Description of Panthera leo
African Lions are large cats with short, tawny coats, and long tails ending in a black tuft. They are sexually dimorphic such that males tend to be larger, averaging 189 kg (416 lbs), compared to 126 kg (227 lbs) for females. Male lions are also distinguished by their manes which vary in color and tend to be fuller in more open habitats.
African lions (Panthera leo) live in most of sub-Saharan Africa except in desert and rainforest habitats. Lions were once exterminated from South Africa, where they remain in Kruger and Kalahari Gemsbok National Parks and possibly some other protected areas. Lions once ranged throughout southwest Asia and north Africa. Asiatic lions (P. l. persica) belong to the single remaining subspecies in this region. Once roaming from Greece to central India, Asiatic lions persist in the Gir forest of northwest India.
In addition to the asiatic subspecies, many taxonomists contend that there are five extant African subspecies. Each subspecies is identified by geographic region. Panthera leo senegalensis (west African or Senegalese lions), P. l. azandica (north east Congo lions), P. l. bleyenberghi (Katanga, Angolan, or south Congo lions), and P. l. krugeri (south African or Transvaal lions). Panthera leo krugeri includes Kalahari lions (sometimes denoted as P. l. verneyi). Lastly, there are East African lions (P. l. nubica). These animals have been categorized as Somali lions (P. l. somaliensis), Masai lions (P. l. massaicus), Serengeti lions (P. l. massaicus), Congo lions (P. l. hollisteri), and Abyssinian lions (P. l. roosevelti). It should be noted, however, that there is some debate as to the validity of the African subspecies classifications, leaving only the Asiatic subspecies, P. l. persica, uncontested.
Biogeographic Regions: palearctic (Native ); oriental (Native ); ethiopian (Native )
- 2001. "Asiatic Lion Information Center" (On-line). Accessed February 09, 2004 at http://www.asiatic-lion.org.
- Alden, P., R. Estes, D. Schlitter, B. McBride. 1998. National Audubon Society Field Guide to African Wildlife. New York, United States of America: Alfred A. Knopf, Inc..
- Estes, R. 1993. The Safari Companion: A Guide to Watching African Mammals. Vermont, United States of America: Chelsea Green Publishing Company.
- O'Brien, S., J. Martenson, C. Packer, L. Herbst, V. de Voss, P. Jocelyn, J. Ott-Jocelyn, D. Wildt, M. Bush. 1987. Biochemical genetic variation in geographically isolated populations of African and Asiatic lions. National Geographic Research, 3/1: 114-124.
- Urban, M., P. West. 2002. "Lion Research Center" (On-line). Accessed February 10, 2004 at http://www.lionresearch.org/.
The Lion formerly ranged from northern Africa through southwest Asia (where it disappeared from most countries within the last 150 years), west into Europe, where it apparently became extinct almost 2,000 years ago, and east into India (Nowell and Jackson 1996, Sunquist and Sunquist 2002). Today, the only remainder of this once widespread population is a single isolated population of the Asiatic Lion P. leo persica in the 1,400 km² Gir Forest National Park and Wildlife Sanctuary. Lions are extinct in North Africa, having perhaps survived in the High Atlas Mountains up to the 1940s (Nowell and Jackson 1996, West and Packer in press).
The map (provided by the Wildlife Conservation Society) shows Lion range as derived from mapping workshops associated with two regional Lion conservation strategies. In West and Central Africa, known and probable range are shown (IUCN 2006a). In Eastern and Southern Africa, known, occasional and possible range are shown (IUCN 2006b). Both strategies are available online from the IUCN SSC Cat Specialist Group website http://www.catsg.org/catsgportal/bulletin-board/20_bulletin-board/home/index_en.htm
Lions are large cats with short, tawny coats, white underparts, and long tails with a black tuft at the end. They are sexually dimorphic and male lions are the only cats with manes. Three year-old male lions grow manes that vary in color from black to blond. Manes tend to be fuller in open habitats. Adult males typically weigh 189 kg; the heaviest male on record weighed 272 kg (Mount Kenya). Females weigh 126 kg on average. The average male height is 1.2 m and the average female height is 1.1 m. Length ranges from 2.4 to 3.3 m and tail length ranges from 0.6 to 1.0 m; the longest male lion recorded was 3.3 m.
Cubs have brown spots on a grayish coat until the age of three months; spots may remain on stomach, especially in east Africa. Albinism does occur in some populations, but there are no published records of melanism (black fur) in lions. Adult lions have 30 total teeth and adult females have four mammae.
