Lion

Lion Temporal range: Pleistocene–Present PreЄ Є O S D C P T J K Pg N ↓
A male Southern African lion photographed in Kruger National Park, South Africa
A Southern African lioness photographed in Etosha National Park, Namibia
Conservation status
Vulnerable (IUCN 3.1)[2]
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Suborder:	Feliformia
Family:	Felidae
Subfamily:	Pantherinae
Genus:	Panthera
Species:	P. leo[1]
Binomial name
Panthera leo[1] (Linnaeus, 1758)[3]
Subspecies
see Subspecies P. l. leo (Sensu stricto) P. l. atrox P. l. europaea P. l. melanochaita (Sensu stricto) P. l. sinhaleyus P. spelaea P. l. vereshchagini
Distribution of Panthera leo in Africa and Eurasia, in the past and present.
Synonyms
Felis leo Linnaeus, 1758

The lion (Panthera leo) is a species in the cat family (Felidae). A muscular, deep-chested cat, it has a short, rounded head, a reduced neck and round ears, and a hairy tuft at the end of its tail, the lion is sexually dimorphic; males are larger than females with a typical weight range of 150 to 250 kg (331 to 551 lb) for the former and 120 to 182 kg (265 to 401 lb) for the latter. In addition, male lions have a prominent mane, which is the most recognisable feature of the species. A lion pride consists of a few adult males, related females and cubs. Groups of female lions typically hunt together, preying mostly on large ungulates. Lions are apex and keystone predators, although they scavenge as opportunity allows. While lions do not typically hunt humans, some have been known to do so.

The lion typically inhabits grasslands and savannas, but is absent in dense forests, it is usually more diurnal than other big cats, but when persecuted adapts to being active at night and at twilight. In the Pleistocene, the lion ranged throughout Eurasia, Africa and North America from the Yukon to Peru. Today, it has been reduced to fragmented populations in Sub-Saharan Africa and one critically endangered population in western India, it has been listed as Vulnerable on the IUCN Red List since 1996, as populations in African countries declined by about 43% since the early 1990s. Lion populations are untenable outside designated protected areas, although the cause of the decline is not fully understood, habitat loss and conflicts with humans are the greatest causes of concern.

One of the most widely recognised animal symbols in human culture, the lion has been extensively depicted in sculptures and paintings, on national flags, and in contemporary films and literature. Lions have been kept in menageries since the time of the Roman Empire, and have been a key species sought for exhibition in zoos across the world since the late 18th century. Cultural depictions of lions were prominent in the Upper Paleolithic period, with carvings and paintings from the Lascaux and Chauvet Caves in France dated to 17,000 years ago, through virtually all ancient and medieval cultures where they once occurred.

Etymology

The lion's name, similar in many Romance languages, is derived from the Latin Latin: leo,^[4] and the Ancient Greek λέων (leon).^[5] The Hebrew word לָבִיא‎ (lavi) may also be related.^[6]

Taxonomy and phylogeny

In 1758, Carl Linnaeus described the lion in his work Systema Naturae and gave it the scientific name Felis leo.^[3] Between the mid 18th and mid 20th centuries, 26 lion specimens were described and proposed as subspecies, of which 11 were recognised as valid in 2005,^[1] they were distinguished on the basis of appearance, size and colour of mane. As these characteristics vary highly between individuals, most of these forms were probably not true subspecies, especially as they were often based upon museum material with "striking, but abnormal" morphological characteristics.^[11]

Based on morphology of 58 lion skulls in three European museums, the subspecies krugeri, nubica, persica, and senegalensis were assessed distinct; but bleyenberghi overlapped with senegalensis and krugeri. The Asiatic lion persica was the most distinctive, and the Cape lion had characteristics allying it more with persica than the other sub-Saharan lions.^[12]

The lion's closest relatives are the other species of the genus Panthera: the tiger, snow leopard, jaguar, and leopard. Results of phylogenetic studies published in 2006 and 2009 indicated that the jaguar and the lion belong to one sister group, which diverged about 2.06 million years ago.^[7][8] Results of later studies published in 2010 and 2011 indicate that the leopard and the lion belong to the same sister group, which diverged 1.95–3.10 million years ago.^[9][10] However, hybridisation between lion and snow leopard populations may have continued until about 2.1 million years ago.^[13]

Subspecies

In the 19th and 20th centuries, several lion type specimen were described and proposed as subspecies.^[14] Between 2008 and 2016, IUCN Red List assessors for lions used only two subspecific names, P. l. leo for African lion populations and P. l. persica for the Asiatic lion population.^[15][16][2] In 2017, the Cat Classification Task Force of the Cat Specialist Group revised classification of the subspecies as follows:^[14]

• P. l. leo (Linnaeus, 1758) is the nominate lion subspecies and includes the regionally extinct North African, as well as the Asian, West and Central African lion populations.^[14] Former synonyms include P. l. persica (Meyer, 1826), P. l. senegalensis (Meyer, 1826), P. l. kamptzi (Matschie, 1900), and P. l. azandica (Allen, 1924).^[1]
• P. l. melanochaita (Smith, 1842) includes the regionally extinct Cape lion, as well as the Southern African and East African lion populations.^[14] Former synonyms include P. l. massaica (Neumann, 1900), P. l. sabakiensis (Lönnberg, 1910), P. l. bleyenberghi (Lönnberg, 1914), P. l. roosevelti (Heller, 1914), P. l. nyanzae (Heller, 1914), P. l. hollisteri (Allen), 1924), P. l. krugeri (Roberts, 1929), P. l. vernayi (Roberts, 1948)^[1] and P. l. webbiensis (Zukowsky, 1964).^[17]

There is phylogenetic evidence that Ethiopia was a contact zone between the two subspecies.^[18][14]

Extinct species and subspecies

Other lion subspecies or sister species to the modern lion existed in prehistoric times:^[19]

• Panthera leo fossilis bone fragments were excavated in Germany, United Kingdom, Italy and Czech Republic, and are estimated at between 680,000 and 600,000 years old.^[20] It was larger than the modern lion.^[21][22]
• Panthera spelaea bone fragments were excavated in Europe, North Asia, Canada and Alaska. It probably became extinct between 14,900 and 11,900 years ago,^[23] the oldest known bone fragments are estimated between 109,000 and 57,000 years old.^[24] It is depicted in Paleolithic cave paintings, ivory carvings, and clay busts, which show it with protruding ears, tufted tails, and faint tiger-like stripes.^[25] A few had a ruff around their necks.^[26]
• P. l. atrox or P. atrox, known as the American lion or American cave lion, existed in the Americas from Canada to Peru in the Pleistocene Epoch until about 10,000 years ago. This form is a sister clade of P. spelaea, and likely arose when an early P. spelaea population became isolated south of the North American continental ice sheet about 0.34 Mya.^[27] It is among the largest purported lion subspecies to have existed, its body length is estimated to have been 1.6–2.5 m (5.2–8.2 ft).^[28]

Dubious subspecies

• P. l. youngi or Panthera youngi, flourished 350,000 years ago.^[29] Its relationship to the extant lion subspecies is obscure, and it probably represents a distinct species.
• P. l. mesopotamica was described on the basis of a relief from the Neo-Assyrian Period, about 1000–600 BC, in ancient Mesopotamia.^[30]
• P. l. europaea was proposed for subfossil remains of lions excavated in Southern Europe that date to the Late Neolith to the Early Iron Age.^[31]
• P. l. maculatus, known as the Marozi or spotted lion, sometimes is thought to be a distinct subspecies, but may be an adult lion that has retained its juvenile spotted pattern. If it was a subspecies in its own right, rather than a small number of aberrantly coloured individuals, it has been extinct since 1931. A less likely identity is a natural leopard-lion hybrid commonly known as a leopon.^[32]

Hybrids

Lions have been known to breed with tigers (most often the Siberian and Bengal tigers) to create hybrids called 'ligers' and 'tiglons' (or 'tigons').^[33][34] They also have been crossed with leopards to produce leopons,^[35] such hybrid breeding is now discouraged due to the emphasis on conserving species and subspecies. Hybrids are still bred in private menageries and in zoos in China.

