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The lion (''Panthera leo'') is a large cat of the genus ''
Panthera ''Panthera'' is a genus within the family (biology), family Felidae that was named and described by Lorenz Oken in 1816 who placed all the spotted cats in this group. Reginald Innes Pocock revised the classification of this genus in 1916 as co ...
'' native to Africa and India. It has a muscular, broad-chested body; 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 Sociality is the degree to which individuals in an animal population tend to associate in social groups (gregariousness) and form cooperative societies. Sociality is a survival response to evolutionary pressures. For example, when a mother wasp ...
, 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 don't actively seek out and prey on humans. The lion inhabits grasslands, savannas and
shrubland Shrubland, scrubland, scrub, brush, or bush is a plant community characterized by vegetation dominated by shrubs, often also including grasses, herbs, and geophytes. Shrubland may either occur naturally or be the result of human activity. It m ...
s. 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 Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the List of sov ...
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
menagerie A menagerie is a collection of captive animals, frequently exotic, kept for display; or the place where such a collection is kept, a precursor to the modern Zoo, zoological garden. The term was first used in 17th-century France, in reference to ...
s since the time of the Roman Empire and have been a key species sought for exhibition in
zoological garden A zoo (short for zoological garden; also called an animal park or menagerie) is a facility in which animals are kept within enclosures for public exhibition and often bred for conservation purposes. The term ''zoological garden'' refers to zool ...
s across the world since the late 18th century.
Cultural depictions of lions Cultural depictions of lions are known in countries of Afro-Eurasia. The lion has been an important symbol to humans for tens of thousands of years. The earliest graphic representations feature lions as organized hunters with great strength, str ...
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 Anglo-Norman may refer to: *Anglo-Normans, the medieval ruling class in England following the Norman conquest of 1066 *Anglo-Norman language **Anglo-Norman literature *Anglo-Norman England, or Norman England, the period in English history from 1066 ...
from Latin (nominative: ), which in turn was a borrowing from Ancient Greek . The Hebrew word may also be related. The generic name ''Panthera'' is traceable to the classical Latin word 'panthēra' and the ancient Greek word πάνθηρ 'panther'.


Taxonomy

''Felis leo'' was the
scientific name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
used by Carl Linnaeus in 1758, who described the lion in his work ''
Systema Naturae ' (originally in Latin written ' with the ligature æ) is one of the major works of the Swedish botanist, zoologist and physician Carl Linnaeus (1707–1778) and introduced the Linnaean taxonomy. Although the system, now known as binomial nomen ...
''. The genus name ''Panthera'' was coined by Lorenz Oken in 1816. Between the mid-18th and mid-20th centuries, 26 lion
specimen Specimen may refer to: Science and technology * Sample (material), a limited quantity of something which is intended to be similar to and represent a larger amount * Biological specimen or biospecimen, an organic specimen held by a biorepository ...
s were described and proposed as subspecies, of which 11 were recognised as valid in 2005. They were distinguished mostly by the size and colour of their manes and skins.


Subspecies

In the 19th and 20th centuries, several lion type specimens were described and proposed as
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
, with about a dozen recognised as valid taxa until 2017. 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. 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: * ''P. l. leo'' − the nominate lion subspecies includes the Asiatic lion, the
regionally extinct In geography, regions, otherwise referred to as zones, lands or territories, are areas that are broadly divided by physical characteristics (physical geography), human impact characteristics (human geography), and the interaction of humanity and t ...
Barbary lion, and lion populations in West and northern parts of Central Africa.
Synonyms A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
include ''P. l. persica'' , ''P. l. senegalensis'' , ''P. l. kamptzi'' , and ''P. l. azandica'' . Multiple authors referred to it as 'northern lion' and 'northern subspecies'. * ''P. l. melanochaita'' − includes the extinct Cape lion and lion populations in East and Southern African regions. Synonyms include ''P. l. somaliensis'' , ''P. l. massaica'' , ''P. l. sabakiensis'' , ''P. l. bleyenberghi'' , ''P. l. roosevelti'' , ''P. l. nyanzae'' , ''P. l. hollisteri'' , ''P. l. krugeri'' , ''P. l. vernayi'' , and ''P. l. webbiensis'' . It has been referred to as 'southern subspecies' and 'southern lion'. 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''. 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. 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.


Fossil records

Other lion subspecies or sister species to the modern lion existed in prehistoric times: *'' P. l. sinhaleyus'' was a fossil carnassial excavated in
Sri Lanka Sri Lanka (, ; si, ශ්‍රී ලංකා, Śrī Laṅkā, translit-std=ISO (); ta, இலங்கை, Ilaṅkai, translit-std=ISO ()), formerly known as Ceylon and officially the Democratic Socialist Republic of Sri Lanka, is an ...
, which was attributed to a lion. It is thought to have become extinct around 39,000 years ago. * ''P. leo 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. *'' P. spelaea'', or the cave lion, lived in Eurasia and
Beringia Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 72 degrees north latitude in the Chukchi Sea; and on the south by the tip ...
during the
Late Pleistocene The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division of ...
. It became extinct due to climate warming or human expansion latest by 11,900 years ago. Bone fragments excavated in European, North Asian, Canadian and Alaskan caves indicate that it ranged from Europe across Siberia into western Alaska. It likely derived from ''P. fossilis'', and was genetically isolated and highly distinct from the modern lion in Africa and Eurasia. It is depicted in
Paleolithic The Paleolithic or Palaeolithic (), also called the Old Stone Age (from Greek: παλαιός ''palaios'', "old" and λίθος ''lithos'', "stone"), is a period in human prehistory that is distinguished by the original development of stone too ...
cave paintings, ivory carvings, and clay busts. * ''P. atrox'', or the American lion, ranged in the Americas from Canada to possibly Patagonia. It arose when a cave lion population in
Beringia Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 72 degrees north latitude in the Chukchi Sea; and on the south by the tip ...
became isolated south of the Cordilleran Ice Sheet about 370,000 years ago. A fossil from Edmonton dates to 11,355 ± 55 years ago.


