lion

 

large carnivore of the cat family, Panthera leo, found in open country in Africa, with a few surviving in India. Lions have short-haired coats of tawny brown, with the tail ending in a dark tuft. Most males have black or tawny manes of varying length growing from the head, neck, and shoulders. The mane may be quite long and magnificent, giving the lion the imposing appearance that has led it to be known as king of the beasts in folklore; studies indicate that long manes are typical mainly of cooler climate lions. Grown males are about 9 ft (2.7 m) long including the 3-ft (90-cm) tail, stand about 3 ft (90 cm) at the shoulder, and weigh up to 400 lb (180 kg). Females are smaller and lack manes. The lion is anatomically very similar to the tiger although it is different in habitat and way of life.

Lions are the only cats that are social rather than solitary. They usually live in groups called prides, which vary in composition but may occasionally include as many as 30 individuals. The lionesses do a considerable part of the hunting. There is no definite breeding season. They inhabit grasslands, scrubland, and semidesert areas, where they hunt antelope, zebra, and other large herbivorous animals, as well as domestic stock. Lions also eat carrion. They do not normally attack humans unless wounded or provoked; under unusual conditions they may prey on humans, but even old and sick animals are more likely to subsist on rodents, insects, and other small prey.

In early historic times lions ranged over Eurasia from E Europe to India and over all of Africa. They were eliminated from Europe and the Middle East by the beginning of the 2d cent. A.D. and from most of the rest of their range in recent times. They are now numerous only in central Africa, although even there they are severely reduced in numbers. At the beginning of the 20th cent. a few pairs remained in India and were preserved as tourist attractions in the Gir forest (now Gir National Park) of Gujarat state in W India. This group had increased to 290 individuals in 1955 but, although still protected, has been somewhat smaller since; they are the only remaining Asiatic lions. In early Christian symbolism the lion represented Jesus and has also represented St. Mark. For the constellation and sign of the zodiac see Leo.

Lions are classified in the phylum Chordata, subphylum Vertebrata, class Mammalia, order Carnivora, family Felidae.

Bibliography

See the many books by J. Adamson; G. B. Schaller, The Serengeti Lion (1972); A. E. Pease, The Book of the Lion (1986).

Lion
Male Female (Lioness)
Conservation status
Vulnerable (IUCN) [1]
Scientific classification
Kingdom: Animalia Phylum: Chordata Class: Mammalia Order: Carnivora Family: Felidae Genus: Panthera Species: P. leo
Binomial name
Panthera leo (Linnaeus, 1758)
Distribution of lions in Africa
Synonyms
Felis leo (Linnaeus, 1758)

The lion (Panthera leo) is a member of the family Felidae and one of four "big cats" in the genus Panthera. Reaching 272 kg (600 lb), it is the second-largest cat after the tiger. They currently exist in the wild in sub-Saharan Africa and in Asia with a critically endangered remnant population in northwest India, having disappeared from North Africa, the Middle East and western Asia in historic times. Until the late Pleistocene (about 10,000 years ago), the lion was the most widespread large land mammal beside man. They were found in most of Africa, much of Eurasia from western Europe to India and the Bering land bridge and in the Americas from Yukon to Peru.

Lions live for approximately 10–14 years in the wild, while in captivity they can live over 20 years. They typically inhabit savanna and grassland, although they may take to bush and forest. Lions are unusually social compared to other cats. A lion pride consists of related females and offspring and a small number of dominant males. Groups of female lions typically hunt together, preying mostly on large ungulates. The lion is an apex and keystone predator, though will resort to scavenging if the opportunity arises. While lions, in general, do not selectively hunt humans, some have been known to become man-eaters and seek human prey.

The lion is a vulnerable species, having seen a possibly irreversible population decline of 30 to 50% over the past two decades in its African range;^[1] populations are untenable outside designated reserves and national parks. Although the cause of the decline is not well-understood, habitat loss and conflicts with humans are currently the greatest causes of concern. Lions have been kept in menageries since Roman times and have been a key species sought after and exhibited in zoos the world over since the late 18th century. Zoos are cooperating worldwide in breeding programs for the endangered Asiatic subspecies.