Asiatic lions (P. l. persica) are slightly smaller than African lions and have shorter manes, thicker elbow and tail tufts, and longitudinal skin folds on their stomachs. Although Asiatic lions are genetically distinct from African lions, the genetic difference between the two species is smaller than that between human races.
Range mass: 126 to 272 kg.
Range length: 2.4 to 3.3 m.
Sexual Dimorphism: male larger; ornamentation
Other Physical Features: endothermic ; bilateral symmetry
Average basal metabolic rate: 94.58 W.
Catalog Number: USNM 181568
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Mammals
Sex/Stage: Male; Adult
Preparation: Skin; Skull
Collector(s): E. Heller
Year Collected: 1911
Locality: Lime Springs, Sotik, Rift Valley, Kenya, Africa
- Type: Allen, J. A. 1924 Apr 11. Bulletin of the American Museum of Natural History. 47: 229.
Catalog Number: USNM 144054
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Mammals
Sex/Stage: Male; Old adult
Preparation: Skin; Skull; Anatomical
Collector(s): K. Menelick
Year Collected: 1906
Locality: Addis Ababa, near, Addis Ababa, Ethiopia, Africa
- Type: Heller, E. 1913 Nov 08. Smithsonian Miscellaneous Collections. 61 (19): 2.
Catalog Number: USNM 164551
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Mammals
Sex/Stage: Male; Adult
Preparation: Skin; Partial Skull
Collector(s): M. Moses
Year Collected: 1909
Locality: Kampala, Uganda, Africa
- Type: Heller, E. 1913 Nov 08. Smithsonian Miscellaneous Collections. 61 (19): 4.
African lions live in plains or savanna habitat with a large prey base (mostly ungulates) and sufficient cover available. In these optimal habitats, lions are the second most abundant large predator, after spotted hyenas Crocuta crocuta. Lions can also live, with wider ranges, in most habitats except in tropical rainforests and in deserts. Lions can live in forested, shrubby, mountainous, and semi-desert habitats. They are capable of living at high altitudes. There is a lion population in the Bale Mountains of Ethiopia at 4,240 m.
Asiatic lions live in scrubland and teak forest in the small Gir Forest preserve of India.
Range elevation: 4240 (high) m.
Habitat Regions: temperate ; tropical ; terrestrial
Terrestrial Biomes: savanna or grassland ; forest ; scrub forest ; mountains
- Cat Specialist Group, 1996. "African Lion Panthera leo (Linnaeus 1758)" (On-line). Accessed February 09, 2004 at http://lynx.uio.no/catfolk/afrleo01.htm.
Habitat and Ecology
The Lion has a broad habitat tolerance, absent only from tropical rainforest and the interior of the Sahara desert (Nowell and Jackson 1996). There are records of Lion to elevations of more than 4,000 m in the Bale Mountains and on Kilimanjaro (West and Packer in press). Although Lions drink regularly when water is available, they are capable of obtaining their moisture requirements from prey and even plants (such as the tsama melon in the Kalahari desert), and thus can survive in very arid environments. Medium- to large-sized ungulates (including antelopes, zebra and wildebeest) are the bulk of their prey, but Lions will take almost any animal, from rodents to a rhino. They also scavenge, displacing other predators (such as the Spotted Hyaena) from their kills.
Lions are the most social of the cats, with related females remaining together in prides, and related and unrelated males forming coalitions competing for tenure over prides. Average pride size (including males and females) is four to six adults; prides generally break into smaller groups when hunting. Lions tend to live at higher densities than most other felids, but with a wide variation from 1.5 adults per 100 km² in southern African semi-desert to 55/100 km² in parts of the Serengeti (Sunquist and Sunquist 2002). Pride ranges can vary widely even in the same region: e.g., from 266-4,532 km² in the Kgalagadi Transfrontier Park of South Africa (Funston 2001), and 20,500 km² in the Serengeti (West and Packer in press).
In India, the habitat of the Asiatic Lion is dry deciduous forest. The Gir National Park and Wildlife Sanctuary is surrounded by cultivated areas and inhabited by the pastoralist Maldharis and their livestock. Domestic cattle have historically been a major part of the Asiatic Lion's diet, although the most common prey is the chital deer. Mean pride size, measured by the number of adult females, tends to be smaller than for African Lions: most Gir prides contain an average of two adult females (Nowell and Jackson 1996).
Lions live in most of sub-Saharan Africa except in desert and rainforest habitats. Optimal habitat for African Lions is savanna where there is sufficient cover to hide from potential prey.