Evolution

The ''Panthera''
lineage Lineage may refer to: Science * Lineage (anthropology), a group that can demonstrate its common descent from an apical ancestor or a direct line of descent from an ancestor * Lineage (evolution), a temporal sequence of individuals, populati ...
is estimated to have genetically diverged from the common ancestor of the
Felidae Felidae () is the family of mammals in the order Carnivora colloquially referred to as cats, and constitutes a clade. A member of this family is also called a felid (). The term "cat" refers both to felids in general and specifically to the ...
around to , and the geographic origin of the genus is most likely northern Central Asia. Results of analyses differ in the phylogenetic relationship of the lion; it was thought to form a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
with the
jaguar The jaguar (''Panthera onca'') is a large cat species and the only living member of the genus '' Panthera'' native to the Americas. With a body length of up to and a weight of up to , it is the largest cat species in the Americas and the th ...
(''P. onca'') that diverged , but also with the
leopard The leopard (''Panthera pardus'') is one of the five extant species in the genus '' Panthera'', a member of the cat family, Felidae. It occurs in a wide range in sub-Saharan Africa, in some parts of Western and Central Asia, Southern Russia, a ...
(''P. pardus'') that diverged to . Hybridisation between lion and snow leopard (''P. uncia'') ancestors possibly continued until about 2.1 million years ago. The lion-leopard clade was distributed in the Asian and African
Palearctic The Palearctic or Palaearctic is the largest of the eight biogeographic realms of the Earth. It stretches across all of Eurasia north of the foothills of the Himalayas, and North Africa. The realm consists of several bioregions: the Euro-Sibe ...
since at least the early Pliocene. The earliest fossils recognisable as lions were found at
Olduvai Gorge The Olduvai Gorge or Oldupai Gorge in Tanzania is one of the most important paleoanthropology, paleoanthropological localities in the world; the many sites exposed by the gorge have proven invaluable in furthering understanding of early human ev ...
in Tanzania and are estimated to be up to 2 million years old. 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 In population genetics, gene flow (also known as gene migration or geneflow and allele flow) is the transfer of genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent a ...
between the two lineages, indicating that they did not share the same geographic area. The Eurasian and American cave lions became extinct at the end of the last glacial period without
mitochondrial A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is use ...
descendants on other continents. 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. They shared a common ancestor probably between 98,000 and 52,000 years ago. 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. 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. 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.


Hybrids

In zoos, lions have been bred with tigers to create hybrids for the curiosity of visitors or for scientific purpose. 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. The leopon is a hybrid between a lion and leopard.


Description

The lion is a muscular, broad-chested cat with a short, rounded head, a reduced neck and round ears. Its 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 Spot or SPOT may refer to: Places * Spot, North Carolina, a community in the United States * The Spot, New South Wales, a locality in Sydney, Australia * South Pole Traverse, sometimes called the South Pole Overland Traverse People * Spot (produ ...
, which fade as the cub reaches adulthood, although faint spots often may still be seen on the legs and underparts. The lion is the only member of the cat family that displays obvious sexual dimorphism. Males have broader heads and a prominent mane that 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. The tail of all lions ends in a dark, hairy tuft that in some lions conceals an approximately -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 months of age. It is readily identifiable by the age of seven months. Schaller, pp. 28–30. Of the living felid species, the lion is rivaled only by the tiger in length, weight, and height at the shoulder. 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. Skeletal muscles of the lion make up 58.8% of its body weight and represents the highest percentage of muscles among mammals.


Size

The size and weight of adult lions varies across global range and habitats. Accounts of a few individuals that were larger than average exist from Africa and India.


Mane

The male lion's mane is the most recognisable feature of the species. It may have evolved around 320,000–190,000 years ago. It starts growing when lions are about a year old. Mane colour varies and darkens with age; research shows its colour and size are influenced by environmental factors such as average ambient temperature. Mane length apparently signals fighting success in male–male relationships; darker-maned individuals may have longer reproductive lives and higher offspring survival, although they suffer in the hottest months of the year. The presence, absence, colour and size of the mane are associated with genetic precondition, sexual maturity, climate and testosterone production; the rule of thumb is that a darker, fuller mane indicates a healthier animal. In Serengeti National Park, female lions favour males with dense, dark manes as mates. Cool ambient temperature in European and North American zoos may result in a heavier mane. Asiatic lions usually have sparser manes than average African lions. Almost all male lions in Pendjari National Park are either maneless or have very short manes.
Maneless lion The term "maneless lion" or "scanty mane lion" often refers to a male lion without a mane, or with a weak one. The purpose of the mane is thought to protect the lion in territorial fights. Although lions are known for their mane, not all males ...
s have also been reported in Senegal, in
Sudan Sudan ( or ; ar, السودان, as-Sūdān, officially the Republic of the Sudan ( ar, جمهورية السودان, link=no, Jumhūriyyat as-Sūdān), is a country in Northeast Africa. It shares borders with the Central African Republic t ...
's
Dinder National Park Dinder National Park is a national park and biosphere reserve in eastern Sudan, and is connected to Ethiopia's Alitash National Park. Location Dinder lies approximately southeast of Khartoum, on either side of the Dinder River bounded to the n ...
and in Tsavo East National Park, Kenya. The original male white lion from
Timbavati The Timbavati Private Nature Reserve is located at the border line between Hoedspruit (in Limpopo) and Acornhoek (in Mpumalanga), north of the Sabi Sand Private Game Reserve on the western edge of Kruger National Park. Geographically and poli ...
in South Africa was also maneless. The hormone testosterone has been linked to mane growth; castrated lions often have little to no mane because the removal of the
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sper ...
s inhibits testosterone production. Increased testosterone may be the cause of maned lionesses reported in northern Botswana.


Colour variation

The white lion is a rare
morph Morph may refer to: Biology * Morph (zoology), a visual or behavioral difference between organisms of distinct populations in a species * Muller's morphs, a classification scheme for genetic mutations * "-morph", a suffix commonly used in tax ...
with a genetic condition called leucism which is caused by a double
recessive allele In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and t ...
. 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 Kruger National Park is a South African National Park and one of the largest game reserves in Africa. It covers an area of in the provinces of Limpopo and Mpumalanga in northeastern South Africa, and extends from north to south and from ea ...
and the adjacent
Timbavati Private Game Reserve The Timbavati Private Nature Reserve is located at the border line between Hoedspruit (in Limpopo) and Acornhoek (in Mpumalanga), north of the Sabi Sand Private Game Reserve on the western edge of Kruger National Park. Geographically and politi ...
in eastern South Africa. They were removed from the wild in the 1970s, thus decreasing the white lion
gene pool The gene pool is the set of all genes, or genetic information, in any population, usually of a particular species. Description A large gene pool indicates extensive genetic diversity, which is associated with robust populations that can surv ...
. Nevertheless, 17 births have been recorded in five prides between 2007 and 2015. White lions are selected for breeding in captivity. They have reportedly been bred in camps in South Africa for use as trophies to be killed during
canned hunt A canned hunt is a trophy hunt which is not " fair chase", typically by having game animals kept in a confined area such as in a fenced ranch (i.e. "canned") to prevent the animals' escape and make tracking easier for the hunter, in order to inc ...
s.


