Giraffe
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For other uses, see Giraffe (disambiguation).
Giraffe[1]
Conservation status
Least Concern (IUCN 3.1)[2]
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Family: Giraffidae
Genus: Giraffa
Species: G. camelopardalis
Binomial name
Giraffa camelopardalis
Linnaeus, 1758
Range map
The giraffe (Giraffa camelopardalis) is an African even-toed ungulate mammal, the tallest of all land-living animal species, and the largest ruminant. It is covered in large, irregular patches of yellow to black fur separated by white, off-white, or dark yellowish brown background. The average mass for an adult male giraffe is 1,191 kilograms (2,630 lb) while the average mass for an adult female is 828 kilograms (1,830 lb).[3][4] It is approximately 4.3 metres (14 ft) to 5.2 metres (17 ft) tall, although the tallest male recorded stood almost 6 metres (20 ft).[3][4]
The giraffe is related to deer and cattle, but is placed in a separate family, the Giraffidae, consisting of only the giraffe and its closest relative, the okapi. Its range extends from Chad in Central Africa to South Africa.
Giraffes usually inhabit savannas, grasslands, or open woodlands. However, when food is scarce they will venture into areas with denser vegetation. They prefer areas with plenty of acacia growth. They will drink large quantities of water when available which enables them to live for extended periods in dry, arid areas.
Contents [hide]
1 Etymology
2 Taxonomy and evolution
2.1 Subspecies
3 Anatomy and morphology
3.1 Horns
3.2 Neck
3.3 Legs and pacing
3.4 Circulatory system
4 Behaviour
4.1 Social structure and breeding habits
4.2 Reproduction
4.3 Necking
4.4 Diet
4.5 Sleeping
4.6 Communication
4.7 Stereotypic behavior
5 Human interactions
5.1 Conservation
5.2 Scientific inspiration
5.3 In art and culture
6 References
7 External links
Etymology
The species name camelopardalis (camelopard) is derived from its early Roman name, where it was described as having characteristics of both a camel and a leopard.[5] The English word camelopard first appeared in the 14th century and survived in common usage well into the 19th century. The Afrikaans language retained it. The Arabic word الزرافة ziraafa or zurapha, meaning "assemblage" (of animals), or just "tall", was used in English from the sixteenth century on, often in the Italianate form giraffa.
Taxonomy and evolution
The giraffe is one of only two living species of the family Giraffidae, along with the okapi. The family was once much more extensive, with numerous other species. The giraffids evolved from a 3 metres (9.8 ft) tall antelope-like mammal that roamed Europe and Asia some 30-50 million years ago.[6]
The earliest known giraffid was Climacoceras, which still resembled deer, having large antler-like ossicones. It first appeared in the early Miocene epoch. Later examples include the genera Palaeotragus and Samotherium, which appeared in the early to mid-Miocene. They were both tall at the shoulder, and had developed the simple, unbranched ossicones of modern giraffids, but still had relatively short necks.[7]
Comparison of the African Miocene giraffids: Palaeotragus (two top) and Climacoceras (two bottom)From the late Pliocene onwards, the variety of giraffids drastically declined, until only the two surviving species remained. The modern genus Giraffa evolved during the Pliocene epoch, and included a number of other long-necked species, such as Giraffa jumae, that do not survive today.[7] Alan Turner proposes, in the 2004 book Evolving Eden, that giraffe ancestors initially had a dark coat with pale spots, and that the spots gradually became star-shaped, before eventually forming the reticulated pattern found today.[8] The modern species, Giraffa camelopardalis, appeared during the Pleistocene 1 million years ago.[citation needed]
The evolution of the long necks of giraffes has been the subject of much debate. The standard story is that they were evolved to allow the giraffes to browse vegetation that was out of the reach of other herbivores in the vicinity, giving them a competitive advantage.[7] However, an alternative theory proposes that the long necks evolved as a secondary sexual characteristic, giving males an advantage in "necking" contests (see below) to establish dominance and obtain access to sexually receptive females. This theory notes that giraffes frequently feed from relatively low-lying shrubs, and that the necks of males are significantly longer than those of females.[9] However, this theory is not universally accepted, and some of the data supporting it has recently been challenged, lending support to the original proposal that neck length is related to browsing habits.[10]
Subspecies
Different authorities recognize different numbers of subspecies, differentiated by colour and pattern variations and range.[1][2][11] Some of these species may prove to in fact be separate species.[2] The subspecies recognized by various authorities include:
Reticulated or Somali Giraffe (G. c. reticulata) — large, polygonal liver-coloured spots outlined by a network of bright white lines. The blocks may sometimes appear deep red and may also cover the legs. Range: northeastern Kenya, Ethiopia, Somalia.
Angolan or Smoky Giraffe (G. c. angolensis) — large spots and some notches around the edges, extending down the entire lower leg. Range: Angola, Zambia.
Kordofan Giraffe (G. c. antiquorum) — smaller, more irregular spots that cover the inner legs. Range: western and southwestern Sudan.
G. c. tippelskirchi has distinctive jagged-edged spotsMasai or Kilimanjaro Giraffe (G. c. tippelskirchi) — jagged-edged, vine-leaf shaped spots of dark chocolate on a yellowish background. Range: central and southern Kenya, Tanzania.
Nubian Giraffe (G. c. camelopardalis) — large, four-sided spots of chestnut brown on an off-white background and no spots on inner sides of the legs or below the hocks. Range: eastern Sudan, northeast Congo.
Rothschild Giraffe or Baringo Giraffe or Ugandan Giraffe (G. c. rothschildi) — deep brown, blotched or rectangular spots with poorly defined cream lines. Hocks may be spotted. Range: Uganda, north-central Kenya.
South African Giraffe (G. c. giraffa) — rounded or blotched spots, some with star-like extensions on a light tan background, running down to the hooves. Range: South Africa, Namibia, Botswana, Zimbabwe, Mozambique.
Thornicroft or Rhodesian Giraffe (G. c. thornicrofti) — star-shaped or leafy spots extend to the lower leg. Range: eastern Zambia.
West African or Nigerian Giraffe (G. c. peralta) — numerous pale, yellowish red spots. Range: Niger, Cameroon.
Some scientists regard Kordofan and West African Giraffes as a single subspecies; similarly with Nubian and Rothschild's Giraffes, and with Angolan and South African Giraffes. Further, some scientists regard all populations except the Masai Giraffes as a single subspecies. By contrast, scientists have proposed four other subspecies — Cape Giraffe (G. c. capensis), Lado Giraffe (G. c. cottoni), Congo Giraffe (G. c. congoensis), and Transvaal Giraffe (G. c. wardi) — but none of these is widely accepted.
Though giraffes of these populations interbreed freely under conditions of captivity, suggesting that they are subspecific populations, genetic testing published in 2007[12] has been interpreted to show that there may be at least six species of giraffe that are reproductively isolated and not interbreeding, even though no natural obstacles, like mountain ranges or impassable rivers block their mutual access. In fact, the study found that the two giraffe populations that live closest to each other— the reticulated giraffe (G. camelopardalis reticulata) of north Kenya, and the Masai giraffe (G. c. tippelskirchi) in south Kenya— separated genetically between 0.13 and 1.62 million years BP, judging from genetic drift in nuclear and mitochondrial DNA.
The implications for conservation of as many as eleven such cryptic species and sub-species were summarised by David Brown for BBC News: "Lumping all giraffes into one species obscures the reality that some kinds of giraffe are on the brink. Some of these populations number only a few hundred individuals and need immediate protection."[13]
Anatomy and morphology
Giraffe skeleton from Royal Natural History by Richard LydekkerMale giraffes are up to 5.5 metres (18 ft) tall at the horn tips, and weigh between 800 and 1,930 kilograms (1,800 and 4,300 lb). Females are between 4 and 4.5 metres (13 and 15 ft) tall and weigh between 550 and 1,180 kilograms (1,200 and 2,600 lb). The coat is made up of brown blotches or patches separated by lighter hair. Each giraffe has a unique coat pattern.[14]
Horns
Both sexes have horns, although the horns of a female are smaller. The prominent horns are formed from ossified cartilage, and are called ossicones. The appearance of horns is a reliable method of identifying the sex of giraffes, with the females displaying tufts of hair on the top of the horns, whereas males' horns tend to be bald on top — an effect of necking in combat. Males sometimes develop calcium deposits which form bumps on their skull as they age, which can give the appearance of up to three additional horns.[15]
Giraffe in captivity at the Melbourne Zoo
Neck
Giraffes have long necks which they use to browse tree leaves. They possess seven vertebrae (although disputed by some zoologists who claim that it has eight[16]) in the neck (the usual number for a mammal) that are elongated. The vertebrae are separated by highly flexible joints. The base of the neck has spines which project upward and form a hump over the shoulders. They have anchor muscles that hold the neck upright.
Legs and pacing
Giraffes also have slightly elongated forelegs, about 10% longer than their hind legs. The pace of the giraffe is an amble, though when pursued it can run extremely fast, up to 55 km/h.[17] It cannot sustain a lengthy chase. Its leg length compels an unusual gait with the left legs moving together followed by right (similar to pacing) at low speed, and the back legs crossing outside the front at high speed. When hunting adult giraffes, lions try to knock the lanky animal off its feet and pull it down. Giraffes are difficult and dangerous prey. The giraffe defends itself with a powerful kick. A single well-placed kick from an adult giraffe can shatter a lion's skull or break its spine. Lions are the only predators which pose a serious threat to an adult giraffe.
Circulatory system
Giraffes bending down to drinkModifications to the giraffe's structure have evolved, particularly to the circulatory system. A giraffe's heart, which can weigh up to 10 kg (22 lb) and measure about 60 cm (2 ft) long, must generate approximately double the normal blood pressure for an average large mammal to maintain blood flow to the brain. In the upper neck, a complex pressure-regulation system called the rete mirabile prevents excess blood flow to the brain when the giraffe lowers its head to drink. Conversely, the blood vessels in the lower legs are under great pressure (because of the weight of fluid pressing down on them). In other animals such pressure would force the blood out through the capillary walls; giraffes, however, have a very tight sheath of thick skin over their lower limbs which maintains high extravascular pressure in the same way as a pilot's g-suit.
Behaviour
A male (bull) with a baby (calf) giraffe at the San Francisco Zoo
Social structure and breeding habits
Female giraffes associate in groups of a dozen or so members, occasionally including a few younger males. Younger males tend to live in "bachelor" herds, with older males often leading solitary lives. Reproduction is polygamous, with a few older males impregnating all the fertile females in a herd. Male giraffes determine female fertility by tasting the female's urine in order to detect estrus, in a multi-step process known as the Flehmen response.
Giraffes will mingle with the other herbivores in the African bush. Their company is beneficial, since they are tall enough to have a much wider scope of an area and will watch for predators.
Mating Angolan Giraffes (G. c. angolensis) at Chudop waterhole, Etosha, Namibia
Reproduction
Giraffe gestation lasts between 400 and 460 days, after which a single calf is normally born, although twins occasionally occur.[18] The mother gives birth standing up and the embryonic sack usually bursts when the baby falls to the ground. Newborn giraffes are about 1.8 m (6 ft) tall.
Within a few hours of being born, calves can run around and are indistinguishable from a week-old calf; however, for the first two weeks, they spend most of their time lying down, guarded by the mother. The young can fall prey to lions, leopards, spotted hyenas, and wild dogs. It has been speculated that their characteristic spotted pattern provides a certain degree of camouflage. Only 25 to 50% of giraffe calves reach adulthood; the life expectancy is between 20 and 25 years in the wild and 28 years in captivity (Encyclopedia of Animals).
Necking
Two male giraffes neckingAs noted above, males often engage in necking, which has been described as having various functions. One of these is combat. Battles can be fatal, but are more often less severe. The longer the neck, and the heavier the head at the end of the neck, the greater the force a giraffe is able to deliver in a blow. It has also been observed that males that are successful in necking have greater access to estrous females, so the length of the neck may be a product of sexual selection.[19]
After a necking duel, a giraffe can land a powerful blow with his head — occasionally knocking a male opponent to the ground. These fights rarely last more than a few minutes or end in physical harm.
Another function of necking is sexual, in which two males caress and court each other, leading up to mounting and climax. Such interactions between males are more frequent than heterosexual coupling.[20] In one study, up to 94% of observed mounting incidents took place between two males. The proportion of same sex activities varied between 30 and 75%, and at any given time one in twenty males were engaged in non-combative necking behaviour with another male. Only 1% of same-sex mounting incidents occurred between females.[21]
Giraffes use their long, prehensile tongues to extend their reach
Diet
The giraffe browses on the twigs of trees, preferring trees of the genera Acacia, Commiphora and Terminalia, and also eats grass and fruit.[11][22] The tongue is tough due to the giraffe's diet, which can include tree thorns. In Southern Africa, giraffes feed on all acacias, especially Acacia erioloba, and possess a specially-adapted tongue and lips that are tough enough to withstand the vicious thorns of this plant. A giraffe can eat 65 pounds (29 kg) of leaves and twigs daily, but can survive on just 15 pounds (6.8 kg).[22] The giraffe requires less food than typical grazing animals because the foliage it eats has more concentrated nutrition and it has a more efficeint digestive system.[11] During the wet season, food is abundant and giraffes disperse widely, but during the dry season they need to congragate around evergreen trees and bushes.[11] As a ruminant, it first chews its food, then swallows for processing and then visibly regurgitates the semi-digested cud up their necks and back into the mouth, in order to chew again. This process is usually repeated several times for each mouthful. The giraffe can surivive without water for extended periods.[22] A giraffe will clean off any bugs that appear on its face with its extremely long tongue (about 45 centimetres (18 in)).
Sleeping
The giraffe has one of the shortest sleep requirements of any mammal, which is between ten minutes and two hours in a 24-hour period, averaging 1.9 hours per day.[23]
Male giraffe making a cough-like mating call at the Roger Williams Park Zoo, Providence, Rhode Island
Communication
Although generally quiet and not vocal, giraffes have been heard to make various sounds. Courting males will emit loud coughs. Females will call their young by whistling or bellowing. Calves will bleat, moo, or make mewing sounds. In addition, giraffes will grunt, snort, hiss, or make strange flute-like sounds. Recent research has shown evidence that the animal communicates at an infrasound level.[24]
Stereotypic behavior
Many animals when kept in captivity, such as in zoos, display abnormal behaviours. Such unnatural behaviours are known as stereotypic behaviours.[25] In particular, giraffes show distinct patterns of stereotypic behaviours when removed from their natural environment. Due to a subconscious response to suckle milk from their mother, something which many human-reared giraffes and other captive animals do not experience, giraffes resort instead to excessive tongue use on inanimate objects.[26]
Due to the obvious social and cultural discomfort associated with the addition of milk delivery devices, animal enclosures are often enriched with other stimuli, such as food and mental distractions (toys, scent markings etc.). This operates as a distraction, removing the giraffe’s focus from its instinctual tendencies towards suckling, resulting in tongue lolling and licking of objects in close proximity.
Human interactions
Conservation
Lone giraffe at Lake Nakuru National Park, Kenya.Overall, the giraffe is regarded as "Least Concern" from a conservation perspective by the International Union for Conservation of Nature (IUCN).[2] However, at least one subspecies, the West African or Nigerian Giraffe (G. c. peralta), has been classified as endangered.[27]
Giraffes are hunted for their tails, hides and meat.[22] The tails are used as good luck charms, thread and flyswatters.[22] In addition, habitat destruction also hurts the giraffe. In the Sahel trees are cut down for firewood and to make way for livestock. Normally, giraffes are able to cope with livestock since they feed in the trees above their heads. The giraffe population is shrinking in West Africa. However, the populations in eastern and southern Africa are stable and, due to the popularity of privately-owned game ranches and sanctuaries (i.e. Bour-Algi Giraffe Sanctuary), are expanding. The giraffe is a protected species in most of its range. The total African giraffe population has been estimated to range from 110,000 to 150,000. Kenya (45,000), Tanzania (30,000), and Botswana (12,000), have the largest national populations.[28]
Scientific inspiration
Giraffes have been used as examples for introducing ideas in evolution, especially to illustrate the ideas of Lamarck. Lamarck believed that the giraffe's long neck developed as a result of ancestral giraffe's reaching to browse on the leaves of tall trees.[29]
The coat patterns of several species of giraffe have been modelled using reaction-diffusion mechanisms.[30]
In art and culture
Painting of a giraffe taken to China from Africa during the Ming DynastyGiraffes can be seen in paintings, including the famous painting of a giraffe which was taken from Africa to China in 1414. The giraffe was placed in a Ming Dynasty zoo.
The Medici giraffe was a giraffe presented to Lorenzo de' Medici in 1486. It caused a great stir on its arrival in Florence, being reputedly the first living giraffe to be seen in Italy since the days of Ancient Rome. Another famous giraffe, called Zarafa, was brought from Africa to Paris in the early 1800s and kept in a menagerie for 18 years.
Giraffe is a novel by the author J. M. Ledgard. The work concerns a true incident in which 49 giraffes were slaughtered in the Czech Republic (then Czechoslovakia) in 1975 following the suspected outbreak of disease amongst the group. The novel contains extensive information about the species, including the long history of European fascination with the beast and its captivity in zoos.
Notable fictional giraffes include:
Toys "R" Us mascot Geoffrey the Giraffe. He was originally portrayed as a cartoon giraffe but in the 2001 commercials he was portrayed as a real-life giraffe who talks; an animatronic version of Geoffrey the Giraffe (created by Stan Winston Studios), was voiced by Jim Hanks in commercials for radio and television.
Longrack of the Transformers universe
Girafarig from the Pokémon franchise
Melman from Madagascar
Tuesday, June 2, 2009
Elephant
From Wikipedia, the free encyclopedia
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For other uses, see Elephant (disambiguation).
Elephant
Fossil range: Pliocene–Recent PreЄЄOSDCPTJKPgN
Asian and African elephants.
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Subphylum: Vertebrata
Class: Mammalia
Order: Proboscidea
Family: Elephantidae
Gray, 1821
Subfamilia
See Classification
Comparative view of the human and elephant frames, c1860.Elephants are large land mammals of the order Proboscidea and the family Elephantidae. There are three living species: the African Bush Elephant, the African Forest Elephant and the Asian Elephant (also known as the Indian Elephant). Other species have become extinct since the last ice age, the Mammoths, dwarf forms of which may have survived as late as 2,000 BC,[1] being the best-known of these. They were once classified along with other thick skinned animals in a now invalid order, Pachydermata.
Elephants are the largest land animals.[2] The elephant's gestation period is 22 months, the longest of any land animal. At birth it is common for an elephant calf to weigh 120 kilograms (260 lb). They typically live for 50 to 70 years, but the oldest recorded elephant lived for 82 years.[3] The largest elephant ever recorded was shot in Angola in 1956. This male weighed about 12,000 kilograms (26,000 lb),[4] with a shoulder height of 4.2 metres (14 ft), a metre (yard) taller than the average male African elephant.[5] The smallest elephants, about the size of a calf or a large pig, were a prehistoric species that lived on the island of Crete during the Pleistocene epoch.[6]
The elephant has appeared in cultures across the world. They are a symbol of wisdom in Asian cultures and are famed for their memory and intelligence, where they are thought to be on par with cetaceans[7] and hominids.[8] Aristotle once said the elephant was "the beast which passeth all others in wit and mind"[9]. The word "elephant" has its origins in the Greek ἐλέφας, meaning "ivory" or "elephant".[10]
Healthy adult elephants have no natural predators[11], although lions may take calves or weak individuals.[12][13] They are, however, increasingly threatened by human intrusion and poaching. Once numbering in the millions, the African elephant population has dwindled to between 470,000 and 690,000 individuals according to a March 2007 estimate.[14] While the elephant is a protected species worldwide, with restrictions in place on capture, domestic use, and trade in products such as ivory, CITES reopening of "one time" ivory stock sales, has resulted in increased poaching. Certain African nations report a decrease of their elephant populations by as much as two-thirds, and populations in certain protected areas are in danger of being eliminated[15] Since recent poaching has increased by as much as 45%, the current population is unknown (2008).[16]
Contents [hide]
1 Taxonomy and evolution
1.1 African Elephant
1.2 Asian Elephant
2 Physical characteristics
2.1 Trunk
2.2 Tusks
2.3 Teeth
2.4 Skin
2.5 Legs and feet
2.6 Ears
3 Biology and behavior
3.1 Social behavior
3.2 Intelligence
3.3 Senses
3.4 Self-awareness
3.5 Communication
3.6 Diet
3.7 Reproduction and life cycle
3.7.1 Elephant calves
3.8 Effect on the environment
4 Threat of extinction
4.1 Hunting
4.2 Habitat loss
4.3 National parks
5 Humanity and elephants
5.1 Harvest from the wild
5.2 Domestication and use
5.2.1 Warfare
5.2.2 Industry
5.2.3 Zoo and circuses
5.2.4 Hybrids
5.3 Elephant rage
5.3.1 Musth
5.3.2 Other causes
5.4 In popular culture
6 See also
7 References
8 Further reading
9 External links
Taxonomy and evolution
See also Elephant classification
Physical difference between an Asian (left) and African (right) elephant.The African Elephant genus contains two (or, arguably, three) living species; whereas the Asian Elephant species is the only surviving member of the Asian Elephant genus, but can be divided into four subspecies.
