Lion
Lion Temporal range:
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Male in Okonjima, Namibia | |
Female (lioness) in Okonjima | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Carnivora |
Suborder: | Feliformia |
Family: | Felidae |
Subfamily: | Pantherinae |
Genus: | Panthera |
Species: | P. leo[1]
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Binomial name | |
Panthera leo[1] | |
Subspecies | |
| |
Historical and present distribution of the lion in Africa, Asia and Europe |
The lion (Panthera leo) is a large
The lion inhabits
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 literature and films. Lions have been kept in
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
Felis leo was the
Subspecies
In the 19th and 20th centuries, several lion
- 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
Other lion subspecies or
- 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]
- 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]
Evolution
The Panthera
Estimates for the divergence time of the modern and cave lion lineages range from 529,000 to 392,000 years ago based on
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
Hybrids
In zoos, lions have been bred with
Description
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
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
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. 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
The white lion is a rare
Distribution and habitat
African lions live in scattered populations across sub-Saharan Africa. The lion prefers grassy plains and
Historical range
In Africa, the range of the lion originally spanned most of the central
In southern Europe and Asia, the lion once ranged in regions where climatic conditions supported an abundance of prey.
Behaviour and ecology
Lions spend much of their time resting; they are inactive for about twenty hours per day.[89] 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.[90]
Group organisation
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 "
Nomadic lions range widely and move around sporadically, either in pairs or alone.
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).[102][103] The area occupied by a pride is called a "pride area" whereas that occupied by a nomad is a "range".[91] 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.[104] Lions tend to assume specific roles in the pride; slower-moving individuals may provide other valuable services to the group.[105] 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.[106] The male or males associated with the pride must defend their relationship with the pride from outside males who may attempt to usurp them.[97] Dominance hierarchies do not appear to exist among individuals of either sex in a pride.[107]
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.[108] 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.[109]
Hunting and diet
The lion is a generalist hypercarnivore and is considered to be both an apex and keystone predator due to its wide prey spectrum.[110][111] Its prey consists mainly of ungulates, particularly blue wildebeest, plains zebra, African buffalo, common warthog, gemsbok and giraffe.[112] In India, chital and sambar deer are the most common wild prey,[46][112][113] while livestock contributes significantly to lion kills outside protected areas.[114] They usually avoid fully grown adult elephants, rhinoceroses and hippopotamus and small prey like dik-dik, hyraxes, hares and monkeys.[112][115] Unusual prey include porcupines and small reptiles. Lions kill other predators but seldom consume them.[116]
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.[117] Single lions are capable of bringing down zebra and wildebeest, while larger prey like buffalo and giraffe are riskier.[97] 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.[118][119] 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.[120] Some evidence suggests, however, that males are just as successful as females; they are typically solo hunters who ambush prey in small bushland.[121] 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.[122]
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.[123] 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.[124] One study in 2018 recorded a lion running at a top speed of 74.1 km/h (46.0 mph).[125] They take advantage of factors that reduce visibility; many kills take place near some form of cover or at night.[126] 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.[127] It has a bite force of 1314.7 Newton at the canine tip and 2023.7 Newton at the carnassial notch.[128]
Lions typically consume prey at the location of the hunt but sometimes drag large prey into cover.[129] 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.[97] Large kills are shared more widely among pride members.[130] An adult lioness requires an average of about 5 kg (11 lb) of meat per day while males require about 7 kg (15 lb).[131] Lions gorge themselves and eat up to 30 kg (66 lb) in one session.[87] 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.[129] Lions defend their kills from scavengers such as vultures and hyenas.[97]
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.[132] 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.[133]
Predatory competition
Lions and
Lions tend to dominate cheetahs and leopards, steal their kills and kill their cubs and even adults when given the chance.[141] Cheetahs often lose their kills to lions or other predators.[142] A study in the Serengeti ecosystem revealed that lions killed at least 17 of 125 cheetah cubs born between 1987 and 1990.[143] Cheetahs avoid their competitors by hunting at different times and habitats.[144] Leopards, by contrast, do not appear to be motivated by an avoidance of lions, as they use heavy vegetation regardless of whether lions are present in an area and both cats are active around the same time of day. In addition, there is no evidence that lions effect leopard abundance.[145] Leopards take refuge in trees, though lionesses occasionally attempt to climb up and retrieve their kills.[146]
Lions similarly dominate African wild dogs, taking their kills and dispatching pups or adult dogs. Population densities of wild dogs are low in areas where lions are more abundant.[147] However, there are a few reported cases of old and wounded lions falling prey to wild dogs.[148][149]
Reproduction and life cycle
Most lionesses reproduce by the time they are four years of age.