Distribution and habitat

African lions live in scattered populations across sub-Saharan Africa. The lion prefers grassy plains and
savannah A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the Canopy (forest), canopy does not close. The open canopy allows sufficient light to rea ...
s, scrub bordering rivers and open woodlands with bushes. It rarely enters closed forests. On
Mount Elgon Mount Elgon is an extinct shield volcano on the border of Uganda and Kenya, north of Kisumu and west of Kitale. The mountain's highest point, named "Wagagai", is located entirely within Uganda.
, the lion has been recorded up to an elevation of and close to the snow line on
Mount Kenya Mount Kenya (Kikuyu: ''Kĩrĩnyaga'', Kamba, ''Ki Nyaa'') is the highest mountain in Kenya and the second-highest in Africa, after Kilimanjaro. The highest peaks of the mountain are Batian (), Nelion () and Point Lenana (). Mount Kenya is locat ...
. Savannahs with an annual rainfall of make up the majority of lion habitat in Africa, estimated at at most, but remnant populations are also present in tropical moist forests in West Africa and montane forests in East Africa. 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.


Historical range

In Africa, the range of the lion originally spanned most of the central African rainforest zone and the
Sahara , photo = Sahara real color.jpg , photo_caption = The Sahara taken by Apollo 17 astronauts, 1972 , map = , map_image = , location = , country = , country1 = , ...
desert. Schaller, p. 5. In the 1960s, it became extinct in North Africa, except in the southern part of Sudan. In southern Europe and Asia, the lion once ranged in regions where climatic conditions supported an abundance of prey. In Greece, it was common as reported by Herodotus in 480 BC; it was considered rare by 300 BC and extirpated by AD 100. It was present in the Caucasus until the 10th century. It lived in
Palestine __NOTOC__ Palestine may refer to: * State of Palestine, a state in Western Asia * Palestine (region), a geographic region in Western Asia * Palestinian territories, territories occupied by Israel since 1967, namely the West Bank (including East ...
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. The last live lion in Iran was sighted in 1942 about northwest of Dezful, although the corpse of a lioness was found on the banks of the Karun river in
Khūzestān Province Khuzestan Province (also spelled Xuzestan; fa, استان خوزستان ''Ostān-e Xūzestān'') is one of the 31 provinces of Iran. It is in the southwest of the country, bordering Iraq and the Persian Gulf. Its capital is Ahvaz and it covers ...
in 1944. It once ranged from
Sind Sindh (; ; ur, , ; historically romanized as Sind) is one of the four provinces of Pakistan. Located in the southeastern region of the country, Sindh is the third-largest province of Pakistan by land area and the second-largest province ...
and Punjab in Pakistan to Bengal and the Narmada River in central India.


Behaviour and ecology

Lions spend much of their time resting; they are inactive for about twenty hours per day. Schaller, p. 122. 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. Schaller, pp. 120–21.


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 " pride". Groups of male lions are called "coalitions". Schaller, p. 33. Females form the stable social unit in a pride and do not tolerate outside females. Schaller, p. 37. Membership changes only with the births and deaths of lionesses, Schaller, p. 39. although some females leave and become nomadic. Schaller, p. 44. 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. The sole exception to this pattern is the
Tsavo lion ''Panthera leo melanochaita'' is a lion subspecies in Southern and East Africa. In this part of Africa, lion populations are regionally extinct in Lesotho, Djibouti and Eritrea, and are threatened by loss of habitat and prey base, killing by local ...
pride that always has just one adult male. Male cubs are excluded from their maternal pride when they reach maturity at around two or three years of age. Some lions are "nomads" that range widely and move around sporadically, either in pairs or alone. Pairs are more frequent among related males who have been excluded from their birth pride. A lion may switch lifestyles; nomads can become residents and vice versa. Interactions between prides and nomads tend to be hostile, although pride females in estrus allow nomadic males to approach them. Males spend years in a nomadic phase before gaining residence in a pride. A study undertaken in the Serengeti National Park revealed that nomadic coalitions gain residency at between 3.5 and 7.3 years of age. In Kruger National Park, dispersing male lions move more than 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. The area occupied by a pride is called a "pride area" whereas that occupied by a nomad is a "range". Males associated with a pride tend to stay on the fringes, patrolling their territory. The reasons for the development of
sociality Sociality is the degree to which individuals in an animal population tend to associate in social groups (gregariousness) and form cooperative societies. Sociality is a survival response to evolutionary pressures. For example, when a mother wasp ...
in lionesses—the most pronounced in any cat species—are the subject of much debate. Increased hunting success appears to be an obvious reason, but this is uncertain upon examination; coordinated hunting allows for more successful predation but also ensures non-hunting members reduce ''per capita'' calorific intake. Some females, however, take a role raising cubs that may be left alone for extended periods. Members of the pride tend to regularly play the same role in hunts and hone their skills. The health of the hunters is the primary need for the survival of the pride; hunters are the first to consume the prey at the site it is taken. Other benefits include possible kin selection, sharing food within the family, protecting the young, maintaining territory, and individual insurance against injury and hunger. Both males and females defend the pride against intruders, but the male lion is better-suited for this purpose due to its stockier, more powerful build. Some individuals consistently lead the defence against intruders, while others lag behind. Lions tend to assume specific roles in the pride; slower-moving individuals may provide other valuable services to the group. 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. The male or males associated with the pride must defend their relationship with the pride from outside males who may attempt to usurp them. 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. 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.


