Thursday, December 27, 2007

Lion

From Wikipedia, the free encyclopedia

For other uses, see Lion (disambiguation).
Lion
Male
Male
Female (Lioness)
Female (Lioness)
Conservation status
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
Synonyms
Felis leo
(Linnaeus, 1758)

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

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

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

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

Contents

Naming and etymology

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

Taxonomy and evolution

Skull of a modern lion at Kruger National Park
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.[8] The closest relatives of the lion are the other Panthera species: the tiger, the jaguar and the leopard. Morphological and genetic studies reveal that the tiger was the first of these recent species to diverge. About 1.9 million years ago the Jaguar branched off the remaining group, which contained ancestors of the leopard and lion. The Lion and leopard subsequently separated about 1 to 1.25 million years ago from each other.[9]

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

Subspecies

Southwest African lion (Panthera leo bleyenberghi)
Southwest African lion (Panthera leo bleyenberghi)

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

Recent

Eight recent subspecies are recognized today:

  • P. l. persica, known as the Asiatic- or South Asian, Persian or Indian lion, was once widespread from Turkey, across the Middle East, to Pakistan, India and even Bangladesh. However, large prides and daylight activity made it easier to poach than tigers or leopards; now around 300 exist in and near the Gir Forest of India.[19]
  • P. l. leo, known as the Barbary lion, is extinct in the wild due to excessive hunting, although captive individuals may still exist. This was the largest of the lion subspecies, at 3–3.5m approx., and weighing over 150 kilograms and more. They ranged from Morocco to Egypt. The last wild Barbary lion was killed in Morocco in 1922.[20]
  • P. l. senegalensis, known as the West African lion, is found in Western Africa, from Senegal to Nigeria.
  • P. l. azandica, known as the North East Congo lion, is found in the Northeastern parts of the Congo.
  • P. l. nubica, known as the East African- or Massai lion, is found in East Africa, from Ethiopia and Kenya to Tanzania and Mozambique.
  • P. l. bleyenberghi, known as the Southwest African- or Katanga lion. It is found in southwestern Africa, Zimbabwe, Angola, Katanga (Zaire).
  • P. l. krugeri, known as the Southeast African- or Transvaal lion, is found in the Transvaal region of South eastern Africa, including Kruger National Park.
  • P. l. melanochaita, known as the Cape lion, became extinct in the wild around 1860. Results of mitochondrial DNA research do not support the status as a distinct subspecies. It seems probable that the Cape lion was only the southernmost population of the extant southern African lion.[16]

Prehistoric

Several additional subspecies of lion existed in prehistoric times:

  • P. l. atrox, known as the American lion or American cave lion, was abundant in the Americas from Alaska to Peru in the Pleistocene Epoch until about 10,000 years ago. This form as well as the cave lion are sometimes considered to represent separate species, but recent phylogenetic studies lead to suggest, that they are in fact subspecies of the lion (Panthera leo).[12] One of the largest lion subspecies to have existed, its body length is estimated to have been 1.6–2.5 m (5–8 ft).[21]
  • P. l. fossilis, known as the Early Middle Pleistocene European cave lion, flourished about 500,000 years ago; fossils have been recovered from Germany and Italy.
Cave lions, Chamber of Felines, Lascaux caves
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.[12] It is known from Paleolithic cave paintings, ivory carvings, and clay busts,[22] indicating it had protruding ears, tufted tails, faint tiger-like stripes, and that at least some males had a "ruff" or primitive mane around their necks.[23]
  • P. l. vereshchagini, known as the East Siberian- or Beringian cave lion, was found in Yakutia (Russia), Alaska (USA), and the Yukon Territory (Canada). Analysis of skulls and mandibles of this lion demonstrate that it is distinct—larger than the European cave lion and smaller than the American cave lion with differing skull proportions.[24][12]

Dubious

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

Physical characteristics

During confrontations with others, the mane makes the lion look bigger than he really is.
During confrontations with others, the mane makes the lion look bigger than he really is.

