The lion (''Panthera leo'') is a large cat of the genus ''

Panthera ''Panthera'' is a genus within the family (biology), family Felidae, and one of two extant genera in the subfamily Pantherinae. It contains the largest living members of the cat family. There are five living species: the jaguar, leopard, lion, ...

'', native to

Sub-Saharan Africa Sub-Saharan Africa is the area and regions of the continent of Africa that lie south of the Sahara. These include Central Africa, East Africa, Southern Africa, and West Africa. Geopolitically, in addition to the list of sovereign states and ...

and

India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...

. It has a muscular, broad-chested body; a short, rounded head; round ears; and a dark, hairy tuft at the tip of its tail. It is

sexually dimorphic Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...

; 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 was ...

, 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 medium-sized and large

ungulate Ungulates ( ) are members of the diverse clade Euungulata ("true ungulates"), which primarily consists of large mammals with Hoof, hooves. Once part of the clade "Ungulata" along with the clade Paenungulata, "Ungulata" has since been determined ...

s. The lion is an apex and keystone predator. The lion inhabits

grassland A grassland is an area where the vegetation is dominance (ecology), dominated by grasses (Poaceae). However, sedge (Cyperaceae) and rush (Juncaceae) can also be found along with variable proportions of legumes such as clover, and other Herbaceo ...

s, savannahs, and

shrubland Shrubland, scrubland, scrub, brush, or bush is a plant community characterized by vegetation dominance (ecology), dominated by shrubs, often also including grasses, herbaceous plant, herbs, and geophytes. Shrubland may either occur naturally o ...

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 The Neolithic or New Stone Age (from Ancient Greek, Greek 'new' and 'stone') is an archaeological period, the final division of the Stone Age in Mesopotamia, Asia, Europe and Africa (c. 10,000 BCE to c. 2,000 BCE). It saw the Neolithic Revo ...

period, the lion ranged throughout Africa and Eurasia, from Southeast Europe to India, but it has been reduced to fragmented populations in sub-Saharan Africa and one population in

western India Western India is a loosely defined region of India consisting of western states of India, Republic of India. The Ministry of Home Affairs (India), Ministry of Home Affairs in its Western Zonal Council Administrative divisions of India, Adminis ...

. It has been listed as Vulnerable on the

IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is an inventory of the global conservation status and extinction risk of biological ...

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 Habitat destruction (also termed habitat loss or habitat reduction) occurs when a natural habitat is no longer able to support its native species. The organisms once living there have either moved elsewhere, or are dead, leading to a decrease ...

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 literature and films. 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 or zoological garden. The term was first used in 17th-century France, referring to ...

s since the time of the

Roman Empire The Roman Empire ruled the Mediterranean and much of Europe, Western Asia and North Africa. The Roman people, Romans conquered most of this during the Roman Republic, Republic, and it was ruled by emperors following Octavian's assumption of ...

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 where animals are kept within enclosures for public exhibition and often bred for Conservation biology, conservation purposes. The term ''zoological garden ...

s across the world since the late 18th century. Cultural depictions of lions have occurred worldwide, particularly as a symbol of power and royalty.

Etymology

The English word ''lion'' is derived via Anglo-Norman from

Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...

(nominative: ), which in turn was a borrowing from

Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...

. The

Hebrew Hebrew (; ''ʿÎbrit'') is a Northwest Semitic languages, Northwest Semitic language within the Afroasiatic languages, Afroasiatic language family. A regional dialect of the Canaanite languages, it was natively spoken by the Israelites and ...

word may also be related. The generic name ''Panthera'' is traceable to the

classical Latin Classical Latin is the form of Literary Latin recognized as a Literary language, literary standard language, standard by writers of the late Roman Republic and early Roman Empire. It formed parallel to Vulgar Latin around 75 BC out of Old Latin ...

word 'panthēra' and the

ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...

word πάνθηρ 'panther'.

Taxonomy

''Felis leo'' was the

scientific name In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...

used by

Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...

in 1758, who described the lion in his work ''

Systema Naturae ' (originally in Latin written ' with the Orthographic ligature, ligature æ) is one of the major works of the Sweden, Swedish botanist, zoologist and physician Carl Linnaeus (1707–1778) and introduced the Linnaean taxonomy. Although the syste ...

''. The genus name ''Panthera'' was coined by

Lorenz Oken Lorenz Oken (1 August 1779 – 11 August 1851) was a Germans, German natural history, naturalist, botany, botanist, biologist, and ornithology, ornithologist. Biography Oken was born Lorenz Okenfuss () in Bohlsbach (now part of Offenburg), Ortena ...

in 1816. Between the mid-18th and mid-20th centuries, 26 lion specimens were described and proposed as subspecies, of which 11 were recognised as valid in 2005. They were distinguished mostly by the size and colour of their manes and skins.

Subspecies

In the 19th and 20th centuries, several lion

type specimen In biology, a type is a particular wikt:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to ancho ...

s were described and proposed as

subspecies In Taxonomy (biology), biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (Morphology (biology), morpholog ...

, with about a dozen recognised as valid

taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...

until 2017. Between 2008 and 2016,

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 image:Hierarchical clustering diagram.png, 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme o ...

, and recognises two subspecies based on results of several phylogeographic studies on lion

evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...