Hunting and diet

The lion is a
generalist A generalist is a person with a wide array of knowledge on a variety of subjects, useful or not. It may also refer to: Occupations * a physician who provides general health care, as opposed to a medical specialist; see also: ** General pract ...
hypercarnivore and is considered to be both an apex and keystone predator due to its wide prey spectrum. Its prey consists mainly of ungulates weighing , particularly blue wildebeest, plains zebra, African buffalo,
gemsbok The gemsbok or South African oryx (''Oryx gazella'') is a large antelope in the genus ''Oryx''. It is native to the extremely dry, arid regions of Southern Africa; notably, the Kalahari Desert. Some authorities formerly classified the East Afric ...
and
giraffe The giraffe is a large African hoofed mammal belonging to the genus ''Giraffa''. It is the tallest living terrestrial animal and the largest ruminant on Earth. Traditionally, giraffes were thought to be one species, ''Giraffa camelopardalis ...
. They also hunt
common warthog The common warthog (''Phacochoerus africanus'') is a wild member of the pig family (Suidae) found in grassland, savanna, and woodland in sub-Saharan Africa. In the past, it was commonly treated as a subspecies of ''P. aethiopicus'', but today th ...
depending on availability, despite weighing less than the preferred weight range. In India, chital and
sambar deer The sambar (''Rusa unicolor'') is a large deer native to the Indian subcontinent and Southeast Asia that is listed as a vulnerable species on the IUCN Red List since 2008. Populations have declined substantially due to severe hunting, local ins ...
are the most common wild prey, while livestock contributes significantly to lion kills outside protected areas. They usually avoid fully grown adult elephants, rhinoceroses and hippopotamus and small prey like dik-dik,
hyrax Hyraxes (), also called dassies, are small, thickset, herbivorous mammals in the order Hyracoidea. Hyraxes are well-furred, rotund animals with short tails. Typically, they measure between long and weigh between . They are superficially simil ...
es,
hare Hares and jackrabbits are mammals belonging to the genus ''Lepus''. They are herbivores, and live solitarily or in pairs. They nest in slight depressions called forms, and their young are able to fend for themselves shortly after birth. The ge ...
s and
monkey Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as the simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes, which constitutes an incomple ...
s. Unusual prey include
porcupine Porcupines are large rodents with coats of sharp spines, or quills, that protect them against predation. The term covers two families of animals: the Old World porcupines of family Hystricidae, and the New World porcupines of family, Erethizont ...
s and small reptiles. Lions kill other predators but seldom consume them. 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. Schaller, p. 153. Single lions are capable of bringing down zebra and wildebeest, while larger prey like buffalo and giraffe are riskier. In
Chobe National Park Chobe National Park is Botswana's first national park, and also the most biologically diverse. Located in the north of the country, it is Botswana's third largest park, after Central Kalahari Game Reserve and Gemsbok National Park, and has one of ...
, large prides have been observed hunting
African bush elephant The African bush elephant (''Loxodonta africana'') is one of two extant African elephant species and one of three extant elephant species. It is the largest living terrestrial animal, with bulls reaching a shoulder height of up to and a body ...
s up to around 15 years old in exceptional cases, with the victims being calves, juveniles, and even subadults. 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. Some evidence suggests, however, that males are just as successful as females; they are typically solo hunters who ambush prey in small bushland. 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. Thus, lions run quickly only in short bursts at about and need to be close to their prey before starting the attack. Schaller, pp. 233, 247–248 One study in 2018 recorded a lion running at a top speed of . They take advantage of factors that reduce visibility; many kills take place near some form of cover or at night. Schaller, p. 237. The lion's attack is short and powerful; they attempt to catch prey with a fast rush and final leap. They usually pull it down by the rump and kill by a strangling bite to the throat. They also kill prey by enclosing its muzzle in their jaws. Male lions usually aim for the backs or hindquarters of rivals, rather than their necks. Lions typically consume prey at the location of the hunt but sometimes drag large prey into cover. 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. Large kills are shared more widely among pride members. Schaller, p. 133. An adult lioness requires an average of about of meat per day while males require about . Lions gorge themselves and eat up to in one session. 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. Schaller, pp. 270–76. Lions defend their kills from scavengers such as vultures and hyenas. Lions scavenge on
carrion Carrion () is the decaying flesh of dead animals, including human flesh. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures, c ...
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. Schaller, p. 213–216. Most carrion on which both hyenas and lions feed upon are killed by hyenas rather than lions. Carrion is thought to provide a large part of lion diet.


Predatory competition

Lions and
spotted hyena The spotted hyena (''Crocuta crocuta''), also known as the laughing hyena, is a hyena species, currently classed as the sole extant member of the genus ''Crocuta'', native to sub-Saharan Africa. It is listed as being of least concern by the IUC ...
s 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%. 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. 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. When confronted on a kill, hyenas may either leave or wait patiently at a distance of until the lions have finished. Schaller, p. 272. Hyenas 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. Schaller, pp. 273–74. Lion predation 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. When the lion population in Kenya's
Masai Mara National Reserve Maasai Mara, also sometimes spelled Masai Mara and locally known simply as The Mara, is a large national game reserve in Narok, Kenya, contiguous with the Serengeti National Park in Tanzania. It is named in honor of the Maasai people, the ancest ...
declined, the spotted hyena population increased rapidly. Lions tend to dominate cheetahs and
leopard The leopard (''Panthera pardus'') is one of the five extant species in the genus '' Panthera'', a member of the cat family, Felidae. It occurs in a wide range in sub-Saharan Africa, in some parts of Western and Central Asia, Southern Russia, a ...
s, steal their kills and kill their cubs and even adults when given the chance. Cheetahs often lose their kills to lions or other predators. A study in the Serengeti ecosystem revealed that lions killed at least 17 of 125 cheetah cubs born between 1987 and 1990. Cheetahs avoid their competitors by hunting at different times and habitats. Leopards take refuge in trees, but lionesses occasionally attempt to climb up and retrieve their kills. Schaller, p. 293. 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. However, there are a few reported cases of old and wounded lion falling prey to wild dogs. Schaller, p. 188. Nile crocodiles may also kill and eat lions, evidenced by the occasional lion claw found in crocodile stomachs.


Reproduction and life cycle

Most lionesses reproduce by the time they are four years of age. Schaller, p. 29. Lions do not mate at a specific time of year and the females are
polyestrous The estrous cycle (, originally ) is the set of recurring physiological changes that are induced by reproductive hormones in most mammalian therian females. Estrous cycles start after sexual maturity in females and are interrupted by anestrous p ...
. Schaller, p. 174. 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. A lioness may mate with more than one male when she is
in heat The estrous cycle (, originally ) is the set of recurring physiological changes that are induced by reproductive hormones in most mammalian therian females. Estrous cycles start after sexual maturity in females and are interrupted by anestrous p ...
. Schaller, p. 142. Generation length of the lion is about seven years. The average gestation period is around 110days; 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.
Scott Scott may refer to: Places Canada * Scott, Quebec, municipality in the Nouvelle-Beauce regional municipality in Quebec * Scott, Saskatchewan, a town in the Rural Municipality of Tramping Lake No. 380 * Rural Municipality of Scott No. 98, Saska ...
, p. 45.
Lion cubs are born blind, their eyes opening around seven days after birth. They weigh at birth and are almost helpless, beginning to crawl a day or two after birth and walking around three weeks of age. Schaller, p. 143. 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. Usually, the mother does not integrate herself and her cubs back into the pride until the cubs are six to eight weeks old. Sometimes the introduction to pride life occurs earlier, particularly if other lionesses have given birth at about the same time. 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. 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. 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. Schaller, p. 147-49. 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. 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. Cubs also die from starvation and abandonment, and predation by leopards, hyenas and wild dogs. Up to 80% of lion cubs will die before the age of two. Both male and female lions may be ousted from prides to become nomads, although most females usually remain with their birth pride. When a pride becomes too large, however, the youngest generation of female cubs may be forced to leave to find their own territory. When a new male lion takes over a pride, adolescents both male and female may be evicted. Lions of both sexes may be involved in group
homosexual Homosexuality is romantic attraction, sexual attraction, or sexual behavior between members of the same sex or gender. As a sexual orientation, homosexuality is "an enduring pattern of emotional, romantic, and/or sexual attractions" to peop ...
and courtship activities. Males will also head-rub and roll around with each other before simulating sex together.