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

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

Mane

A maneless male lion from Tsavo East National Park, Kenya
A maneless male lion from Tsavo East National Park, Kenya
Thermal image of a lion, showing the insulating manes
Thermal image of a lion, showing the insulating manes

The mane of the male lion, unique amongst cats, is one of the most distinctive characteristics of the species. It makes the lion appear larger, providing an excellent intimidation display; this aids the lion during confrontations with other lions and with the species' chief competitor in Africa, the spotted hyena.[32] The presence, absence, color, and size of the mane is associated with genetic precondition, sexual maturity, climate and testosterone production; the rule of thumb is the darker and fuller the mane, the healthier the lion.[33] Research in Tanzania also suggests mane length signals fighting success in male-male relationships. Darker-maned individuals may have longer reproductive lives and higher offspring survival, although they suffer in the hottest months of the year.[34] In prides led by a coalition of two or three males, it is possible that lionesses solicit mating more actively with heavily maned lions.[33]

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

White lions owe their coloring to a recessive gene. They are rare forms of the subspecies Panthera leo krugeri.
White lions owe their coloring to a recessive gene. They are rare forms of the subspecies Panthera leo krugeri.

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

White lions

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

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

Hybrids

A liger is the offspring of a male lion and female tiger.
A liger is the offspring of a male lion and female tiger.
Further information: Panthera hybrid, liger and tigon

Lions have also been known to breed with tigers (most often the Amur and Bengal subspecies) to create hybrids called ligers and tigons.[41] They have also been crossed with leopards to produce leopons,[42] and jaguars to produce jaglions. The marozi is reputedly a spotted lion or a naturally occurring leopon, while the Congolese spotted lion is a complex lion/jaguar/leopard hybrid called a lijagulep. Such hybrids were once commonly bred in zoos, but this is now discouraged due to the emphasis on conserving species and subspecies. Hybrids are still bred in private menageries and in zoos in China.

The liger is a cross between a male lion and a tigress.[43] Because the lion sire passes on a growth-promoting gene, but the corresponding growth-inhibiting gene from the female lion is absent, ligers grow far larger than either parent. They share physical and behavioural qualities of both parent species (spots and stripes on a sandy background). Male ligers are sterile, but female ligers are often fertile. Males have about a 50% chance of having a mane, but if they grow one their manes will be modest: around 50% of a pure lion mane. Ligers are typically between 10 to 12 feet in length, and can be between 800 and 1,000 pounds or more.[43] The less common tigon is a cross between the lioness and the male tiger.[44]

Biology and behaviour

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

Hunting and diet

While a lion's teeth are sharp, prey is usually killed by strangulation.
While a lion's teeth are sharp, prey is usually killed by strangulation.

Lions are powerful animals that usually hunt in groups and stalk their chosen prey. They can reach speeds of 59 km/h (40 mph),[47] though only for short bursts,[48] so they have to be close to their prey before starting the attack. Lions take advantage of factors that reduce visibility; many kills take place near some form of cover or at night.[49] They sneak up to the victim until they reach a distance of about 30 m (98 feet) or less. Typically, several female lions work together and encircle the herd from different points. Once they have closed with a herd, they usually target the closest prey. The attack is short and powerful, the lion attempting to catch the victim with a fast rush and final leap. The prey is usually killed by strangulation.[50]

The lion's prey consists mainly of large mammals, with a preference for wildebeest, impalas, zebras, buffalo and warthogs in Africa and nilgai, wild boars and several deer species in India. Many other species are hunted based on availability, mainly ungulates weighing between 50 and 300 kg such as kudu, hartebeest, gemsbok and eland.[30] Occasionally, lions take relatively small species such as Thomson's gazelle or springbok. Lions, hunting in groups, are capable of taking down most animals, even healthy adults, but they rarely attack very large prey such as buffalo bulls or fully grown male giraffes, due to the danger of injury.[51] They normally feed on mammals no larger than 550 kg, which excludes most adult hippopotamuses, rhinoceroses, elephants, giraffes and buffalos.[52] In some areas, lions specialise in hunting atypical prey-species; this is the case at the Savuti river, where they prey on young elephants.[53] Park guides in the area reported that the lions, driven by extreme hunger, started taking down baby elephants, and then moved on to adolescents and, occasionally, fully grown adults.[54] In the Kruger National Park, giraffes are regularly hunted.[55] Lions also attack domestic livestock; in India cattle contribute significantly to their diet.[36] They are capable of killing other predators such as leopards, cheetahs, hyenas and wild dogs, as well as scavenging animals either dead from natural causes or killed by other predators.[56] A lion may gorge itself and eat up to 30 kg (66 lb) in one sitting;[57] if it is unable to consume all the kill it will rest for a few hours before consuming more. On a hot day, the pride may retreat to shade leaving a male or two to stand guard.[58] An adult female lion requires an average of about 5 kg (11 lb) of meat per day, a male about 7 kg (15.4 lb).[59]

Lion and cub eating a Cape Buffalo
Lion and cub eating a Cape Buffalo

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

Reproduction

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

During a mating bout, a couple may copulate twenty to forty times a day for several days.
During a mating bout, a couple may copulate twenty to forty times a day for several days.

