|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.
Naming and etymology
Taxonomy and evolution
Panthera leo itself evolved in Africa between 1 million and 800,000 years ago before spreading throughout the Holarctic region; 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. Lions died out in northern Eurasia and America at the end of the last glaciation, about 10,000 years ago; this may have been secondary to the extinction of megafauna.
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.
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.
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.
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.
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 behavioral 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
Hunting and diet
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 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, and adult hippopotamuses, rhinoceroses or elephants, due to the danger of injury. In some areas, lions specialize 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 behavior 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.
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.
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. Sub adult males on the other hand, leave the pride when they reach maturity at around 2-3 years of age.
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
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 1412 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
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 coordinated efforts involving lion conservation have been organized 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.
Relation with humans
The lion's proclivity for man-eating has been systematically examined. American and Tanzanian scientists report that man-eating behavior 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.4
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. Mozambicans had for nearly a century before the border was sealed regularly walked across the park in daytime with little harm.
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 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.
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. 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 which was cracked in several places. He further commented that "This lion probably experienced a lot of pain, particularly when it was chewing." GTZ is the German development cooperation agency and has been working with the Tanzanian government on wildlife conservation for nearly two decades. Like 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 the lesser-known incidents in the late 1930s through the late 1940s in what was then Tanganyika (now Tanzania). 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 in what is now Njombe district.
Lions were kept and bred by Assyrian kings as early as 850 BC, and Alexander the Great was said to have been presented with tame lions by the Malhi of northern India. Later in Roman times, lions were kept by emperors to take part in the gladiator arenas. Roman notables, including Sulla, Pompey, and Julius Caesar, often ordered the mass slaughter of hundreds of lions at a time. In the East, Lions were tamed by Indian princes, and Marco Polo reported that Kublai Khan kept lions inside. The first European "zoos" spread amongst 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. 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, probably stocked with animals from an earlier menagerie started in 1125 by Henry I at his palace in Woodstock, near Oxford; where lions had been reported stocked by William of Malmesbury.
Seraglios served as expressions of the nobility's power and wealth. Animals like big cats and elephants, in particular, symbolized power, and would be pitted in fights against each other or domesticated animals. By extension, menageries and seraglios served as demonstrations of the dominance of man 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 would last into the 19th century, at which time it was seen as highly eccentric.
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. 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. A rival menagerie at the Exeter Exchange also exhibited lions until the early 19th century. The Tower menagerie was closed down by William IV, and animals transferred to the London Zoo which opened its gates to the public on 27 April 1828.
Animal species disappear when they cannot peacefully orbit the center of gravity that is man.
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 like the giraffe and hippopotamus, and much less than pandas. 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. The widely reproduced imagery of the heroic hunter chasing lions would dominate a large part of the century. 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."
Lions were kept in cramped and squalid conditions at London Zoo until a larger lion house with roomier cages was built in the 1870s. 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. 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. Lions are now housed in much larger naturalistic areas; modern recommended guidelines more closely approximate conditions in the wild with closer attention to the lions' needs, highlighting the need for dens in separate areas, elevated positions in both sun and shade where lions can sit and adequate ground cover and drainage as well as sufficient space to roam.
Baiting and taming
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 19th century by Frenchman Henri Martin and American Isaac Van Amburgh who both toured widely, and whose techniques were copied by a number of followers. Van Amburgh performed before Queen Victoria of the United Kingdom 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 20th century with cinema. In demonstrating the superiority of man over animal, lion taming served a purpose similar to animal fights of previous centuries. The now iconic lion tamer's chair was possibly first used by American Clyde Beatty (1903-1965).
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