The liger is a cross between a male lion and a tigress,^[33] because the growth-inhibiting gene from the female tiger mother is absent, the growth-promoting gene passed on by the male lion father is unimpeded by a regulating gene and the resulting ligers grow far larger than either parent. They share physical and behavioural qualities of both parent species (spots and stripes on a sandy background). Male ligers are sterile, but female ligers often are fertile. Males have about a 50% chance of having a mane, but if they grow them, their manes will be modest: around 50% the size of a pure lion mane. Ligers are much bigger than normal lions and tigers, typically 3.65 m (12.0 ft) in length, and can weigh up to 500 kg (1,100 lb).^[33][34]

The less common tiglon or tigon is a cross between a lioness and a male tiger;^[36] in contrast to ligers, tigons are often relatively small in comparison to their parents, because of reciprocal gene effects.^[34]

Evolution and genetic diversity

The lion evolved in Africa between 1 million and 800,000 years ago, from where it spread throughout the Holarctic region,^[38] the earliest fossil record in Europe was found near Pakefield in the United Kingdom and is about 680,000 years old.^[20] From this lion the late Pleistocene Eurasian cave lion probably derived about 300,000 years ago.^[39] Fossil remains found in the Cromer Forest Bed suggest that it was of a gigantic size and represented a lineage that was genetically isolated and highly distinct from lions in Africa and Asia,^[40] it was distributed throughout Europe, across Siberia and into western Alaska, via the Beringian landmass. The gradual formation of dense forest likely caused the decline of its geographic range near the end of the Late Pleistocene. Frequently encountered lion bones in cave deposits from Eemian times suggest that the cave lion survived in the Balkans and Asia Minor. There was probably a continuous population extending into India.^[41] Fossil lion remains were found in Pleistocene deposits in West Bengal,^[42] it became extinct about 10,000 years ago at the end of the last glacial period without mitochondrial descendants on other continents.^[21][40]

A fossil carnassial found in the Batadomba Cave indicates that Panthera leo sinhaleyus inhabited Sri Lanka during the late Pleistocene, and is thought to have become extinct around 39,000 years ago. This subspecies was described by Deraniyagala in 1939, it is distinct from the contemporary lion.^[43]

The modern lion probably originated in East and Southern Africa about 100,000 years ago,^[18] during the last glacial maximum until about 20,000 years ago, it was likely distributed throughout most of Southern and Central Africa, and expanded its range northwards during the early Holocene about 10,000 to 4,000 years ago.^[44] Early phylogenetic research was focused on lions from eastern and southern parts of Africa, and already showed that they can possibly be divided in two main clades: one to the west of the Great Rift Valley and the other to the east. Lions in eastern Kenya are genetically much closer to lions in Southern Africa than to lions in the Aberdare National Park in western Kenya.^[45]

In a subsequent study, tissue and bone samples of 32 lion specimens in museums were used. Results indicated that lions form three phylogeographic groups, one each in North Africa and Asia, in Central Africa and in Southern Africa.^[44] Samples of 53 lions, both wild and captive individuals, from 15 countries were used for phylogenetic analysis. Results showed little genetic diversity among lion samples from Asia, West and Central Africa, whereas samples from East and Southern Africa revealed numerous mutations indicating that this group has a longer evolutionary history.^[46] Results of another phylogeographic study indicate that the two groups probably diverged about 186,000–128,000 years ago, it is thought that the Asiatic lion remained connected to North and Central African lions until gene flow was interrupted due to extinction of lion populations in the Middle East and Europe. Haplotypes from the Central African lion group were found in nine of 19 lion samples from Ethiopia, indicating that the Great Rift Valley was not a complete barrier to gene flow, but that this region was a genetic admixture zone between lions in Central and East Africa.^[18]

Approximately 77% of the captive lions registered by the International Species Information System are of unknown origin. Nonetheless, they might carry genes that are extinct in the wild, and might be therefore important to maintain overall genetic variability of the lion,^[47] it is thought that those lions, imported to Europe before the middle of the 19th century, were mainly either Barbary lions from North Africa, or Cape lions from Southern Africa.^[48]

Description

The lion is a muscular, deep-chested cat with a short, rounded head, a reduced neck and round ears, its fur varies from light buff to silvery grey, to yellowish red and dark brown.^[49] The underparts are generally lighter, and cubs are born with dark spots on their bodies, the spots fade as lions reach adulthood, although faint spots often may still be seen on the legs and underparts. The lion is the only member of the cat family that displays obvious sexual dimorphism. Males are more robust than females, have broader heads and a prominent mane, which grows downward and backward and covers most of the head, neck, shoulders, and chest, the mane is typically brownish and tinged with yellow, rust, and black hairs. The tail ends in a dark, hairy tuft; in some lions, the tuft conceals a hard "spine" or "spur", approximately 5 mm (0.20 in) long, formed of the final sections of tail bone fused together. The function of the tuft and spine are unknown. Absent at birth, the tuft develops around ​5 ¹⁄₂ months of age and is readily identifiable at the age of seven months.^[50]

Of the living, non-hybrid felids, the lion is rivalled only by the tiger in length, weight and height at the shoulder,^[51] its skull is very similar to that of the tiger, although the frontal region is usually more depressed and flattened, with a slightly shorter postorbital region and broader nasal openings than that of a tiger. Due to the amount of skull variation in the two species, usually only the structure of the lower jaw can be used as a reliable indicator of species.^[33][34][52] The size and weight of adult lions varies across global range and habitats.^{[53][54][55][56]}

Average	Female lions	Male lions
Head-to-body length	140–175 cm (4 ft 7 in–5 ft 9 in)[56]	170–298 cm (5 ft 7 in–9 ft 9 in)[55]
Tail length	70–100 cm (2 ft 4 in–3 ft 3 in)[56]	90–105 cm (2 ft 11 in–3 ft 5 in)[56]
Weight	120–182 kg (265–401 lb),[56] 124.2–139.8 kg (274–308 lb) in Southern Africa,[53] 119.5 kg (263 lb) in East Africa,[53] 110–120 kg (240–260 lb) in India[54]	150–250 kg (330–550 lb),[56] 187.5–193.3 kg (413–426 lb) in Southern Africa,[53] 174.9 kg (386 lb) in East Africa,[53] 160–190 kg (350–420 lb) in India[54]

Accounts of a few individuals that were larger than average exist from Africa and India.^{[57][58][59][60]} Pleistocene forms like the American lion reached a maximum head-to-body length of 250 cm (8 ft 2 in).^[61]