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

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

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

Health

Male lion in Ngorongoro Crater
Male lion in Ngorongoro Crater

Though adult lions have no natural predators, evidence suggests that the majority die violently from humans or other lions.[72] Various species of tick commonly infest the ears, neck and groin regions of most lions.[73][74] Adult forms of several species of the tapeworm genus Taenia have been isolated from intestines, the lions having ingested larval forms from antelope meat.[75] Lions in the Ngorongoro Crater were afflicted by an outbreak of stable fly (Stomoxys calcitrans) in 1962; this resulted in lions becoming covered in bloody bare patches and emaciated. Lions sought unsuccessfully to evade the biting flies by climbing trees or crawling into hyena burrows; many perished or emigrated as the population dropped from 70 to 15 individuals.[76] A more recent outbreak in 2001 killed six lions.[77] Lions, especially in captivity, are vulnerable to the Canine distemper virus (CDV), feline immunodeficiency virus (FIV) and feline infectious peritonitis (FIP).[15] CDV is spread through domestic dogs and other carnivores; a 1994 outbreak in Serengeti National Park resulted in many lions developing neurological symptoms such as seizures. During the outbreak, several lions died from pneumonia and encephalitis.[78] FIV, which is similar to HIV while not known to adversely affect lions, is worrisome enough in its effect in domestic cats that the Species Survival Plan recommends systematic testing in captive lions. It occurs with high to endemic frequency in several wild lion populations, but is mostly absent from Asiatic and Namibian lions.[15]

Group organization

Lions are predatory carnivores who manifest two types of social organization. Some are residents, living in groups, called prides.[79] The pride consists of usually around five or six related females, their cubs of both sexes, and a group of one to four males known as a coalition who mate with the adult females. Others are nomads, ranging widely and moving sporadically, either singularly or in pairs.[79] Note that a lion may switch lifestyles; nomads may become residents and vice versa. The area a pride occupies is called a pride area, whereas that by a nomad is a range.[79] Why sociality—the most pronounced in any cat species—has developed in lions is the subject of much debate. Increased hunting success appears an obvious reason, but this is less than sure upon examination: coordinated hunting does allow for more successful predation, but also ensures that non-hunting "cheaters" reduce per capita caloric intake. Other benefits include possible kin selection (better to share food with a related lion than with a stranger), protection of the young, maintenance of territory, and individual insurance against injury and hunger.[20]

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

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

Head rubbing and licking are common social behaviors within a pride.
Head rubbing and licking are common social behaviors within a pride.

Communication

When resting, lion socialization occurs through a number of behaviors, and the animal's expressive movements are highly developed. The most common peaceful tactile gestures are head rubbing and social licking,[87] which have been compared with grooming in primates.[88] Head rubbing—nuzzling one's forehead, face and neck against another lion—appears to be a form of greeting,[89] as it is seen often after an animal has been apart from others, or after a fight or confrontation. Males tend to rub other males, while cubs and females rub females.[90] Social licking often occurs in tandem with head rubbing; it is generally mutual and the recipient appears to express pleasure. The head and neck are the most common parts of the body licked, which may have arisen out of utility, as a lion cannot lick these areas individually.[91]

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

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 other. 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 does however encompass more than just 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, seeing as they set boundaries against each other as they would against members of their same 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.[94][95]

Distribution and habitat

The Gir Forest in the State of Gujarat, India is the last natural range of the 300-odd wild Asiatic Lions. Plans are afoot to re-introduce some to Palpur-Kuno Wildlife Sanctuary in the neighboring State of Madhya Pradesh in India.
The Gir Forest in the State of Gujarat, India is the last natural range of the 300-odd wild Asiatic Lions. Plans are afoot to re-introduce some to Palpur-Kuno Wildlife Sanctuary in the neighboring State of Madhya Pradesh in India.