Health and mortality

Although adult lions have no natural predators, evidence suggests most die violently from attacks by humans or other lions. Schaller, p. 183. 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. Schaller, pp. 188–89. 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. Schaller, pp. 189–90. Ticks commonly infest the ears, neck and groin regions of the lions. Schaller, p. 184. Adult forms of several tapeworm species of the genus '' Taenia'' have been isolated from lion intestines, having been ingested as larvae in
antelope The term antelope is used to refer to many species of even-toed ruminant that are indigenous to various regions in Africa and Eurasia. Antelope comprise a wastebasket taxon defined as any of numerous Old World grazing and browsing hoofed mammals ...
meat. Lions in the Ngorongoro Crater were afflicted by an outbreak of stable fly (''
Stomoxys calcitrans ''Stomoxys calcitrans'' is commonly called the stable fly, barn fly, biting house fly, dog fly, or power mower fly. Unlike most members of the family Muscidae, ''Stomoxys calcitrans'' ('sharp mouth' + 'kicking') and others of its genus Hematoph ...
'') 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. A more recent outbreak in 2001 killed six lions. Captive lions have been infected with canine distemper virus (CDV) since at least the mid-1970s. 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 Encephalitis is inflammation of the brain. The severity can be variable with symptoms including reduction or alteration in consciousness, headache, fever, confusion, a stiff neck, and vomiting. Complications may include seizures, hallucinations, ...
.
Feline immunodeficiency virus Feline immunodeficiency virus (FIV) is a '' Lentivirus'' that affects cats worldwide, with 2.5% to 4.4% of felines being infected. FIV was first isolated in 1986, by Niels C Pedersen and Janet K. Yamamoto at the UC Davis School of Veterina ...
and
lentivirus ''Lentivirus'' is a genus of retroviruses that cause chronic and deadly diseases characterized by long incubation periods, in humans and other mammalian species. The genus includes the human immunodeficiency virus (HIV), which causes AIDS. Lent ...
also affect captive lions.


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 head rubbing and
social licking Social grooming is a behavior in which social animals, including humans, clean or maintain one another's body or appearance. A related term, allogrooming, indicates social grooming between members of the same species. Grooming is a major socia ...
, Schaller, p. 85. which have been compared with the role of
allogrooming Social grooming is a behavior in which social animals, including humans, clean or maintain one another's body or appearance. A related term, allogrooming, indicates social grooming between members of the same species. Grooming is a major soci ...
among primates. Head rubbing—nuzzling the forehead, face and neck against another lion—appears to be a form of greeting 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. Schaller, pp. 85–88. 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. Schaller, pp. 88–91. Lions have an array of facial expressions and body postures that serve as visual gestures. Schaller, pp. 103–117. 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. Lions also use chemical and visual marking; males will
spray Spray or spraying commonly refer to: * Spray (liquid drop) ** Aerosol spray ** Blood spray ** Hair spray ** Nasal spray ** Pepper spray ** PAVA spray ** Road spray or tire spray, road debris kicked up from a vehicle tire ** Sea spray, refers to ...
and scrape plots of ground and objects within the territory. 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 Growling is a low, guttural vocalization produced by animals as an aggressive warning but can also be found in other contexts such as playful behaviors or mating. Different animals will use growling in specific contexts as a form of communicati ...
,
snarling A snarl is a sound, often a growl or vicious utterance, often accompanied by a facial expression, where the upper lip is raised, and the nostrils widen, generally indicating hate, anger or pain. In addition to humans, other mammals including ...
, meowing and
roaring A roar is a type of Animal communication#Auditory, animal vocalization that is deep and acoustic resonance, resonating. Many mammals have evolved to produce roars and other roar-like vocals for purposes such as long-distance communication and intim ...
. 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 . Schaller, pp. 103–113. They tend to roar in a very characteristic manner starting with a few deep, long roars that subside into a series of shorter ones.


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.


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. 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. In 1975, it was estimated that since the 1950s, lion numbers had decreased by half to 200,000 or fewer. Estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004. In the
Republic of the Congo The Republic of the Congo (french: République du Congo, ln, Republíki ya Kongó), also known as Congo-Brazzaville, the Congo Republic or simply either Congo or the Congo, is a country located in the western coast of Central Africa to the w ...
,
Odzala-Kokoua National Park Odzala-Kokoua National Park (or Odzala National Park) is a national park in the Republic of the Congo. The park was first protected in 1935, declared a biosphere reserve in 1977, and granted official designation by presidential decree in 2001. Odz ...
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. 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. 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. A population occurs in Cameroon's Waza National Park, where between approximately 14 and 21 animals persisted as of 2009. In addition, 50 to 150 lions are estimated to be present in Burkina Faso's Arly-Singou ecosystem. 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. 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 Alitash National Park, also called Alatish or Alatash National Park, is a national park in North Gondar Zone, Amhara Region, Ethiopia. It is adjacent to Sudan's Dinder National Park. The national park was founded in 2006. Geography Alatish Nation ...
, Ethiopia, close to the Sudanese border. 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. Lion depredation on livestock is significantly reduced in areas where herders keep livestock in improved enclosures. Such measures contribute to mitigating human–lion conflict.


In Asia

The last refuge of the Asiatic lion population is the Gir National Park and surrounding areas in the region of Saurashtra or
Kathiawar Peninsula Kathiawar () is a peninsula, near the far north of India's west coast, of about bordering the Arabian Sea. It is bounded by the Gulf of Kutch in the northwest and by the Gulf of Khambhat (Gulf of Cambay) in the east. In the northeast, it is ...
in Gujarat State, India. The population has risen from approximately 180 lions in 1974 to about 400 in 2010. It is geographically isolated, which can lead to inbreeding and reduced
genetic diversity Genetic diversity is the total number of genetic characteristics in the genetic makeup of a species, it ranges widely from the number of species to differences within species and can be attributed to the span of survival for a species. It is dis ...
. Since 2008, the Asiatic lion has been listed as Endangered on the IUCN Red List. By 2015, the population had grown to 523 individuals inhabiting an area of in Saurashtra. The Asiatic Lion Census conducted in 2017 recorded about 650 individuals. The presence of numerous human habitations close to the National Park results in conflict between lions, local people and their livestock. Some consider the presence of lions a benefit, as they keep populations of crop damaging herbivores in check. 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.


