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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; 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.
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. 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. 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). 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). 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.
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. 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. 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. In prides led by a coalition of two or three males, it is possible that lionesses solicit mating more actively with heavily maned lions.
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. 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. However the males of the Asiatic subspecies are characterized by sparser manes than average African lions.
White lions owe their coloring to a recessive gene. They are rare forms of the subspecies Panthera leo krugeri.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. 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.
The white lion is not a distinct subspecies, but a special morph with a genetic condition, leucism, 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. They have been reportedly bred in camps in South Africa for use as trophies for canned hunts.
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.
Lions have also been known to breed with tigers (most often the Amur and Bengal subspecies) to create hybrids called ligers and tigons. They have also been crossed with leopards to produce leopons, 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. 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. The less common tigon is a cross between the lioness and the male tiger.
Biology and behavior
Lions spend much of their time resting and are inactive for about 20 hours per day. 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.
Hunting and diet
Lions are powerful animals that usually hunt in groups and stalk their chosen prey. They can reach speeds of 59 km/h (40 mph), though only for short bursts, 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. 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.
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. Occasionally, lions take relatively small species such as Thomson's gazelle or springbok. Lions, hunting in groups, are capable of taking down most animals, even healthy adults, but they rarely attack very large prey such as buffalo bulls or fully grown male giraffes, due to the danger of injury. They normally feed on mammals no larger than 550 kg, which excludes most adult hippopotamuses, rhinoceroses, elephants, giraffes and buffalos. In some areas, lions specialise in hunting atypical prey-species; this is the case at the Savuti river, where they prey on young elephants. 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. In the Kruger National Park, giraffes are regularly hunted. Lions also attack domestic livestock; in India cattle contribute significantly to their diet. 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. A lion may gorge itself and eat up to 30 kg (66 lb) in one sitting; 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. 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).
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. 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.
Most lionesses will have reproduced by the time they are four years of age. Lions do not mate at any specific time of year, and the females are polyestrous. 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. A female may mate with more than one male when she is in heat; 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, 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. 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.
When one or more new males take over a pride and oust the previous master(s), the conquerors often kill any remaining cubs; 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. 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.
One scientific study reports that both males and females may interact homosexually. 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.
Lions are predatory carnivores who manifest two types of social organization. Some are residents, living in groups, called prides. 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. 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. 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.
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.
Both males and females defend the pride against intruders. Some individual lions consistently lead the defense against intruders, while others lag behind. These "laggards" are not punished by leaders. Possibly laggards provide other services to the group so that leaders forgive them. An alternative hypothesis is that there is some reward associated with being a leader who fends off intruders. 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; membership only changes with the births and deaths of lionesses, though some females do leave and become nomadic. Subadult males on the other hand, leave the pride when they reach maturity at around 2–3 years of age.
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, which have been compared with grooming in primates. Head rubbing—nuzzling one's forehead, face and neck against another lion—appears to be a form of greeting, 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. 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.
Lions have an array of facial expressions and body postures that serve as visual gestures. 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.
Distribution and habitat
In Africa, lions can be found in savannah grasslands with scattered Acacia trees which serve as shade; their habitat in India is a mixture of dry savannah forest and very dry deciduous scrub forest. In relatively recent times the habitat of lions spanned the southern parts of Eurasia, ranging from Greece to India, and most of Africa except the central rainforest-zone and the Sahara desert. Herodotus reported that lions had been common in Greece around 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 and by 100 AD extirpated. A population of the Asiatic lion survived until the 10th century in the Caucasus, their last European outpost.
The species was eradicated from 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 early 20th century they became extinct in North Africa and the Middle East. By the late 19th century the lion had disappeared from Turkey and most of northern India, while the last sighting of a live Asiatic Lion in Iran was in 1941 (between Shiraz and Jahrom, Fars province), though the corpse of a lioness was found on the banks of Karun river, Khuzestan province in 1944. There are no subsequent reliable reports from Iran. The subspecies now survives only in and around the Gir Forest of northwestern India. About 300 lions live in a 1,412 km² (558 square miles) sanctuary in the state of Gujarat, which covers most of the forest. Their numbers are slowly increasing.
Until the late Pleistocene (about 10,000 years ago), the lion was the most widespread land mammal aside from 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. Parts of this range were occupied by subspecies that are extinct today.
Population and conservation status
Most lions now live in eastern and southern Africa, and their numbers there are rapidly decreasing, with an estimated 30–50% decline over the last two decades. Currently, estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004, down from early 1990s estimates that ranged as high as 100,000 and perhaps 400,000 in 1950. The cause of the decline is not well-understood, and may not be reversible. Currently, habitat loss and conflicts with humans are considered the most significant threats to the species. The remaining populations are often geographically isolated from each other, which can lead to inbreeding, and consequently, a lack of genetic diversity. Therefore the lion is considered a vulnerable species by the International Union for Conservation of Nature and Natural Resources, while the Asiatic subspecies is critically endangered. The lion population in the region of West Africa is isolated from lion populations of Central Africa, with little or no exchange of breeding individuals. The number of mature individuals in West Africa is estimated by two separate recent surveys at 850–1,160 (2002/2004). 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.
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 and Kruger National Park in eastern South Africa. Outside these areas, the issues arising from lions' interaction with livestock and people usually results in the elimination of the former. In India, the last refuge of the Asiatic Lion is the 1,412 km² (558 square miles) Gir Forest National Park in western India which had about 359 lions (as of April 2006). As in Africa, numerous human habitations are close by with the resultant problems between lions, livestock, locals and wildlife officials. The Asiatic Lion Reintroduction Project plans to establish a second independent population of Asiatic Lions at the Kuno Wildlife Sanctuary in the Indian state of Madhya Pradesh. It is important to start a second population to serve as a life insurance for the last surviving Asiatic Lions and to help develop and maintain genetic diversity enabling the species to survive.
The former popularity of the Barbary Lion as a zoo animal has meant that scattered lions in captivity are likely to be descended from Barbary Lion stock. This includes twelve lions at Port Lympne Wild Animal Park in Kent, England that are descended from animals owned by the King of Morocco. Another eleven animals believed to be Barbary lions were found in Addis Ababa zoo, 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.
Following the discovery of the decline of lion population in Africa, several co-ordinated 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 North American lions were not genetically pure, having been hybridized 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 origins, making maintenance of genetic diversity a problem.
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