, namely: * ''P. l. leo'' − the nominate lion subspecies includes the Asiatic lion, the regionally extinct Barbary lion, and lion populations in West and northern parts of Central Africa.

Synonyms A synonym is a word, morpheme, or phrase that means precisely 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 a ...

include ''P. l. persica'' , ''P. l. senegalensis'' , ''P. l. kamptzi'' , and ''P. l. azandica'' . It has been called '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 The Ethiopian Highlands (also called the Abyssinian Highlands) is a rugged mass of mountains in Ethiopia in Northeast Africa. It forms the largest continuous area of its elevation in the continent, with little of its surface falling below , whil ...

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 A genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as ...

-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 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 A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...

carnassial Carnassials are paired upper and lower teeth modified in such a way as to allow enlarged and often self-sharpening edges to pass by each other in a shearing manner. This adaptation is found in carnivorans, where the carnassials are the modified f ...

excavated in

Sri Lanka Sri Lanka, officially the Democratic Socialist Republic of Sri Lanka, also known historically as Ceylon, is an island country in South Asia. It lies in the Indian Ocean, southwest of the Bay of Bengal, separated from the Indian subcontinent, ...

, which was attributed to a lion. It is thought to have become extinct around 39,000 years ago. * ''P. fossilis'' was larger than the modern lion and lived in the

Middle Pleistocene The Chibanian, more widely known as the Middle Pleistocene (its previous informal name), is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocen ...

. Bone fragments were excavated in caves in the United Kingdom, Germany, Italy and Czech Republic. *'' P. spelaea'', or the cave lion, lived in

Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...

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 70th parallel north, 72° north latitude in the Chukchi Sea; and on the south ...

during the

Late Pleistocene The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as the Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division ...

. 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 ( years ago) ( ), also called the Old Stone Age (), is a period in human prehistory that is distinguished by the original development of stone tools, and which represents almost the entire period of human prehist ...

cave paintings, ivory carvings, and clay busts. * ''P. atrox'', or the American lion, ranged in the Americas from Canada to possibly Patagonia during the Late Pleistocene. It diverged from the cave lion around 165,000 years ago. A fossil from

Edmonton Edmonton is the capital city of the Provinces and territories of Canada, Canadian province of Alberta. It is situated on the North Saskatchewan River and is the centre of the Edmonton Metropolitan Region, which is surrounded by Central Alberta ...

dates to 11,355 ± 55 years ago.

Evolution

The ''Panthera'' lineage is estimated to have genetically diverged from the

common ancestor Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. According to modern evolutionary biology, all living beings could be descendants of a unique ancestor commonl ...

of the

Felidae Felidae ( ) is the Family (biology), family of mammals in the Order (biology), order Carnivora colloquially referred to as cats. A member of this family is also called a felid ( ). The 41 extant taxon, extant Felidae species exhibit the gre ...

around to . 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 ...

with the jaguar that diverged , but also with the leopard that diverged to . Hybridisation between lion and snow leopard 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 a biogeographic realm of the Earth, the largest of eight. Confined almost entirely to the Eastern Hemisphere, it stretches across Europe and Asia, north of the foothills of the Himalayas, and North Africa. Th ...

since at least the early

Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58Olduvai Gorge 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 migration and allele flow) is the transfer of genetic variation, genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent ...

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 descendants on other continents. The modern lion was probably widely distributed in Africa during the

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

mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, ...

s 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 The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...

Hybrids

In zoos, lions have been bred with

tiger The tiger (''Panthera tigris'') is a large Felidae, cat and a member of the genus ''Panthera'' native to Asia. It has a powerful, muscular body with a large head and paws, a long tail and orange fur with black, mostly vertical stripes. It is ...

s to create hybrids for the curiosity of visitors or for scientific purpose. The liger is bigger than a lion and a tiger, whereas most

tigon The tigon is a hybrid offspring of a male tiger (''Panthera tigris'') and a female lion, or lioness (''Panthera leo''). 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; 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. The tail of all lions ends in a dark, hairy tuft that, in some lions, conceals an approximately -long, hard "spine" or "spur" composed of dermal papillae. The functions of the spur are unknown. The tuft is absent at birth, develops at around months of age and is readily identifiable at the age of seven months. Schaller, pp. 28–30. 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. The skeletal muscles of the lion make up 58.8% of its body weight and represent the highest percentage of muscles among mammals. The lion has a high concentration of fast twitch muscle fibres, giving them quick bursts of speed but less stamina.

Size

Among felids, the lion is second only to the tiger in size. The size and weight of adult lions vary across its 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 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. Mutations in the genes microphthalmia-associated transcription factor and tyrosinase are possibly responsible for the colour of manes. It starts growing when lions enter adolescence, when

testosterone Testosterone is the primary male sex hormone and androgen in Male, males. In humans, testosterone plays a key role in the development of Male reproductive system, male reproductive tissues such as testicles and prostate, as well as promoting se ...

levels increase, and reach their full size at around four years old. Cool ambient temperatures in European and North American zoos may result in a heavier mane. On average, Asiatic lions have sparser manes than African lions. 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. The core body temperature does apparently not increase regardless of sex, season, feeding time, length and colour of mane, but only surface temperature is affected. Unlike in other felid species, female lions consistently interact with multiple males at once. Another hypothesis suggests that the mane also serves to protect the neck in fights, but this is disputed. 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. Almost all male lions in Pendjari National Park are either maneless or have very short manes. Maneless lions have also been reported in Senegal, in

Sudan Sudan, officially the Republic of the Sudan, is a country in Northeast Africa. It borders the Central African Republic to the southwest, Chad to the west, Libya to the northwest, Egypt to the north, the Red Sea to the east, Eritrea and Ethiopi ...