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. Another 11 animals thought to be Barbary lions kept in
Addis Ababa Zoo Addis Ababa Zoo( Amharic: አንበሳ ጊቢ, romanized: änəbäsa gibi lit. 'lion compound') is a zoological park in Addis Ababa, Ethiopia. History The zoo was founded in 1948 by Emperor Haile Selassie I. Fauna The Addis Ababa Zoo ke ...
are descendants of animals owned by
Emperor Haile Selassie Haile Selassie I ( gez, ቀዳማዊ ኀይለ ሥላሴ, Qädamawi Häylä Səllasé, ; born Tafari Makonnen; 23 July 189227 August 1975) was Emperor of Ethiopia from 1930 to 1974. He rose to power as Regent Plenipotentiary of Ethiopia (' ...
. 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 The Atlas Mountains are a mountain range in the Maghreb in North Africa. It separates the Sahara Desert from the Mediterranean Sea and the Atlantic Ocean; the name "Atlantic" is derived from the mountain range. It stretches around through Moroc ...
of Morocco. 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. 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. Breeding programs need to note origins of the participating animals to avoid cross-breeding different subspecies and thus reducing their conservation value. Captive breeding of lions was halted to eliminate individuals of unknown origin and
pedigree Pedigree may refer to: Breeding * Pedigree chart, a document to record ancestry, used by genealogists in study of human family lines, and in selective breeding of other animals ** Pedigree, a human genealogy (ancestry chart) ** Pedigree (animal ...
. 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. About 77% of the captive lions registered in the
International Species Information System Species360 (formerly International Species Information System or ISIS) founded in 1974, is an international non-profit organization that maintains an online database of wild animals under human care. , the organization serves more than 1,000 zoos ...
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.


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, de Courcy, p. 81-82. 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. 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. 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
menagerie A menagerie is a collection of captive animals, frequently exotic, kept for display; or the place where such a collection is kept, a precursor to the modern Zoo, zoological garden. The term was first used in 17th-century France, in reference to ...
s, an extension of the cabinet of curiosities. They spread from France and Italy during the Renaissance to the rest of Europe. In England, although the seraglio tradition was less developed, lions were kept at the Tower of London in a seraglio established by
King John King John may refer to: Rulers * John, King of England (1166–1216) * John I of Jerusalem (c. 1170–1237) * John Balliol, King of Scotland (c. 1249–1314) * John I of France (15–20 November 1316) * John II of France (1319–1364) * John I o ...
in the 13th century; this was probably stocked with animals from an earlier menagerie started in 1125 by Henry I at his hunting lodge in
Woodstock, Oxfordshire Woodstock is a market town and civil parish, north-west of Oxford in West Oxfordshire in the county of Oxfordshire, England. The 2011 Census recorded a parish population of 3,100. Blenheim Palace, a UNESCO World Heritage Site, is next to Wo ...
, where according to William of Malmesbury lions had been stocked. Blunt, p. 15. Lions were kept in cramped and squalid conditions at
London Zoo London Zoo, also known as ZSL London Zoo or London Zoological Gardens is the world's oldest scientific zoo. It was opened in London on 27 April 1828, and was originally intended to be used as a collection for science, scientific study. In 1831 o ...
until a larger lion house with roomier cages was built in the 1870s. Blunt, p. 208. Further changes took place in the early 20th century when
Carl Hagenbeck Carl Hagenbeck (10 June 1844 – 14 April 1913) was a Germans, German merchant of wild animals who supplied many European zoos, as well as P. T. Barnum. He created the modern zoo with animal enclosures without bars that were closer to their natu ...
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 Melbourne Zoo is a zoo in Melbourne, Australia. It is located within Royal Park in Parkville, approximately north of the centre of Melbourne. It is the primary zoo serving Melbourne. The zoo contains more than 320 animal species from Austra ...
and Sydney's Taronga Zoo; although his designs were popular, the use of bars and caged enclosures prevailed in many zoos until the 1960s. de Courcy, p. 69. 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.
Lion taming Lion taming is the taming and training of lions, either for protection or for use in entertainment, such as the circus. The term often applies to the taming and display of lions and other big cats such as tigers, leopards, jaguars, black panthe ...
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. 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 Equestrianism (from Latin , , , 'horseman', 'horse'), commonly known as horse riding (Commonwealth English) or horseback riding (American English), includes the disciplines of riding, Driving (horse), driving, and Equestrian vaulting, vaulting ...
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. Baratay & Hardouin-Fugier, p. 187. 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).


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 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 Ashurbanipal (Neo-Assyrian language, Neo-Assyrian cuneiform: , meaning "Ashur (god), Ashur is the creator of the heir") was the king of the Neo-Assyrian Empire from 669 BCE to his death in 631. He is generally remembered as the last great king o ...
had one of his lion hunts depicted on a sequence of palace reliefs , known as the Lion Hunt of Ashurbanipal. Lions were also hunted during the Mughal Empire, where Emperor
Jahangir Nur-ud-Din Muhammad Salim (30 August 1569 – 28 October 1627), known by his imperial name Jahangir (; ), was the fourth Mughal Emperor, who ruled from 1605 until he died in 1627. He was named after the Indian Sufi saint, Salim Chishti. Ear ...
is said to have excelled at it. Jackson, p. 156–159. In Ancient Rome, lions were kept by emperors for
hunts Hunt's is the name of a brand of preserved tomato products owned by Conagra Brands. The company was founded in 1888, in Sebastopol, California, as the Hunt Bros. Fruit Packing Co., by Joseph and William Hunt. The brothers relocated to nearby San ...
, gladiator fights and
executions Capital punishment, also known as the death penalty, is the state-sanctioned practice of deliberately killing a person as a punishment for an actual or supposed crime, usually following an authorized, rule-governed process to conclude that t ...
. The
Maasai people The Maasai (; sw, Wamasai) are a Nilotic ethnic group inhabiting northern, central and southern Kenya and northern Tanzania. They are among the best-known local populations internationally due to their residence near the many game parks of t ...
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. 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. The widely reproduced imagery of the heroic hunter chasing lions would dominate a large part of the century. Trophy hunting of lions in recent years has been met with controversy, notably with the
killing of Cecil the lion Cecil ( 2002 – 2 July 2015) was a male African lion who lived primarily in the Hwange National Park in Matabeleland North, Zimbabwe. He was being studied and tracked by a research team of the University of Oxford as part of a long-term study ...
in mid-2015.