Evolution of elephants from the ancient Eocene (bottom) to the modern day (top).Although the fossil evidence is uncertain, scientists discovered genetic evidence that the elephant family shares distant ancestry with the sirenians (sea cows) and the hyraxes through gene comparisons. In the distant past, members of the hyrax family grew to large sizes, and it seems likely that the common ancestor of all three modern families was some kind of amphibious hyracoid. One theory suggests that these animals spent most of their time under water, using their trunks like snorkels for breathing.[17][18] Modern elephants have retained this ability and are known to swim in that manner for up to 6 hours and 50 km (30 miles).
In the past, there was a much wider variety of elephant genera, including the mammoths and stegodons. There was also a much wider variety of species.[19][20]
African Elephant
Main articles: African Elephant, African Bush Elephant, and African Forest Elephant
Elephant crossing a river, Kenya.
African bush (savanna) elephant in Mikumi National Park, Tanzania.The Elephants of the genus Loxodonta, known collectively as African elephants, are currently found in 37 countries in Africa.
African elephants are distinguished from Asian elephants in several ways, the most noticeable being their ears which are much larger. The African elephant is typically larger than the Asian elephant and has a concave back. Both African males and females have external tusks and are usually less hairy than their Asian cousins.
African elephants have traditionally been classified as a single species comprising two distinct subspecies, namely the savanna elephant (Loxodonta africana africana) and the forest elephant (Loxodonta africana cyclotis), but recent DNA analysis suggests that these may actually constitute distinct species.[21] This split is not universally accepted by experts[22] and a third species of African elephant has also been proposed.[23]
This reclassification has important implications for conservation, because it means that where previously it was assumed that a single and endangered species comprised two small populations, if in reality these are two separate species, then as a consequence, both could be more gravely endangered than a more numerous and wide-ranging single species might have been. There is also a potential danger in that, if the forest elephant is not explicitly listed as an endangered species, poachers and smugglers might be able to evade the law forbidding trade in endangered animals and their body parts.
The Forest elephant and the Savanna elephant can also hybridise – that is, breed together – successfully, though their preferences for different terrains reduce such opportunities. As the African elephant has only recently been recognized to comprise two separate species, groups of captive elephants have not been comprehensively classified and some could well be hybrids.
Under the new two species classification, Loxodonta africana refers specifically to the Savanna Elephant, the largest of all elephants. In fact, it is the largest land animal in the world, with the males standing 3.2 metres (10 ft) to 4 metres (13 ft) at the shoulder and weighing 3,500 kilograms (7,700 lb) to a reported 12,000 kilograms (26,000 lb).[24]. The female is smaller, standing about 3 metres (9.8 ft) at the shoulder[25]. Most often, Savanna Elephants are found in open grasslands, marshes, and lakeshores. They range over much of the savanna zone south of the Sahara.
The other putative species, the Forest Elephant (Loxodonta cyclotis), is usually smaller and rounder, and its tusks thinner and straighter compared with the Savanna Elephant. The Forest Elephant can weigh up to 4,500 kilograms (9,900 lb) and stand about 3 metres (10 ft) tall. Much less is known about these animals than their savanna cousins, because environmental and political obstacles make them difficult to study. Normally, they inhabit the dense African rain forests of central and western Africa, although occasionally they roam the edges of forests, thus overlapping the Savanna elephant territories and hybridizing. In 1979, Iain Douglas-Hamilton estimated the continental population of African elephants at around 1.3 million animals.[26] This estimate is controversial and is believed to be a gross overestimate,[27] but it is very widely cited and has become a de facto baseline that continues to be incorrectly used to quantify downward population trends in the species. Through the 1980s, Loxodonta received worldwide attention due to the dwindling numbers of major populations in East Africa, largely as a result of poaching. Today, according to IUCN’s African Elephant Status Report 2007[28] there are approximately between 470,000 and 690,000 African elephants in the wild. Although this estimate only covers about half of the total elephant range, experts do not believe the true figure to be much higher, as it is unlikely that large populations remain to be discovered.[29] By far the largest populations are now found in Southern and Eastern Africa, which together account for the majority of the continental population. According to a recent analysis by IUCN experts, most major populations in Eastern and Southern Africa are stable or have been steadily increasing since the mid-1990s, at an average rate of 4.5% per year.[30][31]
Elephant populations in West Africa, on the other hand, are generally small and fragmented, and only account for a small proportion of the continental total.[32] Much uncertainty remains as to the size of the elephant population in Central Africa, where the prevalence of forest makes population surveys difficult, but poaching for ivory and bushmeat is believed to be intense through much of the region.[33] South Africa elephant population more than doubled, rising from 8,000 to over 20,000, in the thirteen years after a 1995 ban on killing the animals.[34] The ban was lifted in February 2008, sparking controversy among environmental groups.[35]
Asian Elephant
Main article: Asian Elephant
The Asian elephant, Elephas maximus, is smaller than the African. It has smaller ears, and typically, only the males have large external tusks.
An Asian elephant named Sri Hari during Sree Poornathrayesa temple festival, Thrippunithura.
A decorated Indian elephant in Jaipur, India.
Elephant in a Truck, Sri Lanka.
"O Elephante" - Hand-coloured engraving drawn by H.Gobin and engraved by Ramus[36]
Elephant in Sri Lanka
The world population of Asian elephants – also called Indian Elephants – is estimated to be around 60,000, about a tenth of the number of African elephants. More precisely, it is estimated that there are between 38,000 and 53,000 wild elephants and between 14,500 and 15,300 domesticated elephants in Asia with perhaps another 1,000 scattered around zoos in the rest of the world.[37] The Asian elephants' decline has possibly been more gradual than the African and caused primarily by poaching and habitat destruction by human encroachment.
Several subspecies of Elephas maximus have been identified, using morphometric data and molecular markers. Elephas maximus maximus (Sri Lankan Elephant) is found only on the island of Sri Lanka. It is the largest of the Asians. There are an estimated 3,000–4,500 members of this subspecies left today in the wild, although no accurate census has been carried out recently. Large males can weigh upward to 5,400 kg (12,000 lb) and stand over 3.4 m (11 ft) tall. Sri Lankan males have very large cranial bulges, and both sexes have more areas of depigmentation than other Asians. Typically, their ears, face, trunk, and belly have large concentrations of pink-speckled skin. There is an orphanage for elephants in Pinnawala, Sri Lanka, which plays a large role in protecting the Sri Lankan Elephant from extinction.
Elephas maximus indicus (Indian Elephant) makes up the bulk of the Asian elephant population. Numbering approximately 36,000, these elephants are lighter grey in colour, with depigmentation only on the ears and trunk. Large males will ordinarily weigh only about 5,000 kg (11,000 lb), but are as tall as the Sri Lankan. The mainland Asian can be found in 11 Asian countries, from India to Indonesia. They prefer forested areas and transitional zones, between forests and grasslands, where greater food variety is available.
The smallest of all the elephants is the Sumatran Elephant, Elephas maximus sumatranus. Population estimates for this group range from 2,100 to 3,000 individuals. It is very light grey in colour and has less depigmentation than the other Asians, with pink spots only on the ears. Mature Sumatrans will usually only measure 1.7–2.6 m (5.6–8.5 ft) at the shoulder and weigh less than 3,000 kg (6,600 lb). It is considerably smaller than its other Asian (and African) cousins and exists only on the island of Sumatra, usually in forested regions and partially wooded habitats.
In 2003, a further subspecies was identified on Borneo. Named the Borneo pygmy elephant, it is smaller and tamer than any other Asian elephants. It also has relatively larger ears, longer tail and straighter tusks.
Physical characteristics
Trunk
Trunk of African (left) and Asian (right) elephant.
Articulation of elephant trunk
An elephant can use its trunk for a variety of purposes. This one is wiping its eye.
Eyes of an Asian elephant.The proboscis, or trunk, is a fusion of the nose and upper lip, elongated and specialized to become the elephant's most important and versatile appendage. African elephants are equipped with two fingerlike projections at the tip of their trunk, while Asians have only one. According to biologists, the elephant's trunk may have over forty thousand individual muscles in it,[38] making it sensitive enough to pick up a single blade of grass, yet strong enough to rip the branches off a tree. Some sources indicate that the correct number of muscles in an elephant's trunk is closer to one hundred thousand.[39]
Most herbivores (plant eaters, like the elephant) possess teeth adapted for cutting and tearing off plant materials. However, except for the very young or infirm, elephants always use their trunks to tear up their food and then place it in their mouth. They will graze on grass or reach up into trees to grasp leaves, fruit, or entire branches. If the desired food item is too high up, the elephant will wrap its trunk around the tree or branch and shake its food loose or sometimes simply knock the tree down altogether.
The trunk is also used for drinking. Elephants suck water up into the trunk (up to fifteen quarts or fourteen litres at a time) and then blow it into their mouth. Elephants also inhale water to spray on their body during bathing. On top of this watery coating, the animal will then spray dirt and mud, which act as a protective sunscreen. When swimming, the trunk makes an excellent snorkel.[17][18]
This appendage also plays a key role in many social interactions. Familiar elephants will greet each other by entwining their trunks, much like a handshake. They also use them while play-wrestling, caressing during courtship and mother / child interactions, and for dominance displays – a raised trunk can be a warning or threat, while a lowered trunk can be a sign of submission. Elephants can defend themselves very well by flailing their trunk at unwanted intruders or by grasping and flinging them.
An elephant also relies on its trunk for its highly developed sense of smell. Raising the trunk up in the air and swivelling it from side to side, like a periscope, it can determine the location of friends, enemies, and food sources.
Tusks
Tusks of African and Asian elephants.The tusks of an elephant are its second upper incisors. Tusks grow continuously; an adult male's tusks will grow about 18 cm (7 in) a year. Tusks are used to dig for water, salt, and roots; to debark trees, to eat the bark; to dig into baobab trees to get at the pulp inside; and to move trees and branches when clearing a path. In addition, they are used for marking trees to establish territory and occasionally as weapons.
Like humans who are typically right- or left-handed, elephants are usually right- or left-tusked. The dominant tusk, called the master tusk, is generally shorter and more rounded at the tip from wear. Both male and female African elephants have large tusks that can reach over 3 m (10 ft) in length and weigh over 90 kg (200 lb). In the Asian species, only the males have large tusks. Female Asians have tusks which are very small or absent altogether. Asian males can have tusks as long as the much larger Africans, but they are usually much slimmer and lighter; the heaviest recorded is 39 kg (86 lb). The tusk of both species is mostly made of calcium phosphate in the form of apatite. As a piece of living tissue, it is relatively soft (compared with other minerals such as rock), and the tusk, also known as ivory, is strongly favoured by artists for its carvability. The desire for elephant ivory has been one of the major factors in the reduction of the world's elephant population.
Some extinct relatives of elephants had tusks in their lower jaws in addition to their upper jaws, such as Gomphotherium, or only in their lower jaws, such as Deinotherium.
Teeth
Elephants' teeth are very different from those of most other mammals. Over their lives they usually have 28 teeth. These are:
The two upper second incisors: these are the tusks.
The milk precursors of the tusks.
12 premolars, 3 in each side of each jaw.
12 molars, 3 in each side of each jaw.
Replica of an Asian Elephant's molar, showing upper sideThis gives elephants a dental formula of:
Dentition
1.0.3.3
0.0.3.3
Unlike most mammals, which grow baby teeth and then replace them with a permanent set of adult teeth, elephants have cycles of tooth rotation throughout their entire life. The tusks have milk precursors, which fall out quickly and the adult tusks are in place by one year of age, but the molars are replaced five times in an average elephant's lifetime.[40] The teeth do not emerge from the jaws vertically like with human teeth. Instead, they move horizontally, like a conveyor belt. New teeth grow in at the back of the mouth, pushing older teeth toward the front, where they wear down with use and the remains fall out. When an elephant becomes very old, the last set of teeth is worn to stumps, and it must rely on softer foods to chew. Very elderly elephants often spend their last years exclusively in marshy areas where they can feed on soft wet grasses. Eventually, when the last teeth fall out, the elephant will be unable to eat and will die of starvation. Were it not for tooth wearout, their metabolism would allow them to live much longer. Rupert Sheldrake has proposed this as an explanation for the elephant graveyards. However, as more habitat is destroyed, the elephants' living space becomes smaller and smaller; the elderly no longer have the opportunity to roam in search of more appropriate food and will, consequently, die of starvation at an earlier age.
Tusks in the lower jaw are also second incisors. These grew out large in Deinotherium and some mastodons, but in modern elephants they disappear early without erupting.
Skin
Skin of an African (left) and Asian (right) elephant.Elephants are colloquially called pachyderms (from their original scientific classification), which means thick-skinned animals. An elephant's skin is extremely tough around most parts of its body and measures about 2.5 centimetres (1.0 in) thick. However, the skin around the mouth and inside of the ear is paper-thin. Normally, the skin of an Asian is covered with more hair than its African counterpart. This is most noticeable in the young. Asian calves are usually covered with a thick coat of brownish red fuzz. As they get older, this hair darkens and becomes more sparse, but it will always remain on their heads and tails.
The species of elephants are typically greyish in colour, but the Africans very often appear brown or reddish from wallowing in mud holes of coloured soil. Wallowing is an important behaviour in elephant society. Not only is it important for socialization, but the mud acts as a sunscreen, protecting their skin from harsh ultraviolet radiation. Although tough, an elephant's skin is very sensitive. Without regular mud baths to protect it from burning, as well as from insect bites and moisture loss, an elephant's skin would suffer serious damage. After bathing, the elephant will usually use its trunk to blow dirt on its body to help dry and bake on its new protective coat. As elephants are limited to smaller and smaller areas, there is less water available, and local herds will often come too close over the right to use these limited resources.
Wallowing also aids the skin in regulating body temperatures. Elephants have difficulty in releasing heat through the skin because, in proportion to their body size, they have very little of it. The ratio of an elephant's mass to the surface area of its skin is many times that of a human. Elephants have even been observed lifting up their legs to expose the soles of their feet, presumably in an effort to expose more skin to the air. Since wild elephants live in very hot climates, they must have other means of getting rid of excess heat.
Legs and feet
Elephant using its feet to crush a watermelon prior to eating itAn elephant's legs are great straight pillars, as they must be to support its bulk. The elephant needs less muscular power to stand because of its straight legs and large pad-like feet. For this reason an elephant can stand for very long periods of time without tiring. In fact, African elephants rarely lie down unless they are sick or wounded. Indian elephants, in contrast, lie down frequently.
The feet of an elephant are nearly round. African elephants have three nails on each hind foot, and four on each front foot. Indian elephants have four nails on each hind foot and five on each front foot. Beneath the bones of the foot is a tough, gelatinous material that acts as a cushion or shock absorber. Under the elephant's weight the foot swells, but it gets smaller when the weight is removed. An elephant can sink deep into mud, but can pull its legs out readily because its feet become smaller when they are lifted.[citation needed]
Elephants swim well, but cannot trot, jump, or gallop. They do have two gaits: a walk; and a faster gait that is similar to running.
In walking, the legs act as pendulums, with the hips and shoulders rising and falling while the foot is planted on the ground. With no "aerial phase", the faster gait does not meet all the criteria of running, as elephants always have at least one foot on the ground. However, an elephant moving fast uses its legs much like a running animal, with the hips and shoulders falling and then rising while the feet are on the ground. In this gait, an elephant will have three feet off the ground at one time. As both of the hind feet and both of the front feet are off the ground at the same time, this gait has been likened to the hind legs and the front legs taking turns running.[41]
Although they start this "run" at only 8 km/h,[42] elephants can reach speeds up to 40 km/h (25 mph),[43] all the while using the same gait. At this speed, most other four-legged creatures are well into a gallop, even accounting for leg length. Spring-like kinetics could explain the difference between the motion of elephants and other animals.[44]
Ears
Difference between Asian (left) and African (right) elephant ears.The large flapping ears of an elephant are also very important for temperature regulation. Elephant ears are made of a very thin layer of skin stretched over cartilage and a rich network of blood vessels. On hot days, elephants will flap their ears constantly, creating a slight breeze. This breeze cools the surface blood vessels, and then the cooler blood gets circulated to the rest of the animal's body. The hot blood entering the ears can be cooled as much as ten degrees Fahrenheit before returning to the body. Differences in the ear sizes of African and Asian elephants can be explained, in part, by their geographical distribution. Africans originated and stayed near the equator, where it is warmer. Therefore, they have bigger ears. Asians live farther north, in slightly cooler climates, and thus have smaller ears.
The ears are also used in certain displays of aggression and during the males' mating period. If an elephant wants to intimidate a predator or rival, it will spread its ears out wide to make itself look more massive and imposing. During the breeding season, males give off an odour from the musth gland located behind their eyes. Joyce Poole, a well-known elephant researcher, has theorized that the males will fan their ears in an effort to help propel this "elephant cologne" great distances. [45]
Biology and behavior
Social behavior
Elephant footprints (tire tracks for scale)Elephants live in a structured social order. The social lives of male and female elephants are very different. The females spend their entire lives in tightly knit family groups made up of mothers, daughters, sisters, and aunts. These groups are led by the eldest female, or matriarch. Adult males, on the other hand, live mostly solitary lives.
The social circle of the female elephant does not end with the small family unit. In addition to encountering the local males that live on the fringes of one or more groups, the female's life also involves interaction with other families, clans, and subpopulations. Most immediate family groups range from five to fifteen adults, as well as a number of immature males and females. When a group gets too big, a few of the elder daughters will break off and form their own small group. They remain very aware of which local herds are relatives and which are not.
The life of the adult male is very different. As he gets older, he begins to spend more time at the edge of the herd, gradually going off on his own for hours or days at a time. Eventually, days become weeks, and somewhere around the age of fourteen, the mature male, or bull, sets out from his natal group for good. While males do live primarily solitary lives, they will occasionally form loose associations with other males. These groups are called bachelor herds. The males spend much more time than the females fighting for dominance with each other. Only the most dominant males will be permitted to breed with cycling females. The less dominant ones must wait their turn. It is usually the older bulls, forty to fifty years old, that do most of the breeding.
The dominance battles between males can look very fierce, but typically they inflict very little injury. Most of the bouts are in the form of aggressive displays and bluffs. Ordinarily, the smaller, younger, and less confident animal will back off before any real damage can be done. However, during the breeding season, the battles can get extremely aggressive, and the occasional elephant is injured. During this season, known as musth, a bull will fight with almost any other male it encounters, and it will spend most of its time hovering around the female herds, trying to find a receptive mate.
African as well as Asiatic males will engage in same-sex bonding and mounting. Such encounters are often associated with affectionate interactions, such as kissing, trunk intertwining, and placing trunks in each other's mouths. The encounters are analogous to heterosexual bouts, one male often extending his trunk along the other's back and pushing forward with his tusks to signify his intention to mount. Unlike heterosexual relations, which are always of a fleeting nature, those between males result in a "companionship", consisting of an older individual and one or two younger, attendant males. Same-sex relations are common and frequent in both sexes, with Asiatic elephants in captivity devoting roughly 46% of sexual encounters to same-sex activity.[46]
Rogue elephant is a term for a lone, violently aggressive wild elephant. It is a calque of the Sinhala term hora aliya. Its introduction to English has been attributed by the Oxford English Dictionary to Sir James Emerson Tennent, but this usage may have been pre-dated by William Sirr.
Intelligence
Main article: Elephant intelligence
Human, dolphin and elephant brains up to scale. (1)-cerebrum (1a)-temporal lobe and (2)-cerebellumWith a mass just over 5 kg (11 lb), elephant brains are larger than those of any other land animal, and although the largest whales have body masses twentyfold those of a typical elephant, whale brains are barely twice the mass of an elephant's. A wide variety of behaviours, including those associated with grief, making music, art, altruism, allomothering, play, use of tools,[47] compassion and self-awareness [48] evidence a highly intelligent species on par with cetaceans[7] and primates[8]. The largest areas in the elephant brain are those responsible for hearing, smell and movement coordination.
Senses
Elephants have well innervated trunks, and an exceptional sense of hearing and smell. The hearing receptors reside not only in ears, but also in trunks that are sensitive to vibrations, and most significantly feet, which have special receptors for low frequency sound and are exceptionally well innervated. Elephants communicate by sound over large distances of several kilometers partly through the ground, which is important for their social lives. Elephants are observed listening by putting trunks on the ground and carefully positioning their feet.
Their eyesight is relatively poor, and the eyes are aiming down the trunk. An elephant has to raise his head conspicuously to look out horizontally[citation needed].
Self-awareness
Mirror self recognition is a test of self awareness and cognition used in animal studies. A mirror was provided and visible marks were made on the elephant. The elephants investigated these marks, which were visible only via the mirror. The tests also included non-visible marks to rule out the possibility of their using other senses to detect these marks. This shows that elephants recognize the fact that the image in the mirror is their own self and such abilities are considered the basis for empathy, altruism and higher social interactions. This ability has been demonstrated in humans, apes, dolphins,[49] and magpies.[50]
A young elephant in Zimbabwe.
Communication
Elephants make a number of sounds when communicating. Elephant are famous for their trumpet calls which are made when the animal blows though its nostrils. Trumpeting is usually made during excitement. Its use varies from startlement to a cry of help to rage. Elephants also make rumbling growls when greeting each other. The growl become a bellow when the mouth is open and a bellow becomes a moan when prolonged. This can escalate with a roar when threatening another elephant or another animal.