's Dinder National Park and in Tsavo East National Park, Kenya. Castrated lions often have little to no mane because the removal of the

gonad A gonad, sex gland, or reproductive gland is a Heterocrine gland, mixed gland and sex organ that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gon ...

s inhibits testosterone production. Rarely, both wild and captive lionesses have manes. Increased testosterone may be the cause of maned lionesses reported in northern Botswana.

Colour variation

The white lion is a rare morph with a genetic condition called leucism, which is caused by a double recessive allele. It is not albino; it has normal pigmentation in the eyes and skin. White lions have occasionally been encountered in and around Kruger National Park and the adjacent Timbavati Private Game Reserve in eastern South Africa. They were removed from the wild in the 1970s, thus decreasing the white lion gene pool. Nevertheless, 17 births have been recorded in five prides between 2007 and 2015. 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 hunts.

Distribution and habitat

African lions live in scattered populations across sub-Saharan Africa. The lion prefers grassy plains and savannahs, scrub bordering rivers, and open woodlands with bushes. It rarely enters closed forests. On Mount Elgon, the lion has been recorded up to an elevation of and close to the snow line on Mount Kenya. 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 The Sahara (, ) is a desert spanning across North Africa. With an area of , it is the largest hot desert in the world and the list of deserts by area, third-largest desert overall, smaller only than the deserts of Antarctica and the northern Ar ...

desert. Schaller, p. 5. In the 1960s, it became extinct in North Africa, except in the southern part of Sudan. During the mid-

, around 8,000-6,000 years ago, the range of lions expanded into Southeastern and Eastern Europe, partially re-occupying the range of the now extinct cave lion. In

Hungary Hungary is a landlocked country in Central Europe. Spanning much of the Pannonian Basin, Carpathian Basin, it is bordered by Slovakia to the north, Ukraine to the northeast, Romania to the east and southeast, Serbia to the south, Croatia and ...

, the modern lion was present from about 4,500 to 3,200 years

Before Present Before Present (BP) or "years before present (YBP)" is a time scale used mainly in archaeology, geology, and other scientific disciplines to specify when events occurred relative to the origin of practical radiocarbon dating in the 1950s. Because ...

. In

Ukraine Ukraine is a country in Eastern Europe. It is the List of European countries by area, second-largest country in Europe after Russia, which Russia–Ukraine border, borders it to the east and northeast. Ukraine also borders Belarus to the nor ...

, the modern lion was present from about 6,400 to 2,000 years Before Present. In Greece, it was common, as reported by

Herodotus Herodotus (; BC) was a Greek historian and geographer from the Greek city of Halicarnassus (now Bodrum, Turkey), under Persian control in the 5th century BC, and a later citizen of Thurii in modern Calabria, Italy. He wrote the '' Histori ...

in 480 BC; it was considered rare by 300 BC and extirpated by AD 100. In Asia the lion once ranged in regions where climatic conditions supported an abundance of prey. It was present in the

Caucasus The Caucasus () or Caucasia (), is a region spanning Eastern Europe and Western Asia. It is situated between the Black Sea and the Caspian Sea, comprising parts of Southern Russia, Georgia, Armenia, and Azerbaijan. The Caucasus Mountains, i ...

until the 10th century. It lived in the

Levant The Levant ( ) is the subregion that borders the Eastern Mediterranean, Eastern Mediterranean sea to the west, and forms the core of West Asia and the political term, Middle East, ''Middle East''. In its narrowest sense, which is in use toda ...

until the

Middle Ages In the history of Europe, the Middle Ages or medieval period lasted approximately from the 5th to the late 15th centuries, similarly to the post-classical period of global history. It began with the fall of the Western Roman Empire and ...

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

Khuzestan province Khuzestan province () is one of the 31 Provinces of Iran. Located in the southwest of the country, the province borders Iraq and the Persian Gulf, covering an area of . Its capital is the city of Ahvaz. Since 2014, it has been part of Iran's R ...

in 1944. It once ranged from Sind and

Punjab Punjab (; ; also romanised as Panjāb or Panj-Āb) is a geopolitical, cultural, and historical region in South Asia. It is located in the northwestern part of the Indian subcontinent, comprising areas of modern-day eastern Pakistan and no ...

in Pakistan to

Bengal Bengal ( ) is a Historical geography, historical geographical, ethnolinguistic and cultural term referring to a region in the Eastern South Asia, eastern part of the Indian subcontinent at the apex of the Bay of Bengal. The region of Benga ...

and the

Narmada River The Narmada River, previously also known as ''Narbada'' or anglicised as ''Nerbudda'', is the 5th longest river in India and overall the longest west-flowing river in the country. It is also the largest flowing river in the state of Madhya Prade ...

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. 120–121.