Mane

The lion's mane is the most recognisable feature of the species,^[17] the mane starts growing when lions are about a year old. Mane colour varies, and darkens with age. Research results indicate that environmental factors such as average ambient temperature influence the mane's colour and size. Mane length apparently signals fighting success in male–male relationships. Darker-maned individuals may have longer reproductive lives and higher offspring survival, although they suffer in the hottest months of the year, the presence, absence, colour, and size of the mane is associated with genetic precondition, sexual maturity, climate, and testosterone production; the rule of thumb is the darker and fuller the mane, the healthier the lion. In the Serengeti National Park, female lions favour males as mates with dense, dark manes,^[62][63] the main purpose of the mane is thought to protect the lion's neck and throat in territorial fights with rivals.^[64] The cooler ambient temperature in European and North American zoos may result in a heavier mane,^[47] while Asiatic lions usually have sparser manes than average African lions.^[65]

In the area of Pendjari National Park, almost all West African males are maneless or have very weak manes.^[66] Maneless male African lions have also been reported from Senegal, from Sudan's Dinder National Park, and from Tsavo East National Park in Kenya.^[67] The original male white lion from Timbavati was also maneless, the testosterone hormone has been linked to mane growth; therefore, castrated lions often have minimal to no mane, as the removal of the gonads inhibits testosterone production.^[68] Increased testosterone may be the cause of maned lionesses reported from northern Botswana.^[69]

Cave paintings of extinct European cave lions almost exclusively show hunting animals with no manes. Some suggest this as evidence that the males of this species were maneless,^[26] however, since the hunting usually involved groups of lionesses, this presumption remains unproven; in the Chauvet cave, there is a sketchy drawing of two maneless lions, appearing to be walking side by side. One is mostly obscured behind the other, with the former being larger than the latter, and shown with a scrotum.^[38]

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, but has normal pigmentation in eyes and skin. White lion individuals have been occasionally encountered only 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 different prides between 2007 and 2015.^[70] White lions are selected for breeding in captivity.^[71] Reportedly, they have been bred in camps in South Africa for use as trophies to be killed during canned hunts.^[72]

A melanistic Asiatic lion from Khuzestan in Iran was described by Austen Henry Layard, which was dark brown with nearly black patches.^[73]

Behaviour and ecology

Lions spend much of their time resting, and are inactive for about 20 hours per day.^[74] Although lions can be active at any time, their activity generally peaks after dusk with a period of socialising, grooming, and defecating. Intermittent bursts of activity follow through the night hours until dawn, when hunting most often takes place, they spend an average of two hours a day walking, and 50 minutes eating.^[75]

Group organisation

The lion is the most social of all wild cat species, living in groups of related individuals with their offspring, such a group is called a pride. Male lion groups are called a coalition.^[76] Females form the stable social unit in a pride and do not tolerate outside females.^[77] Membership only changes with the births and deaths of lionesses,^[78] although some females leave and become nomadic,^[79] the average pride consists of around 15 lions, including several adult females and up to four males and their cubs of both sexes. Large prides, consisting of up to 30 individuals, have also been observed,^[80] the sole exception to this pattern is the Tsavo lion pride which always has just one adult male.^[81] Male cubs are excluded from their maternal pride when they reach maturity at around 2–3 years of age.^[79]

Another lion behaviour is labeled nomads: lions who range widely and move about sporadically, either singularly or in pairs.^[76] 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.^[82] Interactions between prides and nomads tend to be hostile, although pride females in estrous allow nomad males to approach them.^[83] Males spend years in a nomadic phase before gaining residence in a pride.^[84] A study in the Serengeti National Park revealed that nomadic coalitions gain residency at between 3.5 and 7.3 years of age.^[85]

The area a pride occupies is called a pride area, whereas that by a nomad is a range,^[76] the males associated with a pride tend to stay on the fringes, patrolling their territory. Why sociality – the most pronounced in any cat species – has developed in lionesses is the subject of much debate. Increased hunting success appears an obvious reason, but this is less than sure upon examination: coordinated hunting does allow for more successful predation but also ensures that non-hunting members reduce per capita calorific intake; however, some take a role raising cubs, who may be left alone for extended periods of time. Members of the pride regularly tend to play the same role in hunts and hone their skills, the health of the hunters is the primary need for the survival of the pride, and they are the first to consume the prey at the site it is taken. Other benefits include possible kin selection (better to share food with a related lion than with a stranger), protection of the young, maintenance of territory, and individual insurance against injury and hunger.^[58]

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,^[86] some individuals consistently lead the defence against intruders, while others lag behind.^[87] Lions tend to assume specific roles in the pride, those lagging behind may provide other valuable services to the group.^[88] An alternative hypothesis is that there is some reward associated with being a leader who fends off intruders, and the rank of lionesses in the pride is reflected in these responses,^[89] the male or males associated with the pride must defend their relationship to the pride from outside males who attempt to take over their relationship with the pride.

Asiatic lion prides differ from African prides in group composition. Male Asiatic lions are solitary or associate with up to three males forming a loose pride. Pairs of males rest and feed together, and display marking behaviour at the same sites. Females associate with up to 12 females forming a stronger pride together with their cubs, they share large carcasses among each other, but seldom with males. Female and male lions associate only when mating.^[90] Coalitions of males hold a territory for a longer time than single lions. Males in coalitions of three to four individuals exhibit a pronounced hierarchy with one male dominating the others. Dominant males mate more frequently than their coalition partners, during a study carried out between December 2012 and December 2016, three females were observed switching mating partners in favour of the dominant male.^[91]

Hunting and diet

The lion is a generalist hypercarnivore^[92] and usually hunts in groups. Its prey consists mainly of mammals, particularly ungulates, with a preference for wildebeest, zebras, buffalo, gemsbok, and giraffes in Africa,^[93] and chital, sambar deer, nilgai, wild boar, chinkara and chousingha in India.^[94] Because of its wide prey spectrum, the lion is considered to be an apex and keystone predator.^[95] African lions prefer prey weighing 190–550 kg (420–1,210 lb).^[93] They generally avoid fully grown adult elephants, hippopotamuses and rhinoceroses, as well as very small prey like dik-dik, hyrax, hare and vervet monkey.^[96] However, Thomson's gazelles may be hunted^[97] and warthogs are often taken depending on availability, despite being below the preferred weight range.^[93] In many areas, a small number of species may make up around three-fourths of the lion's diet; in Serengeti National Park, wildebeest, zebras and gazelle are the majority of prey.^[98] In Kruger National Park, giraffes are the most common prey;^[99] in Manyara Park, Cape buffaloes constitute as much as 62% of the lion's diet.^[100] In the Okavango Delta, with its strong seasonal changes in prey, up to eight species may make up three quarters of a lion's diet.^[98] Occasionally adult hippopotamus are taken at Gorongosa National Park and calves are commonly hunted at Virunga National Park; in addition to size, the aquatic nature of hippos makes them normally unavailable as prey.^[92] The lions of Savuti, Botswana, have adapted to hunting young elephants during the dry season, and a pride of 30 lions has been recorded killing individuals between the ages of four and eleven years; in at least one occasion, an adult elephant was observed to have been taken, and due to its immense size compared to lions, the ratio of prey to predator size for the lion is 10–15 : 1, the highest for any known terrestrial mammalian predator.^[101] Lions also attack domestic livestock and in India cattle contribute significantly to their diet;^[65][94] in what was Somaliland in the 20th century, lions were even recorded to have jumped over zaribas that were 10–12 ft (3.0–3.7 m) high or more to obtain livestock.^[102] Unusual prey items include porcupines and small reptiles. Lions will kill other predators such as leopards, cheetahs, and hyenas, but they seldom devour them.^[103]

