Odzala-Kokoua National Park was considered a lion stronghold in the 1990s. By 2014, no lions were recorded in the protected area so the population is considered locally extinct.[189] The West African lion population is isolated from the one in Central Africa, with little or no exchange of breeding individuals. In 2015, it was estimated that this population consists of about 400 animals, including fewer than 250 mature individuals. They persist in three protected areas in the region, mostly in one population in the W A P protected area complex, shared by Benin, Burkina Faso and Niger. This population is listed as Critically Endangered.[13] Field surveys in the WAP ecosystem revealed that lion occupancy is lowest in the W National Park, and higher in areas with permanent staff and thus better protection.[190]

A population occurs in Cameroon's Waza National Park, where between approximately 14 and 21 animals persisted as of 2009.[191] In addition, 50 to 150 lions are estimated to be present in Burkina Faso's Arly-Singou ecosystem.[192] In 2015, an adult male lion and a female lion were sighted in Ghana's Mole National Park. These were the first sightings of lions in the country in 39 years.[193] In the same year, a population of up to 200 lions that was previously thought to have been extirpated was filmed in the Alatash National Park, Ethiopia, close to the Sudanese border.[194][195]

In 2005, Lion Conservation Strategies were developed for West and Central Africa, and or East and Southern Africa. The strategies seek to maintain suitable habitat, ensure a sufficient wild prey base for lions, reduce factors that lead to further fragmentation of populations, and make lion–human coexistence sustainable.[196][197] Lion depredation on livestock is significantly reduced in areas where herders keep livestock in improved enclosures. Such measures contribute to mitigating human–lion conflict.[198]

In Asia

 
A lioness in Gir National Park

The last refuge of the Asiatic lion population is the 1,412 km2 (545 sq mi) Gir National Park and surrounding areas in the region of Saurashtra or Kathiawar Peninsula in Gujarat State, India. The population has risen from approximately 180 lions in 1974 to about 400 in 2010.[199] It is geographically isolated, which can lead to inbreeding and reduced genetic diversity. Since 2008, the Asiatic lion has been listed as Endangered on the IUCN Red List.[12] By 2015, the population had grown to 523 individuals inhabiting an area of 7,000 km2 (2,700 sq mi) in Saurashtra.[200][201][202] The Asiatic Lion Census conducted in 2017 recorded about 650 individuals.[203]

The presence of numerous human habitations close to the National Park results in conflict between lions, local people and their livestock.[204][200] Some consider the presence of lions a benefit, as they keep populations of crop damaging herbivores in check.[205] The establishment of a second, independent Asiatic lion population in Kuno Wildlife Sanctuary, located in Madhya Pradesh was planned but in 2017, the Asiatic Lion Reintroduction Project seemed unlikely to be implemented.[206][207]

Captive breeding

 
Two captive male Asiatic lions in Sanjay Gandhi National Park, India

Lions imported to Europe before the middle of the 19th century were possibly foremost Barbary lions from North Africa, or Cape lions from Southern Africa.[208] Another 11 animals thought to be Barbary lions kept in Addis Ababa Zoo are descendants of animals owned by Emperor Haile Selassie. WildLink International in collaboration with Oxford University launched an ambitious International Barbary Lion Project with the aim of identifying and breeding Barbary lions in captivity for eventual reintroduction into a national park in the Atlas Mountains of Morocco.[209] However, a genetic analysis showed that the captive lions at Addis Ababa Zoo were not Barbary lions, but rather closely related to wild lions in Chad and Cameroon.[210]

In 1982, the Association of Zoos and Aquariums started a Species Survival Plan for the Asiatic lion to increase its chances of survival. In 1987, it was found that most lions in North American zoos were hybrids between African and Asiatic lions.[211] Breeding programs need to note origins of the participating animals to avoid cross-breeding different subspecies and thus reducing their conservation value.[212] Captive breeding of lions was halted to eliminate individuals of unknown origin and pedigree. Wild-born lions were imported to American zoos from Africa between 1989 and 1995. Breeding was continued in 1998 in the frame of an African lion Species Survival Plan.[213]

About 77% of the captive lions registered in the International Species Information System in 2006 were of unknown origin; these animals might have carried genes that are extinct in the wild and may therefore be important to the maintenance of the overall genetic variability of the lion.[62]

Interactions with humans

In zoos and circuses

 
Lion at Melbourne Zoo
 
19th-century etching of a lion tamer in a cage with lions and tigers

Lions are part of a group of exotic animals that have been central to zoo exhibits since the late 18th century. Although many modern zoos are more selective about their exhibits,[214] there are more than 1,000 African and 100 Asiatic lions in zoos and wildlife parks around the world. They are considered an ambassador species and are kept for tourism, education and conservation purposes.[215] Lions can live over twenty years in captivity; for example, three sibling lions at the Honolulu Zoo lived to the age of 22 in 2007.[216][217]

The first European "zoos" spread among noble and royal families in the 13th century, and until the 17th century were called seraglios. At that time, they came to be called menageries, an extension of the cabinet of curiosities. They spread from France and Italy during the Renaissance to the rest of Europe.[218] In England, although the seraglio tradition was less developed, lions were kept at the Tower of London in a seraglio established by King John in the 13th century;[219][220] this was probably stocked with animals from an earlier menagerie started in 1125 by Henry I at his hunting lodge in Woodstock, Oxfordshire, where according to William of Malmesbury lions had been stocked.[221]

Lions were kept in cramped and squalid conditions at London Zoo until a larger lion house with roomier cages was built in the 1870s.[222] Further changes took place in the early 20th century when Carl Hagenbeck designed enclosures with concrete "rocks", more open space and a moat instead of bars, more closely resembling a natural habitat. Hagenbeck designed lion enclosures for both Melbourne Zoo and Sydney's Taronga Zoo; although his designs were popular, the use of bars and caged enclosures prevailed in many zoos until the 1960s.[223] In the late 20th century, larger, more natural enclosures and the use of wire mesh or laminated glass instead of lowered dens allowed visitors to come closer than ever to the animals; some attractions such as the Cat Forest/Lion Overlook of Oklahoma City Zoological Park placed the den on ground level, higher than visitors.[224]

Lion taming has been part of both established circuses and individual acts such as Siegfried & Roy. The practice began in the early 19th century by Frenchman Henri Martin and American Isaac Van Amburgh, who both toured widely and whose techniques were copied by a number of followers.[225] Van Amburgh performed before Queen Victoria in 1838 when he toured Great Britain. Martin composed a pantomime titled Les Lions de Mysore ("the lions of Mysore"), an idea Amburgh quickly borrowed. These acts eclipsed equestrianism acts as the central display of circus shows and entered public consciousness in the early 20th century with cinema. In demonstrating the superiority of human over animal, lion taming served a purpose similar to animal fights of previous centuries.[225] The ultimate proof of a tamer's dominance and control over a lion is demonstrated by the placing of the tamer's head in the lion's mouth. The now-iconic lion tamer's chair was possibly first used by American Clyde Beatty (1903–1965).[226]

Hunting and games

Main article: Lion hunting
See also: Lion baiting
 
Bas-relief of a wounded lioness from Nineveh, c. 645–635 BC

Lion hunting has occurred since ancient times and was often a royal tradition, intended to demonstrate the power of the king over nature. Such hunts took place in a reserved area in front of an audience. The monarch was accompanied by his men and controls were put in place to increase their safety and ease of killing. The earliest surviving record of lion hunting is an ancient Egyptian inscription dated circa 1380 BC that mentions Pharaoh Amenhotep III killing 102 lions in ten years "with his own arrows". The Assyrian emperor Ashurbanipal had one of his lion hunts depicted on a sequence of Assyrian palace reliefs c. 640 BC, known as the Lion Hunt of Ashurbanipal. Lions were also hunted during the Mughal Empire, where Emperor Jahangir is said to have excelled at it.[227] In Ancient Rome, lions were kept by emperors for hunts, gladiator fights and executions.[228]

The Maasai people have traditionally viewed the killing of lions as a rite of passage. Historically, lions were hunted by individuals, however, due to reduced lion populations, elders discourage solo lion hunts.[229] During the European colonisation of Africa in the 19th century, the hunting of lions was encouraged because they were considered pests and lion skins were sold for £1 each.[230] The widely reproduced imagery of the heroic hunter chasing lions would dominate a large part of the century.[231] Trophy hunting of lions in recent years has been met with controversy, notably with the killing of Cecil the lion in mid-2015.[232]

Man-eating

 
The Tsavo maneaters of East Africa on display in the Field Museum of Natural History in Chicago
Further information: Man-eater § Lions

Lions do not usually hunt humans but some (usually males) seem to seek them out. One well-publicised case is the Tsavo maneaters; in 1898, 28 officially recorded railway workers building the Uganda Railway were taken by lions over nine months during the construction of a bridge in Kenya.[233] The hunter who killed the lions wrote a book detailing the animals' predatory behaviour; they were larger than normal and lacked manes, and one seemed to suffer from tooth decay. The infirmity theory, including tooth decay, is not favoured by all researchers; an analysis of teeth and jaws of man-eating lions in museum collections suggests that while tooth decay may explain some incidents, prey depletion in human-dominated areas is a more likely cause of lion predation on humans.[234] Sick or injured animals may be more prone to man-eating but the behaviour is not unusual, nor necessarily aberrant.[235]