Usually, the mother does not integrate herself and her cubs back into the pride until the cubs are six to eight weeks old.[158] Sometimes the introduction to pride life occurs earlier, particularly if other lionesses have given birth at about the same time.[97][160] 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.[160] 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.[161]
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.[97][160] 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.[162]
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.[163] 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,[164] while some females may leave when they reach the age of two.[93] When a new male lion takes over a pride, adolescents both male and female may be evicted.[165]
Health and mortality
Lions may live 12–17 years in the wild.[46] Although adult lions have no natural predators, evidence suggests most die violently from attacks by humans or other lions.[166] 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.[167] 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.[168] Nile crocodiles may also kill and eat lions, evidenced by the occasional lion claw found in crocodile stomachs.[169]
Captive lions have been infected with canine distemper virus (CDV) since at least the mid-1970s.[175] 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.[176] Feline immunodeficiency virus and lentivirus also affect captive lions.[177][178]
Communication
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
Lions have an array of facial expressions and body postures that serve as visual gestures.[184] 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.[185] Lions also use chemical and visual marking;[184] males spray urine[186] and scrape plots of ground and objects within the territory.[184]
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
Conservation
The lion is listed as
In Africa
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.[190] Estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004.[191][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.[192] 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.[193]
A population occurs in Cameroon's
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.[199][200] Lion depredation on livestock is significantly reduced in areas where herders keep livestock in improved enclosures. Such measures contribute to mitigating human–lion conflict.[201]
In Asia
The last refuge of the Asiatic lion population is the 1,412 km2 (545 sq mi) Gir National Park and surrounding areas in the
The presence of numerous human settlements close to Gir National Park resulted in conflict between lions, local people and their livestock.[207][203] Some consider the presence of lions a benefit, as they keep populations of crop damaging herbivores in check.[208]
Captive breeding
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.
In 1982, the
About 77% of the captive lions registered in the
Interactions with humans
In zoos and circuses
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,[215] 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.[216] 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.[217][218]
The first European "zoos" spread among noble and royal families in the 13th century, and until the 17th century were called
Lions were kept in cramped and squalid conditions at London Zoo until a larger lion house with roomier cages was built in the 1870s.[223] 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.[224] 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.[225]
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.[226] 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.[226] 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).[227]
Hunting and games
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
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.[230] 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.[231] The widely reproduced imagery of the heroic hunter chasing lions would dominate a large part of the century.[232] Trophy hunting of lions in recent years has been met with controversy, notably with the killing of Cecil the lion in mid-2015.[233]
Man-eating
Lions do not usually hunt humans but some (usually males) seem to seek them out. One well-publicised case is the
Lions' proclivity for
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.[239]
Cultural significance
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.
In sub-Saharan Africa, the lion has been a common character in stories, proverbs and dances, but rarely featured in visual arts.
The lion was a prominent symbol in ancient Mesopotamia from Sumer up to Assyrian and Babylonian times, where it was strongly associated with kingship.[247] The big cat was a symbol and steed of fertility goddess Inanna.[240] Lions decorate the Processional Way leading to the Ishtar Gate in Babylon which was built by the Nebuchadnezzar II in the 6th century BCE. The Lion of Babylon symbolized the power of the king and protector of the land against enemies, but was also invoked for good luck.[248] The constellation Leo the lion was first recognised by the Sumerians around 4,000 years ago and is considered to be the fifth sign of the zodiac.[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). The power and ferocity of the lion is invoked when describing the anger of God (Amos 3:4–8, Lam 3:10) and the menace of Israel's enemies (Psalm 17:12, Jer 2:30) and Satan (1 Pet 5:8). The book of Isaiah uses the imagery of a lion laying with a calf and child, and eating straw to portray the harmony of creation (Isa 11:6–7). In the Book of Revelation, a lion, an ox, a man and an eagle are seen on a heavenly throne in John's vision;(Rev 4:7).[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] In India, the Lion Capital of Ashoka, erected by Emperor Ashoka in the 3rd century CE, depicts four lions standing back to back. In Hindu mythology, the half-lion Narasimha, an avatar of the deity Vishnu, battles and slays the evil ruler Hiranyakashipu. Though they were never native to the country, lions have played important roles in Chinese culture. Statues of the beast have guarded the entrances to the imperial palace and many religious shrines. The lion dance has been performed in China and beyond for over a thousand years. In Buddhist art, lions are associated with both arhats and bodhisattvas and may be ridden by the Manjushri.[252]
In
See also
Notes
- ^ 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.
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External links
- 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.
- Collier's New Encyclopedia. 1921. .