Young lions first display stalking behaviour around three months of age, although they do not participate in hunting until they are almost a year old, they begin to hunt effectively when nearing the age of two.^[104] Single lions are capable of bringing down prey like zebra and wildebeest, which can be twice their own weight, while hunting larger prey like giraffes and buffalo alone is too much of a risk. Cooperative-hunting lions are usually successful;^[82][97] in prides, lionesses do most of the hunting.^[82] 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 in flight from other lionesses. Males attached to prides do not usually participate in group hunting.^[105] However, some evidence suggests that pride males are just as successful as females; they are solo hunters who ambush prey in small bush.^[106] Lions are not particularly known for their stamina – for instance, a lioness' heart makes up only 0.57% of her body weight (a male's is about 0.45% of his body weight), whereas a hyena's heart is close to 1% of its body weight.^[107] Thus, they only run fast in short bursts,^[108] and need to be close to their prey before starting the attack, they take advantage of factors that reduce visibility; many kills take place near some form of cover or at night.^[109] In addition, since lions are such ambush hunters, humans farming in the vicinity have recently found that lions are easily discouraged if they think their prey has spotted them. To protect their cattle from such attacks with that knowledge in mind, farmers have found that all that they have to do is simply paint eyes on the hindquarters of each cow, which is usually enough for hunting lions to think they are spotted and move to easier prey.^[110]

The attack is short and powerful; they attempt to catch the victim with a fast rush and final leap. The prey usually is killed by strangulation,^[111] which can cause cerebral ischemia or asphyxia (which results in hypoxemic, or "general", hypoxia). The prey also may be killed by the lion enclosing the animal's mouth and nostrils in its jaws (which would also result in asphyxia).^[56] Prey is typically eaten at the location of the hunt, although large prey is sometimes dragged into cover.^[112][113] Lions tend to squabble over a kill, particularly the males. When food is scarce, cubs tend to suffer the most but otherwise all pride members can eat their fill, including old and crippled ones which can live on leftovers.^[82] There is more sharing of larger kills.^[114] An adult lioness requires an average of about 5 kg (11 lb) of meat per day, a male about 7 kg (15 lb).^[115] A lion may gorge itself and eat up to 30 kg (66 lb) in one sitting;^[116] if it is unable to consume all the kill it will rest for a few hours before consuming more. On a hot day, the pride may retreat to shade leaving a male or two to stand guard.^[112] Lions will defend their kills from scavengers like vultures and hyenas.^[82]

Lions scavenge on carrion when the opportunity arises, they scavenge animals either dead from natural causes like diseases, or were killed by other predators, and keep a constant lookout for circling vultures, being keenly aware that they indicate an animal dead or in distress.^[117] In fact, most carrion on which both hyenas and lions feed upon are killed by the hyenas instead of the lions.^[56] Carrion is thought to provide a large part of lion diet.^[118]

Predator competition

African lions and spotted hyenas occupy a similar ecological niche and compete for prey and carrion in the areas where they coexist. A review of data across several studies indicates a dietary overlap of 58.6%.^[119] Lions typically ignore spotted hyenas unless the lions are on a kill or are being harassed by the hyenas, while the latter tend to visibly react to the presence of lions whether there is food or not. Lions seize the kills of spotted hyenas: in the Ngorongoro crater, it is common for lions to subsist largely on kills stolen from hyenas, causing the hyenas to increase their kill rate;^[120] in Botswana's Chobe National Park, the situation is reversed: hyenas frequently challenge lions and steal their kills: they obtain food from 63% of all lion kills.^[121] When confronted on a kill by lions, spotted hyenas may either leave or wait patiently at a distance of 30–100 m (98–328 ft) until the lions have finished,^[122] but they are also bold enough to feed alongside lions, and even force the lions off a kill. The two species may attack one another even when there is no food involved for no apparent reason.^[123][124] Lion predation can account for up to 71% of hyena deaths in Etosha National Park. Spotted hyenas have adapted by frequently mobbing lions that enter their territories.^[125] Where the lion population declined in Kenya's Masai Mara National Reserve, the spotted hyena population increased rapidly.^[126] Experiments on captive spotted hyenas revealed that specimens with no prior experience with lions act indifferently to the sight of them, but will react fearfully to the scent,^[120] the size of male lions allows them occasionally to confront hyenas in otherwise evenly matched brawls and so to tip the balance in favour of the lions.

Lions tend to dominate smaller felids such as African cheetahs and leopards where they co-occur, stealing their kills and killing their cubs and even adults when given the chance,^[127] the cheetah in particular has a 50% chance of losing its kill to lions or other predators.^[128] Lions are major killers of cheetah cubs, accounting for up to 78.2% of predator-killed juveniles in one study.^[129] Cheetahs avoid their competitors using different temporal (time) and spatial (habitat) niches.^[130] Leopards are able to take refuge in trees; however, lionesses will occasionally be successful in climbing to retrieve leopard kills.^[131] Similarly, lions dominate African wild dogs, not only taking their kills but also preying on young and (rarely) adult dogs. Population densities of wild dogs are low in areas where lions are more abundant.^[132] However, there are a few reported cases of old and wounded lions falling prey to wild dogs.^[99][133] African lions may also conflict with Nile crocodiles. Depending on the size of the crocodile and the lion, either can lose kills or carrion to the other. Lions have been known to kill crocodiles venturing onto land,^[134] while the reverse is true for lions entering waterways, as evidenced by the occasional lion claw found in crocodile stomachs.^[135]

As for the Asiatic lion in India, sympatric predators currently include the Indian leopard,^[58][136] mugger crocodile,^[137] Golden jackal^[138] and striped hyena.^[52][139]

Reproduction and life cycle

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Most lionesses will have reproduced by the time they are four years of age.^[140] Lions do not mate at any specific time of year, and the females are polyestrous,^[141] as with other cats' penises, 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.^[142][143] A lioness may mate with more than one male when she is in heat.^[144]

The average gestation period is around 110 days,^[141] the female giving birth to a litter of one to 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 rest of the pride. She will often hunt by herself while the cubs are still helpless, staying relatively close to the thicket or den where the cubs are kept,^[145] the cubs themselves are born blind – their eyes do not open until roughly a week after birth. They weigh 1.2–2.1 kg (2.6–4.6 lb) at birth and are almost helpless, beginning to crawl a day or two after birth and walking around three weeks of age.^[146] 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, to prevent scent from building up at a single den site and thus avoiding the attention of predators that may harm the cubs.^[145]

Usually, the mother does not integrate herself and her cubs back into the pride until the cubs are six to eight weeks old.^[145] Sometimes this introduction to pride life occurs earlier, however, particularly if other lionesses have given birth at about the same time, for instance, lionesses in a pride often synchronise their reproductive cycles and there is communal raising and suckling of the young (once the cubs are past the initial stage of isolation with their mother), who suckle indiscriminately from any or all of the nursing females in the pride. The synchronization of births also has an advantage in that the cubs end up being roughly the same size, and thus have an equal chance of survival, and older cubs do not dominate the sucklings.^[82][147]

When first introduced to the rest of the pride, the cubs initially lack confidence when confronted with adult lions 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.^[147] Lionesses with cubs of their own are more likely to be tolerant of another lioness's cubs than lionesses without cubs, the tolerance of the male lions toward the cubs varies – sometimes, a male will patiently let the cubs play with his tail or his mane, whereas another may snarl and bat the cubs away.^[148]