Lions' proclivity for man-eating has been systematically examined. American and Tanzanian scientists report that man-eating behaviour in rural areas of Tanzania increased greatly from 1990 to 2005. At least 563 villagers were attacked and many eaten over this period. The incidents occurred near Selous Game Reserve in Rufiji River and in Lindi Region near the Mozambican border. While the expansion of villages into bush country is one concern, the authors argue conservation policy must mitigate the danger because in this case, conservation contributes directly to human deaths. Cases in Lindi in which lions seize humans from the centres of substantial villages have been documented.[236] Another study of 1,000 people attacked by lions in southern Tanzania between 1988 and 2009 found that the weeks following the full moon, when there was less moonlight, were a strong indicator of increased night-time attacks on people.[237]

According to Robert R. Frump, Mozambican refugees regularly crossing Kruger National Park, South Africa, at night are attacked and eaten by lions; park officials have said man-eating is a problem there. Frump said thousands may have been killed in the decades after apartheid sealed the park and forced refugees to cross the park at night. For nearly a century before the border was sealed, Mozambicans had regularly crossed the park in daytime with little harm.[238]

Cultural significance

Main article: Cultural depictions of lions
 
Lions carved on a rock weight, Jiroft culture, Iran, 3rd mil. BC

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.[45] It appeared as a symbol for strength and nobility in cultures across Europe, Asia and Africa, despite incidents of attacks on people. The lion has been depicted as "king of the jungle" and "king of beasts", and thus became a popular symbol for royalty and stateliness.[239] The lion is also used as a symbol of sporting teams.[240]

Africa

 
Granite statue of the Egyptian goddess Sekhmet from the Luxor Temple, dated 1403–1365 BC, exhibited in the National Museum of Denmark

In sub-Saharan Africa, the lion has been a common character in stories, proverbs and dances, but rarely featured in visual arts.[241] In some cultures, the lion symbolises power and royalty.[242] In the Swahili language, the lion is known as simba which also means "aggressive", "king" and "strong".[54] Some rulers had the word "lion" in their nickname. Sundiata Keita of the Mali Empire was called "Lion of Mali".[243] The founder of the Waalo kingdom is said to have been raised by lions and returned to his people part-lion to unite them using the knowledge he learned from the lions.[242]

In parts of West Africa, lions symbolised the top class of their social hierarchies.[242] In more heavily forested areas where lions were rare, the leopard represented the top of the hierarchy.[241] In parts of West and East Africa, the lion is associated with healing and provides the connection between seers and the supernatural. In other East African traditions, the lion represents laziness.[242] In much of African folklore, the lion is portrayed as having low intelligence and is easily tricked by other animals.[243]

The ancient Egyptians portrayed several of their war deities as lionesses, which they revered as fierce hunters. Egyptian deities associated with lions include Sekhmet, Bast, Mafdet, Menhit, Pakhet and Tefnut.[239] These deities were often connected with the sun god Ra and his fierce heat, and their dangerous power was invoked to guard people or sacred places. The sphinx, a figure with a lion's body and the head of a human or other creature, represented a pharaoh or deity who had taken on this protective role.[244]

Asia

 
Roaring and striding lion from the Throne Room of Nebuchadnezzar II, 6th century BC, from Babylon, Iraq

The lion was a prominent symbol in ancient Mesopotamia from Sumer up to Assyrian and Babylonian times, where it was strongly associated with kingship.[245] Lions were among the major symbols of the goddess Inanna/Ishtar.[246][247] The Lion of Babylon was the foremost symbol of the Babylonian Empire.[248] The Lion of Judah is the biblical emblem of the tribe of Judah and the later Kingdom of Judah.[249] Lions are frequently mentioned in the Bible, notably in the Book of Daniel, in which the eponymous hero refuses to worship King Darius and is forced to sleep in the lions' den where he is miraculously unharmed (Dan 6). In the Book of Judges, Samson kills a lion as he travels to visit a Philistine woman.(Judg 14).[250]

Indo-Persian chroniclers regarded the lion as keeper of order in the realm of animals. The Sanskrit word mrigendra signifies a lion as king of animals in general or deer in particular.[251] Narasimha, the man-lion, is one of ten avatars of the Hindu god Vishnu.[252] Singh is an ancient Indian vedic name meaning "lion", dating back over 2,000 years. It was originally used only by Rajputs, a Hindu Kshatriya or military caste but is used by millions of Hindu Rajputs and more than twenty million Sikhs today.[253] The Lion Capital of Ashoka, erected by Emperor Ashoka in the 3rd century CE, depicts four lions standing back to back. It was made the National Emblem of India in 1950.[254] The lion is also symbolic for the Sinhalese people, the term derived from the Sanskrit Sinhala, meaning "of lions"[255] while a sword-wielding lion is the central figure on the national flag of Sri Lanka.[256]

The lion is a common motif in Chinese art; it was first used in art during the late Spring and Autumn period (fifth or sixth century BC) and became more popular during the Han dynasty (206 BC – AD 220) when imperial guardian lions started to be placed in front of imperial palaces for protection. Because lions have never been native to China, early depictions were somewhat unrealistic; after the introduction of Buddhist art to China in the Tang dynasty after the sixth century AD, lions were usually depicted wingless with shorter, thicker bodies and curly manes.[257] The lion dance is a traditional dance in Chinese culture in which performers in lion costumes mimic a lion's movements, often with musical accompaniment from cymbals, drums and gongs. They are performed at Chinese New Year, the August Moon Festival and other celebratory occasions for good luck.[258]

Western world

 
Dorothy Gale meets the Cowardly Lion in The Wonderful Wizard of Oz. Art by W. W. Denslow, 1900.

Lion-headed figures and amulets were excavated in tombs in the Greek islands of Crete, Euboea, Rhodes, Paros and Chios. They are associated with the Egyptian deity Sekhmet and date to the early Iron Age between the 9th and 6th centuries BC.[259] Tfhe lion is featured in several of Aesop's fables, notably The Lion and the Mouse.[260] The Nemean lion was symbolic in ancient Greece and Rome, represented as the constellation and zodiac sign Leo, and described in mythology, where it was killed and worn by the hero Heracles,[261] symbolising victory over death.[262] Lancelot and Gawain were also heroes slaying lions in the Middle Ages. In some medieval stories, lions were portrayed as allies and companions.[263] "Lion" was the nickname of several medieval warrior-rulers with a reputation for bravery, such as Richard the Lionheart.[239]

Lions continue to appear in modern literature as characters including the messianic Aslan in the 1950 novel The Lion, the Witch and the Wardrobe and The Chronicles of Narnia series by C. S. Lewis,[264] and the comedic Cowardly Lion in L. Frank Baum's 1900 The Wonderful Wizard of Oz.[265] Lion symbolism was used from the advent of cinema; one of the most iconic and widely recognised lions is Leo, which has been the mascot for Metro-Goldwyn-Mayer studios since the 1920s.[266] The 1966 film Born Free features Elsa the lioness and is based on the book Born Free published in 1960.[267] The lion's role as king of the beasts has been used in the 1994 Disney animated feature film The Lion King.[268]

Lions are frequently depicted on coats of arms, like on the coat of arms of Finland,[269] either as a device on shields or as supporters, but the lioness is used much less frequently.[270] The heraldic lion is particularly common in British arms. It is traditionally depicted in a great variety of attitudes, although within French heraldry only lions rampant are considered to be lions; felined figures in any other position are instead referred to as leopards.[271]

See also

Notes

  1. ^ Populations of India are listed in Appendix I.

References

Citations

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Books

  • Baratay, E. & Hardouin-Fugier, E. (2002). Zoo: A History of Zoological Gardens in the West. London: Reaktion Books. ISBN 978-1-86189-111-2.
  • Blunt, W. (1975). The Ark in the Park: The Zoo in the Nineteenth Century. London: Hamish Hamilton. ISBN 978-0-241-89331-9.
  • de Courcy, C. (1995). The Zoo Story. Ringwood, Victoria: Penguin Books. ISBN 978-0-14-023919-5.
  • Jackson, D. (2010). Lion. London: Reaktion Books. ISBN 978-1861896551.
  • Packer, C. (2023). The Lion: Behavior, Ecology, and Conservation of an Iconic Species. Princeton University Press. ISBN 9780691215297.
  • Schaller, G. B. (1972). The Serengeti Lion: A Study of Predator–Prey Relations. Chicago: University of Chicago Press. ISBN 978-0-226-73639-6.
  • Scott, J.; Scott, A. (2002). Big Cat Diary: Lion. New York: Harper Collins. ISBN 9780007146666.

External links

 
Look up lion in Wiktionary, the free dictionary.
 
Wikispecies has information related to Lion.
 
Wikisource has the text of the 1911 Encyclopædia Britannica article "Lion".
  •   Media related to Lion at Wikimedia Commons
  • IUCN/SSC Cat Specialist Group. "Lion Panthera leo".
  • "Lion Conservation Fund".
  • The Portugal News (2014). . Archived from the original on 2 August 2018. Retrieved 24 May 2018.
  • "Lion" . Collier's New Encyclopedia. 1921.