Lions are predatory carnivores. They usually hunt in groups, but the actual killing is done by an individual lion. They frequently bring down prey much bigger than they are themselves. Showy males have more difficulty hunting than females because of their conspicuousness, therefore females in a pride do the majority of hunting. Males are still more aggressive during feeding than are females, even though they are less likely to have killed the prey.
African lions eat the most common large ungulates in the area (Thompson's gazelles Eudorcas thomsonii, zebras Equus burchellii, impalas Aepyceros melampus, and wildebeests Connochaetes taurinus). Individual prides tend to have their own eating preferences. Some prides tend to target large prey such as cape buffalo Syncerus caffer and giraffe Giraffa camelopardalis. Lions that are not able to capture large prey will eat birds, rodents, fish, ostrich eggs, amphibians and reptiles. Lions also actively scavenge, taking cues from hyenas and vultures.
In Tanzania's Serengeti National Park, local lions subsist on a diet comprized mainly of 7 species: zebras Equus burchellii, wildebeests Connochaetes taurinus, Thompson's gazelles Eudorcas thomsonii, buffalos Syncerus caffer, warthogs Phacochoerus aethiopicus, hartebeests Alcelaphus buselaphus, and topis Damaliscus lunatus provide 90% of their diet.
Hunting effectiveness is increased by hunting in groups. Serengeti research has shown that individual lions succeed in their hunting 17% of the time, whereas group hunts succeed 30% of the time.
Animal Foods: birds; mammals; amphibians; reptiles; fish; eggs; carrion
Primary Diet: carnivore (Eats terrestrial vertebrates)
Lions are the top predators in their range. It is not clear to what extent lions regulate their prey population; some studies have shown that food availability plays a larger role in regulating prey populations than consumption by lions.
- Eltringham, S. 1979. The Ecology and Conservation of Large African Mammals. London, Great Britain: The MacMillan Press Ltd..
Adult lions have no natural predators, excepting persecution by humans. Lions often kill and/or compete with other predators (leopards Panthera pardus and cheetahs Acinonyx jubatus). Spotted hyenas Crocuta crocuta defend kills or scavenged food from immature and female lions, but typically leave the food to a big male lion. Hyenas are known to kill lion cubs, juveniles, or weak and sick adult lions.
Lion cubs, if left alone, can be vulnerable to other large predators. However, infanticide is the primary threat to cubs.
Human poaching is a problem for lions. These animals are poached with wire snares, rifles, and arrows. Since lions are scavengers, they are particularly vulnerable to intentionally poisoned carcasses. There are still poachers that operate within some national parks in Africa. It has been estimated that in the 1960's, poachers were responsible for approximately 20,000 lion deaths per year in Serengeti National Park. Trophy hunting is allowed in 6 African countries.
- spotted hyenas (Crocuta crocuta)
- humans (Homo sapiens)
Known prey organisms
This list may not be complete but is based on published studies.
Predator: prey includes various gazelles, impalas, zebras, and wildebeests.
Life History and Behavior
Lions have the cognitive ability to recognize individuals and to interact with other lions to benefit their own survival. They use visual cues in this communication. For example, the mane is thought to signal to other lions the sex of a male from a distance and to indicate individual fitness. (The rate of mane development is mostly controlled by testosterone.)
Resident male lions regularly mark their territory by spraying vegetation with urine and by scuff-marking. Females spray occasionally. This behavior starts at the age of 2 years. This type of marking uses both chemical and visual communication signals.
Male lions start to roar at 1 year of age and females start shortly after this. The male’s roar is both louder and deeper than the female's. Lions can roar at any time, but they typically stand or crouch while roaring. Roaring serves to advertise territories, to communicate with other pride members, and to demonstrate aggressions toward enemy lions. Lions also roar in chorus; this may be a form of social bonding. This is accoustic communication.
Finally, lions use tactile communication. Males engage in physical aggression during pride take over. There is touching during greetings between pride members. Physical communication passes between lactating females and the cubs they are nursing.
Communication Channels: visual ; tactile ; acoustic ; chemical
Other Communication Modes: choruses ; scent marks
Perception Channels: visual ; tactile ; acoustic ; chemical
- Grinnel, J., C. Packer, A. Pusey. 1995. Cooperation in male lions: kinship, reciprocity, or mutualism?. Animal Behavior, 49/1: 95-105. Accessed February 10, 2004 at http://www.sciencedirect.com/science.
Lions are predatory carnivores that usually feed on hoofed mammals. They typically hunt in groups, but the actual killing is done by an individual lion, usually a female.
Lions are the only truly social cats, with related females residing together in prides led by one or acoalition of several adult males. Females are able to breed at four years, males at five. Females are the predominant care-givers to cubs, which are dependent upon adults until about 16 months old.