Elephants can communicate over long distances by producing and receiving low-frequency sound (infrasound), a sub-sonic rumbling, which can travel in the air and through the ground much farther than higher frequencies. These calls range in frequency from 15-35 Hz and can be as loud as 117 Db, allowing communication for many kilometres, with a possible maximum range of around 10 km.[51] This sound can be felt by the sensitive skin of an elephant's feet and trunk, which pick up the resonant vibrations much as the flat skin on the head of a drum. To listen attentively, every member of the herd will lift one foreleg from the ground, and face the source of the sound, or often lay its trunk on the ground. The lifting presumably increases the ground contact and sensitivity of the remaining legs. This ability is thought also to aid their navigation by use of external sources of infrasound. Discovery of this new aspect of elephant social communication and perception came with breakthroughs in audio technology, which can pick up frequencies outside the range of the human ear. Pioneering research in elephant infrasound communication was done by Katy Payne, of the Elephant Listening Project,[52] and is detailed in her book Silent Thunder. Though this research is still in its infancy, it is helping to solve many mysteries, such as how elephants can find distant potential mates, and how social groups are able to coordinate their movements over extensive range.[51]
Diet
Elephants are herbivores, spending 16 hours a day collecting plant food. Their diet is at least 50% grasses, supplemented with leaves, bamboo, twigs, bark, roots, and small amounts of fruits, seeds and flowers. Because elephants only digest 40% of what they eat, they have to make up for their digestive system's lack of efficiency in volume. An adult elephant can consume 140–270 kg (300–600 lb) of food a day. 60% of that food leaves the elephant's body undigested[citation needed].
Reproduction and life cycle
Elephant calves
Elephant social life revolves around breeding and raising of the calves. A female will usually be ready to breed around the age of thirteen, when she comes into estrus, a short phase of receptiveness lasting a couple of days, for the first time. Females announce their estrus with smell signals and special calls.
Female African elephant with calf, in Kenya.Females prefer bigger, stronger, and, most importantly, older males. Such a reproductive strategy tends to increase their offspring's chances of survival.
After a twenty-two-month pregnancy, the mother will give birth to a calf that will weigh about 113 kg (250 lb) and stand over 76 cm (2.5 ft) tall. Elephants have a very long childhood. They are born with fewer survival instincts than many other animals. Instead, they must rely on their elders to teach them the things they need to know. Today, however, the pressures humans have put on the wild elephant populations, from poaching to habitat destruction, mean that the elderly often die at a younger age, leaving fewer teachers for the young.
A new calf is usually the centre of attention for all herd members. All the adults and most of the other young will gather around the newborn, touching and caressing it with their trunks. The baby is born nearly blind and at first relies, almost completely, on its trunk to discover the world around it.
As everyone in the herd is usually related, all members of the tightly knit female group participate in the care and protection of the young. After the initial excitement, the mother will usually select several full-time baby-sitters, or "allomothers", from her group. According to Cynthia Moss, a well known researcher, these allomothers will help in all aspects of raising the calf.[53] They walk with the young as the herd travels, helping the calves along if they fall or get stuck in the mud. The more allomothers a baby has, the more free time its mother has to feed herself. Providing a calf with nutritious milk means the mother has to eat more nutritious food herself. So, the more allomothers, the better the calf's chances of survival. An elephant is considered an allomother when she is not able to have her own baby. A benefit of being an allomother is that she can gain experience or receive assistance when caring for her own calf.
Effect on the environment
Elephants are a species which many other organisms depend on. One particular example of that are termites mounds: termites eat elephant feces and often begin building their mounds under piles of elephant feces.
Elephants' foraging activities can sometimes greatly affect the areas in which they live. By pulling down trees to eat leaves, breaking branches, and pulling out roots they create clearings in which new young trees and other vegetation can establish themselves. During the dry season, elephants use their tusks to dig into dry river beds to reach underground sources of water. These newly dug water holes may then become the only source of water in the area. Elephants make pathways through their environment which are also used by other animals to access areas normally out of reach. These pathways have sometimes been used by several generations of elephants and today are converted by humans to paved roads.
Threat of extinction
Men with African Elephant tusks, Dar es Salaam, c. 1900
An Elephant resting his head on a tree trunk, Samburu National Reserve, Kenya
An elephant in the Ngorongoro crater, Tanzania
African Savanna Elephant Loxodonta africana, born 1969 (left), and Asian Elephant Elephas maximus, born 1970 (right), at an English zoo
Hunting
The threat to the African elephant presented by the ivory trade is unique to the species. Larger, long-lived, slow-breeding animals, like the elephant, are more susceptible to overhunting than other animals. They cannot hide, and it takes many years for an elephant to grow and reproduce. An elephant needs an average of 140 kg (300 lb) of vegetation a day to survive. As large predators are hunted, the local small grazer populations (the elephant's food competitors) find themselves on the rise. The increased number of herbivores ravage the local trees, shrubs, and grasses. Elephants themselves have few natural predators besides man and, occasionally, lions. However, many African governments legally allow limited hunting. The large amount of money that is charged for the necessary permits is often used to support conservation efforts, and the small number of permits issued (usually for older animals) ensure that populations are not depleted.[1]
Habitat loss
Another threat to elephant's survival in general is the ongoing cultivation of their habitats with increasing risk of conflicts of interest with human cohabitants. These conflicts kill 150 elephants and up to 100 people per year in Sri Lanka.[54] Lacking the massive tusks of its African cousins, the Asian elephant's demise can be attributed mostly to loss of its habitat.
As larger patches of forest disappear, the ecosystem is affected in profound ways. The trees are responsible for anchoring soil and absorbing water runoff. Floods and massive erosion are common results of deforestation. Elephants need massive tracts of land because, much like the slash-and-burn farmers, they are used to crashing through the forest, tearing down trees and shrubs for food and then cycling back later on, when the area has regrown. As forests are reduced to small pockets, elephants become part of the problem, quickly destroying all the vegetation in an area, eliminating all their resources.
National parks
An Elephant sanctuary at Punnathur kotta, Kerala, south India.Africa's first official reserve, Kruger National Park, eventually became one of the world's most famous and successful national parks. [55] There are, however, many problems associated with the establishment of these reserves. For example, elephants range through a wide tract of land with little regard for national borders. Once a reserve is established and fence erected, many animals find themselves cut off from their winter feeding grounds or spring breeding areas. Some animals may die as a result, while others, like the elephants, may just trample over the fences, wreaking havoc in nearby fields. When confined to small territories, elephants can inflict an enormous amount of damage to the local landscapes. [56]
Additionally, some reserves, such as Kruger National Park has, in the opinion of wildlife managers, suffered from elephant overcrowding, at the expense of other species of wildlife within the reserve. On 25 February 2008, the South Africa announced that they would reintroduce culling for the first time since 1994 to control elephant numbers.[57] Nevertheless, as scientists learn more about nature and the environment, it becomes very clear that these parks may be the elephant's last hope against the rapidly changing world around them.
Humanity and elephants
Harvest from the wild
The harvest of elephants, both legal and illegal, has had some unexpected consequences on elephant anatomy as well. African ivory hunters, by killing only tusked elephants, have given a much larger chance of mating to elephants with small tusks or no tusks at all. The propagation of the absent-tusk gene has resulted in the birth of large numbers of tuskless elephants, now approaching 30% in some populations (compare with a rate of about 1% in 1930). Tusklessness, once a very rare genetic abnormality, has become a widespread hereditary trait.
It is possible, if unlikely, that continued selection pressure could bring about a complete absence of tusks in African elephants, a development normally requiring thousands of years of evolution. The effect of tuskless elephants on the environment, and on the elephants themselves, could be dramatic. Elephants use their tusks to root around in the ground for necessary minerals, tear apart vegetation, and spar with one another for mating rights. Without tusks, elephant behaviour could change dramatically.[58]
Domestication and use
Elephants have been working animals used in various capacities by humans. Seals found in the Indus Valley suggest that the elephant was first domesticated in ancient India. However, elephants have never been truly domesticated: the male elephant in his periodic condition of musth is dangerous and difficult to control. Therefore elephants used by humans have typically been female, war elephants being an exception, however: as female elephants in battle will run from a male, only males could be used in war. It is generally more economical to capture wild young elephants and tame them than breeding them in captivity (see also elephant "crushing").
The Judean rebel Eleazar Maccabeus kills a Seleucid war elephant and is crushed under it (Miniature from a manuscript Speculum Humanae Salvationis).The Lao PDR has been domesticating elephant for centuries, and still employs an approximate 500 domesticated elephants, the majority of which work in the Xaignabouli province. These elephants are mainly employed in the logging industry, with ecotourism emerging as a sustainable and environmentally friendly alternative. Elefantasia is a local INGO aiming to reconvert logging elephants into ecotourism practices, thus allowing Asian elephants the ability to supply their mahouts with income whilst still allowed to breed.
Elephants are also commonly exhibited in zoos and wild animal parks. 1200 Elephants are kept in western zoos. A study shows that the lifespan of elephants in European zoos is about half as long as those living in protected areas in Africa and Asia.[59]
Elephants are revered in India (and are worshipped in ceremonies such as the Aanayoottu).
Warfare
Main article: War elephant
War elephants were used by armies in the Indian sub-continent, the Warring States of China, and later by the Persian Empire. This use was adopted by Hellenistic armies after Alexander the Great experienced their worth against king Porus, notably in the Ptolemaic and Seleucid diadoch empires. The Carthaginian general Hannibal took elephants across the Alps when he was fighting the Romans, but brought too few elephants to be of much military use, although his horse cavalry was quite successful; he probably used a now-extinct third African (sub)species, the North African (Forest) elephant, smaller than its two southern cousins, and presumably easier to domesticate. A large elephant in full charge could cause tremendous damage to infantry, and cavalry horses would be afraid of them (see Battle of Hydaspes).
Industry
Elephant work camp in Thailand. Elephants are used for heavy forest work and in circus presentationsThroughout Myanmar (Burma), Siam, India, and most of South Asia elephants were used in the military for heavy labour, especially for uprooting trees and moving logs, and were also commonly used as executioners to crush the condemned underfoot.
Elephants have also been used as mounts for safari-type hunting, especially Indian shikar (mainly on tigers), and as ceremonial mounts for royal and religious occasions, whilst Asian elephants have been used for transport and entertainment.
Zoo and circuses
There is growing resistance[60] against the capture, confinement, and use of wild elephants. Animal rights advocates allege that elephants in zoos and circuses "suffer a life of chronic physical ailments, social deprivation, emotional starvation, and premature death".[61] Zoos argue that standards for treatment of elephants are extremely high and that minimum requirements for such things as minimum space requirements, enclosure design, nutrition, reproduction, enrichment and veterinary care are set to ensure the wellbeing of elephants in captivity. Circuses continue to have a mixed record. Recently, the city of Los Angeles' closed an elephant act with Circus Vazquez due to numerous instances of abuse and neglect (April 2008) [62], and, according to PETA, Ringling Bros. and Barnum & Bailey Circus has lost 25 elephants since 1992.[63]
Elephants have traditionally been a major part of circuses around the world, being intelligent enough to be trained in a variety of acts (see for example P.T. Barnum's Jumbo and John L. Sullivan, the famous "Boxing Elephant"). However, conditions for circus elephants are highly unnatural (confinement in small pens or cages, restraints on their feet, lack of companionship of other elephants, etc) and, perhaps as a result, there are instances of them turning on their keepers or handlers (examples include Black Diamond and "Murderous Mary").
Elephants raised in captivity sometimes show "rocking behavior", a rhythmic and repetitive swaying which is unreported in free ranging wild elephants. Thought to be symptomatic of stress disorders, and probably made worse by a barren environment,[64] rocking behavior may be a precursor to aggressive behavior in captive elephants.[65][66] This link is to an image of Devi (little princess), a 30-year-old Asian Elephant raised in captivity at the San Diego Zoo showing "rocking behavior".
Hybrids
Although successful hybridisation between African and Asian Elephant species is highly unlikely in the wild, in 1978 at Chester Zoo, an Asian elephant cow gave birth to a hybrid calf sired by an African elephant bull (the old terms are used here as these events pre-date the current classifications). "Motty", the resulting hybrid male calf, had an African elephant's cheeks, their ears (large with pointed lobes) and legs (longer and slimmer), but the toenail numbers, (5 for each front foot, 4 hind) and the single trunk finger of an Asian elephant. His wrinkled trunk was like that of an African elephant. His forehead was sloping with one dome and two smaller domes behind it. The body was African in type, but had an Asian-type centre hump and an African-type rear hump. The calf died of infection 12 days later[67]. It is preserved as a mounted specimen at the British Natural History Museum, London. There are unconfirmed rumours of three other hybrid elephants born in zoos or circuses; all are said to have been deformed and none survived.
Elephant rage
Devi (little princess), a 30-year-old Asian Elephant raised in captivity at the San Diego Zoo exhibiting "rocking behavior" (animation), a rhythmic and repetitive swaying which is unreported in free ranging wild elephants. Thought to be symptomatic of stress disorders, and probably made worse by a barren environment,[68] rocking behavior may be a precursor to aggressive behavior in captive elephants.Despite its popularity in zoos, and cuddly portrayal as gentle giants in fiction, elephants are among the world's most potentially dangerous animals. They can crush and kill any other land animal, even the rhinoceros. They can experience unexpected bouts of rage, and can be vindictive.[69] In Africa, groups of young teenage elephants attack human villages in what is thought to be revenge for the destruction of their society by massive cullings done in the 1970s and 80s.[70][71] In India, male elephants attack villages at night, destroying homes and killing people regularly. In the Indian state of Jharkhand, 300 people were killed by elephants between 2000 and 2004, and in Assam, 239 people have been killed by elephants since 2001.[69] In India, elephants kill up to 200 humans every year, and in Sri Lanka around 50 per year.
Musth
Main article: Musth
Adult male elephants naturally periodically enter the state called musth (Hindi for "madness"), sometimes spelt "must" in English.
Other causes
At least a few elephants have been suspected to be drunk during their attacks. In December 1998, a herd of elephants overran a village in India. Although locals reported that nearby elephants had recently been observed drinking beer which rendered them "unpredictable", officials considered it the least likely explanation for the attack.[72] An attack on another Indian village occurred in October 1999, and again locals believed the reason was drunkenness, but the theory was not widely accepted.[73] Purportedly drunk elephants raided yet another Indian village again on December 2002, killing six people, which led to killing of about 200 elephants by locals.[74]
In popular culture
See also: Cultural depictions of elephants
Rudyard Kipling's Elephant's Child.Elephants are ubiquitous in Western popular culture as emblems of the exotic[75] because their unique appearance and size sets them apart from other animals and because, like other African animals such as the giraffe, rhinoceros, and hippopotamus, they are unfamiliar to Western audiences.[76] Popular culture's stock references to elephants rely on this exotic uniqueness.[76] For instance, a "white elephant" is a byword for something expensive, useless and bizarre.[76]
As characters, elephants are relegated largely to children's literature,[75] in which they are generally cast as models of exemplary behaviour,[75] but account for some of this branch of literature's most iconic characters.[75] Many stories stell of isolated young elephants returning to a close-knit community, such as The Elephant’s Child from Rudyard Kipling's Just So Stories (1902), Dumbo (1942) or The Saggy Baggy Elephant (1947).[76] Other elephant heroes given human qualities include Laurent de Brunhoff's anthropomorphic Babar (1935), David McKee's Elmer (1989) and Dr. Seuss's Horton (1940).[76] More than other exotic animals, elephants in fiction are surrogates for humans,[76] with their concern for the community and each other depicted as something to aspire to.[77]
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For other uses, see Elephant (disambiguation).
Elephant
Fossil range: Pliocene–Recent PreЄЄOSDCPTJKPgN
Asian and African elephants.
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Subphylum: Vertebrata
Class: Mammalia
Order: Proboscidea
Family: Elephantidae
Gray, 1821
Subfamilia
See Classification
Comparative view of the human and elephant frames, c1860.Elephants are large land mammals of the order Proboscidea and the family Elephantidae. There are three living species: the African Bush Elephant, the African Forest Elephant and the Asian Elephant (also known as the Indian Elephant). Other species have become extinct since the last ice age, the Mammoths, dwarf forms of which may have survived as late as 2,000 BC,[1] being the best-known of these. They were once classified along with other thick skinned animals in a now invalid order, Pachydermata.
Elephants are the largest land animals.[2] The elephant's gestation period is 22 months, the longest of any land animal. At birth it is common for an elephant calf to weigh 120 kilograms (260 lb). They typically live for 50 to 70 years, but the oldest recorded elephant lived for 82 years.[3] The largest elephant ever recorded was shot in Angola in 1956. This male weighed about 12,000 kilograms (26,000 lb),[4] with a shoulder height of 4.2 metres (14 ft), a metre (yard) taller than the average male African elephant.[5] The smallest elephants, about the size of a calf or a large pig, were a prehistoric species that lived on the island of Crete during the Pleistocene epoch.[6]
The elephant has appeared in cultures across the world. They are a symbol of wisdom in Asian cultures and are famed for their memory and intelligence, where they are thought to be on par with cetaceans[7] and hominids.[8] Aristotle once said the elephant was "the beast which passeth all others in wit and mind"[9]. The word "elephant" has its origins in the Greek ἐλέφας, meaning "ivory" or "elephant".[10]
Healthy adult elephants have no natural predators[11], although lions may take calves or weak individuals.[12][13] They are, however, increasingly threatened by human intrusion and poaching. Once numbering in the millions, the African elephant population has dwindled to between 470,000 and 690,000 individuals according to a March 2007 estimate.[14] While the elephant is a protected species worldwide, with restrictions in place on capture, domestic use, and trade in products such as ivory, CITES reopening of "one time" ivory stock sales, has resulted in increased poaching. Certain African nations report a decrease of their elephant populations by as much as two-thirds, and populations in certain protected areas are in danger of being eliminated[15] Since recent poaching has increased by as much as 45%, the current population is unknown (2008).[16]
Contents [hide]
1 Taxonomy and evolution
1.1 African Elephant
1.2 Asian Elephant
2 Physical characteristics
2.1 Trunk
2.2 Tusks
2.3 Teeth
2.4 Skin
2.5 Legs and feet
2.6 Ears
3 Biology and behavior
3.1 Social behavior
3.2 Intelligence
3.3 Senses
3.4 Self-awareness
3.5 Communication
3.6 Diet
3.7 Reproduction and life cycle
3.7.1 Elephant calves
3.8 Effect on the environment
4 Threat of extinction
4.1 Hunting
4.2 Habitat loss
4.3 National parks
5 Humanity and elephants
5.1 Harvest from the wild
5.2 Domestication and use
5.2.1 Warfare
5.2.2 Industry
5.2.3 Zoo and circuses
5.2.4 Hybrids
5.3 Elephant rage
5.3.1 Musth
5.3.2 Other causes
5.4 In popular culture
6 See also
7 References
8 Further reading
9 External links
Taxonomy and evolution
See also Elephant classification
Physical difference between an Asian (left) and African (right) elephant.The African Elephant genus contains two (or, arguably, three) living species; whereas the Asian Elephant species is the only surviving member of the Asian Elephant genus, but can be divided into four subspecies.
Evolution of elephants from the ancient Eocene (bottom) to the modern day (top).Although the fossil evidence is uncertain, scientists discovered genetic evidence that the elephant family shares distant ancestry with the sirenians (sea cows) and the hyraxes through gene comparisons. In the distant past, members of the hyrax family grew to large sizes, and it seems likely that the common ancestor of all three modern families was some kind of amphibious hyracoid. One theory suggests that these animals spent most of their time under water, using their trunks like snorkels for breathing.[17][18] Modern elephants have retained this ability and are known to swim in that manner for up to 6 hours and 50 km (30 miles).
In the past, there was a much wider variety of elephant genera, including the mammoths and stegodons. There was also a much wider variety of species.[19][20]
African Elephant
Main articles: African Elephant, African Bush Elephant, and African Forest Elephant
Elephant crossing a river, Kenya.
African bush (savanna) elephant in Mikumi National Park, Tanzania.The Elephants of the genus Loxodonta, known collectively as African elephants, are currently found in 37 countries in Africa.
African elephants are distinguished from Asian elephants in several ways, the most noticeable being their ears which are much larger. The African elephant is typically larger than the Asian elephant and has a concave back. Both African males and females have external tusks and are usually less hairy than their Asian cousins.
African elephants have traditionally been classified as a single species comprising two distinct subspecies, namely the savanna elephant (Loxodonta africana africana) and the forest elephant (Loxodonta africana cyclotis), but recent DNA analysis suggests that these may actually constitute distinct species.[21] This split is not universally accepted by experts[22] and a third species of African elephant has also been proposed.[23]
This reclassification has important implications for conservation, because it means that where previously it was assumed that a single and endangered species comprised two small populations, if in reality these are two separate species, then as a consequence, both could be more gravely endangered than a more numerous and wide-ranging single species might have been. There is also a potential danger in that, if the forest elephant is not explicitly listed as an endangered species, poachers and smugglers might be able to evade the law forbidding trade in endangered animals and their body parts.