The male lion is highly distinctive and is easily recognized by its mane. The lion, particularly the face of the male, is one of the most widely recognized animal symbols in human culture. Depictions have existed from the Upper Paleolithic period, with carvings and paintings from the Lascaux and Chauvet Caves, through virtually all ancient and medieval cultures where they historically occurred. It has been extensively depicted in literature, in sculptures, in paintings, on national flags, and in contemporary films and literature.

Naming and etymology

The lion's name, similar in many languages, derives from the Latin leo,^[2] and before that the Ancient Greek leōn/λεων.^[3] The Hebrew word lavi (לָבִיא) may also be related,^[4] as well as the Ancient Egyptian rw.^[5] It was one of the many species originally described, as Felis leo, by Linnaeus in his 18th century work, Systema Naturae.^[6] The generic component of its scientific designation, Panthera leo, is often presumed to derive from Greek pan- ("all") and ther ("beast"), but this may be a folk etymology. Although it came into English through the classical languages, panthera is probably of East Asian origin, meaning "the yellowish animal," or "whitish-yellow".^[7]

Taxonomy and evolution

The oldest lion-like fossil is known from Laetoli in Tanzania and is perhaps 3.5 million years old; some scientists have identified the material as Panthera leo. These records are not well-substantiated, and all that can be said is that they pertain to a Panthera-like felid. The oldest confirmed records of Panthera leo in Africa are about 2 million years younger.^[8] The closest relatives of the lion are the other Panthera species: the tiger, the jaguar and the leopard. Morphological and genetic studies reveal that the tiger was the first of these recent species to diverge. About 1.9 million years ago the Jaguar branched off the remaining group, which contained ancestors of the leopard and lion. The Lion and leopard subsequently separated about 1 to 1.25 million years ago from each other.^[9]

Panthera leo itself evolved in Africa between 1 million and 800,000 years ago before spreading throughout the Holarctic region;^[10] It appeared in Europe for the first time 700,000 years ago with the subspecies Panthera leo fossilis at Isernia in Italy. From this lion derived the later Cave lion (Panthera leo spelaea), which appeared about 300,000 years ago. During the upper Pleistocene the lion spread to North and South America, and developed into Panthera leo atrox, the American lion.^[11] Lions died out in northern Eurasia and America at the end of the last glaciation, about 10,000 years ago;^[12] this may have been secondary to the extinction of megafauna.^[13]

Subspecies

Traditionally 12 recent subspecies of lion were recognized, the largest of which has been recognised as the Barbary Lion.^[14] The major differences between these subspecies are location, mane appearance, size and distribution. Because these characteristics are very insignificant and show a high individual variability, most of these forms were debatable and probably invalid; additionally, they were often based upon zoo material of unknown original who may have had "striking, but abnormal" morphological characteristics.^[15] Today only eight subspecies are usually accepted,^[16][12] but one of these, the Cape lion formerly described as Panthera leo melanochaita is probably invalid.^[16] Even the remaining seven subspecies might be too much; mitochondrial variation in recent African lions is modest, which suggests that all sub-Saharan lions could be considered a single subspecies, possibly divided in two main clades: one to the west of the Great Rift Valley and the other to the east. Lions from Tsavo in Eastern Kenya are much closer genetically to lions in Transvaal (South Africa), than to those in the Aberdare Range in Western Kenya.^[17][18]

Recent

Eight recent subspecies are recognized today:

• P. l. persica, known as the Asiatic- or South Asian, Persian or Indian lion, was once widespread from Turkey, across the Middle East, to Pakistan, India and even Bangladesh. However, large prides and daylight activity made it easier to poach than tigers or leopards; now around 300 exist in and near the Gir Forest of India.^[19]
• P. l. leo, known as the Barbary lion, is extinct in the wild due to excessive hunting, although captive individuals may still exist. This was the largest of the lion subspecies, at 3–3.5m approx., and weighing over 150 kilograms and more. They ranged from Morocco to Egypt. The last wild Barbary lion was killed in Morocco in 1922.^[20]
• P. l. senegalensis, known as the West African lion, is found in Western Africa, from Senegal to Nigeria.
• P. l. azandica, known as the North East Congo lion, is found in the Northeastern parts of the Congo.
• P. l. nubica, known as the East African- or Massai lion, is found in East Africa, from Ethiopia and Kenya to Tanzania and Mozambique.
• P. l. bleyenberghi, known as the Southwest African- or Katanga lion. It is found in southwestern Africa, Zimbabwe, Angola, Katanga (Zaire).
• P. l. krugeri, known as the Southeast African- or Transvaal lion, is found in the Transvaal region of South eastern Africa, including Kruger National Park.
• P. l. melanochaita, known as the Cape lion, became extinct in the wild around 1860. Results of mitochondrial DNA research do not support the status as a distinct subspecies. It seems probable that the Cape lion was only the southernmost population of the extant southern African lion.

"#wp-_note-Conservation-Genetics:Preserving-Genetic-Diversity">[16]

Prehistoric

Several additional subspecies of lion existed in prehistoric times:

• P. l. atrox, known as the American lion or American cave lion, was abundant in the Americas from Alaska to Peru in the Pleistocene Epoch until about 10,000 years ago. This form as well as the cave lion are sometimes considered to represent separate species, but recent phylogenetic studies lead to suggest, that they are in fact subspecies of the lion (Panthera leo).

"#wp-_note-BurgerJ-Molecular-phylogeny">[12] One of the largest lion subspecies to have existed, its body length is estimated to have been 1.6–2.5 m (5–8 ft).^[21]

• P. l. fossilis, known as the Early Middle Pleistocene European cave lion, flourished about 500,000 years ago; fossils have been recovered from Germany and Italy.

• P. l. spelaea, known as the European cave lion, Eurasian cave lion or Upper Pleistocene European cave lion, occurred in Eurasia 300,000 to 10,000 years ago.

"#wp-_note-BurgerJ-Molecular-phylogeny">[12] It is known from Paleolithic cave paintings, ivory carvings, and clay busts,^[22] indicating it had protruding ears, tufted tails, faint tiger-like stripes, and that at least some males had a "ruff" or primitive mane around their necks.^[23]

• P. l. vereshchagini, known as the East Siberian- or Beringian cave lion, was found in Yakutia (Russia), Alaska (USA), and the Yukon Territory (Canada). Analysis of skulls and mandibles of this lion demonstrate that it is distinct—larger than the European cave lion and smaller than the American cave lion with differing skull proportions.^[24]

"#wp-_note-BurgerJ-Molecular-phylogeny">[12]

Dubious

• P. l. sinhaleyus, known as the Sri Lanka lion, appears to have become extinct around 39,000 years ago. It is only known from two teeth found in deposits at Kuruwita. Based on these teeth, P. Deraniyagala erected this subspecies in 1939.^[25]
• P. l. europaea, known as the European lion, was probably identical with Panthera leo persica or Panthera leo spelea; its status as subspecies is unconfirmed. It became extinct around 100 AD due to persecution and over-exploitation. Inhabited the Balkans, the Italian Peninsula, southern France and the Iberian Peninsula. It was a very popular object of hunting among Romans, Greeks and Macedonians.
• P. l. youngi or Panthera youngi , known as the North-Eastern Pleistocene China cave lion, flourished 350,000 years ago.^[26] Its relationship to the extant lion subspecies is obscure, and probably represents a distinct species.
• P. l. maculatus, known as the Marozi or Spotted lion, is sometimes believed 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 colored individuals, it has been extinct since 1931. A less likely identity is a natural leopard/lion hybrid commonly known as a leopon.^[27]

Physical characteristics

The lion is the second largest feline after the tiger. With powerful legs, a strong jaw, and long canine teeth, the lion can bring down and kill large prey.^[28] Lion coloration varies from light buff to yellowish, reddish or dark ochraceous brown. The underparts are generally lighter and the tail tuft is black. The color of the mane varies from blond to black.