Man-eating

Lions do not usually hunt humans but some (usually males) seem to seek them out. One well-publicised case is the
Tsavo maneaters The Tsavo Man-Eaters were a pair of man-eating male lions in the Tsavo region of Kenya, which were responsible for the deaths of many construction workers on the Kenya-Uganda Railway between March and December 1898. The lion pair was said to h ...
; in 1898, 28 officially recorded railway workers building the Kenya-Uganda Railway were taken by lions over nine months during the construction of a bridge in Kenya. 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. Sick or injured animals may be more prone to man-eating but the behaviour is not unusual, nor necessarily aberrant. 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 National Park in
Rufiji District Rufiji is one of the six districts of the Pwani Region of Tanzania. It is bordered to the north by the Kisarawe and Mkuranga Districts, to the east by the Indian Ocean, to the south by the Kilwa District, Lindi Region and to the west by the Morog ...
and in Lindi Province 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. 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. 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.


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. 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. The lion is also used as a symbol of sporting teams.


Africa

In sub-Saharan Africa, the lion has been a common character in stories, proverbs and dances, but rarely featured in visual arts. In some cultures, the lion symbolises power and royalty. In the
Swahili language Swahili, also known by its local name , is the native language of the Swahili people, who are found primarily in Tanzania, Kenya and Mozambique (along the East African coast and adjacent litoral islands). It is a Bantu language, though Swahili ...
, the lion is known as ''simba'' which also means "aggressive", "king" and "strong". Some rulers had the word "lion" in their nickname.
Sundiata Keita Sundiata Keita ( Mandinka, Malinke: ; 1217 – c. 1255) (also known as Manding Diara, Lion of Mali, Sogolon Djata, son of Sogolon, Nare Maghan and Sogo Sogo Simbon Salaba) was a prince and founder of the Mali Empire. He is also the great-uncle ...
of the
Mali Empire The Mali Empire ( Manding: ''Mandé''Ki-Zerbo, Joseph: ''UNESCO General History of Africa, Vol. IV, Abridged Edition: Africa from the Twelfth to the Sixteenth Century'', p. 57. University of California Press, 1997. or Manden; ar, مالي, Māl ...
was called "Lion of Mali". 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. In parts of West Africa, lions symbolised the top class of their social hierarchies. In more heavily forested areas where lions were rare, the leopard represented the top of the hierarchy. Jackson, p. 119. In parts of West and East Africa, the lion is associated with healing and provides the connection between
seers In the United States, the efficiency of air conditioners is often rated by the seasonal energy efficiency ratio (SEER) which is defined by the Air Conditioning, Heating, and Refrigeration Institute, a trade association, in its 2008 standard AHRI ...
and the supernatural. In other East African traditions, the lion represents laziness. In much of African folklore, the lion is portrayed as having low intelligence and is easily tricked by other animals. The ancient Egyptians portrayed several of their
war deities War is an intense armed conflict between states, governments, societies, or paramilitary groups such as mercenaries, insurgents, and militias. It is generally characterized by extreme violence, destruction, and mortality, using regular o ...
as lionesses, which they revered as fierce hunters. Egyptian deities associated with lions include Sekhmet, Bast,
Mafdet Mafdet (also Mefdet, Maftet) was a goddess in the ancient Egyptian religion. She was often depicted wearing a skin of a cheetah, and protected against the bite of snakes and scorpions. She is part of the ancient Egyptian deities during the First ...
, Menhit, Pakhet and Tefnut. 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.


Eastern world

The lion was a prominent symbol in ancient Mesopotamia from
Sumer Sumer () is the earliest known civilization in the historical region of southern Mesopotamia (south-central Iraq), emerging during the Chalcolithic and early Bronze Ages between the sixth and fifth millennium BC. It is one of the cradles of c ...
up to Assyrian and
Babylonia Babylonia (; Akkadian: , ''māt Akkadī'') was an ancient Akkadian-speaking state and cultural area based in the city of Babylon in central-southern Mesopotamia (present-day Iraq and parts of Syria). It emerged as an Amorite-ruled state c. ...
n times, where it was strongly associated with kingship. Lions were among the major symbols of the goddess
Inanna Inanna, also sux, 𒀭𒊩𒌆𒀭𒈾, nin-an-na, label=none is an List of Mesopotamian deities, ancient Mesopotamian goddess of love, war, and fertility. She is also associated with beauty, sex, Divine law, divine justice, and political p ...
/ Ishtar. The Lion of Babylon was the foremost symbol of the Babylonian Empire. The Lion of Judah is the biblical emblem of the tribe of Judah and the later Kingdom of Judah. 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 ''The Lions' Den: Zionism and the Left from Hannah Arendt to Noam Chomsky'' is a 2019 book by associate professor of journalism Susie Linfield, a social and cultural theorist at New York University who self describes as a leftist and a Zionist. ...
where he is miraculously unharmed (). In the Book of Judges, Samson kills a lion as he travels to visit a Philistine woman.(). 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. Narasimha, the man-lion, is one of ten avatars of the
Hindu Hindus (; ) are people who religiously adhere to Hinduism.Jeffery D. Long (2007), A Vision for Hinduism, IB Tauris, , pages 35–37 Historically, the term has also been used as a geographical, cultural, and later religious identifier for ...
god Vishnu. 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 Caste is a form of social stratification characterised by endogamy, hereditary transmission of a style of life which often includes an occupation, ritual status in a hierarchy, and customary social interaction and exclusion based on cultura ...
but is used by millions of Hindu Rajputs and more than twenty million
Sikhs Sikhs ( or ; pa, ਸਿੱਖ, ' ) are people who adhere to Sikhism (Sikhi), a monotheistic religion that originated in the late 15th century in the Punjab region of the Indian subcontinent, based on the revelation of Guru Nanak. The term ...
today. The
Lion Capital of Ashoka The Lion Capital of Ashoka is the Capital (architecture), capital, or head, of a column erected by the Mauryan emperor Ashoka in Sarnath, India, . Its crowning features are four life-sized lions set back to back on a drum-shaped abacus (arch ...
, 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. The lion is also symbolic for the Sinhalese people, the term derived from the Sanskrit ''Sinhala'', meaning "of lions" while a sword-wielding lion is the central figure on the national flag of Sri Lanka. The lion is a common motif in Chinese art; it was first used in art during the late
Spring and Autumn period The Spring and Autumn period was a period in Chinese history from approximately 770 to 476 BC (or according to some authorities until 403 BC) which corresponds roughly to the first half of the Eastern Zhou period. The period's name derives fr ...
(fifth or sixth century BC) and became more popular during the Han Dynasty (206 BCAD 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. 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 Chinese New Year is the festival that celebrates the beginning of a New Year, new year on the traditional lunisolar calendar, lunisolar and solar Chinese calendar. In Sinophone, Chinese and other East Asian cultures, the festival is commonly r ...
, the August Moon Festival and other celebratory occasions for good luck.