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 Pride is a human Emotion, secondary emotion characterized by a sense of satisfaction with one's Identity (philosophy), identity, performance, or accomplishments. It is often considered the opposite of shame or of humility and, depending on conte ...

". 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. The majority of females remain in their birth prides while all males and some females disperse. Packer, p. 33. 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 of up to 30 individuals have been observed. The sole exception to this pattern is the Tsavo lion pride that always had only one adult male. Prides act as fission–fusion societies, and members split into subgroups that keep in contact with roars. Nomadic lions range widely and move around sporadically, either in pairs or alone. Pairs are more frequent among related males. 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 The estrous cycle (, originally ) is a set of recurring physiological changes induced by reproductive hormones in females of mammalian subclass Theria. Estrous cycles start after sexual maturity in females and are interrupted by anestrous phas ...

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 The Serengeti National Park is a large national park in northern Tanzania that stretches over . It is located in eastern Mara Region and northeastern Simiyu Region and contains over of virgin savanna. The park was established in 1940. The Se ...

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 evolution of sociability in lions was likely driven both by high

population density Population density (in agriculture: Standing stock (disambiguation), standing stock or plant density) is a measurement of population per unit land area. It is mostly applied to humans, but sometimes to other living organisms too. It is a key geog ...

and the clumped resources of savannah habitats. The larger the pride, the more high-quality

territory A territory is an area of land, sea, or space, belonging or connected to a particular country, person, or animal. In international politics, a territory is usually a geographic area which has not been granted the powers of self-government, ...

they can defend; "hotspots" are near river

confluence In geography, a confluence (also ''conflux'') occurs where two or more watercourses join to form a single channel (geography), channel. A confluence can occur in several configurations: at the point where a tributary joins a larger river (main ...

s, where they have optimal access to water, prey and vegetation cover. A study on three lion prides in a Zimbabwean wildlife reserve revealed that the dominant pride of 12 lions had the shortest average distance to water and the smallest

home range A home range is the area in which an animal lives and moves on a periodic basis. It is related to the concept of an animal's territory which is the area that is actively defended. The concept of a home range was introduced by W. H. Burt in 1943. ...

of ; the smallest pride of four lions had the longest average distance to water and the largest home range of . 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 patrol the fringes. 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

hypercarnivore A hypercarnivore is an animal that has a diet that is more than 70% meat, either via active predation or by scavenging. The remaining non-meat diet may consist of non-animal foods such as fungi, fruits or other plant material. Some extant exampl ...

and is considered to be both an apex and keystone predator due to its wide prey spectrum. Its prey consists mainly of medium-sized to large

s, particularly

blue wildebeest The blue wildebeest (''Connochaetes taurinus''), also called the common wildebeest, white-bearded gnu or brindled gnu, is a large antelope and one of the two species of wildebeest. It is placed in the genus ''Connochaetes'' and Family (biology), ...

plains zebra The plains zebra (''Equus quagga'', formerly ''Equus burchellii'') is the most common and geographically widespread species of zebra. Its range is fragmented, but spans much of southern and eastern Africa south of the Sahara. Six or seven subspec ...

, African buffalo, gemsbok and

giraffe The giraffe is a large Fauna of Africa, African even-toed ungulate, hoofed mammal belonging to the genus ''Giraffa.'' It is the Largest mammals#Even-toed Ungulates (Artiodactyla), tallest living terrestrial animal and the largest ruminant on ...

. It also frequently takes common warthog despite it being much smaller. In India, chital and sambar deer are the most common wild prey, while livestock contributes significantly to lion kills outside protected areas. It usually avoids adult

elephant Elephants are the largest living land animals. Three living species are currently recognised: the African bush elephant ('' Loxodonta africana''), the African forest elephant (''L. cyclotis''), and the Asian elephant ('' Elephas maximus ...

rhinoceros A rhinoceros ( ; ; ; : rhinoceros or rhinoceroses), commonly abbreviated to rhino, is a member of any of the five extant taxon, extant species (or numerous extinct species) of odd-toed ungulates (perissodactyls) in the family (biology), famil ...

and

hippopotamus The hippopotamus (''Hippopotamus amphibius;'' ; : hippopotamuses), often shortened to hippo (: hippos), further qualified as the common hippopotamus, Nile hippopotamus and river hippopotamus, is a large semiaquatic mammal native to sub-Sahar ...

and small prey like dik-dik, hyraxes,

hare Hares and jackrabbits are mammals belonging to the genus ''Lepus''. They are herbivores and live Solitary animal, solitarily or in pairs. They nest in slight depressions called forms, and their young are precociality, able to fend for themselves ...

s and monkeys, and seldom consumes other predators; unusual prey include small reptiles and

porcupine Porcupines are large rodents with coats of sharp Spine (zoology), spines, or quills, that protect them against predation. The term covers two Family (biology), families of animals: the Old World porcupines of the family Hystricidae, and the New ...

s. 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 Biodiversity, 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. ...