Average listed weights for the lions are between 150–225 kg (330–500 lb) for males, and 120–150 kg (260–330 lb) for females.^[29] Nowell and Jackson report average weights of 181 kg for males and 126 kg for females; one male shot near Mount Kenya was weighed at 272 kg (600 lb).^[20] Head and body length is 170–250 cm (5 ft 7 in–8 ft 2 in) in males and 140–175 cm (4 ft 7 in–5 ft 9 in) in females; shoulder height is about 123 cm (4 ft) in males and 100 cm (3 ft 3 in) in females. The tail length is 70–100 cm (2 ft 3 in–3 ft 3 in).^[30] The tail ends in a hairy tuft. The tuft conceals a spine, approximately 5 mm long, formed of the final sections of tail bone fused together. The lion is the only felid to have a tufted tail and the function of the tuft and spine are unknown. Absent at birth, the tuft develops around 5½ months of age and readily identifiable at 7 months.^[31]

Mane

The mane of the male lion, unique amongst cats, is one of the most distinctive characteristics of the species. It makes the lion appear larger, providing an excellent intimidation display; this aids the lion during confrontations with other lions and with the species' chief competitor in Africa, the spotted hyena.^[32] The presence, absence, color, 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.^[33] Research in Tanzania also suggests mane length 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.^[34] In prides led by a coalition of two or three males, it is possible that lionesses solicit mating more actively with heavily maned lions.^[33]

Scientists once believed that the distinct status of some subspecies could be justified by morphology, including the size of the mane. Morphology was used to identify subspecies such as the Barbary lion and Cape Lion. Research has suggested, however, that environmental factors influence the color and size of a lion's mane, such as the ambient temperature.^[34] The cooler ambient temperature in European and North American zoos, for example, can result in a heavy mane. Thus the mane is an inappropriate marker for identifying subspecies.^[16][35] However the males of the Asiatic subspecies are characterized by sparser manes than average African lions.^[36]

Maneless lions have been reported in Senegal and Tsavo East National Park in Kenya, and the original male white lion from Timbavati was also maneless. Castrated lions have minimal manes. The lack of a mane is found in inbred lion populations; inbreeding also results in poor fertility.^[37] Cave paintings of European cave lions show exclusively animals with no mane or just the hint of a mane, suggesting they were more or less maneless.^[23]

White lions

The white lion is not a distinct subspecies, but a special morph with a genetic condition, leucism,^[15] that causes paler colouration akin to that of the white tiger; the condition is similar to melanism, which causes black panthers. White animals of the Transvaal lion (Panthera leo krugeri) have been occasionally encountered in and around the Kruger National Park and the adjacent Timbavati Private Game Reserve in eastern South Africa, but are more commonly found in captivity, where breeders deliberately select them. The unusual cream color of their coats is due to a recessive gene.^[38] They have been reportedly bred in camps in South Africa for use as trophies for canned hunts.^[39]

Confirmation of the actual existence of the White lion only came in the late 20th century. For hundreds of years prior, the White lion had been a figment of legend circulating through South Africa, the white pelage of the animal said to represent the goodness in all creatures. Claimed sightings were first reported in the early 1900s, and continued, infrequently, for almost 50 years until, in 1975, a litter of white lion cubs were found at Timbavati Game Reserve.^[40]

Black lions

though in few in some parts of africa there have been reports of black lions. most likely this is due to the benificial evolutionary trait of over producing melanin, such as in the case of Black panthers. though since lions with dark manes have a high chance at attracting females it is thought these reports are untrue due to the fact that they're whould be more black lions than the normal tawny brown ones.

Hybrids

Further information: Panthera hybrid, liger and tigon

Lions have also been known to breed with tigers (most often the Amur and Bengal subspecies) to create hybrids called ligers and tigons.^[41] They have also been crossed with leopards to produce leopons,^[42] and jaguars to produce jaglions. The marozi is reputedly a spotted lion or a naturally occurring leopon, while the Congolese spotted lion is a complex lion/jaguar/leopard hybrid called a lijagulep. Such hybrids were once commonly bred in zoos, but this 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.^[43] Because the lion sire passes on a growth-promoting gene, but the corresponding growth-inhibiting gene from the female lion is absent, 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 are often fertile. Males have about a 50% chance of having a mane, but if they grow one their manes will be modest: around 50% of a pure lion mane. Ligers are typically between 10 to 12 feet in length, and can be between 800 and 1,000 pounds or more.^[43] The less common tigon is a cross between the lioness and the male tiger.^[44]