The Forest elephant and the Savanna elephant can also hybridise – that is, breed together – successfully, though their preferences for different terrains reduce such opportunities. As the African elephant has only recently been recognized to comprise two separate species, groups of captive elephants have not been comprehensively classified and some could well be hybrids.
Under the new two species classification, Loxodonta africana refers specifically to the Savanna Elephant, the largest of all elephants. In fact, it is the largest land animal in the world, with the males standing 3.2 metres (10 ft) to 4 metres (13 ft) at the shoulder and weighing 3,500 kilograms (7,700 lb) to a reported 12,000 kilograms (26,000 lb).[24]. The female is smaller, standing about 3 metres (9.8 ft) at the shoulder[25]. Most often, Savanna Elephants are found in open grasslands, marshes, and lakeshores. They range over much of the savanna zone south of the Sahara.
The other putative species, the Forest Elephant (Loxodonta cyclotis), is usually smaller and rounder, and its tusks thinner and straighter compared with the Savanna Elephant. The Forest Elephant can weigh up to 4,500 kilograms (9,900 lb) and stand about 3 metres (10 ft) tall. Much less is known about these animals than their savanna cousins, because environmental and political obstacles make them difficult to study. Normally, they inhabit the dense African rain forests of central and western Africa, although occasionally they roam the edges of forests, thus overlapping the Savanna elephant territories and hybridizing. In 1979, Iain Douglas-Hamilton estimated the continental population of African elephants at around 1.3 million animals.[26] This estimate is controversial and is believed to be a gross overestimate,[27] but it is very widely cited and has become a de facto baseline that continues to be incorrectly used to quantify downward population trends in the species. Through the 1980s, Loxodonta received worldwide attention due to the dwindling numbers of major populations in East Africa, largely as a result of poaching. Today, according to IUCN’s African Elephant Status Report 2007[28] there are approximately between 470,000 and 690,000 African elephants in the wild. Although this estimate only covers about half of the total elephant range, experts do not believe the true figure to be much higher, as it is unlikely that large populations remain to be discovered.[29] By far the largest populations are now found in Southern and Eastern Africa, which together account for the majority of the continental population. According to a recent analysis by IUCN experts, most major populations in Eastern and Southern Africa are stable or have been steadily increasing since the mid-1990s, at an average rate of 4.5% per year.[30][31]
Elephant populations in West Africa, on the other hand, are generally small and fragmented, and only account for a small proportion of the continental total.[32] Much uncertainty remains as to the size of the elephant population in Central Africa, where the prevalence of forest makes population surveys difficult, but poaching for ivory and bushmeat is believed to be intense through much of the region.[33] South Africa elephant population more than doubled, rising from 8,000 to over 20,000, in the thirteen years after a 1995 ban on killing the animals.[34] The ban was lifted in February 2008, sparking controversy among environmental groups.[35]
Asian Elephant
Main article: Asian Elephant
The Asian elephant, Elephas maximus, is smaller than the African. It has smaller ears, and typically, only the males have large external tusks.
An Asian elephant named Sri Hari during Sree Poornathrayesa temple festival, Thrippunithura.
A decorated Indian elephant in Jaipur, India.
Elephant in a Truck, Sri Lanka.
"O Elephante" - Hand-coloured engraving drawn by H.Gobin and engraved by Ramus[36]
Elephant in Sri Lanka
The world population of Asian elephants – also called Indian Elephants – is estimated to be around 60,000, about a tenth of the number of African elephants. More precisely, it is estimated that there are between 38,000 and 53,000 wild elephants and between 14,500 and 15,300 domesticated elephants in Asia with perhaps another 1,000 scattered around zoos in the rest of the world.[37] The Asian elephants' decline has possibly been more gradual than the African and caused primarily by poaching and habitat destruction by human encroachment.
Several subspecies of Elephas maximus have been identified, using morphometric data and molecular markers. Elephas maximus maximus (Sri Lankan Elephant) is found only on the island of Sri Lanka. It is the largest of the Asians. There are an estimated 3,000–4,500 members of this subspecies left today in the wild, although no accurate census has been carried out recently. Large males can weigh upward to 5,400 kg (12,000 lb) and stand over 3.4 m (11 ft) tall. Sri Lankan males have very large cranial bulges, and both sexes have more areas of depigmentation than other Asians. Typically, their ears, face, trunk, and belly have large concentrations of pink-speckled skin. There is an orphanage for elephants in Pinnawala, Sri Lanka, which plays a large role in protecting the Sri Lankan Elephant from extinction.
Elephas maximus indicus (Indian Elephant) makes up the bulk of the Asian elephant population. Numbering approximately 36,000, these elephants are lighter grey in colour, with depigmentation only on the ears and trunk. Large males will ordinarily weigh only about 5,000 kg (11,000 lb), but are as tall as the Sri Lankan. The mainland Asian can be found in 11 Asian countries, from India to Indonesia. They prefer forested areas and transitional zones, between forests and grasslands, where greater food variety is available.
The smallest of all the elephants is the Sumatran Elephant, Elephas maximus sumatranus. Population estimates for this group range from 2,100 to 3,000 individuals. It is very light grey in colour and has less depigmentation than the other Asians, with pink spots only on the ears. Mature Sumatrans will usually only measure 1.7–2.6 m (5.6–8.5 ft) at the shoulder and weigh less than 3,000 kg (6,600 lb). It is considerably smaller than its other Asian (and African) cousins and exists only on the island of Sumatra, usually in forested regions and partially wooded habitats.
In 2003, a further subspecies was identified on Borneo. Named the Borneo pygmy elephant, it is smaller and tamer than any other Asian elephants. It also has relatively larger ears, longer tail and straighter tusks.
Physical characteristics
Trunk
Trunk of African (left) and Asian (right) elephant.
Articulation of elephant trunk
An elephant can use its trunk for a variety of purposes. This one is wiping its eye.
Eyes of an Asian elephant.The proboscis, or trunk, is a fusion of the nose and upper lip, elongated and specialized to become the elephant's most important and versatile appendage. African elephants are equipped with two fingerlike projections at the tip of their trunk, while Asians have only one. According to biologists, the elephant's trunk may have over forty thousand individual muscles in it,[38] making it sensitive enough to pick up a single blade of grass, yet strong enough to rip the branches off a tree. Some sources indicate that the correct number of muscles in an elephant's trunk is closer to one hundred thousand.[39]
Most herbivores (plant eaters, like the elephant) possess teeth adapted for cutting and tearing off plant materials. However, except for the very young or infirm, elephants always use their trunks to tear up their food and then place it in their mouth. They will graze on grass or reach up into trees to grasp leaves, fruit, or entire branches. If the desired food item is too high up, the elephant will wrap its trunk around the tree or branch and shake its food loose or sometimes simply knock the tree down altogether.
The trunk is also used for drinking. Elephants suck water up into the trunk (up to fifteen quarts or fourteen litres at a time) and then blow it into their mouth. Elephants also inhale water to spray on their body during bathing. On top of this watery coating, the animal will then spray dirt and mud, which act as a protective sunscreen. When swimming, the trunk makes an excellent snorkel.[17][18]
This appendage also plays a key role in many social interactions. Familiar elephants will greet each other by entwining their trunks, much like a handshake. They also use them while play-wrestling, caressing during courtship and mother / child interactions, and for dominance displays – a raised trunk can be a warning or threat, while a lowered trunk can be a sign of submission. Elephants can defend themselves very well by flailing their trunk at unwanted intruders or by grasping and flinging them.
An elephant also relies on its trunk for its highly developed sense of smell. Raising the trunk up in the air and swivelling it from side to side, like a periscope, it can determine the location of friends, enemies, and food sources.
Tusks
Tusks of African and Asian elephants.The tusks of an elephant are its second upper incisors. Tusks grow continuously; an adult male's tusks will grow about 18 cm (7 in) a year. Tusks are used to dig for water, salt, and roots; to debark trees, to eat the bark; to dig into baobab trees to get at the pulp inside; and to move trees and branches when clearing a path. In addition, they are used for marking trees to establish territory and occasionally as weapons.
Like humans who are typically right- or left-handed, elephants are usually right- or left-tusked. The dominant tusk, called the master tusk, is generally shorter and more rounded at the tip from wear. Both male and female African elephants have large tusks that can reach over 3 m (10 ft) in length and weigh over 90 kg (200 lb). In the Asian species, only the males have large tusks. Female Asians have tusks which are very small or absent altogether. Asian males can have tusks as long as the much larger Africans, but they are usually much slimmer and lighter; the heaviest recorded is 39 kg (86 lb). The tusk of both species is mostly made of calcium phosphate in the form of apatite. As a piece of living tissue, it is relatively soft (compared with other minerals such as rock), and the tusk, also known as ivory, is strongly favoured by artists for its carvability. The desire for elephant ivory has been one of the major factors in the reduction of the world's elephant population.
Some extinct relatives of elephants had tusks in their lower jaws in addition to their upper jaws, such as Gomphotherium, or only in their lower jaws, such as Deinotherium.
Teeth
Elephants' teeth are very different from those of most other mammals. Over their lives they usually have 28 teeth. These are:
The two upper second incisors: these are the tusks.
The milk precursors of the tusks.
12 premolars, 3 in each side of each jaw.
12 molars, 3 in each side of each jaw.
Replica of an Asian Elephant's molar, showing upper sideThis gives elephants a dental formula of:
Dentition
1.0.3.3
0.0.3.3
Unlike most mammals, which grow baby teeth and then replace them with a permanent set of adult teeth, elephants have cycles of tooth rotation throughout their entire life. The tusks have milk precursors, which fall out quickly and the adult tusks are in place by one year of age, but the molars are replaced five times in an average elephant's lifetime.[40] The teeth do not emerge from the jaws vertically like with human teeth. Instead, they move horizontally, like a conveyor belt. New teeth grow in at the back of the mouth, pushing older teeth toward the front, where they wear down with use and the remains fall out. When an elephant becomes very old, the last set of teeth is worn to stumps, and it must rely on softer foods to chew. Very elderly elephants often spend their last years exclusively in marshy areas where they can feed on soft wet grasses. Eventually, when the last teeth fall out, the elephant will be unable to eat and will die of starvation. Were it not for tooth wearout, their metabolism would allow them to live much longer. Rupert Sheldrake has proposed this as an explanation for the elephant graveyards. However, as more habitat is destroyed, the elephants' living space becomes smaller and smaller; the elderly no longer have the opportunity to roam in search of more appropriate food and will, consequently, die of starvation at an earlier age.
Tusks in the lower jaw are also second incisors. These grew out large in Deinotherium and some mastodons, but in modern elephants they disappear early without erupting.
Skin
Skin of an African (left) and Asian (right) elephant.Elephants are colloquially called pachyderms (from their original scientific classification), which means thick-skinned animals. An elephant's skin is extremely tough around most parts of its body and measures about 2.5 centimetres (1.0 in) thick. However, the skin around the mouth and inside of the ear is paper-thin. Normally, the skin of an Asian is covered with more hair than its African counterpart. This is most noticeable in the young. Asian calves are usually covered with a thick coat of brownish red fuzz. As they get older, this hair darkens and becomes more sparse, but it will always remain on their heads and tails.
The species of elephants are typically greyish in colour, but the Africans very often appear brown or reddish from wallowing in mud holes of coloured soil. Wallowing is an important behaviour in elephant society. Not only is it important for socialization, but the mud acts as a sunscreen, protecting their skin from harsh ultraviolet radiation. Although tough, an elephant's skin is very sensitive. Without regular mud baths to protect it from burning, as well as from insect bites and moisture loss, an elephant's skin would suffer serious damage. After bathing, the elephant will usually use its trunk to blow dirt on its body to help dry and bake on its new protective coat. As elephants are limited to smaller and smaller areas, there is less water available, and local herds will often come too close over the right to use these limited resources.
Wallowing also aids the skin in regulating body temperatures. Elephants have difficulty in releasing heat through the skin because, in proportion to their body size, they have very little of it. The ratio of an elephant's mass to the surface area of its skin is many times that of a human. Elephants have even been observed lifting up their legs to expose the soles of their feet, presumably in an effort to expose more skin to the air. Since wild elephants live in very hot climates, they must have other means of getting rid of excess heat.
Legs and feet
Elephant using its feet to crush a watermelon prior to eating itAn elephant's legs are great straight pillars, as they must be to support its bulk. The elephant needs less muscular power to stand because of its straight legs and large pad-like feet. For this reason an elephant can stand for very long periods of time without tiring. In fact, African elephants rarely lie down unless they are sick or wounded. Indian elephants, in contrast, lie down frequently.
The feet of an elephant are nearly round. African elephants have three nails on each hind foot, and four on each front foot. Indian elephants have four nails on each hind foot and five on each front foot. Beneath the bones of the foot is a tough, gelatinous material that acts as a cushion or shock absorber. Under the elephant's weight the foot swells, but it gets smaller when the weight is removed. An elephant can sink deep into mud, but can pull its legs out readily because its feet become smaller when they are lifted.[citation needed]
Elephants swim well, but cannot trot, jump, or gallop. They do have two gaits: a walk; and a faster gait that is similar to running.
In walking, the legs act as pendulums, with the hips and shoulders rising and falling while the foot is planted on the ground. With no "aerial phase", the faster gait does not meet all the criteria of running, as elephants always have at least one foot on the ground. However, an elephant moving fast uses its legs much like a running animal, with the hips and shoulders falling and then rising while the feet are on the ground. In this gait, an elephant will have three feet off the ground at one time. As both of the hind feet and both of the front feet are off the ground at the same time, this gait has been likened to the hind legs and the front legs taking turns running.[41]
Although they start this "run" at only 8 km/h,[42] elephants can reach speeds up to 40 km/h (25 mph),[43] all the while using the same gait. At this speed, most other four-legged creatures are well into a gallop, even accounting for leg length. Spring-like kinetics could explain the difference between the motion of elephants and other animals.[44]
Ears
Difference between Asian (left) and African (right) elephant ears.The large flapping ears of an elephant are also very important for temperature regulation. Elephant ears are made of a very thin layer of skin stretched over cartilage and a rich network of blood vessels. On hot days, elephants will flap their ears constantly, creating a slight breeze. This breeze cools the surface blood vessels, and then the cooler blood gets circulated to the rest of the animal's body. The hot blood entering the ears can be cooled as much as ten degrees Fahrenheit before returning to the body. Differences in the ear sizes of African and Asian elephants can be explained, in part, by their geographical distribution. Africans originated and stayed near the equator, where it is warmer. Therefore, they have bigger ears. Asians live farther north, in slightly cooler climates, and thus have smaller ears.
The ears are also used in certain displays of aggression and during the males' mating period. If an elephant wants to intimidate a predator or rival, it will spread its ears out wide to make itself look more massive and imposing. During the breeding season, males give off an odour from the musth gland located behind their eyes. Joyce Poole, a well-known elephant researcher, has theorized that the males will fan their ears in an effort to help propel this "elephant cologne" great distances. [45]
Biology and behavior
Social behavior
Elephant footprints (tire tracks for scale)Elephants live in a structured social order. The social lives of male and female elephants are very different. The females spend their entire lives in tightly knit family groups made up of mothers, daughters, sisters, and aunts. These groups are led by the eldest female, or matriarch. Adult males, on the other hand, live mostly solitary lives.
The social circle of the female elephant does not end with the small family unit. In addition to encountering the local males that live on the fringes of one or more groups, the female's life also involves interaction with other families, clans, and subpopulations. Most immediate family groups range from five to fifteen adults, as well as a number of immature males and females. When a group gets too big, a few of the elder daughters will break off and form their own small group. They remain very aware of which local herds are relatives and which are not.
The life of the adult male is very different. As he gets older, he begins to spend more time at the edge of the herd, gradually going off on his own for hours or days at a time. Eventually, days become weeks, and somewhere around the age of fourteen, the mature male, or bull, sets out from his natal group for good. While males do live primarily solitary lives, they will occasionally form loose associations with other males. These groups are called bachelor herds. The males spend much more time than the females fighting for dominance with each other. Only the most dominant males will be permitted to breed with cycling females. The less dominant ones must wait their turn. It is usually the older bulls, forty to fifty years old, that do most of the breeding.
The dominance battles between males can look very fierce, but typically they inflict very little injury. Most of the bouts are in the form of aggressive displays and bluffs. Ordinarily, the smaller, younger, and less confident animal will back off before any real damage can be done. However, during the breeding season, the battles can get extremely aggressive, and the occasional elephant is injured. During this season, known as musth, a bull will fight with almost any other male it encounters, and it will spend most of its time hovering around the female herds, trying to find a receptive mate.
African as well as Asiatic males will engage in same-sex bonding and mounting. Such encounters are often associated with affectionate interactions, such as kissing, trunk intertwining, and placing trunks in each other's mouths. The encounters are analogous to heterosexual bouts, one male often extending his trunk along the other's back and pushing forward with his tusks to signify his intention to mount. Unlike heterosexual relations, which are always of a fleeting nature, those between males result in a "companionship", consisting of an older individual and one or two younger, attendant males. Same-sex relations are common and frequent in both sexes, with Asiatic elephants in captivity devoting roughly 46% of sexual encounters to same-sex activity.[46]
Rogue elephant is a term for a lone, violently aggressive wild elephant. It is a calque of the Sinhala term hora aliya. Its introduction to English has been attributed by the Oxford English Dictionary to Sir James Emerson Tennent, but this usage may have been pre-dated by William Sirr.
Intelligence
Main article: Elephant intelligence
Human, dolphin and elephant brains up to scale. (1)-cerebrum (1a)-temporal lobe and (2)-cerebellumWith a mass just over 5 kg (11 lb), elephant brains are larger than those of any other land animal, and although the largest whales have body masses twentyfold those of a typical elephant, whale brains are barely twice the mass of an elephant's. A wide variety of behaviours, including those associated with grief, making music, art, altruism, allomothering, play, use of tools,[47] compassion and self-awareness [48] evidence a highly intelligent species on par with cetaceans[7] and primates[8]. The largest areas in the elephant brain are those responsible for hearing, smell and movement coordination.
Senses
Elephants have well innervated trunks, and an exceptional sense of hearing and smell. The hearing receptors reside not only in ears, but also in trunks that are sensitive to vibrations, and most significantly feet, which have special receptors for low frequency sound and are exceptionally well innervated. Elephants communicate by sound over large distances of several kilometers partly through the ground, which is important for their social lives. Elephants are observed listening by putting trunks on the ground and carefully positioning their feet.
Their eyesight is relatively poor, and the eyes are aiming down the trunk. An elephant has to raise his head conspicuously to look out horizontally[citation needed].
Self-awareness
Mirror self recognition is a test of self awareness and cognition used in animal studies. A mirror was provided and visible marks were made on the elephant. The elephants investigated these marks, which were visible only via the mirror. The tests also included non-visible marks to rule out the possibility of their using other senses to detect these marks. This shows that elephants recognize the fact that the image in the mirror is their own self and such abilities are considered the basis for empathy, altruism and higher social interactions. This ability has been demonstrated in humans, apes, dolphins,[49] and magpies.[50]
A young elephant in Zimbabwe.
Communication
Elephants make a number of sounds when communicating. Elephant are famous for their trumpet calls which are made when the animal blows though its nostrils. Trumpeting is usually made during excitement. Its use varies from startlement to a cry of help to rage. Elephants also make rumbling growls when greeting each other. The growl become a bellow when the mouth is open and a bellow becomes a moan when prolonged. This can escalate with a roar when threatening another elephant or another animal.
Elephants can communicate over long distances by producing and receiving low-frequency sound (infrasound), a sub-sonic rumbling, which can travel in the air and through the ground much farther than higher frequencies. These calls range in frequency from 15-35 Hz and can be as loud as 117 Db, allowing communication for many kilometres, with a possible maximum range of around 10 km.[51] This sound can be felt by the sensitive skin of an elephant's feet and trunk, which pick up the resonant vibrations much as the flat skin on the head of a drum. To listen attentively, every member of the herd will lift one foreleg from the ground, and face the source of the sound, or often lay its trunk on the ground. The lifting presumably increases the ground contact and sensitivity of the remaining legs. This ability is thought also to aid their navigation by use of external sources of infrasound. Discovery of this new aspect of elephant social communication and perception came with breakthroughs in audio technology, which can pick up frequencies outside the range of the human ear. Pioneering research in elephant infrasound communication was done by Katy Payne, of the Elephant Listening Project,[52] and is detailed in her book Silent Thunder. Though this research is still in its infancy, it is helping to solve many mysteries, such as how elephants can find distant potential mates, and how social groups are able to coordinate their movements over extensive range.[51]
Diet
Elephants are herbivores, spending 16 hours a day collecting plant food. Their diet is at least 50% grasses, supplemented with leaves, bamboo, twigs, bark, roots, and small amounts of fruits, seeds and flowers. Because elephants only digest 40% of what they eat, they have to make up for their digestive system's lack of efficiency in volume. An adult elephant can consume 140–270 kg (300–600 lb) of food a day. 60% of that food leaves the elephant's body undigested[citation needed].
Reproduction and life cycle
Elephant calves
Elephant social life revolves around breeding and raising of the calves. A female will usually be ready to breed around the age of thirteen, when she comes into estrus, a short phase of receptiveness lasting a couple of days, for the first time. Females announce their estrus with smell signals and special calls.