February 16, 2024
lion, other, uses, disambiguation, lion, panthera, large, genus, panthera, native, africa, india, muscular, broad, chested, body, short, rounded, head, round, ears, hairy, tuft, tail, sexually, dimorphic, adult, male, lions, larger, than, females, have, promin. For other uses see Lion disambiguation The lion Panthera leo is a large cat of the genus Panthera native to Africa and India It has a muscular broad chested body a short rounded head round ears and a hairy tuft at the end of its tail It is sexually dimorphic adult male lions are larger than females and have a prominent mane It is a social species forming groups called prides A lion s pride consists of a few adult males related females and cubs Groups of female lions usually hunt together preying mostly on large ungulates The lion is an apex and keystone predator although some lions scavenge when opportunities occur and have been known to hunt humans lions typically do not actively seek out and prey on humans LionTemporal range Pleistocene Present PreꞒ Ꞓ O S D C P T J K Pg N Male in Okonjima NamibiaFemale lioness in OkonjimaConservation statusVulnerable IUCN 3 1 2 CITES Appendix II CITES a 2 Scientific classificationDomain EukaryotaKingdom AnimaliaPhylum ChordataClass MammaliaOrder CarnivoraSuborder FeliformiaFamily FelidaeSubfamily PantherinaeGenus PantheraSpecies P leo 1 Binomial namePanthera leo 1 Linnaeus 1758 3 SubspeciesP l leo P l melanochaita P l sinhaleyusHistorical and present distribution of the lion in Africa Asia and EuropeThe lion inhabits grasslands savannahs and shrublands It is usually more diurnal than other wild cats but when persecuted it adapts to being active at night and at twilight During the Neolithic period the lion ranged throughout Africa and Eurasia from Southeast Europe to India but it has been reduced to fragmented populations in sub Saharan Africa and one population in western India It has been listed as Vulnerable on the IUCN Red List since 1996 because populations in African countries have 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 for concern One of the most widely recognised animal symbols in human culture the lion 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 zoological gardens across the world since the late 18th century Cultural depictions of lions were prominent in Ancient Egypt and depictions have occurred in virtually all ancient and medieval cultures in the lion s historic and current range Contents 1 Etymology 2 Taxonomy 2 1 Subspecies 2 2 Fossil records 2 3 Evolution 2 4 Hybrids 3 Description 3 1 Size 3 2 Mane 3 3 Colour variation 4 Distribution and habitat 4 1 Historical range 5 Behaviour and ecology 5 1 Group organisation 5 2 Hunting and diet 5 3 Predatory competition 5 4 Reproduction and life cycle 5 5 Health and mortality 5 6 Communication 6 Conservation 6 1 In Africa 6 2 In Asia 6 3 Captive breeding 7 Interactions with humans 7 1 In zoos and circuses 7 2 Hunting and games 7 3 Man eating 8 Cultural significance 8 1 Africa 8 2 Asia 8 3 Western world 9 See also 10 Notes 11 References 11 1 Citations 11 2 Books 12 External linksEtymologyThe English word lion is derived via Anglo Norman liun from Latin leōnem nominative leō which in turn was a borrowing from Ancient Greek lewn leōn The Hebrew word ל ב יא lavi may also be related 4 The generic name Panthera is traceable to the classical Latin word panthera and the ancient Greek word pan8hr panther 5 Taxonomy nbsp The upper cladogram is based on the 2006 study 6 7 the lower one on the 2010 8 and 2011 9 studies Felis leo was the scientific name used by Carl Linnaeus in 1758 who described the lion in his work Systema Naturae 3 The genus name Panthera was coined by Lorenz Oken in 1816 10 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 1 They were distinguished mostly by the size and colour of their manes and skins 11 Subspecies nbsp Range map showing distribution of subspecies and cladesIn the 19th and 20th centuries several lion type specimens were described and proposed as subspecies with about a dozen recognised as valid taxa until 2017 1 Between 2008 and 2016 IUCN Red List assessors used only two subspecific names P l leo for African lion populations and P l persica for the Asiatic lion population 2 12 13 In 2017 the Cat Classification Task Force of the Cat Specialist Group revised lion taxonomy and recognises two subspecies based on results of several phylogeographic studies on lion evolution namely 14 P l leo Linnaeus 1758 the nominate lion subspecies includes the Asiatic lion the regionally extinct Barbary lion and lion populations in West and northern parts of Central Africa 14 Synonyms include P l persica Meyer 1826 P l senegalensis Meyer 1826 P l kamptzi Matschie 1900 and P l azandica Allen 1924 1 Multiple authors referred to it as northern lion and northern subspecies 15 16 P l melanochaita Smith 1842 includes the extinct Cape lion and lion populations in East and Southern African regions 14 Synonyms include P l somaliensis Noack 1891 P l massaica Neumann 1900 P l sabakiensis Lonnberg 1910 P l bleyenberghi Lonnberg 1914 P l roosevelti Heller 1914 P l nyanzae Heller 1914 P l hollisteri Allen 1924 P l krugeri Roberts 1929 P l vernayi Roberts 1948 and P l webbiensis Zukowsky 1964 1 11 It has been referred to as southern subspecies and southern lion 16 However there seems to be some degree of overlap between both groups in northern Central Africa DNA analysis from a more recent study indicates that Central African lions are derived from both northern and southern lions as they cluster with P leo leo in mtDNA based phylogenies whereas their genomic DNA indicates a closer relationship with P leo melanochaita 17 Lion samples from some parts of the Ethiopian Highlands cluster genetically with those from Cameroon and Chad while lions from other areas of Ethiopia cluster with samples from East Africa Researchers therefore assume Ethiopia is a contact zone between the two subspecies 18 Genome wide data of a wild born historical lion sample from Sudan showed that it clustered with P l leo in mtDNA based phylogenies but with a high affinity to P l melanochaita This result suggested that the taxonomic position of lions in Central Africa may require revision 19 Fossil records nbsp Skull of an American lion on display at the National Museum of Natural HistoryOther lion subspecies or sister species to the modern lion existed in prehistoric times 20 P l sinhaleyus was a fossil carnassial excavated in Sri Lanka which was attributed to a lion It is thought to have become extinct around 39 000 years ago 21 P fossilis was larger than the modern lion and lived in the Middle Pleistocene Bone fragments were excavated in caves in the United Kingdom Germany Italy and Czech Republic 22 23 P spelaea or the cave lion lived in Eurasia and Beringia during the Late Pleistocene It became extinct due to climate warming or human expansion latest by 11 900 years ago 24 Bone fragments excavated in European North Asian Canadian and Alaskan caves indicate that it ranged from Europe across Siberia into western Alaska 25 It likely derived from P fossilis 26 and was genetically isolated and highly distinct from the modern lion in Africa and Eurasia 27 26 It is depicted in Paleolithic cave paintings ivory carvings and clay busts 28 P atrox or the American lion ranged in the Americas from Canada to possibly Patagonia 29 It arose when a cave lion population in Beringia became isolated south of the Cordilleran Ice Sheet about 370 000 years ago 30 31 A fossil from Edmonton dates to 11 355 55 years ago 32 Evolution nbsp red Panthera spelaeablue Panthera atroxgreen Panthera leoMaximal range of the modern lionand its prehistoric relativesin the late PleistoceneThe Panthera lineage is estimated to have genetically diverged from the common ancestor of the Felidae around 9 32 to 4 47 million years ago to 11 75 to 0 97 million years ago 6 33 34 Results of analyses differ in the phylogenetic relationship of the lion it was thought to form a sister group with the jaguar P onca that diverged 3 46 to 1 22 million years ago 6 but also with the leopard P pardus that diverged 3 1 to 1 95 million years ago 8 9 to 4 32 to 0 02 million years ago Hybridisation between lion and snow leopard P uncia ancestors possibly continued until about 2 1 million years ago 34 The lion leopard clade was distributed in the Asian and African Palearctic since at least the early Pliocene 35 The earliest fossils recognisable as lions were found at Olduvai Gorge in Tanzania and are estimated to be up to 2 million years old 33 Estimates for the divergence time of the modern and cave lion lineages range from 529 000 to 392 000 years ago based on mutation rate per generation time of the modern lion There is no evidence for gene flow between the two lineages indicating that they did not share the same geographic area 19 The Eurasian and American cave lions became extinct at the end of the last glacial period without mitochondrial descendants on other continents 27 36 37 The modern lion was probably widely distributed in Africa during the Middle Pleistocene and started to diverge in sub Saharan Africa during the Late Pleistocene Lion populations in East and Southern Africa became separated from populations in West and North Africa when the equatorial rainforest expanded 183 500 to 81 800 years ago 38 They shared a common ancestor probably between 98 000 and 52 000 years ago 19 Due to the expansion of the Sahara between 83 100 and 26 600 years ago lion populations in West and North Africa became separated As the rainforest decreased and thus gave rise to more open habitats lions moved from West to Central Africa Lions from North Africa dispersed to southern Europe and Asia between 38 800 and 8 300 years ago 38 Extinction of lions in southern Europe North Africa and the Middle East interrupted gene flow between lion populations in Asia and Africa Genetic evidence revealed numerous mutations in lion samples from East and Southern Africa which indicates that this group has a longer evolutionary history than genetically less diverse lion samples from Asia and West and Central Africa 39 A whole genome wide sequence of lion samples showed that samples from West Africa shared alleles with samples from Southern Africa and samples from Central Africa shared alleles with samples from Asia This phenomenon indicates that Central Africa was a melting pot of lion populations after they had become isolated possibly migrating through corridors in the Nile Basin during the early Holocene 19 Hybrids Further information Panthera hybrid In zoos lions have been bred with tigers to create hybrids for the curiosity of visitors or for scientific purpose 40 41 The liger is bigger than a lion and a tiger whereas most tigons are relatively small compared to their parents because of reciprocal gene effects 42 43 The leopon is a