, 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. In typical group 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. 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. 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 They take advantage of factors that reduce visibility; many kills take place near some form of cover or at night. Schaller, p. 237. One study in 2018 recorded a lion running at a top speed of . The lion accelerates at the start of the chase by a rate of per second, whereas zebras, wildebeest and Thomson's gazelle accelerate by a rate of per second, per second, and per second respectively; acceleration appears to be more important than steady displacement speed in lion hunts. 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 In vertebrate anatomy, the throat is the front part of the neck, internally positioned in front of the vertebrae. It contains the Human pharynx, pharynx and larynx. An important section of it is the epiglottis, separating the esophagus from the t ...

or muzzle. It can hold the prey's throat for up to 13 minutes, until the prey stops moving. It has a bite force from 1593.8 to 1768 Newtons at the canine tip and up 4167.6 Newtons at the

notch. 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 (), also known as a carcass, is the decaying flesh of dead animals. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures ...

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 hyenas occupy a similar ecological niche and compete for prey and carrion; a review of data across several studies indicates a dietary overlap of 58.6%. 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 may feed alongside lions and force them off a kill. The two species attack one another even when there is no food involved for no apparent reason. Schaller, pp. 273–74. Lions can account for up to 71% of hyena deaths in

Etosha National Park Etosha National Park is a national park in northwestern Namibia and one of the largest national parks in Africa. It was proclaimed a game reserve in March 1907 in Ordinance 88 by the Governor of German South West Africa, Friedrich von Lindequist. ...

. Hyenas have adapted by frequently mobbing lions that enter their home ranges. When the lion population in Kenya's Masai Mara National Reserve declined, the spotted hyena population increased rapidly. Lions tend to dominate

cheetah The cheetah (''Acinonyx jubatus'') is a large Felidae, cat and the Fastest animals, fastest land animal. It has a tawny to creamy white or pale buff fur that is marked with evenly spaced, solid black spots. The head is small and rounded, wit ...

s and leopards, 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, 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 affect leopard abundance. Leopards take refuge in trees, though lionesses occasionally attempt to climb up and retrieve their kills. Schaller, p. 293. 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. However, there are a few reported cases of old and wounded lions falling prey to wild dogs. Schaller, p. 188.

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. 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 Ovulation is an important part of the menstrual cycle in female vertebrates where the egg cells are released from the ovaries as part of the ovarian cycle. In female humans ovulation typically occurs near the midpoint in the menstrual cycle and ...

. A lioness may mate with more than one male when she is in heat. Schaller, p. 142. Lions of both sexes may be involved in group homosexual and courtship activities. Males will also head-rub and roll around with each other before mounting each other. 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, 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. Male cubs are excluded from their maternal pride when they reach maturity at around two or three years of age, Schaller, p. 44. while some females may leave when they reach the age of two. When a new male lion takes over a pride, adolescents both male and female may be evicted.

Health and mortality

Lions may live 12–17 years in the wild. 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. 188–189. 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. 189–190.

Nile crocodile The Nile crocodile (''Crocodylus niloticus'') is a large crocodilian native to freshwater habitats in Africa, where it is present in 26 countries. It is widely distributed in sub-Saharan Africa, occurring mostly in the eastern, southern, and ce ...

s may also kill and eat lions, evidenced by the occasional lion claw found in crocodile stomachs. 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 refers to numerous extant or recently extinct species of the ruminant artiodactyl family Bovidae that are indigenous to most of Africa, India, the Middle East, Central Asia, and a small area of Eastern Europe. Antelopes do ...

meat. Lions in the Ngorongoro Crater were afflicted by an outbreak of stable fly ('' Stomoxys calcitrans'') in 1962, resulting in lions becoming emaciated and covered in bloody, bare patches. Lions sought unsuccessfully to evade the biting flies by climbing trees or crawling into hyena burrows; many died or migrated and the local population dropped from 70 to 15 individuals. 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 Pneumonia is an Inflammation, inflammatory condition of the lung primarily affecting the small air sacs known as Pulmonary alveolus, alveoli. Symptoms typically include some combination of Cough#Classification, productive or dry cough, ches ...

and

encephalitis Encephalitis is inflammation of the Human brain, brain. The severity can be variable with symptoms including reduction or alteration in consciousness, aphasia, headache, fever, confusion, a stiff neck, and vomiting. Complications may include se ...

. Feline immunodeficiency virus and lentivirus 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, Schaller, p. 85. which have been compared with the role of allogrooming 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 spray urine Schaller
p. 116.
/ref> 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 roaring,

growling Growling is a low, guttural Animal communication, vocalization produced by animals as an aggression, aggressive warning but can also be found in other contexts such as playful behaviors or mating. Different animals will use growling in specific ...

and snarling. Other sounds produced include puffing, humming, and a soft sound similar to purring, while cubs communicate by

meow A meow or miaow is a cat vocalization. ''Meows'' may have diverse tones in terms of their sound, and what is heard can vary from being chattered to calls, murmurs, and whispers. Adult cats rarely meow to each other. Thus, an adult cat meowin ...

ing and bleating. Roaring is used to advertise the animal's 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 grunts.

Conservation

The lion is listed as Vulnerable on the

. 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, also known as Congo-Brazzaville, the Congo Republic or simply the Congo (the last ambiguously also referring to the neighbouring Democratic Republic of the Congo), is a country located on the western coast of Central ...

, 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. 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 Benin, officially the Republic of Benin, is a country in West Africa. It was formerly known as Dahomey. It is bordered by Togo to the west, Nigeria to the east, Burkina Faso to the north-west, and Niger to the north-east. The majority of its po ...