Female African elephant with calf, in Kenya.Females prefer bigger, stronger, and, most importantly, older males. Such a reproductive strategy tends to increase their offspring's chances of survival.
After a twenty-two-month pregnancy, the mother will give birth to a calf that will weigh about 113 kg (250 lb) and stand over 76 cm (2.5 ft) tall. Elephants have a very long childhood. They are born with fewer survival instincts than many other animals. Instead, they must rely on their elders to teach them the things they need to know. Today, however, the pressures humans have put on the wild elephant populations, from poaching to habitat destruction, mean that the elderly often die at a younger age, leaving fewer teachers for the young.
A new calf is usually the centre of attention for all herd members. All the adults and most of the other young will gather around the newborn, touching and caressing it with their trunks. The baby is born nearly blind and at first relies, almost completely, on its trunk to discover the world around it.
As everyone in the herd is usually related, all members of the tightly knit female group participate in the care and protection of the young. After the initial excitement, the mother will usually select several full-time baby-sitters, or "allomothers", from her group. According to Cynthia Moss, a well known researcher, these allomothers will help in all aspects of raising the calf.[53] They walk with the young as the herd travels, helping the calves along if they fall or get stuck in the mud. The more allomothers a baby has, the more free time its mother has to feed herself. Providing a calf with nutritious milk means the mother has to eat more nutritious food herself. So, the more allomothers, the better the calf's chances of survival. An elephant is considered an allomother when she is not able to have her own baby. A benefit of being an allomother is that she can gain experience or receive assistance when caring for her own calf.
Effect on the environment
Elephants are a species which many other organisms depend on. One particular example of that are termites mounds: termites eat elephant feces and often begin building their mounds under piles of elephant feces.
Elephants' foraging activities can sometimes greatly affect the areas in which they live. By pulling down trees to eat leaves, breaking branches, and pulling out roots they create clearings in which new young trees and other vegetation can establish themselves. During the dry season, elephants use their tusks to dig into dry river beds to reach underground sources of water. These newly dug water holes may then become the only source of water in the area. Elephants make pathways through their environment which are also used by other animals to access areas normally out of reach. These pathways have sometimes been used by several generations of elephants and today are converted by humans to paved roads.
Threat of extinction
Men with African Elephant tusks, Dar es Salaam, c. 1900
An Elephant resting his head on a tree trunk, Samburu National Reserve, Kenya
An elephant in the Ngorongoro crater, Tanzania
African Savanna Elephant Loxodonta africana, born 1969 (left), and Asian Elephant Elephas maximus, born 1970 (right), at an English zoo
Hunting
The threat to the African elephant presented by the ivory trade is unique to the species. Larger, long-lived, slow-breeding animals, like the elephant, are more susceptible to overhunting than other animals. They cannot hide, and it takes many years for an elephant to grow and reproduce. An elephant needs an average of 140 kg (300 lb) of vegetation a day to survive. As large predators are hunted, the local small grazer populations (the elephant's food competitors) find themselves on the rise. The increased number of herbivores ravage the local trees, shrubs, and grasses. Elephants themselves have few natural predators besides man and, occasionally, lions. However, many African governments legally allow limited hunting. The large amount of money that is charged for the necessary permits is often used to support conservation efforts, and the small number of permits issued (usually for older animals) ensure that populations are not depleted.[1]
Habitat loss
Another threat to elephant's survival in general is the ongoing cultivation of their habitats with increasing risk of conflicts of interest with human cohabitants. These conflicts kill 150 elephants and up to 100 people per year in Sri Lanka.[54] Lacking the massive tusks of its African cousins, the Asian elephant's demise can be attributed mostly to loss of its habitat.
As larger patches of forest disappear, the ecosystem is affected in profound ways. The trees are responsible for anchoring soil and absorbing water runoff. Floods and massive erosion are common results of deforestation. Elephants need massive tracts of land because, much like the slash-and-burn farmers, they are used to crashing through the forest, tearing down trees and shrubs for food and then cycling back later on, when the area has regrown. As forests are reduced to small pockets, elephants become part of the problem, quickly destroying all the vegetation in an area, eliminating all their resources.
National parks
An Elephant sanctuary at Punnathur kotta, Kerala, south India.Africa's first official reserve, Kruger National Park, eventually became one of the world's most famous and successful national parks. [55] There are, however, many problems associated with the establishment of these reserves. For example, elephants range through a wide tract of land with little regard for national borders. Once a reserve is established and fence erected, many animals find themselves cut off from their winter feeding grounds or spring breeding areas. Some animals may die as a result, while others, like the elephants, may just trample over the fences, wreaking havoc in nearby fields. When confined to small territories, elephants can inflict an enormous amount of damage to the local landscapes. [56]
Additionally, some reserves, such as Kruger National Park has, in the opinion of wildlife managers, suffered from elephant overcrowding, at the expense of other species of wildlife within the reserve. On 25 February 2008, the South Africa announced that they would reintroduce culling for the first time since 1994 to control elephant numbers.[57] Nevertheless, as scientists learn more about nature and the environment, it becomes very clear that these parks may be the elephant's last hope against the rapidly changing world around them.
Humanity and elephants
Harvest from the wild
The harvest of elephants, both legal and illegal, has had some unexpected consequences on elephant anatomy as well. African ivory hunters, by killing only tusked elephants, have given a much larger chance of mating to elephants with small tusks or no tusks at all. The propagation of the absent-tusk gene has resulted in the birth of large numbers of tuskless elephants, now approaching 30% in some populations (compare with a rate of about 1% in 1930). Tusklessness, once a very rare genetic abnormality, has become a widespread hereditary trait.
It is possible, if unlikely, that continued selection pressure could bring about a complete absence of tusks in African elephants, a development normally requiring thousands of years of evolution. The effect of tuskless elephants on the environment, and on the elephants themselves, could be dramatic. Elephants use their tusks to root around in the ground for necessary minerals, tear apart vegetation, and spar with one another for mating rights. Without tusks, elephant behaviour could change dramatically.[58]
Domestication and use
Elephants have been working animals used in various capacities by humans. Seals found in the Indus Valley suggest that the elephant was first domesticated in ancient India. However, elephants have never been truly domesticated: the male elephant in his periodic condition of musth is dangerous and difficult to control. Therefore elephants used by humans have typically been female, war elephants being an exception, however: as female elephants in battle will run from a male, only males could be used in war. It is generally more economical to capture wild young elephants and tame them than breeding them in captivity (see also elephant "crushing").
The Judean rebel Eleazar Maccabeus kills a Seleucid war elephant and is crushed under it (Miniature from a manuscript Speculum Humanae Salvationis).The Lao PDR has been domesticating elephant for centuries, and still employs an approximate 500 domesticated elephants, the majority of which work in the Xaignabouli province. These elephants are mainly employed in the logging industry, with ecotourism emerging as a sustainable and environmentally friendly alternative. Elefantasia is a local INGO aiming to reconvert logging elephants into ecotourism practices, thus allowing Asian elephants the ability to supply their mahouts with income whilst still allowed to breed.
Elephants are also commonly exhibited in zoos and wild animal parks. 1200 Elephants are kept in western zoos. A study shows that the lifespan of elephants in European zoos is about half as long as those living in protected areas in Africa and Asia.[59]
Elephants are revered in India (and are worshipped in ceremonies such as the Aanayoottu).
Warfare
Main article: War elephant
War elephants were used by armies in the Indian sub-continent, the Warring States of China, and later by the Persian Empire. This use was adopted by Hellenistic armies after Alexander the Great experienced their worth against king Porus, notably in the Ptolemaic and Seleucid diadoch empires. The Carthaginian general Hannibal took elephants across the Alps when he was fighting the Romans, but brought too few elephants to be of much military use, although his horse cavalry was quite successful; he probably used a now-extinct third African (sub)species, the North African (Forest) elephant, smaller than its two southern cousins, and presumably easier to domesticate. A large elephant in full charge could cause tremendous damage to infantry, and cavalry horses would be afraid of them (see Battle of Hydaspes).
Industry
Elephant work camp in Thailand. Elephants are used for heavy forest work and in circus presentationsThroughout Myanmar (Burma), Siam, India, and most of South Asia elephants were used in the military for heavy labour, especially for uprooting trees and moving logs, and were also commonly used as executioners to crush the condemned underfoot.
Elephants have also been used as mounts for safari-type hunting, especially Indian shikar (mainly on tigers), and as ceremonial mounts for royal and religious occasions, whilst Asian elephants have been used for transport and entertainment.
Zoo and circuses
There is growing resistance[60] against the capture, confinement, and use of wild elephants. Animal rights advocates allege that elephants in zoos and circuses "suffer a life of chronic physical ailments, social deprivation, emotional starvation, and premature death".[61] Zoos argue that standards for treatment of elephants are extremely high and that minimum requirements for such things as minimum space requirements, enclosure design, nutrition, reproduction, enrichment and veterinary care are set to ensure the wellbeing of elephants in captivity. Circuses continue to have a mixed record. Recently, the city of Los Angeles' closed an elephant act with Circus Vazquez due to numerous instances of abuse and neglect (April 2008) [62], and, according to PETA, Ringling Bros. and Barnum & Bailey Circus has lost 25 elephants since 1992.[63]
Elephants have traditionally been a major part of circuses around the world, being intelligent enough to be trained in a variety of acts (see for example P.T. Barnum's Jumbo and John L. Sullivan, the famous "Boxing Elephant"). However, conditions for circus elephants are highly unnatural (confinement in small pens or cages, restraints on their feet, lack of companionship of other elephants, etc) and, perhaps as a result, there are instances of them turning on their keepers or handlers (examples include Black Diamond and "Murderous Mary").
Elephants raised in captivity sometimes show "rocking behavior", a rhythmic and repetitive swaying which is unreported in free ranging wild elephants. Thought to be symptomatic of stress disorders, and probably made worse by a barren environment,[64] rocking behavior may be a precursor to aggressive behavior in captive elephants.[65][66] This link is to an image of Devi (little princess), a 30-year-old Asian Elephant raised in captivity at the San Diego Zoo showing "rocking behavior".
Hybrids
Although successful hybridisation between African and Asian Elephant species is highly unlikely in the wild, in 1978 at Chester Zoo, an Asian elephant cow gave birth to a hybrid calf sired by an African elephant bull (the old terms are used here as these events pre-date the current classifications). "Motty", the resulting hybrid male calf, had an African elephant's cheeks, their ears (large with pointed lobes) and legs (longer and slimmer), but the toenail numbers, (5 for each front foot, 4 hind) and the single trunk finger of an Asian elephant. His wrinkled trunk was like that of an African elephant. His forehead was sloping with one dome and two smaller domes behind it. The body was African in type, but had an Asian-type centre hump and an African-type rear hump. The calf died of infection 12 days later[67]. It is preserved as a mounted specimen at the British Natural History Museum, London. There are unconfirmed rumours of three other hybrid elephants born in zoos or circuses; all are said to have been deformed and none survived.
Elephant rage
Devi (little princess), a 30-year-old Asian Elephant raised in captivity at the San Diego Zoo exhibiting "rocking behavior" (animation), a rhythmic and repetitive swaying which is unreported in free ranging wild elephants. Thought to be symptomatic of stress disorders, and probably made worse by a barren environment,[68] rocking behavior may be a precursor to aggressive behavior in captive elephants.Despite its popularity in zoos, and cuddly portrayal as gentle giants in fiction, elephants are among the world's most potentially dangerous animals. They can crush and kill any other land animal, even the rhinoceros. They can experience unexpected bouts of rage, and can be vindictive.[69] In Africa, groups of young teenage elephants attack human villages in what is thought to be revenge for the destruction of their society by massive cullings done in the 1970s and 80s.[70][71] In India, male elephants attack villages at night, destroying homes and killing people regularly. In the Indian state of Jharkhand, 300 people were killed by elephants between 2000 and 2004, and in Assam, 239 people have been killed by elephants since 2001.[69] In India, elephants kill up to 200 humans every year, and in Sri Lanka around 50 per year.
Musth
Main article: Musth
Adult male elephants naturally periodically enter the state called musth (Hindi for "madness"), sometimes spelt "must" in English.
Other causes
At least a few elephants have been suspected to be drunk during their attacks. In December 1998, a herd of elephants overran a village in India. Although locals reported that nearby elephants had recently been observed drinking beer which rendered them "unpredictable", officials considered it the least likely explanation for the attack.[72] An attack on another Indian village occurred in October 1999, and again locals believed the reason was drunkenness, but the theory was not widely accepted.[73] Purportedly drunk elephants raided yet another Indian village again on December 2002, killing six people, which led to killing of about 200 elephants by locals.[74]
In popular culture
See also: Cultural depictions of elephants
Rudyard Kipling's Elephant's Child.Elephants are ubiquitous in Western popular culture as emblems of the exotic[75] because their unique appearance and size sets them apart from other animals and because, like other African animals such as the giraffe, rhinoceros, and hippopotamus, they are unfamiliar to Western audiences.[76] Popular culture's stock references to elephants rely on this exotic uniqueness.[76] For instance, a "white elephant" is a byword for something expensive, useless and bizarre.[76]
As characters, elephants are relegated largely to children's literature,[75] in which they are generally cast as models of exemplary behaviour,[75] but account for some of this branch of literature's most iconic characters.[75] Many stories stell of isolated young elephants returning to a close-knit community, such as The Elephant’s Child from Rudyard Kipling's Just So Stories (1902), Dumbo (1942) or The Saggy Baggy Elephant (1947).[76] Other elephant heroes given human qualities include Laurent de Brunhoff's anthropomorphic Babar (1935), David McKee's Elmer (1989) and Dr. Seuss's Horton (1940).[76] More than other exotic animals, elephants in fiction are surrogates for humans,[76] with their concern for the community and each other depicted as something to aspire to.[77]
Lion
From Wikipedia, the free encyclopedia
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For other uses, see Lion (disambiguation).
Semi-protected
Featured article
Lion[1]
Fossil range: Early Pleistocene to recent
Male
Male
Female (Lioness)
Female (Lioness)
Conservation status
Vulnerable (IUCN 3.1)[2]
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
Distribution of lions in Africa
Distribution of lions in India. The Gir Forest, in the State of Gujarat, is the last natural range of approximately 300 wild Asiatic Lions. There are plans to reintroduce some lions to Kuno Wildlife Sanctuary in the neighboring State of Madhya Pradesh.
Distribution of lions in India. The Gir Forest, in the State of Gujarat, is the last natural range of approximately 300 wild Asiatic Lions. There are plans to reintroduce some lions to Kuno Wildlife Sanctuary in the neighboring State of Madhya Pradesh.
Synonyms
Felis leo
Linnaeus, 1758[3]
The lion (Panthera leo) is one of four big cats in the genus Panthera, and a member of the family Felidae. With some males exceeding 250 kg (550 lb) in weight,[4] it is the second-largest living cat after the tiger. Wild lions currently exist 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[specify]. Until the late Pleistocene, which was about 10,000 years ago, the lion was the most widespread large land mammal after humans. They were found in most of Africa, much of Eurasia from western Europe to India, and in the Americas from the Yukon to Peru.
Lions live for around 10–14 years in the wild, while in captivity they can live over 20 years. In the wild, males seldom live longer than ten years as fights with rivals occasionally cause injuries.[5] They typically inhabit savanna and grassland, although they may take to bush and forest. Lions are unusually social compared to other cats. A pride of lions consists of related females and offspring and a small number of adult males. Groups of female lions typically hunt together, preying mostly on large ungulates. The lion is an apex and keystone predator, although they will scavenge if the opportunity arises. While lions do not typically hunt humans selectively, 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 percent over the past two decades in its African range.[6] Lion populations are untenable outside of designated reserves and national parks. Although the cause of the decline is not fully 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 for exhibition in zoos the world over since the late eighteenth century. Zoos are cooperating worldwide in breeding programs for the endangered Asiatic subspecies.
Visually, 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.
Contents
[hide]
* 1 Etymology
* 2 Taxonomy and evolution
o 2.1 Subspecies
+ 2.1.1 Recent
+ 2.1.2 Prehistoric
+ 2.1.3 Dubious
o 2.2 Hybrids
* 3 Physical characteristics
o 3.1 Mane
o 3.2 White lions
* 4 Biology and behavior
o 4.1 Group organization
o 4.2 Hunting and diet
o 4.3 Reproduction and life cycle
o 4.4 Health
o 4.5 Communication
o 4.6 Interspecific predatory relationships
* 5 Distribution and habitat
* 6 Population and conservation status
o 6.1 Man-eaters
o 6.2 In captivity
o 6.3 Baiting and taming
* 7 Cultural depictions
* 8 Notes
* 9 References
* 10 External links
Etymology
The lion's name, similar in many Romance languages, derives from the Latin leo;[7] cf. the Ancient Greek λέων (leon).[8] The Hebrew word lavi (לָבִיא) may also be related,[9] as well as the Ancient Egyptian rw.[10] It was one of the many species originally described, as Felis leo, by Linnaeus in his eighteenth century work, Systema Naturae.[3] The generic component of its scientific designation, Panthera leo, often is 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".[11]
Taxonomy and evolution
Skull of a modern lion at Kruger National Park
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.[12] 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.[13]
Panthera leo itself evolved in Africa between 1 million and 800,000 years ago, before spreading throughout the Holarctic region.[14] 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.[15] Lions died out in northern Eurasia and America at the end of the last glaciation, about 10,000 years ago;[16] this may have been secondary to the extinction of Pleistocene megafauna.[17]
Subspecies
Southwest African lions (Panthera leo bleyenberghi)
Traditionally, twelve recent subspecies of lion were recognized, the largest of which has been recognized as the Barbary Lion.[18] The major differences separating 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 often were based upon zoo material of unknown origin that may have had "striking, but abnormal" morphological characteristics.[19] Today only eight subspecies usually are accepted,[16][20] but one of these (the Cape Lion formerly described as Panthera leo melanochaita) probably is invalid.[20] Even the remaining seven subspecies might be too many; 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.[21][22]
Recent
Eight recent subspecies are recognized today:
* P. l. persica, known as the Asiatic Lion or South Asian, Persian, or Indian Lion, once was widespread from Turkey, across the Middle East, to Pakistan, India, and even to Bangladesh. However, large prides and daylight activity made them easier to poach than tigers or leopards; now around 300 exist in and near the Gir Forest of India.[23]
* 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 one of the largest of the lion subspecies, with reported lengths of 3–3.3 metres (10–10.8 ft) and weights of more than 200 kilograms (440 lb) for males. They ranged from Morocco to Egypt. The last wild Barbary lion was killed in Morocco in 1922.[24]
* P. l. senegalensis, known as the West African Lion, is found in western Africa, from Senegal to Nigeria.
* P. l. azandica, known as the Northeast 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, is found in southwestern Africa, Namibia, Botswana, Angola, Katanga (Zaire), Zambia, and Zimbabwe.
* P. l. krugeri, known as the Southeast African Lion or Transvaal Lion, is found in the Transvaal region of southeastern 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 P. l. krugeri.[20]
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 sometimes are considered to represent separate species, but recent phylogenetic studies suggest that they are in fact, subspecies of the lion (Panthera leo).[16] 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).[25]
* 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. It was larger than today's African lions, reaching the American cave lion in size[26][16]
Cave Lions, Chamber of Felines, Lascaux caves
* 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.[16] This species is known from Paleolithic cave paintings (such as the one displayed to the right), ivory carvings, and clay busts, [27] indicating it had protruding ears, tufted tails, perhaps faint tiger-like stripes, and that at least some males had a ruff or primitive mane around their necks.[28] With this example being a hunting scene it is likely that it depicts females hunting for the pride using the same strategy as their contemporary relatives and males may not be part of the subject.
* 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 distinctly—larger than the European cave lion and smaller than the American cave lion with differing skull proportions.[16][29]
Dubious
* P. l. sinhaleyus, known as the Sri Lanka Lion, appears to have become extinct approximately 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.[30]
* P. l. europaea, known as the European Lion, probably was identical with Panthera leo persica or Panthera leo spelea; its status as a subspecies is unconfirmed. It became extinct around 100 AD due to persecution and over-exploitation. It 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, flourished 350,000 years ago.[31] Its relationship to the extant lion subspecies is obscure, and it probably represents a distinct species.
* P. l. maculatus, known as the Marozi or Spotted lion, sometimes is 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.[32]
Hybrids
Further information: Panthera hybrid, liger, and tigon
Lions have been known to breed with tigers (most often the Siberian and Bengal subspecies) to create hybrids called ligers and tigons.[33] They also have been crossed with leopards to produce leopons,[34] 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 once commonly were 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.[35] Because the growth-inhibiting gene from the female tiger is absent, a growth-promoting gene is passed on by the male lion, 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 are often fertile. Males have about a 50 percent chance of having a mane, but if they grow one, their manes will be modest: around 50 percent of a pure lion mane. Ligers are typically between 3.0 and 3.7 m (10 to 12 feet) in length, and can be between 360 and 450 kg (800 to 1,000 pounds) or more.[35] The less common tigon is a cross between the lioness and the male tiger.[36]
Physical characteristics
During confrontations with others, the mane makes the lion look larger
The lion is the tallest (at the shoulder) of the felines, and also is the second-heaviest feline after the tiger. With powerful legs, a strong jaw, and 8 cm (3.1 in) long canine teeth, the lion can bring down and kill large prey.[37] The skull of the lion is very similar to that of the tiger, though the frontal region is usually more depressed and flattened, with a slightly shorter postorbital region. The lion's skull has broader nasal openings than the tiger. However, 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.[38] Lion coloration varies from light buff to yellowish, reddish, or dark ochraceous brown. The underparts are generally lighter and the tail tuft is black. Lion cubs are born with brown rosettes (spots) on their body, rather like those of a leopard. Although these fade as lions reach adulthood, faint spots often may still be seen on the legs and underparts, particularly on lionesses.