hybrid between a lion and leopard 44 Description nbsp A tuft at the end of the tail is a distinct characteristic of the lion nbsp Skeleton The lion is a muscular broad chested cat with a short rounded head a reduced neck and round ears males have broader heads The fur varies in colour from light buff to silvery grey yellowish red and dark brown The colours of the underparts are generally lighter A new born lion has dark spots which fade as the cub reaches adulthood although faint spots may still be seen on the legs and underparts 45 46 The tail of all lions ends in a dark hairy tuft that in some lions conceals an approximately 5 mm 0 20 in long hard spine or spur that is formed from the final fused sections of tail bone The functions of the spur are unknown The tuft is absent at birth and develops at around 5 1 2 months of age It is readily identifiable at the age of seven months 47 Its skull is very similar to that of the tiger although the frontal region is usually more depressed and flattened and has a slightly shorter postorbital region and broader nasal openings than those of the 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 48 49 The skeletal muscles of the lion make up 58 8 of its body weight and represent the highest percentage of muscles among mammals 50 51 Size Among felids the lion is second only to the tiger in size 46 The size and weight of adult lions vary across its range and habitats 52 53 54 55 Accounts of a few individuals that were larger than average exist from Africa and India 45 56 57 58 Average Female lions Male lionsHead and body length 160 184 cm 63 72 in 59 184 208 cm 72 82 in 59 Tail length 72 89 5 cm 28 3 35 2 in 59 82 5 93 5 cm 32 5 36 8 in 59 Weight 118 37 143 52 kg 261 0 316 4 lb in Southern Africa 52 119 5 kg 263 lb in East Africa 52 110 120 kg 240 260 lb in India 53 186 55 225 kg 411 3 496 0 lb in Southern Africa 52 174 9 kg 386 lb in East Africa 52 160 190 kg 350 420 lb in India 53 Mane nbsp A six year old male in Phinda Private Game Reserve nbsp Male in Pendjari National Park The male lion s mane is the most recognisable feature of the species 11 It may have evolved around 320 000 190 000 years ago 60 It grows downwards and backwards covering most of the head neck shoulders and chest The mane is typically brownish and tinged with yellow rust and black hairs 46 It starts growing when lions enter adolescence when testosterone levels increase and reach their full size at around four years old 61 Cool ambient temperatures in European and North American zoos may result in a heavier mane 62 On average Asiatic lions have sparser manes than African lions 63 This feature likely evolved to signal the fitness of males to females The mane of the lion is thought to serve mating purposes Males with darker manes appear to have greater reproductive success and are more likely to remain in a pride for longer They have longer and thicker hair and higher testosterone levels but they are also more vulnerable to heat stress 64 65 Unlike in other felid species female lions consistently interact with multiple males at once 66 Another hypothesis suggests that the mane also serves to protect the neck in fights but this is disputed 67 68 During fights including those involving maneless females and adolescents the neck is not targeted as much as the face back and hindquarters Injured lions also begin to lose their manes 69 Almost all male lions in Pendjari National Park are either maneless or have very short manes 70 Maneless lions have also been reported in Senegal in Sudan s Dinder National Park and in Tsavo East National Park Kenya 71 Castrated lions often have little to no mane because the removal of the gonads inhibits testosterone production 72 Rarely lionesses in the wild and in captivity have been observed to grow manes 73 74 Increased testosterone may be the cause of maned lionesses reported in northern Botswana 75 Colour variation Further information White lion The white lion is a rare morph with a genetic condition called leucism which is caused by a double recessive allele It is not albino it has normal pigmentation in the eyes and skin White lions have occasionally been encountered in and around Kruger National Park and the adjacent Timbavati Private Game Reserve in eastern South Africa They were removed from the wild in the 1970s thus decreasing the white lion gene pool Nevertheless 17 births have been recorded in five prides between 2007 and 2015 76 White lions are selected for breeding in captivity 77 They have reportedly been bred in camps in South Africa for use as trophies to be killed during canned hunts 78 Distribution and habitat nbsp Lion in Gir National ParkAfrican lions live in scattered populations across sub Saharan Africa The lion prefers grassy plains and savannahs scrub bordering rivers and open woodlands with bushes It rarely enters closed forests On Mount Elgon the lion has been recorded up to an elevation of 3 600 m 11 800 ft and close to the snow line on Mount Kenya 45 Savannahs with an annual rainfall of 300 to 1 500 mm 12 to 59 in make up the majority of lion habitat in Africa estimated at 3 390 821 km2 1 309 203 sq mi at most but remnant populations are also present in tropical moist forests in West Africa and montane forests in East Africa 79 The Asiatic lion now survives only in and around Gir National Park in Gujarat western India Its habitat is a mixture of dry savannah forest and very dry deciduous scrub forest 12 Historical range In Africa the range of the lion originally spanned most of the central African rainforest zone and the Sahara desert 80 In the 1960s it became extinct in North Africa except in the southern part of Sudan 81 79 82 In southern Europe and Asia the lion once ranged in regions where climatic conditions supported an abundance of prey 83 In Greece it was common as reported by Herodotus in 480 BC it was considered rare by 300 BC and extirpated by AD 100 45 It was present in the Caucasus until the 10th century 49 It lived in Palestine until the Middle Ages and in Southwest Asia until the late 19th century By the late 19th century it had been extirpated in most of Turkey 84 The last live lion in Iran was sighted in 1942 about 65 km 40 mi northwest of Dezful 85 although the corpse of a lioness was found on the banks of the Karun river in Khuzestan province in 1944 86 It once ranged from Sind and Punjab in Pakistan to Bengal and the Narmada River in central India 87 Behaviour and ecologyLions spend much of their time resting they are inactive for about twenty hours per day 88 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 continue until dawn when hunting most often takes place They spend an average of two hours a day walking and fifty minutes eating 89 Group organisation nbsp Lion pride in Etosha National Park nbsp A lioness left and two males in Masai Mara The lion is the most social of all wild felid species living in groups of related individuals with their offspring Such a group is called a pride Groups of male lions are called coalitions 90 Females form the stable social unit in a pride and do not tolerate outside females 91 The majority of females remain in their birth prides while all males and some females will disperse 92 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 been observed 93 The sole exception to this pattern is the Tsavo lion pride that always has just one adult male 94 Prides act as fission fusion societies and members will split into subgroups that keep in contact with roars 95 Nomadic lions range widely and move around sporadically either in pairs or alone 90 Pairs are more frequent among related males A lion may switch lifestyles nomads can become residents and vice versa 96 Interactions between prides and nomads tend to be hostile although pride females in estrus allow nomadic males to approach them 97 Males spend years in a nomadic phase before gaining residence in a pride 98 A study undertaken in the Serengeti National Park revealed that nomadic coalitions gain residency at between 3 5 and 7 3 years of age 99 In Kruger National Park dispersing male lions move more than 25 km 16 mi away from their natal pride in search of their own territory Female lions stay closer to their natal pride Therefore female lions in an area are more closely related to each other than male lions in the same area 100 The evolution of sociability in lions was likely driven both by high population density and the clumped resources of savannah habitats The larger the pride the more high quality territory they can defend hotspots being near river confluences where the cats have better access to water prey and shelter via vegetation 101 102 The area occupied by a pride is called a pride area whereas that occupied by a nomad is a range 90 Males associated with a pride patrol the fringes 46 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 defense against intruders while others lag behind 103 Lions tend to assume specific roles in the pride slower moving individuals may provide other valuable services to the group 104 Alternatively there may be rewards associated with being a leader that fends off intruders the rank of lionesses in the pride is reflected in these responses 105 The male or males associated with the pride must defend their relationship with the pride from outside males who may attempt to usurp them 96 Dominance hierarchies do not appear to exist among individuals of either sex in a pride 106 Asiatic lion prides differ in group composition Male Asiatic lions are solitary or associate with up to three males forming a loose pride while females associate with up to 12 other females forming a stronger pride together with their cubs Female and male lions associate only when mating 107 Coalitions of males hold territory for a longer time than single lions Males in coalitions of three or four individuals exhibit a pronounced hierarchy in which one male dominates the others and mates more frequently 108 Hunting and diet nbsp Male lion and cub feeding on a Cape buffalo in Sabi Sand Game Reserve nbsp Four lionesses catching a buffalo in the Serengeti nbsp A skeletal mount of a lion attacking a common eland on display at The Museum of Osteology The lion is a generalist hypercarnivore and is considered to be both an apex and keystone predator due to its wide prey spectrum 109 110 Its prey consists mainly of ungulates particularly blue wildebeest plains zebra African buffalo common warthog gemsbok and giraffe 111 In India chital and sambar deer are the most common wild prey 46 111 112 while livestock contributes significantly to lion kills outside protected areas 113 They usually avoid fully grown adult elephants rhinoceroses and hippopotamus and small prey like dik dik hyraxes hares and monkeys 111 114 Unusual prey include porcupines and small reptiles Lions kill other predators but seldom consume them 115 Young lions