In Africa, lions can be found in savannah grasslands with scattered Acacia trees which serve as shade;[96] their habitat in India is a mixture of dry savannah forest and very dry deciduous scrub forest.[97] 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.[98] A population of the Asiatic lion survived until the 10th century in the Caucasus, their last European outpost.[99]

The species was eradicated from Palestine by the Middle Ages and from most of the rest of Asia after the arrival of readily available firearms in the 18th century. Between the late 19th and early 20th century they became extinct in North Africa and the Middle East. By the late 19th century the lion had disappeared from Turkey and most of northern India,[15][100] while the last sighting of a live Asiatic Lion in Iran was in 1941 (between Shiraz and Jahrom, Fars province), though the corpse of a lioness was found on the banks of Karun river, Khuzestan province in 1944. There are no subsequent reliable reports from Iran.[57] The subspecies now survives only in and around the Gir Forest of northwestern India.[19] 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.[101]

Until the late Pleistocene (about 10,000 years ago), the lion was the most widespread land mammal aside from man. They were found in most of Africa, much of Eurasia from western Europe to India and the Bering land bridge, and in the Americas from Yukon to Peru. Parts of this range were occupied by subspecies that are extinct today.

Population and conservation status

Lion cubs playing in the Serengeti
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% decline over the last two decades.[1] Currently, estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004,[102][103] 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.[1] Currently, habitat loss and conflicts with humans are considered the most significant threats to the species.[104][105] 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.[1]

An Asiatic Lioness Panthera leo persica, named MOTI, was born in Helsinki Zoo (Finland) in October 1994; she arrived at Bristol Zoo (England) in January 1996. The Gir Forest in India is the natural home of the Asiatic lion but this animal was born in captivity.
An Asiatic Lioness Panthera leo persica, named MOTI, was born in Helsinki Zoo (Finland) in October 1994; she arrived at Bristol Zoo (England) in January 1996. The Gir Forest in India is the natural home of the Asiatic lion but this animal was born in captivity.

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.[106] 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.[107] 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.[108] It is important to start a second population to serve as a life insurance for the last surviving Asiatic Lions and to help develop and maintain genetic diversity enabling the species to survive.

The former popularity of the Barbary Lion as a zoo animal has meant that scattered lions in captivity are likely to be descended from Barbary Lion stock. This includes twelve lions at Port Lympne Wild Animal Park in Kent, England that are descended from animals owned by the King of Morocco.[109] 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.[35]

Following the discovery of the decline of lion population in Africa, several co-ordinated efforts involving lion conservation have been organised in an attempt to stem this decline. Lions are one species included in the Species Survival Plan, a coordinated attempt by the Association of Zoos and Aquariums to increase its chances of survival. The plan was originally started in 1982 for the Asiatic Lion, but was suspended when it was found that most North American lions were not genetically pure, having been hybridized with African lions. The African lion plan started in 1993, focusing especially on the South African subspecies, although there are difficulties in assessing the genetic diversity of captive lions, since most individuals are of unknown origins, making maintenance of genetic diversity a problem.[15]

Relation with humans

Man-eaters

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 were taken by lions over nine months during the construction of a railway 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.[110] 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.[111] In their analysis of Tsavo and man-eating generally, 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.[112]

The Tsavo maneaters on display in the Field Museum of Natural History
The Tsavo maneaters on display in the Field Museum of Natural History

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.[113]

Author Robert R. Frump wrote in The Man-eaters of Eden that Mozambican refugees regularly crossing Kruger National Park at night in South Africa are attacked and eaten by the lions; park officials have conceded that man-eating is a problem there. Frump believes thousands may have been killed in the decades after apartheid sealed the park and forced the refugees to cross the park at night. Mozambicans had for nearly a century before the border was sealed regularly walked across the park in daytime with little harm.[114]

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.[113]

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.[115][116] 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."[117] GTZ is the German development cooperation agency and has been working with the Tanzanian government on wildlife conservation for nearly two decades. Like in other cases this lion was large, lacked a mane, and had a tooth problem.