Burkina Faso Burkina Faso is a landlocked country in West Africa, bordered by Mali to the northwest, Niger to the northeast, Benin to the southeast, Togo and Ghana to the south, and Ivory Coast to the southwest. It covers an area of 274,223 km2 (105,87 ...

and

Niger Niger, officially the Republic of the Niger, is a landlocked country in West Africa. It is a unitary state Geography of Niger#Political geography, bordered by Libya to the Libya–Niger border, north-east, Chad to the Chad–Niger border, east ...

. This population is listed as

Critically Endangered An IUCN Red List critically endangered (CR or sometimes CE) species is one that has been categorized by the International Union for Conservation of Nature as facing an extremely high risk of extinction in the wild. As of December 2023, of t ...

. 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, Ethiopia, close to the Sudanese border. In 2005, Lion Conservation Strategies were developed for West and Central Africa, and 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 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 Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely genetic distance, related genetically. By analogy, the term is used in human reproduction, but more commonly refers to the genet ...

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 correlated to the span of survival for a species. It is d ...

. Since 2008, the Asiatic lion has been listed as

Endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching, inv ...

on the

. By 2015, the population had grown to 523 individuals inhabiting an area of in Saurashtra. In 2017, about 650 individuals were recorded during the Asiatic Lion Census. The presence of numerous human settlements close to Gir National Park resulted 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.

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 are descendants of animals owned by Emperor Haile Selassie. WildLink International in collaboration with

Oxford University The University of Oxford is a collegiate research university in Oxford, England. There is evidence of teaching as early as 1096, making it the oldest university in the English-speaking world and the second-oldest continuously operating u ...

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. They separate the Sahara Desert from the Mediterranean Sea and the Atlantic Ocean; the name "Atlantic" is derived from the mountain range, which stretches around through M ...

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. 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 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, pp. 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

s, an extension of the

cabinet of curiosities Cabinets of curiosities ( and ), also known as wonder-rooms ( ), were encyclopedic collections of objects whose categorical boundaries were, in Renaissance Europe, yet to be defined. Although more rudimentary collections had preceded them, t ...

. They spread from France and Italy during the

Renaissance The Renaissance ( , ) is a Periodization, period of history and a European cultural movement covering the 15th and 16th centuries. It marked the transition from the Middle Ages to modernity and was characterized by an effort to revive and sur ...

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 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 parishes in England, civil parish, north-west of Oxford in West Oxfordshire in the county of Oxfordshire, England. The United Kingdom Census 2021, 2021 census recorded a parish population of 3,521, up from t ...

, where according to William of Malmesbury lions had been stocked. Blunt, p. 15. Lions were kept in cramped and squalid conditions at London Zoo 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 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, Melbourne, Royal Park in Parkville, Victoria, Parkville, approximately north of the centre of Melbourne. It is the primary zoo serving Melbourne. As of 2021, the ...

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, cheetahs, ...

has been part of both established

circus A circus is a company of performers who put on diverse entertainment shows that may include clowns, acrobats, trained animals, trapeze acts, musicians, dancers, hoopers, tightrope walkers, jugglers, magicians, ventriloquists, and unicy ...

es 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. Martin composed a

pantomime Pantomime (; informally panto) is a type of musical comedy stage production designed for family entertainment, generally combining gender-crossing actors and topical humour with a story more or less based on a well-known fairy tale, fable or ...

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, and vaulting. This broad description includes the ...

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 Egypt Ancient Egypt () was a cradle of civilization concentrated along the lower reaches of the Nile River in Northeast Africa. It emerged from prehistoric Egypt around 3150BC (according to conventional Egyptian chronology), when Upper and Lower E ...

ian inscription dated circa 1380 BC that mentions Pharaoh Amenhotep III killing 102 lions in ten years "with his own arrows". The

Assyria Assyria (Neo-Assyrian cuneiform: , ''māt Aššur'') was a major ancient Mesopotamian civilization that existed as a city-state from the 21st century BC to the 14th century BC and eventually expanded into an empire from the 14th century BC t ...

n emperor Ashurbanipal had one of his lion hunts depicted on a sequence of Assyrian palace reliefs , known as the Lion Hunt of Ashurbanipal. Lions were also hunted during the

Mughal Empire The Mughal Empire was an Early modern period, early modern empire in South Asia. At its peak, the empire stretched from the outer fringes of the Indus River Basin in the west, northern Afghanistan in the northwest, and Kashmir in the north, to ...

, where Emperor

Jahangir Nur-ud-din Muhammad Salim (31 August 1569 – 28 October 1627), known by his imperial name Jahangir (; ), was List of emperors of the Mughal Empire, Emperor of Hindustan from 1605 until his death in 1627, and the fourth Mughal emperors, Mughal ...

is said to have excelled at it. Jackson, pp. 156–159. In

Ancient Rome In modern historiography, ancient Rome is the Roman people, Roman civilisation from the founding of Rome, founding of the Italian city of Rome in the 8th century BC to the Fall of the Western Roman Empire, collapse of the Western Roman Em ...

, lions were kept by emperors for hunts, gladiator fights and executions. The

Maasai people The Maasai (;) are a Nilotic peoples, Nilotic ethnic group inhabiting northern, central and southern Kenya and northern Tanzania, near the African Great Lakes region.