Lions are the only members of the cat family to display obvious sexual dimorphism—that is, males and females look distinctly different. They also have specialized roles that each gender plays in the pride. For instance, the lioness, the hunter, lacks the male's thick cumbersome mane. It seems to impede the male's ability to be camouflaged when stalking the prey and create overheating in chases. The color of the male's mane varies from blond to black, generally becoming darker as the lion grows older.
Two lionesses in Masai Mara, Kenya
Weights for adult lions generally lie between 150–250 kg (330–550 lb) for males and 120–182 kg (264–400 lb) for females.[4] 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).[24] Lions tend to vary in size depending on their environment and area, resulting in a wide spread in recorded weights. For instance, lions in southern Africa tend to be about 5 percent heavier than those in East Africa, in general.[39]
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 107 cm (3 ft 6 in) in females. The tail length is 90-105 cm (2 ft 11 in - 3 ft 5 in) in males and 70–100 cm in females (2 ft 4 in – 3 ft 3 in).[4] The longest known lion was a black-maned male shot near Mucsso, southern Angola in October 1973; the heaviest known lion was a man-eater shot in 1936 just outside Hectorspruit in eastern Transvaal, South Africa and weighed 313 kg (690 lb).[40] Lions in captivity tend to be larger than lions in the wild—the heaviest lion on record is a male at Colchester Zoo in England named Simba in 1970, who weighed 375 kg (826 lb).[41]
The most distinctive characteristic shared by both females and males is that the tail ends in a hairy tuft. In some lions, the tuft conceals a hard "spine" or "spur", 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—the function of the tuft and spine are unknown. Absent at birth, the tuft develops around 5½ months of age and is readily identifiable at 7 months.[42]
Mane
Thermographic image of a lion, showing the insulating mane
The mane of the adult male lion, unique among 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.[43] 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. Sexual selection of mates by lionesses favors males with the densest, darkest mane. [44] 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.[45] In prides including a coalition of two or three males, it is possible that lionesses solicit mating more actively with the males who are more heavily maned.[44]
A maneless male lion, who also has little body hair—from Tsavo East National Park, Kenya
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.[45] The cooler ambient temperature in European and North American zoos, for example, may result in a heavier mane. Thus the mane is not an appropriate marker for identifying subspecies.[20][46] The males of the Asiatic subspecies, however, are characterized by sparser manes than average African lions.[47]
Maneless male lions have been reported in Senegal and Tsavo East National Park in Kenya, and the original male white lion from Timbavati also was maneless. Castrated lions have minimal manes. The lack of a mane sometimes is found in inbred lion populations; inbreeding also results in poor fertility.[48]
Lioness showing the ruff that sometimes leads to misidentification as a male
Many lionesses have a ruff that may be apparent in certain poses. Sometimes it is indicated in sculptures and drawings, especially ancient artwork, and is misinterpreted as a male mane. It differs from a mane, however, in being at the jaw line below the ears, of much less hair length, and frequently not noticeable, whereas a mane extends above the ears of males, often obscuring their outline entirely.
Cave paintings of extinct European Cave Lions exclusively show animals with no mane, or just the hint of a mane, suggesting to some that they were more or less maneless;[28] however, females hunting for a pride are the likely subjects of the drawings—since they are shown in a group related to hunting—so these images do not enable a reliable judgment about whether the males had manes. The drawings do suggest that the extinct species used the same social organization and hunting strategies as contemporary lions.
White lions
White lions owe their coloring to a recessive gene; they are rare forms of the subspecies Panthera leo krugeri
The white lion is not a distinct subspecies, but a special morph with a genetic condition, leucism,[19] that causes paler colouration akin to that of the white tiger; the condition is similar to melanism, which causes black panthers. They are not albinos, having normal pigmentation in the eyes and skin. White Transvaal lion (Panthera leo krugeri) individuals occasionally have been encountered in and around 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.[49] Reportedly, they have been bred in camps in South Africa for use as trophies to be killed during canned hunts.[50]
Confirmation of the existence of white lions only came in the late twentieth century. For hundreds of years prior, the white lion had been thought to be a figment of legend circulating in South Africa, the white pelage of the animal said to represent the goodness in all creatures. Sightings were first reported in the early 1900s, and continued, infrequently, for almost fifty years until, in 1975, a litter of white lion cubs was found at Timbavati Game Reserve.[51]
Biology and behavior
Lions spend much of their time resting and are inactive for about 20 hours per day.[52] 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 until dawn, when hunting most often takes place. They spend an average of two hours a day walking and 50 minutes eating.[53]
Group organization
Lions are predatory carnivores who manifest two types of social organization. Some are residents, living in groups, called prides.[54] The pride usually consists of approximately five or six related females, their cubs of both sexes, and one or two males (known as a coalition if more than one) who mate with the adult females (although extremely large prides, consisting of up to 30 individuals, have been observed). The coalition of males associated with a pride usually amounts to two, but may increase to four and decrease again over time. Male cubs are excluded from their maternal pride when they reach maturity.
A pride on the move near Governors Camp, in the Massai Mara, Kenya
The second organizational behaviour is labeled nomads, who range widely and move about sporadically, either singularly or in pairs.[54] Pairs are more frequent among related males who have been excluded from their birth pride. Note that a lion may switch lifestyles; nomads may become residents and vice versa. Males have to go through this lifestyle and some never are able to join another pride. A female who becomes a nomad has much greater difficulty joining a new pride, as the females in a pride are related, and they reject most attempts by an unrelated female to join their family group.
The area a pride occupies is called a pride area, whereas that by a nomad is a range.[54] 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 caloric 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. 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.[24]
Lioness in a burst of speed while hunting in the Serengeti
Lionesses do the majority of the hunting for their pride, being smaller, swifter and more agile than the males, and unencumbered by the heavy and conspicuous mane, which causes overheating during exertion. They act as a co-ordinated group in order to stalk and bring down the prey successfully. However, if nearby the hunt, males have a tendency to dominate the kill once the lionesses have succeeded and eaten. They are more likely to share with the cubs than with the lionesses, but rarely share food they have killed by themselves. Smaller prey is eaten at the location of the hunt, thereby being shared among the hunters; when the kill is larger it often is dragged to the pride area. There is more sharing of larger kills, [55] although pride members often behave aggressively toward each other as each tries to consume as much food as possible.
Both males and females defend the pride against intruders. Some individual lions consistently lead the defense against intruders, while others lag behind.[56] Lions tend to assume specific roles in the pride. Those lagging behind may provide other valuable services to the group.[57] 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.[58] 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. Females form the stable social unit in a pride and do not tolerate outside females; [59] membership only changes with the births and deaths of lionesses,[60] although some females do leave and become nomadic.[61] Subadult males on the other hand, must leave the pride when they reach maturity at around 2–3 years of age.[61]
Hunting and diet
While a lioness such as this has very sharp teeth, prey is usually killed by strangulation
Lions are powerful animals who usually hunt in coordinated groups and stalk their chosen prey. However, they are not particularly known for their stamina - for instance, a lioness' heart makes up only 0.57 percent of her body weight (a male's is about 0.45 percent of his body weight), whereas a hyena's heart is close to 1 percent of its body weight.[60] Thus, although lionesses can reach speeds of 60 km/h (40 mph),[62] they only can do so for short bursts[63] so they have 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.[64] They sneak up to the victim until they reach a distance of approximately 30 metres (98 ft) or less. Typically, several lionesses 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; they attempt to catch the victim with a fast rush and final leap. The prey usually is killed by strangulation,[65] 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[4] (which would also result in asphyxia). Smaller prey, though, may simply be killed by a swipe of a lion's paw.[4]
The prey consists mainly of large mammals, with a preference for wildebeest, impalas, zebras, buffalo, and warthogs in Africa and nilgai, wild boar, and several deer species in India. Many other species are hunted, based on availability. Mainly this will include ungulates weighing between 50 and 300 kg (110–660 lb) such as kudu, hartebeest, gemsbok, and eland.[4] Occasionally, they take relatively small species such as Thomson's Gazelle or springbok. Lions living near the Namib coast feed extensively on seals.[66] Lions hunting in groups are capable of taking down most animals, even healthy adults, but in most parts of their range they rarely attack very large prey such as fully grown male giraffes due to the danger of injury.
Seven lions along the road in the Masai Mara park reserve in Kenya
Extensive statistics collected over various studies show that lions normally feed on mammals in the range 190–550 kg (420–1210 lb). Wildebeest rank at the top of preferred prey (making nearly half of the lion prey in the Serengeti) followed by zebra.[67] Most adult hippopotamuses, rhinoceroses, elephants, and smaller gazelles, impala, and other agile antelopes are generally excluded. However giraffes and buffalos are often taken in certain regions. For instance, in Kruger National Park, giraffes are regularly hunted.[68], and in Manyara Pack, Cape buffaloes constitute as much as 62% of the lion's diet,[69] due to the high number density of buffaloes. Occasionally hippopotamus is also taken, but adult rhinoceroses are generally avoided. Even though smaller than 190 kg (420 lb), warthogs are often taken depending on availability.[70] In some areas, they specialise in hunting atypical prey species; this is the case at the Savuti river, where they prey on elephants.[71] 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 during the night when elephants' vision is poor.[72] Lions also attack domestic livestock; in India cattle contribute significantly to their diet.[47] They are capable of killing other predators such as leopards, cheetahs, hyenas, and wild dogs, though (unlike most felids) they seldom devour the competitors after killing them. They also scavenge animals either dead from natural causes or killed by other predators, and keep a constant lookout for circling vultures, being keenly aware that they indicate an animal dead or in distress.[73] A lion may gorge itself and eat up to 30 kg (66 lb) in one sitting;[74] 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.[75] An adult lioness requires an average of about 5 kg (11 lb) of meat per day, a male about 7 kg (15.4 lb).[76]
The hunters of a pride sharing a zebra where the kill occurred
Because lionesses 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 kills more easily against other large predators such as hyenas, which may be attracted by vultures from kilometers away in open savannas. Lionesses do most of the hunting; males attached to prides do not usually participate in hunting, except in the case of larger quarry such as giraffe and buffalo. In typical 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.[77]
Young lions first display stalking behavior 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.[78]
Reproduction and life cycle
Most lionesses will have reproduced by the time they are four years of age.[79] Lions do not mate at any specific time of year, and the females are polyestrous.[80] As with 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.[81] A lioness may mate with more than one male when she is in heat;[82] during a mating bout, which could last several days, the couple copulates twenty to forty times a day and are likely to forgo eating. Lions reproduce very well in captivity.
During a mating bout, a couple may copulate 20 to 40 times a day for several days
The average gestation period is around 110 days,[80] 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 whilst the cubs are still helpless, staying relatively close to the thicket or den where the cubs are kept. [83] 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.[84] 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.[83]
Usually, the mother does not integrate herself and her cubs back into the pride until the cubs are six to eight weeks old. [85] However, sometimes this introduction to pride life occurs earlier, particularly if other lionesses have given birth at about the same time. For instance, lionesses in a pride often synchronize their reproductive cycles so that they cooperate in the 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. In addition to greater protection, 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. If one lioness gives birth to a litter of cubs a couple of months after another lioness, for instance, then the younger cubs, being much smaller than their older brethren, are usually dominated by larger cubs at mealtimes—consequently, death by starvation is more common amongst the younger cubs.
A pregnant lioness (right)
In addition to starvation, cubs also face many other dangers, such as predation by jackals, hyenas, leopards, martial eagles and snakes. Even buffaloes, should they catch the scent of lion cubs, often stampede towards the thicket or den where they are being kept, doing their best to trample the cubs to death whilst warding off the lioness. 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, [86] perhaps because females do not become fertile and receptive until their cubs mature or die. All in all, as many as 80 percent of the cubs will die before the age of two.[87]
When first introduced to the rest of the pride, the cubs initially lack confidence when confronted with adult lions other than their mother. However, they soon begin to immerse themselves in the pride life, playing amongst themselves or attempting to initiate play with the adults. Lionesses with cubs of their own are more likely to be tolerant of another lioness's cubs than lionesses without cubs. The tolerance of the male lions towards 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. [88]
The tolerance of male lions towards the cubs varies. They are, however, generally more likely to share food with the cubs than with the lionesses.
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,[89] if they have not already been critically injured whilst defending the pride (once ousted from a pride by rival males, male lions rarely manage a second take-over). This leaves a short window for their own offspring to be born and mature. If they are able to procreate as soon as they take over a pride, potentially, they may have more offspring reaching maturity before they also are displaced. A lioness often will attempt to defend her cubs fiercely from a usurping male, but such actions are rarely successful. He usually kills all of the existing cubs who are less than two years old. A lioness is weaker and much lighter than a male; success is more likely when a group of three or four mothers within a pride join forces against one male.[86]
Contrary to popular belief, it is not only males that are ousted from their pride to become nomads, although the majority of 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, subadult lions, both male and female, may be evicted. [90] Life is harsh for a female nomad. Nomadic lionesses rarely manage to raise their cubs to maturity, without the protection of other pride members.
One scientific study reports that both males and females may interact homosexually.[91][92] 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 percent of mountings have been observed to occur with other males. 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.[93] This is particularly true of male lions, who, as the main defenders of the pride, are more likely to come into aggressive contact with rival males. In fact, even though a male lion may reach an age of 15 or 16 years if he manages to avoid being ousted by other males, the majority of adult males do not live to be more than 10 years old. This is why the average lifespan of a male lion tends to be significantly less than that of a lioness in the wild. However, members of both sexes can be injured or even killed by other lions when two prides with overlapping territories come into conflict.
Various species of tick commonly infest the ears, neck and groin regions of most lions.[94][95] Adult forms of several species of the tapeworm genus Taenia have been isolated from intestines, the lions having ingested larval forms from antelope meat.[96] 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.[97] A more recent outbreak in 2001 killed six lions.[98] Lions, especially in captivity, are vulnerable to the Canine distemper virus (CDV), feline immunodeficiency virus (FIV), and feline infectious peritonitis (FIP).[19] 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.[99] 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.[19]
Communication
Head rubbing and licking are common social behaviors within a pride
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,[100] which have been compared with grooming in primates.[101] Head rubbing—nuzzling one's forehead, face and neck against another lion—appears to be a form of greeting,[102] 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.[103] 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.[104]
Lions have an array of facial expressions and body postures that serve as visual gestures.[105] 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 tend to roar in a very characteristic manner, starting with a few deep, long roars that trail off into a series of shorter ones. 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.[106] Lions have the loudest roar of any big cat.
Interspecific predatory relationships
The relationship between lions and spotted hyenas in areas where they coexist is unique in its complexity and intensity. Lions and spotted hyenas are both apex predators which feed on the same prey, and are therefore in direct competition with one another. As such, they will often fight over and steal each others' kills. Though hyenas are popularly assumed to be opportunistic scavengers profiting from the lion's hunting abilities, it is quite often the case that the reverse is true. In Tanzania's Ngorongoro Crater, the spotted hyena population greatly exceeds that of the resident lions, which obtain a large proportion of their food by stealing hyena prey. The feud between the two species, however, extends beyond battles over food. In animals, it is usually the case that territorial boundaries of another species are disregarded. Hyenas and lions are an exception to this; they set boundaries against each other as they would against members of their own species. Male lions in particular are extremely aggressive toward hyenas, and have been observed to hunt and kill hyenas without eating them. Conversely, hyenas are major predators of lion cubs, and will harass lionesses over kills.[107][108] However, healthy adult males, even single ones, are generally avoided at all costs.
Lions tend to dominate smaller felines such as cheetahs and leopards in areas where they are sympatric. They will steal their kills and will kill their cubs and even adults when given the chance. The cheetah has a 50 percent chance of losing its kill to lions or other predators.[109] Lions are major killers of cheetah cubs, up to 90 percent of which are lost in their first weeks of life due to attacks by other predators. Cheetahs avoid competition by hunting at different times of the day and hide their cubs in thick brush. Leopards also use such tactics, but have the advantage of being able to subsist much better on small prey than either lions or cheetahs. Also, unlike cheetahs, leopards can climb trees and use them to keep their cubs and kills away from lions. However, lionesses will occasionally be successful in climbing to retrieve leopard kills.[110] Similarly, lions dominate African wild dogs, not only taking their kills but also preying on both young and adult dogs (although the latter are rarely caught).[111]
The Nile crocodile is the only sympatric predator (besides humans) that can singly threaten the lion. 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,[112] while the reverse is true for lions entering waterways containing crocodiles, as evidenced by the fact that lion claws have on occasion been found in crocodile stomachs.[113]
Distribution and habitat
In Africa, lions can be found in savanna grasslands with scattered Acacia trees which serve as shade;[114] their habitat in India is a mixture of dry savanna forest and very dry deciduous scrub forest.[115] 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.[116] A population of the Asiatic Lion survived until the tenth century in the Caucasus, their last European outpost.[117]
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 eighteenth century. Between the late nineteenth and early twentieth century they became extinct in North Africa and the Middle East. By the late nineteenth century the lion had disappeared from Turkey and most of northern India,[19][118] 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, Khūzestān Province in 1944. There are no subsequent reliable reports from Iran.[74] The subspecies now survives only in and around the Gir Forest of northwestern India.[23] 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.[119]
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.[31] Parts of this range were occupied by subspecies that are extinct today.
Population and conservation status
Main article: Lion hunting
Lion cubs playing in the Serengeti
Most lions now live in eastern and southern Africa, and their numbers there are rapidly decreasing, with an estimated 30–50 percent decline over the last two decades.[6] Currently, estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004,[120][121] 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.[6] Currently, habitat loss and conflicts with humans are considered the most significant threats to the species.[122][123] 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.[6]
An Asiatic Lioness Panthera leo persica, named Moti, born in captivity in Helsinki Zoo (Finland) in October 1994; she arrived at Bristol Zoo (England) in January 1996.
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.[124] 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.[125] 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.[126] It is important to start a second population to serve as a gene pool 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.[127] 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.[46]
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 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 origin, making maintenance of genetic diversity a problem.[19]
Man-eaters
Main article: Man-eater
While lions do not usually hunt people, some (usually males) seem to seek out human prey; well-publicized cases include the Tsavo maneaters, where 28 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, and the 1991 Mfuwe man-eater, which killed six people in the Laungwa River Valley in Zambia.[128] In both, the hunters who killed the lions wrote books detailing the animals' predatory behavior. The Mfuwe and Tsavo incidents bear similarities: the lions in both incidents were larger than normal, lacked manes, and seemed to suffer from tooth decay. The infirmity theory, including tooth decay, is not favored 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.[129] In their analysis of Tsavo and man-eating generally, Kerbis Peterhans and Gnoske acknowledge that sick or injured animals may be more prone to man-eating, but that the behavior 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 amongst other pantherines and primates in the paleontological record.[130] 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.[131]
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.[132]
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.[131]
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.[133][134] 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."[135] 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 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.[136]
In captivity
Widely seen in captivity,[137] 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.[138] Though many modern zoos are more selective about their exhibits,[139] there are over 1000 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.[140] 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, are still living.[141] A zoo-based lion breeding programme usually takes into account the separation of the various lion subspecies, while mitigating the inbreeding that is likely to occur when animals are divided by subspecies.[142]
A lion at Paignton Zoo
Lions were kept and bred by Assyrian kings as early as 850 BC,[116] and Alexander the Great was said to have been presented with tame lions by the Malhi of northern India.[143] 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.[144] In the East, lions were tamed by Indian princes, and Marco Polo reported that Kublai Khan kept lions inside.[145] The first European "zoos" spread amongst noble and royal families in the thirteenth century, and until the seventeenth 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.[146] 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 thirteenth century,[147][148] 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.[149]
Seraglios served as expressions of the nobility's power and wealth. Animals such as 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 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 seventeenth 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 nineteenth century, at which time it was seen as highly eccentric.[150]
Albrecht Dürer, Lions sketch. Circa 1520
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 seventeenth century, in contrast to more open conditions in Florence at the time.[151] The menagerie was open to the public by the eighteenth century; admission was a sum of three half-pence or the supply of a cat or dog for feeding to the lions.[152] A rival menagerie at the Exeter Exchange also exhibited lions until the early nineteenth century.[153] The Tower menagerie was closed down by William IV,[152] and animals transferred to the London Zoo which opened its gates to the public on 27 April 1828.[154]
Animal species disappear when they cannot peacefully orbit the center of gravity that is man.