first display stalking behaviour at around three months of age although they do not participate in hunting until they are almost a year old and begin to hunt effectively when nearing the age of two 116 Single lions are capable of bringing down zebra and wildebeest while larger prey like buffalo and giraffe are riskier 96 In Chobe National Park large prides have been observed hunting African bush elephants up to around 15 years old in exceptional cases with the victims being calves juveniles and even subadults 117 118 In typical hunts each lioness has a favoured 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 fleeing from other lionesses Males attached to prides do not usually participate in group hunting 119 Some evidence suggests however that males are just as successful as females they are typically solo hunters who ambush prey in small bushland 120 They may join in the hunting of large slower moving prey like buffalo and even hunt them on their own Moderately sized hunting groups generally have higher success rates than lone females and larger groups 121 Lions are not particularly known for their stamina For instance a lioness s heart comprises only 0 57 of her body weight and a male s is about 0 45 of his body weight whereas a hyena s heart comprises almost 1 of its body weight 122 Thus lions run quickly only in short bursts at about 48 59 km h 30 37 mph and need to be close to their prey before starting the attack 123 One study in 2018 recorded a lion running at a top speed of 74 1 km h 46 0 mph 124 They take advantage of factors that reduce visibility many kills take place near some form of cover or at night 125 The lion s attack is short and powerful it attempts to catch prey with a fast rush and final leap usually pulls it down by the rump and kills with a clamping bite to the throat or muzzle It can hold the prey s throat for up to 13 minutes until the prey stops moving 126 Lions typically consume prey at the location of the hunt but sometimes drag large prey into cover 127 They tend to squabble over kills particularly the males Cubs suffer most when food is scarce but otherwise all pride members eat their fill including old and crippled lions which can live on leftovers 96 Large kills are shared more widely among pride members 128 An adult lioness requires an average of about 5 kg 11 lb of meat per day while males require about 7 kg 15 lb 129 Lions gorge themselves and eat up to 30 kg 66 lb in one session 86 If it is unable to consume all of the kill it rests for a few hours before continuing to eat On hot days the pride retreats to shade with one or two males standing guard 127 Lions defend their kills from scavengers such as vultures and hyenas 96 Lions scavenge on carrion when the opportunity arises scavenging animals dead from natural causes such as disease or those that were killed by other predators Scavenging lions keep a constant lookout for circling vultures which indicate the death or distress of an animal 130 Most carrion on which both hyenas and lions feed upon are killed by hyenas rather than lions 55 Carrion is thought to provide a large part of lion diet 131 Predatory competition nbsp Lioness chasing a spotted hyena in Kruger National Park nbsp Lioness stealing a kill from a leopard in Kruger National Park Lions and spotted hyenas occupy a similar ecological niche and compete for prey and carrion a review of data across several studies indicates a dietary overlap of 58 6 132 Lions typically ignore hyenas unless they are on a kill or are being harassed while the latter tend to visibly react to the presence of lions with or without the presence of food In the Ngorongoro crater lions subsist largely on kills stolen from hyenas causing them to increase their kill rate 133 In Botswana s Chobe National Park the situation is reversed as hyenas there frequently challenge lions and steal their kills obtaining food from 63 of all lion kills 134 When confronted on a kill hyenas may either leave or wait patiently at a distance of 30 100 m 98 328 ft until the lions have finished 135 Hyenas may feed alongside lions and force them off a kill The two species attack one another even when there is no food involved for no apparent reason 136 Lions can account for up to 71 of hyena deaths in Etosha National Park Hyenas have adapted by frequently mobbing lions that enter their home ranges 137 When the lion population in Kenya s Masai Mara National Reserve declined the spotted hyena population increased rapidly 138 Lions tend to dominate cheetahs and leopards steal their kills and kill their cubs and even adults when given the chance 139 Cheetahs often lose their kills to lions or other predators 140 A study in the Serengeti ecosystem revealed that lions killed at least 17 of 125 cheetah cubs born between 1987 and 1990 141 Cheetahs avoid their competitors by hunting at different times and habitats 142 Leopards take refuge in trees but lionesses occasionally attempt to climb up and retrieve their kills 143 Lions similarly dominate African wild dogs taking their kills and slaying pups or adult dogs Population densities of wild dogs are low in areas where lions are more abundant 144 However there are a few reported cases of old and wounded lions falling prey to wild dogs 145 146 Reproduction and life cycle nbsp Lions mating at Masai Mara nbsp A lion cub in Masai Mara Most lionesses reproduce by the time they are four years of age 147 Lions do not mate at a specific time of year and the females are polyestrous 148 Like those of other cats the male lion s penis has spines that point backward During withdrawal of the penis the spines rake the walls of the female s vagina which may cause ovulation 149 150 A lioness may mate with more than one male when she is in heat 151 Lions of both sexes may be involved in group homosexual and courtship activities Males will also head rub and roll around with each other before mounting each other 152 153 Generation length of the lion is about seven years 154 The average gestation period is around 110 days 148 the female gives birth to a litter of between one and four cubs in a secluded den which may be a thicket a reed bed a cave or some other sheltered area usually away from the pride She will often hunt alone while the cubs are still helpless staying relatively close to the den 155 Lion cubs are born blind their eyes opening around seven days after birth They weigh 1 2 2 1 kg 2 6 4 6 lb at birth and are almost helpless beginning to crawl a day or two after birth and walking around three weeks of age 156 To avoid a buildup of scent attracting the attention of predators the lioness moves her cubs to a new den site several times a month carrying them one by one by the nape of the neck 155 Usually the mother does not integrate herself and her cubs back into the pride until the cubs are six to eight weeks old 155 Sometimes the introduction to pride life occurs earlier particularly if other lionesses have given birth at about the same time 96 157 When first introduced to the rest of the pride lion cubs lack confidence when confronted with adults other than their mother They soon begin to immerse themselves in the pride life however playing among themselves or attempting to initiate play with the adults 157 Lionesses with cubs of their own are more likely to be tolerant of another lioness s cubs than lionesses without cubs Male tolerance of the cubs varies one male could patiently let the cubs play with his tail or his mane while another may snarl and bat the cubs away 158 source source source source source Video of a lioness and her cubs in Phinda ReservePride lionesses often synchronise their reproductive cycles and communal rearing and suckling of the young which suckle indiscriminately from any or all of the nursing females in the pride The synchronisation of births is advantageous because the cubs grow to being roughly the same size and have an equal chance of survival and sucklings are not dominated by older cubs 96 157 Weaning occurs after six or seven months Male lions reach maturity at about three years of age and at four to five years are capable of challenging and displacing adult males associated with another pride They begin to age and weaken at between 10 and 15 years of age at the latest 159 When one or more new males oust the previous males associated with a pride the victors often kill any existing young cubs perhaps because females do not become fertile and receptive until their cubs mature or die Females often fiercely defend their cubs from a usurping male but are rarely successful unless a group of three or four mothers within a pride join forces against the male 160 Cubs also die from starvation and abandonment and predation by leopards hyenas and wild dogs Male cubs are excluded from their maternal pride when they reach maturity at around two or three years of age 161 while some females may leave when they reach the age of two 92 When a new male lion takes over a pride adolescents both male and female may be evicted 162 Health and mortality nbsp Lions in a tree near Lake NakuruLions may live 12 17 years 46 Although adult lions have no natural predators evidence suggests most die violently from attacks by humans or other lions 163 Lions often inflict serious injuries on members of other prides they encounter in territorial disputes or members of the home pride when fighting at a kill 164 Crippled lions and cubs may fall victim to hyenas and leopards or be trampled by buffalo or elephants Careless lions may be maimed when hunting prey 165 Nile crocodiles may also kill and eat lions evidenced by the occasional lion claw found in crocodile stomachs 166 Ticks commonly infest the ears neck and groin regions of the lions 167 168 Adult forms of several tapeworm species of the genus Taenia have been isolated from lion intestines having been ingested as larvae in antelope meat 169 Lions in the Ngorongoro Crater were afflicted by an outbreak of stable fly Stomoxys calcitrans in 1962 resulting in lions becoming emaciated and covered in bloody bare patches Lions sought unsuccessfully to evade the biting flies by climbing trees or crawling into hyena burrows many died or migrated and the local population dropped from 70 to 15 individuals 170 A more recent outbreak in 2001 killed six lions 171 Captive lions have been infected with canine distemper virus CDV since at least the mid 1970s 172 CDV is spread by 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 173 Feline immunodeficiency virus and lentivirus also affect captive lions 174 175 Communication nbsp Head rubbing among pride members is a common social behaviour When resting lion socialisation occurs through a number of behaviours the animal s expressive movements are highly developed The most common peaceful tactile gestures are head rubbing and social licking 176 which have been compared with the role of