Weaning occurs after six to seven months. Male lions reach maturity at about 3 years of age and, at 4–5 years of age, are capable of challenging and displacing the adult male(s) associated with another pride, they begin to age and weaken between 10 and 15 years of age at the latest,^[149] Furthermore, when one or more new males oust the previous male(s) associated with a pride, the conqueror(s) often kill any existing young cubs, perhaps because females do not become fertile and receptive until their cubs mature or die. A lioness often will attempt to defend her cubs fiercely from a usurping male, but such actions are rarely successful. Success is more likely when a group of three or four mothers within a pride join forces against one male.^[150] Other sources of mortality for cubs include starvation and abandonment, as well as predation by leopards, hyenas and wild dogs.^[133][82] All in all, as many as 80% of the cubs will die before the age of two.^[151]

Contrary to popular belief, it is not only males that are ousted from their pride to become nomads, although most females certainly do remain with their birth pride. However, when the pride becomes too large, the next generation of female cubs may be forced to leave to eke out their own territory. Furthermore, when a new male lion takes over the pride, adolescent lions, both male and female, may be evicted.^[152]

Both males and females may interact homosexually. Lions are shown to be involved in group homosexual and courtship activities. Male lions will also head rub and roll around with each other before simulating sex together.^[153][154]

Health

Although adult lions have no natural predators, evidence suggests that the majority die violently from humans or other lions.^[155] Lions often inflict serious injuries on each other, either members of different prides encountering each other in territorial disputes, or members of the same pride fighting at a kill.^[156] Crippled lions and lion cubs may fall victim to hyenas, leopards, or be trampled by buffalo or elephants, and careless lions may be maimed when hunting prey.^[157]

Various species of tick commonly infest the ears, neck and groin regions of most lions.^[158][159] Adult forms of several species of the tapeworm genus Taenia have been isolated from intestines, the lions having ingested larval forms from antelope meat.^[160] Lions in the Ngorongoro Crater were afflicted by an outbreak of stable fly (Stomoxys calcitrans) in 1962; this resulted in lions becoming covered in bloody bare patches and emaciated. Lions sought unsuccessfully to evade the biting flies by climbing trees or crawling into hyena burrows; many perished or emigrated as the population dropped from 70 to 15 individuals.^[161] A more recent outbreak in 2001 killed six lions.^[162] Lions, especially in captivity, are vulnerable to the canine distemper virus (CDV), feline immunodeficiency virus (FIV), and feline infectious peritonitis (FIP).^[11] CDV is spread through 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.^[163] FIV, which is similar to HIV while not known to adversely affect lions, is worrisome enough in its effect in domestic cats that the Species Survival Plan recommends systematic testing in captive lions, it occurs with high to endemic frequency in several wild lion populations, but is mostly absent from Asiatic and Namibian lions.^[11]

Communication

When resting, lion socialisation occurs through a number of behaviours, and the animal's expressive movements are highly developed, the most common peaceful tactile gestures are head rubbing and social licking,^[164] which have been compared with grooming in primates.^[165] Head rubbing – nuzzling one's forehead, face and neck against another lion – appears to be a form of greeting,^[166] as it 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.^[167] 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, which may have arisen out of utility, as a lion cannot lick these areas individually.^[168]

	Lion roar A lion in captivity roaring
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Lions have an array of facial expressions and body postures that serve as visual gestures.^[169] 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.^[170] Lions also use chemical and visual marking; male lions will spray and scrape both plots of ground and objects within their territory.^[169]

Their repertoire of vocalisations is also large; variations in intensity and pitch, rather than discrete signals, appear central to communication. Most lion vocals are variations of growling/snarling, miaowing and roaring. Other sounds produced include purring, puffing, bleating and humming.^[171] Lions tend to roar in a very characteristic manner, starting with a few deep, long roars that trail off into a series of shorter ones,^[172][173] they most often roar at night; the sound, which can be heard from a distance of 8 kilometres (5.0 mi), is used to advertise the animal's presence.^[171] Lions have the loudest roar of any big cat.

Distribution and habitat

The lion prefers grassy plains and savannahs, open woodlands with bushes and scrub bordering rivers, it is absent in rainforest and rarely enters closed forest. On Mount Elgon, it has been recorded up to an elevation of 3,600 m (11,800 ft) and close to the snow line on Mount Kenya.^[57]

Lion range in Africa originally spanned most of the central rainforest-zone and the Sahara desert.^[174] Lions occur in savanna grasslands with scattered Acacia trees, which serve as shade,^[175] the species became extinct in North Africa in the 1960s.^[176]

In Eurasia, the lion once ranged from Greece to India. Herodotus reported that lions had been common in Greece in 480 BC; they attacked the baggage camels of the Persian king Xerxes on his march through the country. Aristotle considered them rare by 300 BC. By 100 CE, they were extirpated.^[174] A population of Asiatic lion survived until the 10th century in the Caucasus, their last European outpost,^[52] the species was eradicated in Palestine by the Middle Ages, and from most of the rest of Asia after the arrival of readily available firearms in the 18th century. Between the late 19th and late 20th centuries, they became extinct in Southwest Asia. By the late 19th century, the lion had been extirpated in most of northern India and Turkey,^[73][177] the last live lion in Iran was sighted in 1942, about 65 km (40 mi) northwest of Dezful.^[178] The corpse of a lioness was found on the banks of the Karun river, Khūzestān Province in 1944. There are no subsequent reliable reports from Iran.^[116]

The Asiatic lion now survives only in and around Gir Forest National Park in Gujarat, western India, its habitat is a mixture of dry savanna forest and very dry deciduous scrub forest.^[15]

Population and conservation status

Conservation of both African and Asian lions has required the setup and maintenance of national parks and game reserves; among the best known are Etosha National Park in Namibia, Serengeti National Park in Tanzania, Kruger National Park in South Africa, and Gir National Park in India.

In Africa

Most lions now live in East and Southern Africa, and their numbers are rapidly decreasing, with an estimated 30–50% decline per 20 years in the late half of the 20th century. Therefore, the species is listed as Vulnerable on the IUCN Red List;^[2] in 1975, it was estimated that since the 1950s, lion numbers decreased by half to 200,000 and perhaps even less.^[179] Estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004.^[180][181] Primary causes of the decline include disease and human interference. Habitat loss and conflicts with humans are considered the most significant threats to the species.^[2]

The Ewaso Lions Project protects lions in the Samburu National Reserve, Buffalo Springs National Reserve and Shaba National Reserve of the Ewaso Ng'iro ecosystem in Northern Kenya.^[182] Outside these areas, the issues arising from lions' interaction with livestock and people usually results in the elimination of the lions.^[183]

Zambia's Kafue National Park is a key refuge for lions, where frequent, uncontrolled bushfires combined with hunting of lions and prey species limits the ability of the lion population to recover. When favourable habitat is inundated in the wet season, lions expand home ranges and travel greater distances, and cub mortality is high.^[184]

In 2015, a population of lions that was previously thought extirpated was filmed in the Alatash National Park, Ethiopia, close to the Sudanese border, this population might number up to 200 animals.^[185][186]

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 less than 250 mature individuals. They persist in only three protected areas in the region, with a majority in one population in the tri-national WAP protected area complex, this population listed as Critically Endangered.^[16] 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.^[187] A population occurs in Cameroon's Waza National Park, where approximately 14–21 animals persisted as of 2009.^[188] There is disagreement over the size of the largest individual population in West Africa: the estimates range from 100 to 400 lions in Burkina Faso's Arly-Singou ecosystem;^[2] 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.^[189]