—Pierre-Amédée Pichot, 1891[155]
The wild animals trade flourished alongside improved colonial trade of the nineteenth 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 pandas.[156] 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.[157] The widely reproduced imagery of the heroic hunter chasing lions would dominate a large part of the century.[158] 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."[159]
Lion at Melbourne Zoo enjoying an elevated grassy area with some tree shelter
Lions were kept in cramped and squalid conditions at London Zoo until a larger lion house with roomier cages was built in the 1870s.[160] Further changes took place in the early twentieth 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 twentieth century. Though his designs were popular, the old bars and cage enclosures prevailed until the 1960s in many zoos.[161] In the later decades of the twentieth 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.[19] 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.[140]
There have also been instances where a lion was kept by a private individual, such as the lioness Elsa, who was raised by George Adamson and his wife Joy Adamson and came to develop a strong bonds with them, particularly the latter. The lioness later achieved fame, her life being documented in a series of books and films.
Baiting and taming
Main articles: Lion-baiting and Lion taming
Nineteenth century etching of a lion tamer in a cage of lions
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 1825.[162][163]
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.[164] 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 twentieth century with cinema. In demonstrating the superiority of human over animal, lion taming served a purpose similar to animal fights of previous centuries.[164] The now iconic lion tamer's chair was possibly first used by American Clyde Beatty (1903–1965).[165]
Cultural depictions
For more details on this topic, see Cultural depictions of lions.
The Lion Capital of Asoka, originally erected around 250 BCE atop an Ashoka Pillar at Sarnath is the national emblem of India.
The lion has been an icon for humanity for thousands of years, appearing in cultures across Europe, Asia, and Africa. Despite incidents of attacks on humans, lions have enjoyed a positive depiction in culture as strong but noble. A common depiction is their representation as "king of the jungle" or "king of the beasts"; hence, the lion has been a popular symbol of royalty and stateliness,[166] as well as a symbol of bravery; it is featured in several fables of the sixth century BC Greek storyteller Aesop.[167]
Representations of lions date back 32,000 years; the lion-headed ivory carving from Vogelherd cave in the Swabian Alb in southwestern Germany has been determined to be about 32,000 years old from the Aurignacian culture.[16] Two lions were depicted mating in the Chamber of Felines in 15,000-year-old Paleolithic cave paintings in the Lascaux caves. Cave lions are also depicted in the Chauvet Cave, discovered in 1994; this has been dated at 32,000 years of age,[27] though it may be of similar or younger age to Lascaux.[168]
Ancient Egypt venerated the lioness (the fierce hunter) as their war deities and among those in the Egyptian pantheon are, Bast, Mafdet, Menhit, Pakhet, Sekhmet, Tefnut, and the Sphinx; [166] Among the Egyptian pantheon also are sons of these goddesses such as, Maahes, and, as attested by Egyptians as a Nubian deity, Dedun.[169][170]
The Lion Gate of Mycenae (detail)—two lionesses flank the central column that represents a goddess—c. 1300 BC renovation of an existing structure that was demolished to build the new
Careful examination of the lion deities noted in many ancient cultures reveal that many are lioness also. Admiration for the co-operative hunting strategies of lionesses was evident in very ancient times. Most of the lion gates depict lionesses. 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.[171]
The emblem of Jerusalem is a lion standing in front of the Western Wall and flanked by olive branches.
The lion is the biblical emblem of the tribe of Judah and later the Kingdom of Judah. It is contained within Jacob's blessing to his fourth son in the penultimate chapter of the Book of Genesis, "Judah is a lion's whelp; On prey, my son have you grown. He crouches, lies down like a lion, like the king of beasts—who dare rouse him?" (Genesis 49:9[172]). In the modern state of Israel, the lion remains the symbol of the capital city of Jerusalem, emblazoned on both the flag and coat of arms of the city.
The lion was a prominent symbol in both the Old Babylonian and Neo-Babylonian Empire periods. The classic Babylonian lion motif, found as a statue, carved or painted on walls, is often referred to as the striding lion of Babylon. It is in Babylon that the biblical Daniel is said to have been delivered from the lion's den.[173] Such symbolism was appropriated by Saddam Hussein's regime in Iraq for their Lion of Babylon tank, with the technology adapted from a Russian model.
In the Puranic texts of Hinduism, Narasimha ("man-lion") a half-lion, half-man incarnation or (avatara) of Vishnu, is worshipped by his devotees and saved the child devotee Prahlada from his father, the evil demon king Hiranyakashipu;[174] 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.[175] Narasimha is worshiped as "Lion God."
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.[176][177]
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.[178].
"Bharat Mata" ("Mother India"), National personification of India, depicted with an Asiatic/Indian lion at her side
Farther south on the Indian subcontinent, the Asiatic lion is symbolic for the Sinhalese,[179] 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.[180]
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 were usually depicted without wings, their bodies became thicker and shorter, and their manes became curly.[181] 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.[182]
The island nation of Singapore (Singapura) derives its name from the Malay words singa (lion) and pura (city), which in turn is from the Tamil-Sanskrit சிங்க singa सिंह siṃha and पुर புர pura, which is cognate to the Greek πόλις, pólis.[183] According to the Malay Annals, this name was given by a fourteenth century Sumatran Malay prince named Sang Nila Utama, who, on alighting the island after a thunderstorm, spotted an auspicious beast on shore that his chief minister identified as a lion (Asiatic lion).[184]
Flag of Sri Lanka
"Aslan" or "Arslan (Ottoman ارسلان arslān and اصلان aṣlān) is the Turkish and Mongolian word for "lion". It was used as a title by a number of Seljuk and Ottoman rulers, including Alp Arslan and Ali Pasha, and is a Turkic/Iranian name.
"Lion" was the nickname of medieval warrior rulers with a reputation for bravery, such as Richard I of England, known as Richard the Lionheart,[166], Henry the Lion (German: Heinrich der Löwe), Duke of Saxony 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. (The lioness[185] is much more infrequent.) 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.[186] 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[187] of the NFL, Chelsea[188] and Aston Villa of the English Premier League,[189] (and the Premiership itself) to a host of smaller clubs around the world. Villa sport a Scottish Lion Rampant on their crest, as do Rangers and Dundee United of the Scottish Premier League.
The lion is a popular symbol and mascot of high schools, colleges and universities throughout the United States. This statue is on the campus of the University of North Alabama.
Lions continue to feature in modern literature, from the messianic Aslan in The Lion, the Witch and the Wardrobe and following books from the Narnia series written by C. S. Lewis,[190] to the comedic Cowardly Lion in The Wonderful Wizard of Oz.[191] 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, the mascot for Metro-Goldwyn-Mayer (MGM) studios, which has been in use since the 1920s.[192] The 1960s saw the appearance of what is possibly the most famous lioness, the Kenyan animal Elsa in the movie Born Free,[193] based on the true-life international bestselling book of the same title.[194] The lion's role as King of the Beasts has been used in cartoons, from the 1950s manga which gave rise to the first Japanese colour TV animation series, Kimba the White Lion, Leonardo Lion of King Leonardo and his Short Subjects, both from the 1960s, up to the 1994 Disney animated feature film The Lion King,[195][196] which also featured the popular song "The Lion Sleeps Tonight" in its soundtrack. A lion appears on the South African 50-Rand banknotes (see South African rand).
From Wikipedia, the free encyclopedia
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For other uses, see Lion (disambiguation).
Semi-protected
Featured article
Lion[1]
Fossil range: Early Pleistocene to recent
Male
Male
Female (Lioness)
Female (Lioness)
Conservation status
Vulnerable (IUCN 3.1)[2]
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
Distribution of lions in Africa
Distribution of lions in India. The Gir Forest, in the State of Gujarat, is the last natural range of approximately 300 wild Asiatic Lions. There are plans to reintroduce some lions to Kuno Wildlife Sanctuary in the neighboring State of Madhya Pradesh.
Distribution of lions in India. The Gir Forest, in the State of Gujarat, is the last natural range of approximately 300 wild Asiatic Lions. There are plans to reintroduce some lions to Kuno Wildlife Sanctuary in the neighboring State of Madhya Pradesh.
Synonyms
Felis leo
Linnaeus, 1758[3]
The lion (Panthera leo) is one of four big cats in the genus Panthera, and a member of the family Felidae. With some males exceeding 250 kg (550 lb) in weight,[4] it is the second-largest living cat after the tiger. Wild lions currently exist 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[specify]. Until the late Pleistocene, which was about 10,000 years ago, the lion was the most widespread large land mammal after humans. They were found in most of Africa, much of Eurasia from western Europe to India, and in the Americas from the Yukon to Peru.
Lions live for around 10–14 years in the wild, while in captivity they can live over 20 years. In the wild, males seldom live longer than ten years as fights with rivals occasionally cause injuries.[5] They typically inhabit savanna and grassland, although they may take to bush and forest. Lions are unusually social compared to other cats. A pride of lions consists of related females and offspring and a small number of adult males. Groups of female lions typically hunt together, preying mostly on large ungulates. The lion is an apex and keystone predator, although they will scavenge if the opportunity arises. While lions do not typically hunt humans selectively, 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 percent over the past two decades in its African range.[6] Lion populations are untenable outside of designated reserves and national parks. Although the cause of the decline is not fully 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 for exhibition in zoos the world over since the late eighteenth century. Zoos are cooperating worldwide in breeding programs for the endangered Asiatic subspecies.
Visually, 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.
Contents
[hide]
* 1 Etymology
* 2 Taxonomy and evolution
o 2.1 Subspecies
+ 2.1.1 Recent
+ 2.1.2 Prehistoric
+ 2.1.3 Dubious
o 2.2 Hybrids
* 3 Physical characteristics
o 3.1 Mane
o 3.2 White lions
* 4 Biology and behavior
o 4.1 Group organization
o 4.2 Hunting and diet
o 4.3 Reproduction and life cycle
o 4.4 Health
o 4.5 Communication
o 4.6 Interspecific predatory relationships
* 5 Distribution and habitat
* 6 Population and conservation status
o 6.1 Man-eaters
o 6.2 In captivity
o 6.3 Baiting and taming
* 7 Cultural depictions
* 8 Notes
* 9 References
* 10 External links
Etymology
The lion's name, similar in many Romance languages, derives from the Latin leo;[7] cf. the Ancient Greek λέων (leon).[8] The Hebrew word lavi (לָבִיא) may also be related,[9] as well as the Ancient Egyptian rw.[10] It was one of the many species originally described, as Felis leo, by Linnaeus in his eighteenth century work, Systema Naturae.[3] The generic component of its scientific designation, Panthera leo, often is 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".[11]
Taxonomy and evolution
Skull of a modern lion at Kruger National Park
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.[12] 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.[13]
Panthera leo itself evolved in Africa between 1 million and 800,000 years ago, before spreading throughout the Holarctic region.[14] 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.[15] Lions died out in northern Eurasia and America at the end of the last glaciation, about 10,000 years ago;[16] this may have been secondary to the extinction of Pleistocene megafauna.[17]
Subspecies
Southwest African lions (Panthera leo bleyenberghi)
Traditionally, twelve recent subspecies of lion were recognized, the largest of which has been recognized as the Barbary Lion.[18] The major differences separating 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 often were based upon zoo material of unknown origin that may have had "striking, but abnormal" morphological characteristics.[19] Today only eight subspecies usually are accepted,[16][20] but one of these (the Cape Lion formerly described as Panthera leo melanochaita) probably is invalid.[20] Even the remaining seven subspecies might be too many; 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.[21][22]
Recent
Eight recent subspecies are recognized today:
* P. l. persica, known as the Asiatic Lion or South Asian, Persian, or Indian Lion, once was widespread from Turkey, across the Middle East, to Pakistan, India, and even to Bangladesh. However, large prides and daylight activity made them easier to poach than tigers or leopards; now around 300 exist in and near the Gir Forest of India.[23]
* 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 one of the largest of the lion subspecies, with reported lengths of 3–3.3 metres (10–10.8 ft) and weights of more than 200 kilograms (440 lb) for males. They ranged from Morocco to Egypt. The last wild Barbary lion was killed in Morocco in 1922.[24]
* P. l. senegalensis, known as the West African Lion, is found in western Africa, from Senegal to Nigeria.
* P. l. azandica, known as the Northeast 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, is found in southwestern Africa, Namibia, Botswana, Angola, Katanga (Zaire), Zambia, and Zimbabwe.
* P. l. krugeri, known as the Southeast African Lion or Transvaal Lion, is found in the Transvaal region of southeastern 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 P. l. krugeri.[20]
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 sometimes are considered to represent separate species, but recent phylogenetic studies suggest that they are in fact, subspecies of the lion (Panthera leo).[16] 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).[25]
* 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. It was larger than today's African lions, reaching the American cave lion in size[26][16]
Cave Lions, Chamber of Felines, Lascaux caves
* 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.[16] This species is known from Paleolithic cave paintings (such as the one displayed to the right), ivory carvings, and clay busts, [27] indicating it had protruding ears, tufted tails, perhaps faint tiger-like stripes, and that at least some males had a ruff or primitive mane around their necks.[28] With this example being a hunting scene it is likely that it depicts females hunting for the pride using the same strategy as their contemporary relatives and males may not be part of the subject.
* 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 distinctly—larger than the European cave lion and smaller than the American cave lion with differing skull proportions.[16][29]
Dubious
* P. l. sinhaleyus, known as the Sri Lanka Lion, appears to have become extinct approximately 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.[30]
* P. l. europaea, known as the European Lion, probably was identical with Panthera leo persica or Panthera leo spelea; its status as a subspecies is unconfirmed. It became extinct around 100 AD due to persecution and over-exploitation. It 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, flourished 350,000 years ago.[31] Its relationship to the extant lion subspecies is obscure, and it probably represents a distinct species.
* P. l. maculatus, known as the Marozi or Spotted lion, sometimes is 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.[32]
Hybrids
Further information: Panthera hybrid, liger, and tigon
Lions have been known to breed with tigers (most often the Siberian and Bengal subspecies) to create hybrids called ligers and tigons.[33] They also have been crossed with leopards to produce leopons,[34] 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 once commonly were 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.[35] Because the growth-inhibiting gene from the female tiger is absent, a growth-promoting gene is passed on by the male lion, 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 are often fertile. Males have about a 50 percent chance of having a mane, but if they grow one, their manes will be modest: around 50 percent of a pure lion mane. Ligers are typically between 3.0 and 3.7 m (10 to 12 feet) in length, and can be between 360 and 450 kg (800 to 1,000 pounds) or more.[35] The less common tigon is a cross between the lioness and the male tiger.[36]
Physical characteristics
During confrontations with others, the mane makes the lion look larger
The lion is the tallest (at the shoulder) of the felines, and also is the second-heaviest feline after the tiger. With powerful legs, a strong jaw, and 8 cm (3.1 in) long canine teeth, the lion can bring down and kill large prey.[37] The skull of the lion is very similar to that of the tiger, though the frontal region is usually more depressed and flattened, with a slightly shorter postorbital region. The lion's skull has broader nasal openings than the tiger. However, 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.[38] Lion coloration varies from light buff to yellowish, reddish, or dark ochraceous brown. The underparts are generally lighter and the tail tuft is black. Lion cubs are born with brown rosettes (spots) on their body, rather like those of a leopard. Although these fade as lions reach adulthood, faint spots often may still be seen on the legs and underparts, particularly on lionesses.
Lions are the only members of the cat family to display obvious sexual dimorphism—that is, males and females look distinctly different. They also have specialized roles that each gender plays in the pride. For instance, the lioness, the hunter, lacks the male's thick cumbersome mane. It seems to impede the male's ability to be camouflaged when stalking the prey and create overheating in chases. The color of the male's mane varies from blond to black, generally becoming darker as the lion grows older.
Two lionesses in Masai Mara, Kenya
Weights for adult lions generally lie between 150–250 kg (330–550 lb) for males and 120–182 kg (264–400 lb) for females.[4] 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).[24] Lions tend to vary in size depending on their environment and area, resulting in a wide spread in recorded weights. For instance, lions in southern Africa tend to be about 5 percent heavier than those in East Africa, in general.[39]
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 107 cm (3 ft 6 in) in females. The tail length is 90-105 cm (2 ft 11 in - 3 ft 5 in) in males and 70–100 cm in females (2 ft 4 in – 3 ft 3 in).[4] The longest known lion was a black-maned male shot near Mucsso, southern Angola in October 1973; the heaviest known lion was a man-eater shot in 1936 just outside Hectorspruit in eastern Transvaal, South Africa and weighed 313 kg (690 lb).[40] Lions in captivity tend to be larger than lions in the wild—the heaviest lion on record is a male at Colchester Zoo in England named Simba in 1970, who weighed 375 kg (826 lb).[41]
The most distinctive characteristic shared by both females and males is that the tail ends in a hairy tuft. In some lions, the tuft conceals a hard "spine" or "spur", 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—the function of the tuft and spine are unknown. Absent at birth, the tuft develops around 5½ months of age and is readily identifiable at 7 months.[42]
Mane
Thermographic image of a lion, showing the insulating mane
The mane of the adult male lion, unique among 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.[43] 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. Sexual selection of mates by lionesses favors males with the densest, darkest mane. [44] 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.[45] In prides including a coalition of two or three males, it is possible that lionesses solicit mating more actively with the males who are more heavily maned.[44]
A maneless male lion, who also has little body hair—from Tsavo East National Park, Kenya
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.[45] The cooler ambient temperature in European and North American zoos, for example, may result in a heavier mane. Thus the mane is not an appropriate marker for identifying subspecies.[20][46] The males of the Asiatic subspecies, however, are characterized by sparser manes than average African lions.[47]
Maneless male lions have been reported in Senegal and Tsavo East National Park in Kenya, and the original male white lion from Timbavati also was maneless. Castrated lions have minimal manes. The lack of a mane sometimes is found in inbred lion populations; inbreeding also results in poor fertility.[48]
Lioness showing the ruff that sometimes leads to misidentification as a male
Many lionesses have a ruff that may be apparent in certain poses. Sometimes it is indicated in sculptures and drawings, especially ancient artwork, and is misinterpreted as a male mane. It differs from a mane, however, in being at the jaw line below the ears, of much less hair length, and frequently not noticeable, whereas a mane extends above the ears of males, often obscuring their outline entirely.
Cave paintings of extinct European Cave Lions exclusively show animals with no mane, or just the hint of a mane, suggesting to some that they were more or less maneless;[28] however, females hunting for a pride are the likely subjects of the drawings—since they are shown in a group related to hunting—so these images do not enable a reliable judgment about whether the males had manes. The drawings do suggest that the extinct species used the same social organization and hunting strategies as contemporary lions.
White lions
White lions owe their coloring to a recessive gene; they are rare forms of the subspecies Panthera leo krugeri
The white lion is not a distinct subspecies, but a special morph with a genetic condition, leucism,[19] that causes paler colouration akin to that of the white tiger; the condition is similar to melanism, which causes black panthers. They are not albinos, having normal pigmentation in the eyes and skin. White Transvaal lion (Panthera leo krugeri) individuals occasionally have been encountered in and around 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.[49] Reportedly, they have been bred in camps in South Africa for use as trophies to be killed during canned hunts.[50]
Confirmation of the existence of white lions only came in the late twentieth century. For hundreds of years prior, the white lion had been thought to be a figment of legend circulating in South Africa, the white pelage of the animal said to represent the goodness in all creatures. Sightings were first reported in the early 1900s, and continued, infrequently, for almost fifty years until, in 1975, a litter of white lion cubs was found at Timbavati Game Reserve.[51]
Biology and behavior
Lions spend much of their time resting and are inactive for about 20 hours per day.[52] 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 until dawn, when hunting most often takes place. They spend an average of two hours a day walking and 50 minutes eating.[53]
Group organization
Lions are predatory carnivores who manifest two types of social organization. Some are residents, living in groups, called prides.[54] The pride usually consists of approximately five or six related females, their cubs of both sexes, and one or two males (known as a coalition if more than one) who mate with the adult females (although extremely large prides, consisting of up to 30 individuals, have been observed). The coalition of males associated with a pride usually amounts to two, but may increase to four and decrease again over time. Male cubs are excluded from their maternal pride when they reach maturity.
A pride on the move near Governors Camp, in the Massai Mara, Kenya
The second organizational behaviour is labeled nomads, who range widely and move about sporadically, either singularly or in pairs.[54] Pairs are more frequent among related males who have been excluded from their birth pride. Note that a lion may switch lifestyles; nomads may become residents and vice versa. Males have to go through this lifestyle and some never are able to join another pride. A female who becomes a nomad has much greater difficulty joining a new pride, as the females in a pride are related, and they reject most attempts by an unrelated female to join their family group.
The area a pride occupies is called a pride area, whereas that by a nomad is a range.[54] 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 caloric 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. 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.[24]
Lioness in a burst of speed while hunting in the Serengeti
Lionesses do the majority of the hunting for their pride, being smaller, swifter and more agile than the males, and unencumbered by the heavy and conspicuous mane, which causes overheating during exertion. They act as a co-ordinated group in order to stalk and bring down the prey successfully. However, if nearby the hunt, males have a tendency to dominate the kill once the lionesses have succeeded and eaten. They are more likely to share with the cubs than with the lionesses, but rarely share food they have killed by themselves. Smaller prey is eaten at the location of the hunt, thereby being shared among the hunters; when the kill is larger it often is dragged to the pride area. There is more sharing of larger kills, [55] although pride members often behave aggressively toward each other as each tries to consume as much food as possible.