Western world

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. The lion is featured in several of Aesop's fables, notably
The Lion and the Mouse The Lion and the Mouse is one of Aesop's Fables, numbered 150 in the Perry Index. There are also Eastern variants of the story, all of which demonstrate mutual dependence regardless of size or status. In the Renaissance the fable was provided w ...
. The Nemean lion was symbolic in ancient Greece and Rome, represented as the constellation and zodiac sign
Leo Leo or Léo may refer to: Acronyms * Law enforcement officer * Law enforcement organisation * ''Louisville Eccentric Observer'', a free weekly newspaper in Louisville, Kentucky * Michigan Department of Labor and Economic Opportunity Arts an ...
, and described in mythology, where it was killed and worn by the hero Heracles, symbolising victory over death.
Lancelot Lancelot du Lac (French for Lancelot of the Lake), also written as Launcelot and other variants (such as early German ''Lanzelet'', early French ''Lanselos'', early Welsh ''Lanslod Lak'', Italian ''Lancillotto'', Spanish ''Lanzarote del Lago' ...
and Gawain were also heroes slaying lions in the Middle Ages. In some medieval stories, lions were portrayed as allies and companions. "Lion" was the nickname of several medieval warrior-rulers with a reputation for bravery, such as Richard the Lionheart. Lions continue to appear in modern literature as characters including the messianic Aslan in the 1950 novel ''
The Lion, the Witch and the Wardrobe ''The Lion, the Witch and the Wardrobe'' is a fantasy novel for children by C. S. Lewis, published by Geoffrey Bles in 1950. It is the first published and best known of seven novels in ''The Chronicles of Narnia'' (1950–1956). Among all the ...
'' and '' The Chronicles of Narnia'' series by
C. S. Lewis Clive Staples Lewis (29 November 1898 – 22 November 1963) was a British writer and Anglican lay theologian. He held academic positions in English literature at both Oxford University (Magdalen College, 1925–1954) and Cambridge Univers ...
, and the comedic
Cowardly Lion The Cowardly Lion is a character in the fictional Land of Oz created by American author L. Frank Baum. He is depicted as an African lion, but like all animals in Oz, he can speak. Since lions are supposed to be "The Kings of Beasts," the Cowardly ...
in
L. Frank Baum Lyman Frank Baum (; May 15, 1856 – May 6, 1919) was an American author best known for his children's books, particularly ''The Wonderful Wizard of Oz'' and its sequels. He wrote 14 novels in the ''Oz'' series, plus 41 other novels (not includ ...
's 1900 '' The Wonderful Wizard of Oz''. Lion symbolism was used from the advent of cinema; one of the most iconic and widely recognised lions is
Leo Leo or Léo may refer to: Acronyms * Law enforcement officer * Law enforcement organisation * ''Louisville Eccentric Observer'', a free weekly newspaper in Louisville, Kentucky * Michigan Department of Labor and Economic Opportunity Arts an ...
, which has been the mascot for Metro-Goldwyn-Mayer studios since the 1920s. The 1966 film '' Born Free'' features Elsa the lioness and is based on the 1960 non-fiction book with the same title. The lion's role as king of the beasts has been used in the 1994
Disney animated feature This list of theatrical animated feature films consists of animated films produced or released by The Walt Disney Studios, the film division of The Walt Disney Company.This list does not include films or specials streamed on Disney+ as Disne ...
film '' The Lion King''. Lions are frequently depicted on
coats of arms A coat of arms is a heraldic visual design on an escutcheon (i.e., shield), surcoat, or tabard (the latter two being outer garments). The coat of arms on an escutcheon forms the central element of the full heraldic achievement, which in its wh ...
, like on the coat of arms of Finland, either as a device on shields or as
supporters In heraldry, supporters, sometimes referred to as ''attendants'', are figures or objects usually placed on either side of the Escutcheon (heraldry), shield and depicted holding it up. Early forms of supporters are found in medieval seals. H ...
, but the lioness is used much less frequently. 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; feline figures in any other position are instead referred to as leopards.
Arthur Fox-Davies Arthur Charles Fox-Davies (28 February 1871 – 19 May 1928) was a British expert on heraldry. His ''Complete Guide to Heraldry'', published in 1909, has become a standard work on heraldry in England. A barrister by profession, Fox-Davies worke ...
, ''A Complete Guide to Heraldry'', T.C. and E.C. Jack, London, 1909, Chapter XI, https://archive.org/details/completeguidetoh00foxduoft.


See also

*
List of largest cats This list of largest cats shows the 10 largest extant Felidae species, ordered by maximum reported weight and size of wild individuals on record. The list does not contain cat hybrids, such as the liger or tigon A tigon (), tiglon () (portman ...
*
Mapogo lion coalition The Mapogo lion coalition was a band of male South African lions that controlled the Sabi Sand region in Kruger National Park. The coalition became infamous for their sheer power and strength in taking over and dominating an area of approximate ...
* '' Roar'' (1981 film) *
Tiger versus lion Historically, a comparison of the tiger (''Panthera tigris'') versus the lion (''Panthera leo'') has been a popular topic of discussion by hunters, naturalists, artists and poets, and has inspired the popular imagination. In the past, lions and ...


Explanatory notes


References


Citations


Books

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External links

* * * * {{Authority control Apex predators Articles containing video clips Big cats Extant Pliocene first appearances Felids of Africa Mammals described in 1758 Mammals of Sub-Saharan Africa Mammals of South Asia National symbols of Burundi National symbols of Chad National symbols of Eswatini National symbols of Kenya National symbols of Malawi National symbols of Morocco National symbols of Sierra Leone National symbols of Singapore National symbols of South Africa National symbols of Sri Lanka National symbols of the Republic of the Congo National symbols of Togo National symbols of Tunisia Panthera Taxa named by Carl Linnaeus Vulnerable animals Vulnerable biota of Africa