In Gabon's Batéké Plateau National Park, a single male lion was repeatedly recorded by camera-traps between January 2015 and September 2017. Five hair samples of this lion were collected and compared with samples from museum specimens that had been shot in the area in 1959. Genetic analysis revealed that the Batéké lion is closely related to lions killed in this region in the past, the samples grouped with lion samples from Namibia and Botswana. Thus it is possible that the Batéké lion either dispersed from a Southern African lion population, or is a survivor of the ancestral Batéké lion population that was considered to be extinct since the late 1990s.^[190]

In the Republic of the Congo, the Odzala-Kokoua National Park was considered a lion stronghold in the 1990s. By 2014, no lions were recorded in the protected area, so that the population is considered locally extinct in the country;^[191] in the Democratic Republic of the Congo, there are about 175 lions in Garamba National Park and Bili-Uere Hunting Reserve, and 60 in Virunga National Park. The latter form a contiguous population with lions in Uganda;^[192][193] in 2010, the lion population in Uganda was estimated at 408 ± 46 individuals in three protected areas including Queen Elizabeth National Park.^[194] Little is known about lion distribution and population sizes in adjacent South Sudan; in Sudan, lions were reported in Southern Darfur and Southern Kordofan provinces in the 1980s.^[181][193]

In Asia

The last refuge of the Asiatic lion is the 1,412 km² (545 sq mi) Gir Forest National Park and surrounding areas in India. The population has risen from approximately 180 lions in 1974 to about 400 in 2010,^[195] it is geographically isolated, which can lead to inbreeding, and consequently, reduced genetic diversity. Since 2008, it has been listed as Endangered on the IUCN Red List.^[15] By 2015, the population had grown to 523 individuals inhabiting an area of 7,000 km² (2,700 sq mi) in the Saurashtra region of Gujerat state.^{[196][197][198]} The Asiatic Lion Census conducted in 2017 revealed about 650 individuals.^[199]

Numerous human habitations are close by with the resultant conflict between lions, local people and their livestock,^[200][196] the establishment of a second independent Asiatic lion population was planned in Kuno Wildlife Sanctuary located in Madhya Pradesh. But in 2017, the Asiatic Lion Reintroduction Project seemed unlikely to be implemented.^[201][202]

In captivity

Following the discovery of the decline of lion population in Africa, several coordinated efforts involving lion conservation have been organised in an attempt to stem this decline. Lions are one species included in the Species Survival Plan, a coordinated attempt by the Association of Zoos and Aquariums to increase its chances of survival. The plan was originally started in 1982 for the Asiatic lion, but was suspended when it was found that most Asiatic lions in North American zoos were not genetically pure, having been hybridised with African lions, the African lion plan started in 1993, focusing especially on the South African subspecies, although there are difficulties in assessing the genetic diversity of captive lions, since most individuals are of unknown origin, making maintenance of genetic diversity a problem.^[11]

Lions are part of a group of exotic animals that are the core of zoo exhibits since the late eighteenth century; members of this group are invariably large vertebrates and include elephants, rhinoceroses, hippopotamuses, large primates, and other big cats; zoos sought to gather as many of these species as possible.^[203] Although many modern zoos are more selective about their exhibits,^[203] 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.^[204] Lions can reach an age of over 20 years in captivity; Apollo, a resident lion of Honolulu Zoo in Honolulu, Hawaii, died at age 22 in August 2007. His two sisters, born in 1986, were still alive in August 2007.^[205] Breeding programs need to note origins to avoid breeding different subspecies and thus reducing conservation value.^[206] However, several Asiatic-African lion crosses have been bred.^[207]

The former popularity of the Barbary lion as a zoo animal has meant that captive lions likely descended from Barbary lion stock, this includes lions at Port Lympne Wild Animal Park in Kent, England that are descended from animals owned by the King of Morocco.^[208] Another eleven animals thought to be Barbary lions are kept in Addis Ababa Zoo, and are descendants of animals owned by Emperor Haile Selassie. WildLink International, in collaboration with Oxford University, launched their ambitious International Barbary Lion Project with the aim of identifying and breeding Barbary lions in captivity for eventual reintroduction into a national park in the Atlas Mountains of Morocco.^[209]

At the ancient Egyptian cities of Taremu and Per-Bast were temples dedicated to the lion goddesses of Egypt, Sekhmet and Bast, and at Taremu there was a temple dedicated to the son of the deity, Maahes the lion prince, where live lions were kept and allowed to roam within the temple. The Greeks called the city Leontopolis the "City of Lions" and documented that practice. Lions were kept and bred by Assyrian kings as early as 850 BC,^[174] and Alexander the Great was said to have been presented with tame lions by the Malhi of northern India.^[210] In Ancient Rome, lions were kept by emperors to take part in the gladiator arenas or for executions (see bestiarii, damnatio ad bestias, and venatio). Roman notables, including Sulla, Pompey, and Julius Caesar, often ordered the mass slaughter of hundreds of lions at a time.^[211] In India, lions were tamed by Indian princes. Marco Polo reported that Kublai Khan kept lions.^[212]

The first European "zoos" spread among noble and royal families in the 13th century, and until the 17th century were called seraglios; at that time, they came to be called menageries, an extension of the cabinet of curiosities. They spread from France and Italy during the Renaissance to the rest of Europe;^[213] 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,^[214][215] probably stocked with animals from an earlier menagerie started in 1125 by Henry I at his hunting lodge in Woodstock, near Oxford, where lions had been stocked according to William of Malmesbury.^[216]

Seraglios served as expressions of the nobility's power and wealth. Animals such as big cats and elephants, in particular, symbolised power, and were pitted in fights against each other or domesticated animals. By extension, menageries and seraglios served as demonstrations of the dominance of humanity over nature. Consequently, the defeat of such natural "lords" by a cow in 1682 astonished the spectators, and the flight of an elephant before a rhinoceros drew jeers, such fights would slowly fade out in the 17th century with the spread of the menagerie and their appropriation by the commoners. The tradition of keeping big cats as pets lasted into the 19th century, at which time it was seen as highly eccentric.^[217]

The presence of lions at the Tower of London was intermittent, being restocked when a monarch or his consort, such as Margaret of Anjou the wife of Henry VI, either sought or were given animals. Records indicate they were kept in poor conditions there in the 17th century, in contrast to more open conditions in Florence at the time,^[218] the menagerie was open to the public by the 18th century; admission was a sum of three half-pence or the supply of a cat or dog for feeding to the lions.^[219] A rival menagerie at the Exeter Exchange also exhibited lions until the early 19th century,^[220] the Tower menagerie was closed by William IV,^[219] and animals transferred to the London Zoo, which opened its gates to the public on 27 April 1828.^[221]

The wild animals trade flourished alongside improved colonial trade of the 19th century. Lions were considered fairly common and inexpensive, although they would barter higher than tigers, they were less costly than larger, or more difficult to transport animals such as the giraffe and hippopotamus, and much less than giant pandas.^[222] Like other animals, lions were seen as little more than a natural, boundless commodity that was mercilessly exploited with terrible losses in capture and transportation,^[223] the widely reproduced imagery of the heroic hunter chasing lions would dominate a large part of the century.^[224] Explorers and hunters exploited a popular Manichean division of animals into "good" and "evil" to add thrilling value to their adventures, casting themselves as heroic figures, this resulted in big cats always suspected of being man-eaters, representing "both the fear of nature and the satisfaction of having overcome it."^[225]