allogrooming among primates 177 Head rubbing nuzzling the forehead face and neck against another lion appears to be a form of greeting 178 and is seen often after an animal has been apart from others or after a fight or confrontation Males tend to rub other males while cubs and females rub females 179 Social licking often occurs in tandem with head rubbing it is generally mutual and the recipient appears to express pleasure The head and neck are the most common parts of the body licked this behaviour may have arisen out of utility because lions cannot lick these areas themselves 180 nbsp Lion roar source source A captive lion roaring Problems playing this file See media help Lions have an array of facial expressions and body postures that serve as visual gestures 181 A common facial expression is the grimace face or flehmen response which a lion makes when sniffing chemical signals and involves an open mouth with bared teeth raised muzzle wrinkled nose closed eyes and relaxed ears 182 Lions also use chemical and visual marking 181 males spray urine 183 and scrape plots of ground and objects within the territory 181 The lion s repertoire of vocalisations is large variations in intensity and pitch appear to be central to communication Most lion vocalisations are variations of growling snarling meowing and roaring Other sounds produced include purring puffing bleating and humming Roaring is used to advertise its presence Lions most often roar at night a sound that can be heard from a distance of 8 kilometres 5 mi 184 They tend to roar in a very characteristic manner starting with a few deep long roars that subside into grunts 185 186 ConservationThe lion is listed as Vulnerable on the IUCN Red List The Indian population is listed on CITES Appendix I and the African population on CITES Appendix II 2 In Africa source source source source Video of a wild lionessSeveral large and well managed protected areas in Africa host large lion populations Where an infrastructure for wildlife tourism has been developed cash revenue for park management and local communities is a strong incentive for lion conservation 2 Most lions now live in East and Southern Africa their numbers are rapidly decreasing and fell by an estimated 30 50 in the late half of the 20th century Primary causes of the decline include disease and human interference 2 In 1975 it was estimated that since the 1950s lion numbers had decreased by half to 200 000 or fewer 187 Estimates of the African lion population range between 16 500 and 47 000 living in the wild in 2002 2004 188 81 In the Republic of the Congo Odzala Kokoua National Park was considered a lion stronghold in the 1990s By 2014 no lions were recorded in the protected area so the population is considered locally extinct 189 The West African lion population is isolated from the one in Central Africa with little or no exchange of breeding individuals In 2015 it was estimated that this population consists of about 400 animals including fewer than 250 mature individuals They persist in three protected areas in the region mostly in one population in the W A P protected area complex shared by Benin Burkina Faso and Niger This population is listed as Critically Endangered 13 Field surveys in the WAP ecosystem revealed that lion occupancy is lowest in the W National Park and higher in areas with permanent staff and thus better protection 190 A population occurs in Cameroon s Waza National Park where between approximately 14 and 21 animals persisted as of 2009 191 In addition 50 to 150 lions are estimated to be present in Burkina Faso s Arly Singou ecosystem 192 In 2015 an adult male lion and a female lion were sighted in Ghana s Mole National Park These were the first sightings of lions in the country in 39 years 193 In the same year a population of up to 200 lions that was previously thought to have been extirpated was filmed in the Alatash National Park Ethiopia close to the Sudanese border 194 195 In 2005 Lion Conservation Strategies were developed for West and Central Africa and or East and Southern Africa The strategies seek to maintain suitable habitat ensure a sufficient wild prey base for lions reduce factors that lead to further fragmentation of populations and make lion human coexistence sustainable 196 197 Lion depredation on livestock is significantly reduced in areas where herders keep livestock in improved enclosures Such measures contribute to mitigating human lion conflict 198 In Asia nbsp A lioness in Gir National ParkThe last refuge of the Asiatic lion population is the 1 412 km2 545 sq mi Gir National Park and surrounding areas in the region of Saurashtra or Kathiawar Peninsula in Gujarat State India The population has risen from approximately 180 lions in 1974 to about 400 in 2010 199 It is geographically isolated which can lead to inbreeding and reduced genetic diversity Since 2008 the Asiatic lion has been listed as Endangered on the IUCN Red List 12 By 2015 the population had grown to 523 individuals inhabiting an area of 7 000 km2 2 700 sq mi in Saurashtra 200 201 202 The Asiatic Lion Census conducted in 2017 recorded about 650 individuals 203 The presence of numerous human habitations close to the National Park results in conflict between lions local people and their livestock 204 200 Some consider the presence of lions a benefit as they keep populations of crop damaging herbivores in check 205 The establishment of a second independent Asiatic lion population in Kuno Wildlife Sanctuary located in Madhya Pradesh was planned but in 2017 the Asiatic Lion Reintroduction Project seemed unlikely to be implemented 206 207 Captive breeding nbsp Two captive male Asiatic lions in Sanjay Gandhi National Park IndiaLions imported to Europe before the middle of the 19th century were possibly foremost Barbary lions from North Africa or Cape lions from Southern Africa 208 Another 11 animals thought to be Barbary lions kept in Addis Ababa Zoo are descendants of animals owned by Emperor Haile Selassie WildLink International in collaboration with Oxford University launched an ambitious International Barbary Lion Project with the aim of identifying and breeding Barbary lions in captivity for eventual reintroduction into a national park in the Atlas Mountains of Morocco 209 However a genetic analysis showed that the captive lions at Addis Ababa Zoo were not Barbary lions but rather closely related to wild lions in Chad and Cameroon 210 In 1982 the Association of Zoos and Aquariums started a Species Survival Plan for the Asiatic lion to increase its chances of survival In 1987 it was found that most lions in North American zoos were hybrids between African and Asiatic lions 211 Breeding programs need to note origins of the participating animals to avoid cross breeding different subspecies and thus reducing their conservation value 212 Captive breeding of lions was halted to eliminate individuals of unknown origin and pedigree Wild born lions were imported to American zoos from Africa between 1989 and 1995 Breeding was continued in 1998 in the frame of an African lion Species Survival Plan 213 About 77 of the captive lions registered in the International Species Information System in 2006 were of unknown origin these animals might have carried genes that are extinct in the wild and may therefore be important to the maintenance of the overall genetic variability of the lion 62 Interactions with humansIn zoos and circuses nbsp Lion at Melbourne Zoo nbsp 19th century etching of a lion tamer in a cage with lions and tigers Lions are part of a group of exotic animals that have been central to zoo exhibits since the late 18th century Although many modern zoos are more selective about their exhibits 214 there are more than 1 000 African and 100 Asiatic lions in zoos and wildlife parks around the world They are considered an ambassador species and are kept for tourism education and conservation purposes 215 Lions can live over twenty years in captivity for example three sibling lions at the Honolulu Zoo lived to the age of 22 in 2007 216 217 The first European zoos spread among noble and royal families in the 13th century and until the 17th century were called seraglios At that time they came to be called menageries an extension of the cabinet of curiosities They spread from France and Italy during the Renaissance to the rest of Europe 218 In England although the seraglio tradition was less developed lions were kept at the Tower of London in a seraglio established by King John in the 13th century 219 220 this was probably stocked with animals from an earlier menagerie started in 1125 by Henry I at his hunting lodge in Woodstock Oxfordshire where according to William of Malmesbury lions had been stocked 221 Lions were kept in cramped and squalid conditions at London Zoo until a larger lion house with roomier cages was built in the 1870s 222 Further changes took place in the early 20th century when Carl Hagenbeck designed enclosures with concrete rocks more open space and a moat instead of bars more closely resembling a natural habitat Hagenbeck designed lion enclosures for both Melbourne Zoo and Sydney s Taronga Zoo although his designs were popular the use of bars and caged enclosures prevailed in many zoos until the 1960s 223 In the late 20th century larger more natural enclosures and the use of wire mesh or laminated glass instead of lowered dens allowed visitors to come closer than ever to the animals some attractions such as the Cat Forest Lion Overlook of Oklahoma City Zoological Park placed the den on ground level higher than visitors 224 Lion taming has been part of both established circuses and individual acts such as Siegfried amp Roy The practice began in the early 19th century by Frenchman Henri Martin and American Isaac Van Amburgh who both toured widely and whose techniques were copied by a number of followers 225 Van Amburgh performed before Queen Victoria in 1838 when he toured Great Britain Martin composed a pantomime titled Les Lions de Mysore the lions of Mysore an idea Amburgh quickly borrowed These acts eclipsed equestrianism acts as the central display of circus shows and entered public consciousness in the early 20th century with cinema In demonstrating the superiority of human over animal lion taming served a purpose similar to animal fights of previous centuries 225 The ultimate proof of a tamer s dominance and control over a lion is demonstrated by the placing of the tamer s head in the lion s mouth The now iconic lion tamer s chair was possibly first used by American Clyde Beatty 1903 1965 226 Hunting and games Main article Lion hunting See also Lion baiting nbsp Bas relief of a wounded lioness from Nineveh c 645 635 BCLion hunting has occurred since ancient times and was often a royal tradition intended to demonstrate the power of the king over nature Such hunts took place in a