Both males and females defend the pride against intruders. Some individual lions consistently lead the defense against intruders, while others lag behind.[56] Lions tend to assume specific roles in the pride. Those lagging behind may provide other valuable services to the group.[57] 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.[58] 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. Females form the stable social unit in a pride and do not tolerate outside females; [59] membership only changes with the births and deaths of lionesses,[60] although some females do leave and become nomadic.[61] Subadult males on the other hand, must leave the pride when they reach maturity at around 2–3 years of age.[61]
Hunting and diet
While a lioness such as this has very sharp teeth, prey is usually killed by strangulation
Lions are powerful animals who usually hunt in coordinated groups and stalk their chosen prey. However, they are not particularly known for their stamina - for instance, a lioness' heart makes up only 0.57 percent of her body weight (a male's is about 0.45 percent of his body weight), whereas a hyena's heart is close to 1 percent of its body weight.[60] Thus, although lionesses can reach speeds of 60 km/h (40 mph),[62] they only can do so for short bursts[63] so they have 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.[64] They sneak up to the victim until they reach a distance of approximately 30 metres (98 ft) or less. Typically, several lionesses 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; they attempt to catch the victim with a fast rush and final leap. The prey usually is killed by strangulation,[65] 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[4] (which would also result in asphyxia). Smaller prey, though, may simply be killed by a swipe of a lion's paw.[4]
The prey consists mainly of large mammals, with a preference for wildebeest, impalas, zebras, buffalo, and warthogs in Africa and nilgai, wild boar, and several deer species in India. Many other species are hunted, based on availability. Mainly this will include ungulates weighing between 50 and 300 kg (110–660 lb) such as kudu, hartebeest, gemsbok, and eland.[4] Occasionally, they take relatively small species such as Thomson's Gazelle or springbok. Lions living near the Namib coast feed extensively on seals.[66] Lions hunting in groups are capable of taking down most animals, even healthy adults, but in most parts of their range they rarely attack very large prey such as fully grown male giraffes due to the danger of injury.
Seven lions along the road in the Masai Mara park reserve in Kenya
Extensive statistics collected over various studies show that lions normally feed on mammals in the range 190–550 kg (420–1210 lb). Wildebeest rank at the top of preferred prey (making nearly half of the lion prey in the Serengeti) followed by zebra.[67] Most adult hippopotamuses, rhinoceroses, elephants, and smaller gazelles, impala, and other agile antelopes are generally excluded. However giraffes and buffalos are often taken in certain regions. For instance, in Kruger National Park, giraffes are regularly hunted.[68], and in Manyara Pack, Cape buffaloes constitute as much as 62% of the lion's diet,[69] due to the high number density of buffaloes. Occasionally hippopotamus is also taken, but adult rhinoceroses are generally avoided. Even though smaller than 190 kg (420 lb), warthogs are often taken depending on availability.[70] In some areas, they specialise in hunting atypical prey species; this is the case at the Savuti river, where they prey on elephants.[71] 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 during the night when elephants' vision is poor.[72] Lions also attack domestic livestock; in India cattle contribute significantly to their diet.[47] They are capable of killing other predators such as leopards, cheetahs, hyenas, and wild dogs, though (unlike most felids) they seldom devour the competitors after killing them. They also scavenge animals either dead from natural causes or killed by other predators, and keep a constant lookout for circling vultures, being keenly aware that they indicate an animal dead or in distress.[73] A lion may gorge itself and eat up to 30 kg (66 lb) in one sitting;[74] 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.[75] An adult lioness requires an average of about 5 kg (11 lb) of meat per day, a male about 7 kg (15.4 lb).[76]
The hunters of a pride sharing a zebra where the kill occurred
Because lionesses 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 kills more easily against other large predators such as hyenas, which may be attracted by vultures from kilometers away in open savannas. Lionesses do most of the hunting; males attached to prides do not usually participate in hunting, except in the case of larger quarry such as giraffe and buffalo. In typical 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.[77]
Young lions first display stalking behavior 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.[78]
Reproduction and life cycle
Most lionesses will have reproduced by the time they are four years of age.[79] Lions do not mate at any specific time of year, and the females are polyestrous.[80] As with 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.[81] A lioness may mate with more than one male when she is in heat;[82] during a mating bout, which could last several days, the couple copulates twenty to forty times a day and are likely to forgo eating. Lions reproduce very well in captivity.
During a mating bout, a couple may copulate 20 to 40 times a day for several days
The average gestation period is around 110 days,[80] 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 whilst the cubs are still helpless, staying relatively close to the thicket or den where the cubs are kept. [83] 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.[84] 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.[83]
Usually, the mother does not integrate herself and her cubs back into the pride until the cubs are six to eight weeks old. [85] However, sometimes this introduction to pride life occurs earlier, particularly if other lionesses have given birth at about the same time. For instance, lionesses in a pride often synchronize their reproductive cycles so that they cooperate in the 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. In addition to greater protection, 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. If one lioness gives birth to a litter of cubs a couple of months after another lioness, for instance, then the younger cubs, being much smaller than their older brethren, are usually dominated by larger cubs at mealtimes—consequently, death by starvation is more common amongst the younger cubs.
A pregnant lioness (right)
In addition to starvation, cubs also face many other dangers, such as predation by jackals, hyenas, leopards, martial eagles and snakes. Even buffaloes, should they catch the scent of lion cubs, often stampede towards the thicket or den where they are being kept, doing their best to trample the cubs to death whilst warding off the lioness. 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, [86] perhaps because females do not become fertile and receptive until their cubs mature or die. All in all, as many as 80 percent of the cubs will die before the age of two.[87]
When first introduced to the rest of the pride, the cubs initially lack confidence when confronted with adult lions other than their mother. However, they soon begin to immerse themselves in the pride life, playing amongst themselves or attempting to initiate play with the adults. Lionesses with cubs of their own are more likely to be tolerant of another lioness's cubs than lionesses without cubs. The tolerance of the male lions towards 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. [88]
The tolerance of male lions towards the cubs varies. They are, however, generally more likely to share food with the cubs than with the lionesses.
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,[89] if they have not already been critically injured whilst defending the pride (once ousted from a pride by rival males, male lions rarely manage a second take-over). This leaves a short window for their own offspring to be born and mature. If they are able to procreate as soon as they take over a pride, potentially, they may have more offspring reaching maturity before they also are displaced. A lioness often will attempt to defend her cubs fiercely from a usurping male, but such actions are rarely successful. He usually kills all of the existing cubs who are less than two years old. A lioness is weaker and much lighter than a male; success is more likely when a group of three or four mothers within a pride join forces against one male.[86]
Contrary to popular belief, it is not only males that are ousted from their pride to become nomads, although the majority of 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, subadult lions, both male and female, may be evicted. [90] Life is harsh for a female nomad. Nomadic lionesses rarely manage to raise their cubs to maturity, without the protection of other pride members.
One scientific study reports that both males and females may interact homosexually.[91][92] 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 percent of mountings have been observed to occur with other males. 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.[93] This is particularly true of male lions, who, as the main defenders of the pride, are more likely to come into aggressive contact with rival males. In fact, even though a male lion may reach an age of 15 or 16 years if he manages to avoid being ousted by other males, the majority of adult males do not live to be more than 10 years old. This is why the average lifespan of a male lion tends to be significantly less than that of a lioness in the wild. However, members of both sexes can be injured or even killed by other lions when two prides with overlapping territories come into conflict.
Various species of tick commonly infest the ears, neck and groin regions of most lions.[94][95] Adult forms of several species of the tapeworm genus Taenia have been isolated from intestines, the lions having ingested larval forms from antelope meat.[96] 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.[97] A more recent outbreak in 2001 killed six lions.[98] Lions, especially in captivity, are vulnerable to the Canine distemper virus (CDV), feline immunodeficiency virus (FIV), and feline infectious peritonitis (FIP).[19] 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.[99] 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.[19]
Communication
Head rubbing and licking are common social behaviors within a pride
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,[100] which have been compared with grooming in primates.[101] Head rubbing—nuzzling one's forehead, face and neck against another lion—appears to be a form of greeting,[102] 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.[103] 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.[104]
Lions have an array of facial expressions and body postures that serve as visual gestures.[105] 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 tend to roar in a very characteristic manner, starting with a few deep, long roars that trail off into a series of shorter ones. 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.[106] Lions have the loudest roar of any big cat.
Interspecific predatory relationships
The relationship between lions and spotted hyenas in areas where they coexist is unique in its complexity and intensity. Lions and spotted hyenas are both apex predators which feed on the same prey, and are therefore in direct competition with one another. As such, they will often fight over and steal each others' kills. Though hyenas are popularly assumed to be opportunistic scavengers profiting from the lion's hunting abilities, it is quite often the case that the reverse is true. In Tanzania's Ngorongoro Crater, the spotted hyena population greatly exceeds that of the resident lions, which obtain a large proportion of their food by stealing hyena prey. The feud between the two species, however, extends beyond battles over food. In animals, it is usually the case that territorial boundaries of another species are disregarded. Hyenas and lions are an exception to this; they set boundaries against each other as they would against members of their own species. Male lions in particular are extremely aggressive toward hyenas, and have been observed to hunt and kill hyenas without eating them. Conversely, hyenas are major predators of lion cubs, and will harass lionesses over kills.[107][108] However, healthy adult males, even single ones, are generally avoided at all costs.
Lions tend to dominate smaller felines such as cheetahs and leopards in areas where they are sympatric. They will steal their kills and will kill their cubs and even adults when given the chance. The cheetah has a 50 percent chance of losing its kill to lions or other predators.[109] Lions are major killers of cheetah cubs, up to 90 percent of which are lost in their first weeks of life due to attacks by other predators. Cheetahs avoid competition by hunting at different times of the day and hide their cubs in thick brush. Leopards also use such tactics, but have the advantage of being able to subsist much better on small prey than either lions or cheetahs. Also, unlike cheetahs, leopards can climb trees and use them to keep their cubs and kills away from lions. However, lionesses will occasionally be successful in climbing to retrieve leopard kills.[110] Similarly, lions dominate African wild dogs, not only taking their kills but also preying on both young and adult dogs (although the latter are rarely caught).[111]
The Nile crocodile is the only sympatric predator (besides humans) that can singly threaten the lion. 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,[112] while the reverse is true for lions entering waterways containing crocodiles, as evidenced by the fact that lion claws have on occasion been found in crocodile stomachs.[113]
Distribution and habitat
In Africa, lions can be found in savanna grasslands with scattered Acacia trees which serve as shade;[114] their habitat in India is a mixture of dry savanna forest and very dry deciduous scrub forest.[115] 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.[116] A population of the Asiatic Lion survived until the tenth century in the Caucasus, their last European outpost.[117]
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 eighteenth century. Between the late nineteenth and early twentieth century they became extinct in North Africa and the Middle East. By the late nineteenth century the lion had disappeared from Turkey and most of northern India,[19][118] 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, Khūzestān Province in 1944. There are no subsequent reliable reports from Iran.[74] The subspecies now survives only in and around the Gir Forest of northwestern India.[23] 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.[119]
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.[31] Parts of this range were occupied by subspecies that are extinct today.
Population and conservation status
Main article: Lion hunting
Lion cubs playing in the Serengeti
Most lions now live in eastern and southern Africa, and their numbers there are rapidly decreasing, with an estimated 30–50 percent decline over the last two decades.[6] Currently, estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004,[120][121] 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.[6] Currently, habitat loss and conflicts with humans are considered the most significant threats to the species.[122][123] 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.[6]
An Asiatic Lioness Panthera leo persica, named Moti, born in captivity in Helsinki Zoo (Finland) in October 1994; she arrived at Bristol Zoo (England) in January 1996.
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.[124] 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.[125] 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.[126] It is important to start a second population to serve as a gene pool 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.[127] 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.[46]
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 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 origin, making maintenance of genetic diversity a problem.[19]
Man-eaters
Main article: Man-eater
While lions do not usually hunt people, some (usually males) seem to seek out human prey; well-publicized cases include the Tsavo maneaters, where 28 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, and the 1991 Mfuwe man-eater, which killed six people in the Laungwa River Valley in Zambia.[128] In both, the hunters who killed the lions wrote books detailing the animals' predatory behavior. The Mfuwe and Tsavo incidents bear similarities: the lions in both incidents were larger than normal, lacked manes, and seemed to suffer from tooth decay. The infirmity theory, including tooth decay, is not favored 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.[129] In their analysis of Tsavo and man-eating generally, Kerbis Peterhans and Gnoske acknowledge that sick or injured animals may be more prone to man-eating, but that the behavior 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 amongst other pantherines and primates in the paleontological record.[130] 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.[131]
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.[132]
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.[131]
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.[133][134] 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."[135] 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 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.[136]
In captivity
Widely seen in captivity,[137] 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.[138] Though many modern zoos are more selective about their exhibits,[139] there are over 1000 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.[140] 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, are still living.[141] A zoo-based lion breeding programme usually takes into account the separation of the various lion subspecies, while mitigating the inbreeding that is likely to occur when animals are divided by subspecies.[142]
A lion at Paignton Zoo
Lions were kept and bred by Assyrian kings as early as 850 BC,[116] and Alexander the Great was said to have been presented with tame lions by the Malhi of northern India.[143] 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.[144] In the East, lions were tamed by Indian princes, and Marco Polo reported that Kublai Khan kept lions inside.[145] The first European "zoos" spread amongst noble and royal families in the thirteenth century, and until the seventeenth 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.[146] 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 thirteenth century,[147][148] 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.[149]
Seraglios served as expressions of the nobility's power and wealth. Animals such as 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 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 seventeenth 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 nineteenth century, at which time it was seen as highly eccentric.[150]
Albrecht Dürer, Lions sketch. Circa 1520
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 seventeenth century, in contrast to more open conditions in Florence at the time.[151] The menagerie was open to the public by the eighteenth century; admission was a sum of three half-pence or the supply of a cat or dog for feeding to the lions.[152] A rival menagerie at the Exeter Exchange also exhibited lions until the early nineteenth century.[153] The Tower menagerie was closed down by William IV,[152] and animals transferred to the London Zoo which opened its gates to the public on 27 April 1828.[154]
Animal species disappear when they cannot peacefully orbit the center of gravity that is man.
—Pierre-Amédée Pichot, 1891[155]
The wild animals trade flourished alongside improved colonial trade of the nineteenth 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 pandas.[156] 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.[157] The widely reproduced imagery of the heroic hunter chasing lions would dominate a large part of the century.[158] 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."[159]
Lion at Melbourne Zoo enjoying an elevated grassy area with some tree shelter
Lions were kept in cramped and squalid conditions at London Zoo until a larger lion house with roomier cages was built in the 1870s.[160] Further changes took place in the early twentieth 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 twentieth century. Though his designs were popular, the old bars and cage enclosures prevailed until the 1960s in many zoos.[161] In the later decades of the twentieth 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.[19] 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.[140]
There have also been instances where a lion was kept by a private individual, such as the lioness Elsa, who was raised by George Adamson and his wife Joy Adamson and came to develop a strong bonds with them, particularly the latter. The lioness later achieved fame, her life being documented in a series of books and films.
Baiting and taming
Main articles: Lion-baiting and Lion taming
Nineteenth century etching of a lion tamer in a cage of lions
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 1825.[162][163]
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.[164] 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 twentieth century with cinema. In demonstrating the superiority of human over animal, lion taming served a purpose similar to animal fights of previous centuries.[164] The now iconic lion tamer's chair was possibly first used by American Clyde Beatty (1903–1965).[165]
Cultural depictions
For more details on this topic, see Cultural depictions of lions.
The Lion Capital of Asoka, originally erected around 250 BCE atop an Ashoka Pillar at Sarnath is the national emblem of India.
The lion has been an icon for humanity for thousands of years, appearing in cultures across Europe, Asia, and Africa. Despite incidents of attacks on humans, lions have enjoyed a positive depiction in culture as strong but noble. A common depiction is their representation as "king of the jungle" or "king of the beasts"; hence, the lion has been a popular symbol of royalty and stateliness,[166] as well as a symbol of bravery; it is featured in several fables of the sixth century BC Greek storyteller Aesop.[167]
Representations of lions date back 32,000 years; the lion-headed ivory carving from Vogelherd cave in the Swabian Alb in southwestern Germany has been determined to be about 32,000 years old from the Aurignacian culture.[16] Two lions were depicted mating in the Chamber of Felines in 15,000-year-old Paleolithic cave paintings in the Lascaux caves. Cave lions are also depicted in the Chauvet Cave, discovered in 1994; this has been dated at 32,000 years of age,[27] though it may be of similar or younger age to Lascaux.[168]
Ancient Egypt venerated the lioness (the fierce hunter) as their war deities and among those in the Egyptian pantheon are, Bast, Mafdet, Menhit, Pakhet, Sekhmet, Tefnut, and the Sphinx; [166] Among the Egyptian pantheon also are sons of these goddesses such as, Maahes, and, as attested by Egyptians as a Nubian deity, Dedun.[169][170]
The Lion Gate of Mycenae (detail)—two lionesses flank the central column that represents a goddess—c. 1300 BC renovation of an existing structure that was demolished to build the new
Careful examination of the lion deities noted in many ancient cultures reveal that many are lioness also. Admiration for the co-operative hunting strategies of lionesses was evident in very ancient times. Most of the lion gates depict lionesses. 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.[171]
The emblem of Jerusalem is a lion standing in front of the Western Wall and flanked by olive branches.
The lion is the biblical emblem of the tribe of Judah and later the Kingdom of Judah. It is contained within Jacob's blessing to his fourth son in the penultimate chapter of the Book of Genesis, "Judah is a lion's whelp; On prey, my son have you grown. He crouches, lies down like a lion, like the king of beasts—who dare rouse him?" (Genesis 49:9[172]). In the modern state of Israel, the lion remains the symbol of the capital city of Jerusalem, emblazoned on both the flag and coat of arms of the city.
The lion was a prominent symbol in both the Old Babylonian and Neo-Babylonian Empire periods. The classic Babylonian lion motif, found as a statue, carved or painted on walls, is often referred to as the striding lion of Babylon. It is in Babylon that the biblical Daniel is said to have been delivered from the lion's den.[173] Such symbolism was appropriated by Saddam Hussein's regime in Iraq for their Lion of Babylon tank, with the technology adapted from a Russian model.
In the Puranic texts of Hinduism, Narasimha ("man-lion") a half-lion, half-man incarnation or (avatara) of Vishnu, is worshipped by his devotees and saved the child devotee Prahlada from his father, the evil demon king Hiranyakashipu;[174] 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.[175] Narasimha is worshiped as "Lion God."
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.[176][177]
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.[178].
"Bharat Mata" ("Mother India"), National personification of India, depicted with an Asiatic/Indian lion at her side
Farther south on the Indian subcontinent, the Asiatic lion is symbolic for the Sinhalese,[179] 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.[180]
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 were usually depicted without wings, their bodies became thicker and shorter, and their manes became curly.[181] 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.[182]
The island nation of Singapore (Singapura) derives its name from the Malay words singa (lion) and pura (city), which in turn is from the Tamil-Sanskrit சிங்க singa सिंह siṃha and पुर புர pura, which is cognate to the Greek πόλις, pólis.[183] According to the Malay Annals, this name was given by a fourteenth century Sumatran Malay prince named Sang Nila Utama, who, on alighting the island after a thunderstorm, spotted an auspicious beast on shore that his chief minister identified as a lion (Asiatic lion).[184]
Flag of Sri Lanka
"Aslan" or "Arslan (Ottoman ارسلان arslān and اصلان aṣlān) is the Turkish and Mongolian word for "lion". It was used as a title by a number of Seljuk and Ottoman rulers, including Alp Arslan and Ali Pasha, and is a Turkic/Iranian name.
"Lion" was the nickname of medieval warrior rulers with a reputation for bravery, such as Richard I of England, known as Richard the Lionheart,[166], Henry the Lion (German: Heinrich der Löwe), Duke of Saxony 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. (The lioness[185] is much more infrequent.) 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.[186] 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[187] of the NFL, Chelsea[188] and Aston Villa of the English Premier League,[189] (and the Premiership itself) to a host of smaller clubs around the world. Villa sport a Scottish Lion Rampant on their crest, as do Rangers and Dundee United of the Scottish Premier League.
The lion is a popular symbol and mascot of high schools, colleges and universities throughout the United States. This statue is on the campus of the University of North Alabama.
Lions continue to feature in modern literature, from the messianic Aslan in The Lion, the Witch and the Wardrobe and following books from the Narnia series written by C. S. Lewis,[190] to the comedic Cowardly Lion in The Wonderful Wizard of Oz.[191] 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, the mascot for Metro-Goldwyn-Mayer (MGM) studios, which has been in use since the 1920s.[192] The 1960s saw the appearance of what is possibly the most famous lioness, the Kenyan animal Elsa in the movie Born Free,[193] based on the true-life international bestselling book of the same title.[194] The lion's role as King of the Beasts has been used in cartoons, from the 1950s manga which gave rise to the first Japanese colour TV animation series, Kimba the White Lion, Leonardo Lion of King Leonardo and his Short Subjects, both from the 1960s, up to the 1994 Disney animated feature film The Lion King,[195][196] which also featured the popular song "The Lion Sleeps Tonight" in its soundtrack. A lion appears on the South African 50-Rand banknotes (see South African rand).
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