The "All-Africa" record of man-eating generally is considered to be not Tsavo, but the lesser-known incidents in the late 1930s through the late 1940s in what was then Tanganyika (now Tanzania). George Rushby, game warden and professional hunter, eventually dispatched the pride, which over three generations is thought to have killed and eaten 1,500 to 2,000 in what is now Njombe district.[118]

In captivity

Widely seen in captivity,[119] lions are part of a group of exotic animals which have formed the core of zoo exhibits the world over since the late 18th century; members of this group are invariably large vertebrates and include elephants, rhinoceroses, hippopotamuses and larger primates as well as other big cats; zoos sought to gather as many of these species as possible.[120] Though many modern zoos are more selective about their exhibits,[121] 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.[122] 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.[123] A zoo-based lion breeding programme usually involves the consideration of several matters, such as the separation of the various lion subspecies, while, at the same time, attempting to prevent the often negative effects of inbreeding, which can often occur when lions are divided in this manner.[124]

Lions were kept and bred by Assyrian kings as early as 850 BC,[98] and Alexander the Great was said to have been presented with tame lions by the Malhi of northern India.[125] 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.[126] In the East, Lions were tamed by Indian princes, and Marco Polo reported that Kublai Khan kept lions inside.[127] The first European "zoos" spread amongst noble and royal families in the 13th century, and until the 17th century were called seraglios; at that time, they came to be called menageries, an extension of the cabinet of curiosities. They spread from France and Italy during the Renaissance to the rest of Europe.[128] In England, although the seraglio tradition was less developed, Lions were kept at the Tower of London in a seraglio established by King John in the 13th century,[129][130] 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.[131]

Albrecht Dürer, Lions sketch. Circa 1520.
Albrecht Dürer, Lions sketch. Circa 1520.

Seraglios served as expressions of the nobility's power and wealth. Animals like big cats and elephants, in particular, symbolized power, and would be pitted in fights against each other or domesticated animals. By extension, menageries and seraglios served as demonstrations of the dominance of man over nature. Consequently, the defeat of such natural "lords" by a cow in 1682 astonished the spectators, and the flight of an elephant before a rhinoceros drew jeers. Such fights would slowly fade out in the 17th century with the spread of the menagerie and their appropriation by the commoners. The tradition of keeping big cats as pets would last into the 19th century, at which time it was seen as highly eccentric.[132]

The presence of lions at the Tower of London was intermittent, being restocked when a monarch or his consort, such as Margaret of Anjou the wife of Henry VI, either sought or were given animals. Records indicate they were kept in poor conditions there in the 17th century, in contrast to more open conditions in Florence at the time.[133] The menagerie was open to the public by the 18th century; admission was a sum of three half-pence or the supply of a cat or dog for feeding to the lions.[134] A rival menagerie at the Exeter Exchange also exhibited lions until the early 19th century.[135] The Tower menagerie was closed down by William IV,[134] and animals transferred to the London Zoo which opened its gates to the public on 27 April 1828.[136]

Animal species disappear when they cannot peacefully orbit the center of gravity that is man.
—Pierre-Amédée Pichot, 1891[137]

The wild animals trade flourished alongside improved colonial trade of the 19th century. Lions were considered fairly common and inexpensive. Although they would barter higher than tigers, they were less costly than larger, or more difficult to transport animals like the giraffe and hippopotamus, and much less than pandas.[138] 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.[139] The widely reproduced imagery of the heroic hunter chasing lions would dominate a large part of the century.[140] 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."[141]

Lion at Melbourne Zoo enjoying an elevated grassy area with some tree shelter
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.[142] Further changes took place in the early 20th century, when Carl Hagenbeck designed enclosures more closely resembling a natural habitat, with concrete 'rocks', more open space and a moat instead of bars. He designed lion enclosures for both Melbourne Zoo and Sydney's Taronga Zoo, among others, in the early 20th century. Though his designs were popular, the old bars and cage enclosures prevailed until the 1960s in many zoos.[143] In the later decades of the 20th century, larger, more natural enclosures and the use of wire mesh or laminated glass instead of lowered dens allowed visitors to come closer than ever to the animals, with some attractions even placing the den on ground higher than visitors, such as the Cat Forest/Lion Overlook of Oklahoma City Zoological Park.[15] 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.[122]

Baiting and taming

19th century Etching of a lion tamer in a cage of lions.
19th century Etching of a lion tamer in a cage of lions.
Main articles: Lion-baiting and Lion taming

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 17th century. It was finally banned in Vienna by 1800 and England in 1825.[144][145]