reserved area in front of an audience The monarch was accompanied by his men and controls were put in place to increase their safety and ease of killing The earliest surviving record of lion hunting is an ancient Egyptian inscription dated circa 1380 BC that mentions Pharaoh Amenhotep III killing 102 lions in ten years with his own arrows The Assyrian emperor Ashurbanipal had one of his lion hunts depicted on a sequence of Assyrian palace reliefs c 640 BC known as the Lion Hunt of Ashurbanipal Lions were also hunted during the Mughal Empire where Emperor Jahangir is said to have excelled at it 227 In Ancient Rome lions were kept by emperors for hunts gladiator fights and executions 228 The Maasai people have traditionally viewed the killing of lions as a rite of passage Historically lions were hunted by individuals however due to reduced lion populations elders discourage solo lion hunts 229 During the European colonisation of Africa in the 19th century the hunting of lions was encouraged because they were considered pests and lion skins were sold for 1 each 230 The widely reproduced imagery of the heroic hunter chasing lions would dominate a large part of the century 231 Trophy hunting of lions in recent years has been met with controversy notably with the killing of Cecil the lion in mid 2015 232 Man eating nbsp The Tsavo maneaters of East Africa on display in the Field Museum of Natural History in ChicagoFurther information Man eater Lions Lions do not usually hunt humans but some usually males seem to seek them out One well publicised case is the Tsavo maneaters in 1898 28 officially recorded railway workers building the Uganda Railway were taken by lions over nine months during the construction of a bridge in Kenya 233 The hunter who killed the lions wrote a book detailing the animals predatory behaviour they were larger than normal and lacked manes and one seemed to suffer from tooth decay The infirmity theory including tooth decay is not favoured by all researchers an analysis of teeth and jaws of man eating lions in museum collections suggests that while tooth decay may explain some incidents prey depletion in human dominated areas is a more likely cause of lion predation on humans 234 Sick or injured animals may be more prone to man eating but the behaviour is not unusual nor necessarily aberrant 235 Lions proclivity for man eating has been systematically examined American and Tanzanian scientists report that man eating behaviour in rural areas of Tanzania increased greatly from 1990 to 2005 At least 563 villagers were attacked and many eaten over this period The incidents occurred near Selous Game Reserve in Rufiji River and in Lindi Region near the Mozambican border While the expansion of villages into bush country is one concern the authors argue conservation policy must mitigate the danger because in this case conservation contributes directly to human deaths Cases in Lindi in which lions seize humans from the centres of substantial villages have been documented 236 Another study of 1 000 people attacked by lions in southern Tanzania between 1988 and 2009 found that the weeks following the full moon when there was less moonlight were a strong indicator of increased night time attacks on people 237 According to Robert R Frump Mozambican refugees regularly crossing Kruger National Park South Africa at night are attacked and eaten by lions park officials have said man eating is a problem there Frump said thousands may have been killed in the decades after apartheid sealed the park and forced refugees to cross the park at night For nearly a century before the border was sealed Mozambicans had regularly crossed the park in daytime with little harm 238 Cultural significanceMain article Cultural depictions of lions nbsp Lions carved on a rock weight Jiroft culture Iran 3rd mil BCThe 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 45 It appeared as a symbol for strength and nobility in cultures across Europe Asia and Africa despite incidents of attacks on people The lion has been depicted as king of the jungle and king of beasts and thus became a popular symbol for royalty and stateliness 239 The lion is also used as a symbol of sporting teams 240 Africa nbsp Granite statue of the Egyptian goddess Sekhmet from the Luxor Temple dated 1403 1365 BC exhibited in the National Museum of DenmarkIn sub Saharan Africa the lion has been a common character in stories proverbs and dances but rarely featured in visual arts 241 In some cultures the lion symbolises power and royalty 242 In the Swahili language the lion is known as simba which also means aggressive king and strong 54 Some rulers had the word lion in their nickname Sundiata Keita of the Mali Empire was called Lion of Mali 243 The founder of the Waalo kingdom is said to have been raised by lions and returned to his people part lion to unite them using the knowledge he learned from the lions 242 In parts of West Africa lions symbolised the top class of their social hierarchies 242 In more heavily forested areas where lions were rare the leopard represented the top of the hierarchy 241 In parts of West and East Africa the lion is associated with healing and provides the connection between seers and the supernatural In other East African traditions the lion represents laziness 242 In much of African folklore the lion is portrayed as having low intelligence and is easily tricked by other animals 243 The ancient Egyptians portrayed several of their war deities as lionesses which they revered as fierce hunters Egyptian deities associated with lions include Sekhmet Bast Mafdet Menhit Pakhet and Tefnut 239 These deities were often connected with the sun god Ra and his fierce heat and their dangerous power was invoked to guard people or sacred places The sphinx a figure with a lion s body and the head of a human or other creature represented a pharaoh or deity who had taken on this protective role 244 Asia nbsp Roaring and striding lion from the Throne Room of Nebuchadnezzar II 6th century BC from Babylon IraqThe lion was a prominent symbol in ancient Mesopotamia from Sumer up to Assyrian and Babylonian times where it was strongly associated with kingship 245 Lions were among the major symbols of the goddess Inanna Ishtar 246 247 The Lion of Babylon was the foremost symbol of the Babylonian Empire 248 The Lion of Judah is the biblical emblem of the tribe of Judah and the later Kingdom of Judah 249 Lions are frequently mentioned in the Bible notably in the Book of Daniel in which the eponymous hero refuses to worship King Darius and is forced to sleep in the lions den where he is miraculously unharmed Dan 6 In the Book of Judges Samson kills a lion as he travels to visit a Philistine woman Judg 14 250 Indo Persian chroniclers regarded the lion as keeper of order in the realm of animals The Sanskrit word mrigendra signifies a lion as king of animals in general or deer in particular 251 Narasimha the man lion is one of ten avatars of the Hindu god Vishnu 252 Singh is an ancient Indian vedic name meaning lion dating back over 2 000 years It was originally used only by Rajputs a Hindu Kshatriya or military caste but is used by millions of Hindu Rajputs and more than twenty million Sikhs today 253 The Lion Capital of Ashoka erected by Emperor Ashoka in the 3rd century CE depicts four lions standing back to back It was made the National Emblem of India in 1950 254 The lion is also symbolic for the Sinhalese people the term derived from the Sanskrit Sinhala meaning of lions 255 while a sword wielding lion is the central figure on the national flag of Sri Lanka 256 The lion is a common motif in Chinese art it was first used in art during the late Spring and Autumn period fifth or sixth century BC and became more popular during the Han dynasty 206 BC AD 220 when imperial guardian lions started to be placed in front of imperial palaces for protection Because lions have never been native to China early depictions were somewhat unrealistic after the introduction of Buddhist art to China in the Tang dynasty after the sixth century AD lions were usually depicted wingless with shorter thicker bodies and curly manes 257 The lion dance is a traditional dance in Chinese culture in which performers in lion costumes mimic a lion s movements often with musical accompaniment from cymbals drums and gongs They are performed at Chinese New Year the August Moon Festival and other celebratory occasions for good luck 258 Western world nbsp Dorothy Gale meets the Cowardly Lion in The Wonderful Wizard of Oz Art by W W Denslow 1900 Lion headed figures and amulets were excavated in tombs in the Greek islands of Crete Euboea Rhodes Paros and Chios They are associated with the Egyptian deity Sekhmet and date to the early Iron Age between the 9th and 6th centuries BC 259 Tfhe lion is featured in several of Aesop s fables notably The Lion and the Mouse 260 The Nemean lion was symbolic in ancient Greece and Rome represented as the constellation and zodiac sign Leo and described in mythology where it was killed and worn by the hero Heracles 261 symbolising victory over death 262 Lancelot and Gawain were also heroes slaying lions in the Middle Ages In some medieval stories lions were portrayed as allies and companions 263 Lion was the nickname of several medieval warrior rulers with a reputation for bravery such as Richard the Lionheart 239 Lions continue to appear in modern literature as characters including the messianic Aslan in the 1950 novel The Lion the Witch and the Wardrobe and The Chronicles of Narnia series by C S Lewis 264 and the comedic Cowardly Lion in L Frank Baum s 1900 The Wonderful Wizard of Oz 265 Lion symbolism was used from the advent of cinema one of the most iconic and widely recognised lions is Leo which has been the mascot for Metro Goldwyn Mayer studios since the 1920s 266 The 1966 film Born Free features Elsa the lioness and is based on the book Born Free published in 1960 267 The lion s role as king of the beasts has been used in the 1994 Disney animated feature film The Lion King 268 Lions are frequently depicted on coats of arms like on the coat of arms of Finland 269 either as a device on shields or as supporters but the lioness is used much less frequently 270 The heraldic lion is particularly common in British arms It is traditionally depicted in a great variety of attitudes although within French heraldry only lions rampant are considered to be lions felined figures in any other position are instead referred to as leopards 271 See alsoList of largest cats Mapogo lion coalition Roar film from 1981Notes Populations of India are listed in Appendix I ReferencesCitations a b c d e Wozencraft W C 2005 Species Panthera leo In Wilson D E Reeder D M eds Mammal Species of the World A Taxonomic and Geographic Reference 3rd ed Johns Hopkins University Press p 546 ISBN 978 0 