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 Trophy hunting is a form of hunting for field sports, sport in which parts of the hunted wild animals are kept and displayed as trophies. The animal being targeted, known as the "game (hunting), game", is typically a mature male specimen from a p ...

of lions in recent years has been met with controversy, notably with the killing of Cecil the lion 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; in 1898, 28 officially recorded workers building the 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 Game Reserve in Rufiji River and in

Lindi Region Lindi Region (''Mkoa wa Lindi'' in Swahili language, Swahili) is one of Tanzania's 31 administrative Regions of Tanzania, regions. The region covers an area of . The region is comparable in size to the combined land area of the nation state of Sr ...

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. Frump said thousands may have been killed in the decades after

apartheid Apartheid ( , especially South African English: , ; , ) was a system of institutionalised racial segregation that existed in South Africa and South West Africa (now Namibia) from 1948 to the early 1990s. It was characterised by an ...

sealed the park and forced refugees to cross the park at night.

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 is considered to be the 'King of Beasts' and has symbolised power, royalty and protection. Several leaders have had "lion" in their name including Sundiata Keita of the

Mali Empire The Mali Empire (Manding languages, 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 ...

, who was called "Lion of Mali", and Richard the Lionheart of England. The male's mane makes it a particularly recognisable feature and thus has been represented more than the female. Nevertheless, the lioness has also had importance as a guardian. In sub-Saharan Africa, the lion has been a common character in stories, proverbs and dances, but rarely featured in visual arts. In the

Swahili language Swahili, also known as as it is referred to endonym and exonym, in the Swahili language, is a Bantu languages, Bantu language originally spoken by the Swahili people, who are found primarily in Tanzania, Kenya, and Mozambique (along the East Af ...

, the lion is known as ''simba'' which also means "aggressive", "king" and "strong". In parts of West and East Africa, the lion is associated with healing and provides the connection between seers and the supernatural. In other East African traditions, the lion represents laziness. In much of African folklore, the lion is portrayed as having low intelligence and is easily tricked. In

Nubia Nubia (, Nobiin language, Nobiin: , ) is a region along the Nile river encompassing the area between the confluence of the Blue Nile, Blue and White Nile, White Niles (in Khartoum in central Sudan), and the Cataracts of the Nile, first cataract ...

, the lion-god Apedemak was associated with the flooding of the Nile. In

Ancient Egypt Ancient Egypt () was a cradle of civilization concentrated along the lower reaches of the Nile River in Northeast Africa. It emerged from prehistoric Egypt around 3150BC (according to conventional Egyptian chronology), when Upper and Lower E ...

, lions were linked both with the sun and the waters of the Nile. Several gods were conceived as being part lion, including the war deities

Sekhmet In Egyptian mythology, Sekhmet ( or Sachmis , from ; ) is a warrior goddess as well as goddess of medicine. Sekhmet is also a solar deity, sometimes given the epithet "the Eye of Ra, eye of Ra". She is often associated with the goddesses Hatho ...

and Maahes, and Tefnut, the goddess of moisture.. The lion was a prominent symbol in ancient

Mesopotamia Mesopotamia is a historical region of West Asia situated within the Tigris–Euphrates river system, in the northern part of the Fertile Crescent. Today, Mesopotamia is known as present-day Iraq and forms the eastern geographic boundary of ...

from

Sumer Sumer () is the earliest known civilization, located in the historical region of southern Mesopotamia (now south-central Iraq), emerging during the Chalcolithic and Early Bronze Age, early Bronze Ages between the sixth and fifth millennium BC. ...

up to

n and

Babylonia Babylonia (; , ) was an Ancient history, ancient Akkadian language, Akkadian-speaking state and cultural area based in the city of Babylon in central-southern Mesopotamia (present-day Iraq and parts of Kuwait, Syria and Iran). It emerged as a ...

n times, where it was strongly associated with kingship. The big cat was a symbol and steed of fertility goddess

Inanna Inanna is the List of Mesopotamian deities, ancient Mesopotamian goddess of war, love, and fertility. She is also associated with political power, divine law, sensuality, and procreation. Originally worshipped in Sumer, she was known by the Akk ...

. Lions decorate the Processional Way leading to the Ishtar Gate in Babylon which was built by

Nebuchadnezzar II Nebuchadnezzar II, also Nebuchadrezzar II, meaning "Nabu, watch over my heir", was the second king of the Neo-Babylonian Empire, ruling from the death of his father Nabopolassar in 605 BC to his own death in 562 BC. Often titled Nebuchadnezzar ...

in the 6th century BCE. The Lion of Babylon symbolised the power of the king and protection of the land against enemies, but was also invoked for good luck. The constellation Leo the lion was first recognised by the Sumerians around 4,000 years ago and is the fifth sign of the

zodiac The zodiac is a belt-shaped region of the sky that extends approximately 8° north and south celestial latitude of the ecliptic – the apparent path of the Sun across the celestial sphere over the course of the year. Within this zodiac ...

. In ancient Israel, a lion represented the

tribe of Judah According to the Hebrew Bible, the tribe of Judah (, ''Shevet Yehudah'') was one of the twelve Tribes of Israel, named after Judah (son of Jacob), Judah, the son of Jacob. Judah was one of the tribes to take its place in Canaan, occupying it ...