Lion taming refers to the practice of taming lions for entertainment, either as part of an established circus or as an individual act, such as Siegfried & Roy. The term is also often used for the taming and display of other big cats such as tigers, leopards and cougars. The practice was pioneered in the first half of the 19th century by Frenchman Henri Martin and American Isaac Van Amburgh who both toured widely, and whose techniques were copied by a number of followers.[146] Van Amburgh performed before Queen Victoria of the United Kingdom in 1838 when he toured Great Britain. Martin composed a pantomime titled Les Lions de Mysore ("the lions of Mysore"), an idea that Amburgh quickly borrowed. These acts eclipsed equestrianism acts as the central display of circus shows, but truly entered public consciousness in the early 20th century with cinema. In demonstrating the superiority of man over animal, lion taming served a purpose similar to animal fights of previous centuries.[146] The now iconic lion tamer's chair was possibly first used by American Clyde Beatty (1903–1965).[147]

Cultural depictions

For more details on this topic, see Cultural depictions of lions.
Bronze Lion from Nineveh.
Bronze Lion from Nineveh.
The warrior goddess Sekhmet, shown with her sun disk and cobra crown
The warrior goddess Sekhmet, shown with her sun disk and cobra crown

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,[148] as well as a symbol of bravery; it is featured in several fables of the sixth century BC Greek storyteller Aesop.[149]

Representations of lions date back 32,000 years; the Lion man 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.[12] 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,[22] though it may be of similar or younger age to Lascaux.[150]

Ancient Egypt venerated the lion, with the Sphinx and the lion-headed deity Sekhmet;[148] Maahes and Dedun were Egyptian deities in full lion form.[151][152] 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.[153]

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.[154] 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. Narasimha ("man-lion") is described as an incarnation (avatara) of Vishnu within the Puranic texts of Hinduism;[155] who takes the form of half-man / half-lion, having a human torso and lower body, but with a lion-like face and claws.[156] it is worshiped as "Lion God" thus Indian or Asiatic Lions which were commonly found throughout most of India in ancient times are considered sacred by all Hindus in India.

"Bharat Mata" ("Mother India"), National personification of India, depicted with an Asiatic / Indian lion at her side
"Bharat Mata" ("Mother India"), National personification of India, depicted with an Asiatic / Indian lion at her side

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.[157][158]

Found famously on numerous Flags and Coat of Arms all across Asia and Europe, the Asiatic Lions also stand firm on the National Emblem of India.[159].

Further south on the Indian subcontinent, the Asiatic lion is symbolic for the Sinhalese,[160] 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.[161]

The Asiatic Lion is a common motif in Chinese art. They were first used in art during the late Spring and Autumn Period (5th or 6th 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 6th century AD), lions were usually depicted without wings, their bodies became thicker and shorter, and their manes became curly.[162] 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.[163]

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.[164] According to the Malay Annals, this name was given by a 14th 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).[165] Recent studies of Singapore indicate that lions have never lived there, and the beast seen by Sang Nila Utama was more likely to have been a tiger.

"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 rampant on the royal coat of arms of Scotland
Lion rampant on the royal coat of arms of Scotland

"Lion" was the nickname of medieval warrior rulers with a reputation for bravery, such as Richard I of England, known as Richard the Lionheart,[148], Henry the Lion (German: Heinrich der Löwe), Duke of Saxony and Robert III of Flanders nicknamed "The Lion of Flanders" - a major Flemish national hero up to the present. Lions were frequently depicted on coats of arms, either as a device on shields themselves, or as supporters. 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.[166] As a heraldic symbol or epithet, the lion continues to be used for modern sporting teams, from national soccer teams such as England, Scotland and Singapore to famous clubs such as the Detroit Lions[167] of the NFL, Chelsea[168] and Aston Villa of the English Premier League,[169] (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.
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,[170] to the comedic Cowardly Lion in The Wonderful Wizard of Oz.[171] 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.[172] The 1960s saw the appearance of what is possibly the most famous lioness, the Kenyan animal Elsa in the movie Born Free,[173] based on the true-life international bestselling book of the same title.[174] 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,[175][176] which also featured the popular song "The Lion Sleeps Tonight" in its soundtrack.

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References

  • Baratay, Eric; Elisabeth Hardouin-Fugier (2002). Zoo : a history of zoological gardens in the West. London: Reaktion Books. ISBN 1-861891-11-3.
  • Blunt, Wilfred (1975). The Ark in the Park: The Zoo in the Nineteenth Century. London: Hamish Hamilton. ISBN 0-241-89331-3.
  • de Courcy, Catherine (1995). The Zoo Story. Ringwood, Victoria: Penguin Books. ISBN 0-14-023919-7.
  • Schaller, George B. (1972). The Serengeti lion: A study of predator-prey relations. Chicago: University of Chicago Press. ISBN 0-226-73639-3.

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