8018 8221 0 OCLC 62265494 a b c d e f Bauer H Packer C Funston P F Henschel P amp Nowell K 2017 errata version of 2016 assessment Panthera leo IUCN Red List of Threatened Species 2016 e T15951A115130419 doi 10 2305 IUCN UK 2016 3 RLTS T15951A107265605 en Retrieved 15 January 2022 a b Linnaeus C 1758 Felis leo Caroli Linnaei Systema naturae per regna tria naturae secundum classes ordines genera species cum characteribus differentiis synonymis locis Vol Tomus I decima reformata ed Holmiae Laurentius Salvius p 41 in Latin lion Oxford English Dictionary Online ed Oxford University Press Retrieved 20 March 2022 Subscription or participating institution membership required Liddell H G amp Scott R 1940 pan8hr A Greek English Lexicon Revised and augmented ed Oxford Clarendon Press a b c Johnson W E Eizirik E Pecon Slattery J Murphy W J Antunes A Teeling E amp O Brien S J 2006 The late miocene radiation of modern Felidae A genetic assessment Science 311 5757 73 77 Bibcode 2006Sci 311 73J doi 10 1126 science 1122277 PMID 16400146 S2CID 41672825 Werdelin L Yamaguchi N Johnson W E amp O Brien S J 2010 Phylogeny and evolution of cats Felidae Biology and Conservation of Wild Felids 59 82 a b Davis B W Li G amp Murphy W J 2010 Supermatrix and species tree methods resolve phylogenetic relationships within the big cats Panthera Carnivora Felidae Molecular Phylogenetics and Evolution 56 1 64 76 doi 10 1016 j ympev 2010 01 036 PMID 20138224 a b Mazak J H Christiansen P Kitchener A C amp Goswami A 2011 Oldest known pantherine skull and evolution of the tiger PLOS ONE 6 10 e25483 Bibcode 2011PLoSO 625483M doi 10 1371 journal pone 0025483 PMC 3189913 PMID 22016768 Oken L 1816 1 Art Panthera Lehrbuch der Zoologie 2 Abtheilung Jena August Schmid amp Comp p 1052 a b c Hemmer H 1974 Untersuchungen zur Stammesgeschichte der Pantherkatzen Pantherinae Teil 3 Zur Artgeschichte des Lowen Panthera Panthera leo Linnaeus 1758 Veroffentlichungen der Zoologischen Staatssammlung 17 167 280 a b c Breitenmoser U Mallon D P Ahmad Khan J and Driscoll C 2008 Panthera leo ssp persica IUCN Red List of Threatened Species 2008 e T15952A5327221 doi 10 2305 IUCN UK 2008 RLTS T15952A5327221 en a b Henschel P Bauer H Sogbohoussou E amp Nowell K 2015 Panthera leo West Africa subpopulation IUCN Red List of Threatened Species 2015 doi 10 2305 IUCN UK 2015 2 RLTS T68933833A54067639 en a b c Kitchener A C Breitenmoser Wursten C Eizirik E Gentry A Werdelin L Wilting A Yamaguchi N Abramov A V Christiansen P Driscoll C Duckworth J W Johnson W Luo S J Meijaard E O Donoghue P Sanderson J Seymour K Bruford M Groves C Hoffmann M Nowell K Timmons Z amp Tobe S 2017 A revised taxonomy of the Felidae The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group PDF Cat News Special Issue 11 71 73 Wood J G 1865 Felidae or the Cat Tribe The Illustrated Natural History Mammalia Volume 1 London Routledge p 129 148 a b Hunter L Barrett P 2018 Lion Panthera leo The Field Guide to Carnivores of the World 2 ed London Oxford New York New Delhi Sydney Bloomsbury pp 46 47 ISBN 978 1 4729 5080 2 de Manuel M Barnett R Sandoval Velasco M Yamaguchi N Vieira F G Mendoza M L Z Liu S Martin M D Sinding M S S Mak S S T Caroe C Liu S Guo C Zheng J Zazula G Baryshnikov G Eizirik E Koepfli K P Johnson W E Antunes A Sicheritz Ponten T Gopalakrishnan S Larson G Yang H O Brien S J Hansen A J Zhang G Marques Bonet T Gilbert M T P 2020 The evolutionary history of extinct and living lions Proceedings of the National Academy of Sciences 117 20 10927 10934 Bibcode 2020PNAS 11710927D doi 10 1073 pnas 1919423117 PMC 7245068 PMID 32366643 Bertola L D Jongbloed H Van Der Gaag K J De Knijff P Yamaguchi N Hooghiemstra H Bauer H Henschel P White P A Driscoll C A Tende T Ottosson U Saidu Y Vrieling K de Iongh H H 2016 Phylogeographic patterns in Africa and High Resolution Delineation of genetic clades in the Lion Panthera leo Scientific Reports 6 30807 Bibcode 2016NatSR 630807B doi 10 1038 srep30807 PMC 4973251 PMID 27488946 a b c d Manuel M d Ross B Sandoval Velasco M Yamaguchi N Vieira F G Mendoza M L Z Liu S Martin M D Sinding M H S Mak S S T Caroe C Liu S Guo C Zheng J Zazula G Baryshnikov G Eizirik E Koepfli K P Johnson W E Antunes A Sicheritz Ponten T Gopalakrishnan S Larson G Yang H O Brien S J Hansen A J Zhang G Marques Bonet T amp Gilbert M T P 2020 The evolutionary history of extinct and living lions Proceedings of the National Academy of Sciences of the United States of America 117 20 10927 10934 Bibcode 2020PNAS 11710927D doi 10 1073 pnas 1919423117 PMC 7245068 PMID 32366643 Christiansen P 2008 Phylogeny of the great cats Felidae Pantherinae and the influence of fossil taxa and missing characters Cladistics 24 6 977 992 doi 10 1111 j 1096 0031 2008 00226 x PMID 34892880 S2CID 84497516 Manamendra Arachchi K Pethiyagoda R Dissanayake R amp Meegaskumbura M 2005 A second extinct big cat from the late Quaternary of Sri Lanka PDF The Raffles Bulletin of Zoology Supplement 12 423 434 Archived from the original PDF on 7 August 2007 Marciszak A amp Stefaniak K 2010 Two forms of cave lion Middle Pleistocene Panthera spelaea fossilis Reichenau 1906 and Upper Pleistocene Panthera spelaea spelaea Goldfuss 1810 from the Bisnik Cave Poland Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen 258 3 339 351 doi 10 1127 0077 7749 2010 0117 Sabol M 2014 Panthera fossilis Reichenau 1906 Felidae Carnivora from Za Hajovnou Cave Moravia The Czech Republic A Fossil Record from 1987 2007 Acta Musei Nationalis Pragae Series B Historia Naturalis 70 1 2 59 70 doi 10 14446 AMNP 2014 59 Stuart A J amp Lister A M 2011 Extinction chronology of the cave lion Panthera spelaea Quaternary Science Reviews 30 17 2329 40 Bibcode 2011QSRv 30 2329S doi 10 1016 j quascirev 2010 04 023 Hemmer H 2011 The story of the cave lion Panthera Leo Spelaea Goldfuss 1810 A review Quaternaire 4 201 208 a b Barnett R Mendoza M L Z Soares A E R Ho S Y W Zazula G Yamaguchi N Shapiro B Kirillova I V Larson G amp Gilbert M T P 2016 Mitogenomics of the Extinct Cave Lion Panthera spelaea Goldfuss 1810 resolve its position within the Panthera cats Open Quaternary 2 4 doi 10 5334 oq 24 hdl 10576 22920 a b Burger J Rosendahl W Loreille O Hemmer H Eriksson T Gotherstrom A Hiller J Collins M J Wess T amp Alt K W 2004 Molecular phylogeny of the extinct cave lion Panthera leo spelaea PDF Molecular Phylogenetics and Evolution 30 3 841 849 doi 10 1016 j ympev 2003 07 020 PMID 15012963 Archived from the original PDF on 25 September 2007 Packer C amp Clottes J 2000 When Lions Ruled France PDF Natural History 109 9 52 57 Archived PDF from the original on 29 December 2015 Chimento N R amp Agnolin F L 2017 The fossil American lion Panthera atrox in South America Palaeobiogeographical implications Comptes Rendus Palevol 16 8 850 864 Bibcode 2017CRPal 16 850C doi 10 1016 j crpv 2017 06 009 Harington C R 1969 Pleistocene remains of the lion like cat Panthera atrox from the Yukon Territory and northern Alaska Canadian Journal of Earth Sciences 6 5 1277 1288 Bibcode 1969CaJES 6 1277H doi 10 1139 e69 127 Christiansen P amp Harris J M 2009 Craniomandibular morphology and phylogenetic affinities of Panthera atrox implications for the evolution and paleobiology of the lion lineage Journal of Vertebrate Paleontology 29 3 934 945 Bibcode 2009JVPal 29 934C doi 10 1671 039 029 0314 S2CID 85975640 King L M amp Wallace S C 2014 Phylogenetics of Panthera including Panthera atrox based on craniodental characters Historical Biology 26 6 827 833 Bibcode 2014HBio 26 827K doi 10 1080 08912963 2013 861462 S2CID 84229141 a b Werdelin L Yamaguchi N Johnson W E amp O Brien S J 2010 Phylogeny and evolution of cats Felidae In Macdonald D W amp Loveridge A J eds Biology and Conservation of Wild Felids Oxford UK Oxford University Press pp 59 82 ISBN 978 0 19 923445 5 a b Li G Davis B W Eizirik E amp Murphy W J 2016 Phylogenomic evidence for ancient hybridization in the genomes of living cats Felidae Genome Research 26 1 1 11 doi 10 1101 gr 186668 114 PMC 4691742 PMID 26518481 Tseng Z J Wang X Slater G J Takeuchi G T Li Q Liu J amp Xie G 2014 Himalayan fossils of the oldest known pantherine establish ancient origin of big cats Proceedings of the Royal Society B Biological Sciences 281 1774 20132686 doi 10 1098 rspb 2013 2686 PMC 3843846 PMID 24225466 Barnett R Shapiro B Barnes I Ho S Y W Burger J Yamaguchi N Higham T F G Wheeler H T Rosendahl W Sher A V Sotnikova M Kuznetsova T Baryshnikov G F Martin L D Harington C R Burns J A amp Cooper A 2009 Phylogeography of lions Panthera leo ssp reveals three distinct taxa and a late Pleistocene reduction in genetic diversity PDF Molecular Ecology 18 8 1668 1677 Bibcode 2009MolEc 18 1668B doi 10 1111 j 1365 294X 2009 04134 x PMID 19302360 S2CID 46716748 Archived PDF from the original on 8 August 2017 Argant A amp Brugal J P 2017 The cave lion Panthera Leo spelaea and its evolution Panthera spelaea intermedia nov subspecies Acta Zoologica Cracoviensia 60 2 58 103 doi 10 3409 azc 60 2 59 a b Barnett R Yamaguchi N Shapiro B Ho S Y Barnes I Sabin R Werdelin L Cuisin J amp Larson G 2014 Revealing the maternal demographic history of Panthera leo using ancient DNA and a spatially explicit 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Hybrid Dysgenesis PhD Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology Uppsala Acta Universitatis Upsaliensis p 8 10 Archived PDF from the original on 18 May 2019 Rafferty J P 2011 The Liger Carnivores Meat eating Mammals New York The Rosen Publishing Group p 120 ISBN 978 1 61530 340 3 Retrieved 4 July 2014 Zhang Z Chen J Li L Tao M Zhang C Qin Q Xiao J Liu Y Liu S 2014 Research advances in animal distant hybridization PDF Science China Life Sciences 57 9 889 902 doi 10 1007 s11427 014 4707 1 PMID 25091377 S2CID 18179301 Archived PDF from the original on 30 October 2018 a b c d e Guggisberg C A W 1975 Lion Panthera leo Linnaeus 1758 Wild Cats of the World New York Taplinger Publishing pp 138 179 ISBN 978 0 8008 8324 9 a b c d e f Haas S K Hayssen V Krausman P R 2005 Panthera leo PDF Mammalian Species 762 1 11 doi 10 1644 1545 1410 2005 762 0001 PL 2 0 CO 2 S2CID 198968757 Archived from the original PDF on 28 July 2017 Schaller pp 28 30 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Wikisource has the text of the 1911 Encyclopaedia Britannica article Lion nbsp Media related to Lion at Wikimedia Commons IUCN SSC Cat Specialist Group Lion Panthera leo Lion Conservation Fund The Portugal News 2014 Rare desert lion killed in Angola after supplying unprecedented data Archived from the original on 2 August 2018 Retrieved 24 May 2018 Lion Collier s New Encyclopedia 1921 Portals nbsp Cats nbsp Mammals nbsp Animals nbsp Biology nbsp Africa nbsp India Retrieved from https en wikipedia org w index php title Lion amp oldid 1204049338, wikipedia, wiki, book, books, library,

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