Biology and behaviour

Lions spend much of their time resting and are inactive for about 20 hours per day.^[45] Although lions can be active at any time, their activity generally peaks after dusk with a period of socializing, grooming and defecating. Intermittent bursts of activity follow through the night hours to dawn, when hunting most often takes place. They spend an average two hours a day walking and 50 minutes eating.^[46]

Hunting and diet

Lions are powerful animals that usually hunt in groups and stalk their chosen prey. They can reach speeds of  mph ( km/h),^[47] though only for short bursts,^[48] so they have to be close to their prey before starting the attack. Lions take advantage of factors that reduce visibility; many kills take place near some form of cover or at night.^[49] They sneak up to the victim until they reach a distance of about 30 m (98 feet) or less. Typically, several female lions work together and encircle the herd from different points. Once they have closed with a herd, they usually target the closest prey. The attack is short and powerful, the lion attempting to catch the victim with a fast rush and final leap. The prey is usually killed by strangulation.^[50]

The lion's prey consists mainly of large mammals, with a preference for wildebeest, impalas, zebras, buffalo and warthogs in Africa and nilgai, wild boars and several deer species in India. Many other species are hunted based on availability, mainly ungulates weighing between 50 and 300 kg such as kudu, hartebeest, gemsbok and eland.^[30] Occasionally, lions take relatively small species such as Thomson's gazelle or springbok. Lions are capable of taking down most animals, even healthy adults. However, they rarely attack very large animals such as buffalo bulls, fully grown male giraffes,^[51] and adult hippopotamuses, rhinoceroses or elephants, due to the danger of injury.^[52] In some areas, lions specialise in hunting atypical prey-species; this is the case at the Savuti river, where they prey on young elephants.^[53] Park guides in the area reported that the lions, driven by extreme hunger, started taking down baby elephants, and then moved on to adolescents and, occasionally, fully grown adults.^[54] In the Kruger National Park, giraffes are regularly hunted.^[55] Lions also attack domestic livestock; in India cattle contribute significantly to their diet.^[36] They are capable of killing other predators such as leopards, cheetahs, hyenas and wild dogs, as well as scavenging animals either dead from natural causes or killed by other predators.^[56] A lion may gorge itself and eat up to 30 kg (66 lb) in one sitting;^[57] 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.^[58] An adult female lion requires an average of about 5 kg (11 lb) of meat per day, a male about 7 kg (15.4 lb).^[59]

Because lions hunt in open spaces where they are easily seen by their prey, cooperative hunting increases the likelihood of a successful hunt; this is especially true with larger species. Teamwork also enables them to defend their prey more easily against other large predators such as hyenas, which can be attracted by vultures over kilometers in open savannas. Lionesses do most of the hunting. Males attached to prides do not usually participate, except when hunting large animals such as buffalo and giraffe. In group hunts, each lioness has a favored 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.^[60] Young lions first display stalking behaviour around three months of age, although they do not participate in actual hunting until they are almost a year old. They begin to hunt effectively when nearing the age of two.^[61]

Reproduction

Most lionesses will have reproduced by the time they are four years of age.^[62] Lions do not mate at any specific time of year, and the females are polyestrous.^[63] Like other cats, the male lion's penis has spines which point backwards. Upon withdrawal of the penis, the spines rake the walls of the female's vagina, which may cause ovulation.^[64] A female may mate with more than one male when she is in heat;^[65] during a mating bout, which could last several days, the couple copulates twenty to forty times a day and are likely to forgo hunting. Lions reproduce very well in captivity.