. Lions are frequently mentioned in the

Bible The Bible is a collection of religious texts that are central to Christianity and Judaism, and esteemed in other Abrahamic religions such as Islam. The Bible is an anthology (a compilation of texts of a variety of forms) originally writt ...

, notably in the

Book of Daniel The Book of Daniel is a 2nd-century BC biblical apocalypse with a 6th-century BC setting. It is ostensibly a narrative detailing the experiences and Prophecy, prophetic visions of Daniel, a Jewish Babylonian captivity, exile in Babylon ...

, in which the eponymous hero is forced to sleep in the lions' den. Cowardly lion2

Indo-Persian chroniclers regarded the lion as keeper of order in the realm of animals. The

Sanskrit Sanskrit (; stem form ; nominal singular , ,) is a classical language belonging to the Indo-Aryan languages, Indo-Aryan branch of the Indo-European languages. It arose in northwest South Asia after its predecessor languages had Trans-cultural ...

word ''mrigendra'' signifies a lion as king of animals. 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 Narasimha (, , or , ), is the fourth avatara of the Hindu god Vishnu in the Satya Yuga. He incarnated as a part-lion, part-man and killed Hiranyakashipu, ended religious persecution and calamity on earth, and restored dharma. Narasimha has th ...

, an avatar of the deity

Vishnu Vishnu (; , , ), also known as Narayana and Hari, is one of the Hindu deities, principal deities of Hinduism. He is the supreme being within Vaishnavism, one of the major traditions within contemporary Hinduism, and the god of preservation ( ...

, battles and slays the evil ruler Hiranyakashipu. In Buddhist art, lions are associated with both '' arhats'' and

bodhisattva In Buddhism, a bodhisattva is a person who has attained, or is striving towards, '' bodhi'' ('awakening', 'enlightenment') or Buddhahood. Often, the term specifically refers to a person who forgoes or delays personal nirvana or ''bodhi'' in ...

s and may be ridden by the Manjushri. Though they were never native to the country, lions have played important roles in

Chinese culture Chinese culture () is one of the Cradle of civilization#Ancient China, world's earliest cultures, said to originate five thousand years ago. The culture prevails across a large geographical region in East Asia called the Sinosphere as a whole ...

. Statues of the beast have guarded the entrances to the imperial palace and many religious shrines. The

lion dance Lion dance ( zh, s=舞狮, t=舞獅, p=wǔshī, c=, first=t) is a form of traditional dance in Culture of China, Chinese culture and other Asian countries in which performers mimic a Asiatic lion, lion's movements in a lion costume to bring good l ...

has been performed for over a thousand years. In

ancient Greece Ancient Greece () was a northeastern Mediterranean civilization, existing from the Greek Dark Ages of the 12th–9th centuries BC to the end of classical antiquity (), that comprised a loose collection of culturally and linguistically r ...

, the lion is featured in several of

Aesop's fables Aesop's Fables, or the Aesopica, is a collection of fables credited to Aesop, a Slavery in ancient Greece, slave and storyteller who lived in ancient Greece between 620 and 564 Before the Common Era, BCE. Of varied and unclear origins, the stor ...

, notably The Lion and the Mouse. In

Greek mythology Greek mythology is the body of myths originally told by the Ancient Greece, ancient Greeks, and a genre of ancient Greek folklore, today absorbed alongside Roman mythology into the broader designation of classical mythology. These stories conc ...

, the Nemean lion is slain by the hero

Heracles Heracles ( ; ), born Alcaeus (, ''Alkaios'') or Alcides (, ''Alkeidēs''), was a Divinity, divine hero in Greek mythology, the son of ZeusApollodorus1.9.16/ref> and Alcmene, and the foster son of Amphitryon.By his adoptive descent through ...

who wears its skin.

Lancelot Lancelot du Lac (French for Lancelot of the Lake), alternatively written as Launcelot and other variants, is a popular character in the Matter of Britain, Arthurian legend's chivalric romance tradition. He is typically depicted as King Arthu ...

and

Gawain Gawain ( ), also known in many other forms and spellings, is a character in Matter of Britain, Arthurian legend, in which he is King Arthur's nephew and one of the premier Knights of the Round Table. The prototype of Gawain is mentioned und ...

were also heroes slaying lions in

medieval Europe In the history of Europe, the Middle Ages or medieval period lasted approximately from the 5th to the late 15th centuries, similarly to the post-classical period of World history (field), global history. It began with the fall of the West ...

. Lions continue to appear in modern literature such as the Cowardly Lion in L. Frank Baum's 1900 '' The Wonderful Wizard of Oz'', and in

C. S. Lewis Clive Staples Lewis (29 November 1898 – 22 November 1963) was a British writer, literary scholar and Anglican lay theologian. He held academic positions in English literature at both Magdalen College, Oxford (1925–1954), and Magdalen ...

's '' The Lion, the Witch and the Wardrobe''. For cinema, the lion was portrayed as the ruler of animals in the 1994 Disney animated feature film '' The Lion King''. Jackson, pp. 7, 96, 99, 103–105, 128, 135, 150, 197.

Notes

References

Citations

Books

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External links