Lions were kept in cramped and squalid conditions at London Zoo until a larger lion house with roomier cages was built in the 1870s.^[226] Further changes took place in the early 20th century, when Carl Hagenbeck designed enclosures more closely resembling a natural habitat, with concrete 'rocks', more open space and a moat instead of bars, he designed lion enclosures for both Melbourne Zoo and Sydney's Taronga Zoo, among others, in the early 20th century. Though his designs were popular, the old bars and cage enclosures prevailed until the 1960s in many zoos;^[227] in the later decades of the 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, with some attractions even placing the den on ground higher than visitors, such as the Cat Forest/Lion Overlook of Oklahoma City Zoological Park.^[11]

Interactions with humans

In entertainment

Lion-baiting is a blood sport involving the baiting of lions in combat with other animals, usually dogs. Records of it exist in ancient times through until the seventeenth century, it was finally banned in Vienna by 1800 and England in 1835.^[228][229]

Lion taming refers to the practice of taming lions for entertainment, either as part of an established circus or as an individual act, such as Siegfried & Roy. The term is also often used for the taming and display of other big cats such as tigers, leopards, and cougars, the practice was pioneered in the first half of the nineteenth century by Frenchman Henri Martin and American Isaac Van Amburgh who both toured widely, and whose techniques were copied by a number of followers.^[230] 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 that Amburgh quickly borrowed, these acts eclipsed equestrianism acts as the central display of circus shows, but truly entered public consciousness in the early twentieth century with cinema. In demonstrating the superiority of human over animal, lion taming served a purpose similar to animal fights of previous centuries,^[230] the ultimate proof of a tamer's dominance and control over a lion is demonstrated by placing his head in the lion's mouth. The now iconic lion tamer's chair was possibly first used by American Clyde Beatty (1903–1965).^[231]

Man-eating

While lions do not usually hunt people, some (usually males) seem to seek out human prey; one well-publicised case includes the Tsavo maneaters, where 28 officially recorded railway workers building the Kenya-Uganda Railway were taken by lions over nine months during the construction of a bridge over the Tsavo River in Kenya in 1898.^[232] The hunter who killed the lions wrote a book detailing the animals' predatory behaviour, the lions were larger than normal, 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.^[233]

In their analysis of Tsavo and general man-eating, Kerbis Peterhans and Gnoske acknowledge that sick or injured animals may be more prone to man-eating, but that the behaviour is "not unusual, nor necessarily 'aberrant'" where the opportunity exists; if inducements such as access to livestock or human corpses are present, lions will regularly prey upon human beings. The authors note that the relationship is well-attested among other pantherines and primates in the paleontological record.^[234]

The lion's 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 – a number far exceeding the more famed "Tsavo" incidents of a century earlier. The incidents occurred near Selous National Park in Rufiji District and in Lindi Province near the Mozambican border. While the expansion of villagers into bush country is one concern, the authors argue that conservation policy must mitigate the danger because, in this case, conservation contributes directly to human deaths. Cases in Lindi have been documented where lions seize humans from the center of substantial villages.^[235] 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 attacks on people.^[236]

Author Robert R. Frump wrote in The Man-eaters of Eden that Mozambican refugees regularly crossing Kruger National Park at night in South Africa are attacked and eaten by the lions; park officials have conceded that man-eating is a problem there. Frump believes thousands may have been killed in the decades after apartheid sealed the park and forced the refugees to cross the park at night, for nearly a century before the border was sealed, Mozambicans had regularly walked across the park in daytime with little harm.^[237]

Packer estimates more than 200 Tanzanians are killed each year by lions, crocodiles, elephants, hippos, and snakes, and that the numbers could be double that amount, with lions thought to kill at least 70 of those. Packer has documented that between 1990 and 2004, lions attacked 815 people in Tanzania, killing 563. Packer and Ikanda are among the few conservationists who believe western conservation efforts must take account of these matters not just because of ethical concerns about human life, but also for the long term success of conservation efforts and lion preservation.^[235]

A man-eating lion was killed by game scouts in Southern Tanzania in April 2004, it is believed to have killed and eaten at least 35 people in a series of incidents covering several villages in the Rufiji Delta coastal region.^[238] Dr Rolf D. Baldus, the GTZ wildlife programme coordinator, commented that it was likely that the lion preyed on humans because it had a large abscess underneath a molar that was cracked in several places. He further commented that "This lion probably experienced a lot of pain, particularly when it was chewing."^[239] GTZ is the German development cooperation agency and has been working with the Tanzanian government on wildlife conservation for nearly two decades, as in other cases this lion was large, lacked a mane, and had a tooth problem.

The "All-Africa" record of man-eating generally is considered to be not Tsavo, but incidents in the early 1930s through the late 1940s in what was then Tanganyika (now Tanzania), inflicted by a pride of lions commonly referred to as the "Njombe lions". George Rushby, game warden and professional hunter, eventually dispatched the pride, which over three generations is thought to have killed and eaten 1,500 to 2,000 people in what is now Njombe district.^[240]

Sometimes, even Asiatic lions may become man-eaters, the area of the Gir sanctuary is now insufficient to sustain their large number,^[136] and consequently, lions have moved outside, making them a potential threat to people not only within the park, but also in surrounding places. Two attacks on humans were reported in 2012, including in area about 50–60 km (31–37 mi) from the sanctuary.^{[58][241][242]}

Cultural depictions

The lion is one of the most widely recognised animal symbols in human culture, it has been extensively depicted in sculptures and paintings, on national flags, and in contemporary films and literature. Lions have been kept in menageries since the time of the Roman Empire, and have been a key species sought for exhibition in zoos over the world since the late 18th century.^[57] It is featured in several of Aesop's fables written in the sixth century BC,^[244] it appeared as a symbol for strength and nobility in cultures across Europe, Asia, and Africa, despite incidents of attacks on people. It has been depicted as "king of the jungle" or "king of beasts"; hence, a popular symbol for royalty and stateliness.^[245] Depictions of lions are known from the Upper Paleolithic period. Carvings and paintings were discovered in the Lascaux and Chauvet Caves in France and dated to 15,000 to 17,000 years old.^[246][243] A lioness-headed ivory carving from Vogelherd cave in the Swabian Alb in southwestern Germany is dubbed Löwenmensch (lion-human) in German. The sculpture has been determined to be at least 32,000 years old and from the Aurignacian culture.^[21] But it may date to as early as 40,000 years ago.^[247]

In Africa, cultural views of the lion have varied by region; in some cultures, the lion symbolises power and royalty and some powerful rulers had the word "lion" in their nickname. For example, Marijata of the Mali Empire (c. 1235 – c. 1600) was given the name "Lion of Mali".^[248] Njaay, the legendary founder of the Waalo kingdom (1287–1855), 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 beasts. In parts of West Africa, to be compared to a lion was considered to be one of the greatest compliments, the social hierarchies of their societies where connected to the animal kingdom and the lion represented the top class.^[249] However, in more forested areas where lions were rare, the more numerous leopard represented the top of the hierarchy;^[250] in parts of West and East Africa, the lion is associated with healing and is seen as the link between the seers and the supernatural. In other East African traditions, the lion is the symbol of laziness;^[249] in many folktales, lions are portrayed as having low intelligence and are easily tricked by other animals.^[248] Although lions were commonly used in stories, proverbs and dances, they rarely featured in visual arts.^[250]