The average gestation period is around 110 days,^[63] the female giving birth to a litter of one to four cubs. Lionesses in a pride will synchronize their reproductive cycles so that they cooperate in the raising and suckling of the young, who suckle indiscriminately from any or all of the nursing females in the pride. Cubs are usually born and initially kept hidden from view in thickets or sheltered areas. They weigh 1.2–2.1 kg at birth and are almost helpless, beginning to crawl a day or two after birth and walking around three weeks of age.^[66] Weaning occurs after six to seven months. In the wild, competition for food is fierce, and as many as 80% of the cubs will die before the age of two.^[67]

When one or more new males take over a pride and oust the previous master(s), the conquerors often kill any remaining cubs;^[68] females do not again become fertile and receptive until the cubs grow up or die. The male lions reach maturity at about 3 years of age and are capable of taking over another pride at 4–5 years old. They begin to age and weaken between 10 and 15 years of age at the latest.^[69] This leaves a short window for their own offspring to be born and mature—the fathers have to procreate as soon as they take over the pride. The lioness will often attempt to defend her cubs fiercely from a usurping male, but such actions are rarely successful, as he usually kills all the previous top male's cubs that are less than two years old and the female is much lighter and has less strength than the male. However, success is more likely when a group of 3 or 4 mothers within the pride join forces against one male.^[68]

One scientific study reports that both males and females may interact homosexually.^[70][71] Male lions pair-bond for a number of days and initiate homosexual activity with affectionate nuzzling and caressing, leading to mounting and thrusting. A study found that about 8% of mountings have been observed to occur with other males, while female pairings are held to be fairly common in captivity but have not been observed in the wild.

Health

Though adult lions have no natural predators, evidence suggests that the majority die violently from humans or other lions.^[72] Various species of tick commonly infest the ears, neck and groin regions of most lions.^[73][74] Adult forms of several species of the tapeworm genus Taenia have been isolated from intestines, the lions having ingested larval forms from antelope meat.^[75] 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.^[76] A more recent outbreak in 2001 killed six lions.^[77] Lions, especially in captivity, are vulnerable to the Canine distemper virus (CDV), feline immunodeficiency virus (FIV) and feline infectious peritonitis (FIP).^[15] 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.^[78] 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.^[15]

Group organization

Lions are predatory carnivores who manifest two types of social organization. Some are residents, living in groups, called prides.^[79] The pride consists of usually around five or six related females, their cubs of both sexes, and a group of one to four males known as a coalition who mate with the adult females. Others are nomads, ranging widely and moving sporadically, either singularly or in pairs.^[79] Note that a lion may switch lifestyles; nomads may become residents and vice versa. The area a pride occupies is called a pride area, whereas that by a nomad is a range.^[79] Why sociality—the most pronounced in any cat species—has developed in lions 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 "cheaters" reduce per capita caloric intake. 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.^[20]

Being smaller and more agile than males, and lacking the conspicuous mane, lionesses do the pride's hunting, while the stronger males patrol the territory and protect the pride. There is no clear hierarchy with food: male lions often eat animals killed by lionesses but will never share food they have killed themselves; they will take food from cubs but are more likely to share with cubs than lionesses, which are more likely to share with each other. There is more sharing with larger kills.^[80]

Both males and females defend the pride against intruders. Some individual lions consistently lead the defense against intruders, while others lag behind.^[81] These "laggards" are not punished by leaders. Possibly laggards provide other services to the group so that leaders forgive them.^[82] An alternative hypothesis is that there is some reward associated with being a leader who fends off intruders.^[83] The leading male or males often have to defend against outside males attempting to take over the pride. Females form a stable social unit in a pride and will not tolerate outside females; ^[84] membership only changes with the births and deaths of lionesses,^[85] though some females do leave and become nomadic.^[86] Subadult males on the other hand, leave the pride when they reach maturity at around 2–3 years of age.^[86]

Communication

When resting, lion socialization occurs through a number of behaviors, and the animal's expressive movements are highly developed. The most common peaceful tactile gestures are head rubbing and social licking,^[87] which have been compared with grooming in primates.^[88] Head rubbing—nuzzling one's forehead, face and neck against another lion—appears to be a form of greeting,^[89] 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.^[90] 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.^[91]

Lions have an array of facial expressions and body postures that serve as visual gestures.^[92] Their repertoire of vocalizations is also large; variations in intensity and pitch, rather than discrete signals, appear central to communication. Lion sounds include snarling, purring, hissing, coughing, miaowing, woofing and roaring. Lions most often roar at night; the sound, which can be heard from a distance of 8 km, is used to advertise the animal's presence.^[93]

Distribution and habitat