The ancient Egyptians portrayed several of their war deities as lionesses,^[245] whom they revered as fierce hunters.^[245] Egyptian deities associated with lions include: Bast, Mafdet, Menhit, Pakhet, Sekhmet, Tefnut, and the Sphinx.^[245] In Egypt, the avenging goddess Sekhmet, represented as a lioness, symbolized the ferocious heat of the sun, the lion was also believed to act as a guide to the underworld, through which the sun was believed to pass each night. The presence of lion -footed tombs found in Egypt and images of mummies carried on the backs of lions suggests this close association of the lions with the underworld.^[251]

The lion was a prominent symbol in ancient Mesopotamia (from Sumer up to Assyrian and Babylonian times), where it was strongly associated with kingship.^[252] Lions were among the major symbols of the goddess Inanna/Ishtar,^[253][254] the Lion of Babylon was the foremost symbol of the Babylonian Empire.^[255] The Lion Hunt of Ashurbanipal is a famous sequence of Assyrian palace reliefs from c. 640 BC, now in the British Museum.^[256] In Meopotamia, the lion was linked with both the fertility goddess Ishtar and the supreme Mesopotamian god Marduk, the theme of the royal lion hunt, a common motif in the early iconography in West Asia, symbolized death and resurrection, as the continuation of life was ensured by killing a god like animal. In some stone reliefs depicting the Royal hunt of lions, the divinity and the courage of the lion is equated with the divinity and courage of the king.^[257]

The Nemean lion was symbolic in ancient Greece and Rome, represented as the constellation and zodiac sign Leo, and described in mythology, where its skin was borne by the hero Heracles.^[258] Myths which have a hero killing a lion,such as the one in which Herakles slays the Nemean lion, symbolize victory over death. Similarly the wearing of lion skin such as the lion skin worn by Herackles also symbolizes victory over death.^[251]

The lion is the biblical emblem of the tribe of Judah and the later Kingdom of Judah.^[259] Lions are frequently mentioned in the Bible, notably in the Book of Daniel, in which the eponymous hero refuses to worship King Darius and is forced to sleep in the lions' den where he is miraculously unharmed (Dan 6). In the Book of Judges, Samson kills a lion as he travels to visit a Philistine woman.(Judg 14). The power and ferocity of the lion is invoked when describing both the anger of God (Amos 3:4–8, Lam 3:10) and the menace of Israel's enemies (Psm 17:12, Jer 2:30) and Satan (1 Pet 5:8). The book of Isaiah uses the imagery of a lion laying with a calf and child and eating straw to portray the harmony of creation (Isa 11:6–7); in the Book of Revelation, a lion, an ox, a man, and an eagle are on a heavenly throne in John's vision (Rev 4:7). The early Christian Church used this image to symbolise the four gospels, the lion symbolising the Gospel of Mark.^[260]

In the Puranic texts of Hinduism, Narasimha ("man-lion") a half-lion, half-man incarnation or (avatar) of Vishnu, is worshipped by his devotees and saved the child devotee Prahlada from his father, the evil demon king Hiranyakashipu;^[261] Vishnu takes the form of half-man/half-lion, in Narasimha, having a human torso and lower body, but with a lion-like face and claws.^[262] Singh is an ancient Indian vedic name meaning "lion" (Asiatic lion), dating back over 2000 years to ancient India. It was originally only used by Rajputs a Hindu Kshatriya or military caste in India. After the birth of the Khalsa brotherhood in 1699, the Sikhs also adopted the name "Singh" due to the wishes of Guru Gobind Singh. Along with millions of Hindu Rajputs today, it is also used by over 20 million Sikhs worldwide.^[263] Found famously on numerous flags and coats of arms all across Asia and Europe, the Asiatic lions also stand firm on the National Emblem of India.^[264] Farther south in South Asia, the Asiatic lion is symbolic for the Sinhalese,^[265] Sri Lanka's ethnic majority; the term derived from the Indo-Aryan Sinhala, meaning the "lion people" or "people with lion blood", while a sword-wielding lion is the central figure on the national flag of Sri Lanka.^[266]

The Asiatic lion is a common motif in Chinese art, they were first used in art during the late Spring and Autumn period (fifth or sixth century BC), and became much more popular during the Han Dynasty (206 BC – AD 220), when imperial guardian lions started to be placed in front of imperial palaces for protection. Because lions have never been native to China, early depictions were somewhat unrealistic; after the introduction of Buddhist art to China in the Tang Dynasty (after the sixth century AD), lions usually were wingless, with shorter, thicker bodies, and curly manes.^[267] The lion dance is a form of traditional dance in Chinese culture in which performers mimic a lion's movements in a lion costume, often with musical accompaniment from cymbals, drums, and gongs, they are performed at Chinese New Year, the August Moon Festival and other celebratory occasions for good luck.^[268]

The island nation of Singapore derives its name from the Malay words singa (lion) and pora (city/fortress), which in turn is from the Tamil-Sanskrit சிங்க singa सिंह siṃha and पुर புர pura, which is cognate to the Greek πόλις, pólis.^[269] According to the Malay Annals, this name was given by a fourteenth-century Sumatran Malay prince Sang Nila Utama, who, on alighting the island after a thunderstorm, spotted an auspicious beast on shore that appeared to be a lion.^[270]

"Lion" was the nickname of several medieval warrior rulers with a reputation for bravery, such as the English King Richard the Lionheart,^[245] Henry the Lion, (German: Heinrich der Löwe), Duke of Saxony, William the Lion, King of Scotland, and Robert III of Flanders nicknamed "The Lion of Flanders"—a major Flemish national icon up to the present. Lions are frequently depicted on coats of arms, either as a device on shields themselves, or as supporters, but the lioness is much more infrequent,^[271] the formal language of heraldry, called blazon, employs French terms to describe the images precisely. Such descriptions specified whether lions or other creatures were "rampant" or "passant", that is whether they were rearing or crouching.^[272]

The lion is used as a symbol of sporting teams, from national association football teams such as England, Scotland and Singapore to famous clubs such as the Detroit Lions^[273] of the NFL, Chelsea^[274] and Aston Villa of the English Premier League,^[275] (and the Premiership itself), Eintracht Braunschweig of the Bundesliga, and to a host of smaller clubs around the world.

Lions continue to be featured in modern literature, from the messianic Aslan in The Lion, the Witch and the Wardrobe and following books from The Chronicles of Narnia series written by C. S. Lewis,^[276] to the comedic Cowardly Lion in The Wonderful Wizard of Oz.^[277] The advent of moving pictures saw the continued presence of lion symbolism; one of the most iconic and widely recognised lions is Leo the Lion, which has been the mascot for Metro-Goldwyn-Mayer (MGM) studios since the 1920s.^[278] The 1960s saw the appearance of what is possibly the most famous lioness, the Kenyan animal Elsa in the movie Born Free,^[279] based on the true-life book of the same title.^[280] The lion's role as king of the beasts has been used in cartoons, such as the 1994 Disney animated feature film The Lion King.^[281]

See also

• Bokator
• Lion lights (lights used to repel lions)
• Okonjima
• Northeast African lion
• Physical comparison of tigers and lions
• Tiger versus lion

Notes

References

Cited texts

External links

Look up lion in Wiktionary, the free dictionary.

Wikispecies has information related to Lion

Wikisource has the text of the 1921 Collier's Encyclopedia article Lion.

Wikimedia Commons has media related to Lion.

• Species portrait Lion; IUCN/SSC Cat Specialist Group
• "Lion Conservation Fund".  Example of a fund and its projects about the research and conservation of the lion.
• A lion that traveled almost 1,300 km (810 mi) between Angola and Namibia