Female lions typically live longer than males. Males reach their prime between five and nine years but few males survive past ten years of age. Some males have survived until 16 in the wild. Females normally live until 15 or 16 years. In the Serengeti, females live up to 18 years. In captivity, lions live approximately 13 years. The oldest known lion was 30 years old.
Adult lions have no predators, but are vulnerable to humans, starvation, and attacks from other lions. Infanticide is an important contributor to cub mortality and cub mortality increases when prey is scarce.
Female Asiatic lions live an average of 17 to 18 years, with a maximum of 21 years. Male Asiatic lions generally live for 16 years. Adult Asiatic lions have a less than 10% mortality rate. In the Gir forest, 33% of cubs die during their first year of life.
Status: wild: 18 (high) years.
Status: captivity: 30 (high) years.
Status: wild: 14 years.
Status: captivity: 30 (high) years.
Status: captivity: 13 years.
Status: wild: 15.0 years.
Status: wild: 29.0 years.
Status: wild: 15.0 years.
Lifespan, longevity, and ageing
Lions breed year-round and are usually polygynous. It is estimated that lions copulate 3,000 times for every cub that survives over one year. One estrus out of every 5 results in a litter and lions mate approximately 2.2 times per hour for the 4 day estrus period. The first male member of a pride that reaches a female in heat has the mating priority over her. Fighting between pride members over females does not normally occur.
Male lions are conspicuously large and showy because they have the opportunity to control the reproduction of many females when they rule over a pride. Males form coalitions with each other to increase their chances of pride takeover. The fierce competition among males and the social structure of a pride have led to infanticide by both males and females. Successful males that takeover a pride have about 2 years before another younger, stronger coalition will replace them. Pride takeover battles are often violent leading to severe injury or death of the losing lions.
It is to the successful male’s reproductive advantage to kill the suckling cubs of the defeated males. A nursing lioness that loses her cubs will come back into estrous within 2 to 3 weeks. The normal time between births is 2 years, which is the typical time for a male to rule a pride. Therfore, by killing all unweaned cubs at the time of pride takeover, males can ensure that they have some opportunity to father offspring of females who would otherwise not be available to them during their tenure as pride leaders. Females vigorously defend their cubs during a takeover and are sometimes killed also.
Mating System: polygynous ; cooperative breeder
Female lions are polyestrous, breeding throughout the year and peaking in the rainy season. Female lions tend to have cubs every 2 years. However, if a female's cubs are killed (usually by an intruding male lion), then the female comes into estrus early and has more cubs. Females are able to breed at 4 years of age and males at 5 years.
One to six cubs are born after a 3.5 month gestation period. There is an interbirth interval of approximately twenty to thirty months. Newborn cubs weigh 1 to 2 kg. Eyes typically open by 11 days, cubs can walk by 15 days and are able to run by 1 month of age. Mother lions keep their cubs in hiding until they reach about 8 weeks of age. The cubs are weaned between 7 and 10 months, however they are dependent upon adults in the pride until they are at least 16 months old.
Breeding interval: Female lions tend to have cubs every 2 years. However, if a female's cubs are killed (usually by an intruding male lion), then she will come into estrus early and have more cubs.
Breeding season: Breeding peaks during the rainy season, but is year-round.
Range number of offspring: 1 to 6.
Average number of offspring: 3.
Average gestation period: 3.5 months.
Range weaning age: 7 to 10 months.
Range time to independence: 16 (low) months.
Average age at sexual or reproductive maturity (female): 4 years.
Average age at sexual or reproductive maturity (male): 5 years.
Key Reproductive Features: iteroparous ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization
Average birth mass: 1300 g.
Average gestation period: 109 days.
Average number of offspring: 3.
Average age at sexual or reproductive maturity (male)
Sex: male: 1095 days.
Average age at sexual or reproductive maturity (female)
Sex: female: 1095 days.
Females are mainly responsible for care of the young. Females nurse their young, but will also nurse the young of their female relatives in the pride if litters are born close together. Cub mortality is lowest when related females in the same pride synchronously reproduce and cross-suckle. Since synchronous reproduction is common in prides, cubs are often raised in crèches where the entire pride helps to raise several litters.
Cubs are often left alone for more than one full day by the time they are 5 to 7 months old. This is a particularly vulnerable time for the cubs to be attacked by predators (often hyenas). Hungry mothers occasionally abandon weak cubs that can not keep up with the pride.
Although males do not directly provide care for the young in a pride, they are important in the protection of the cubs from rival males. So long as a male maintains control over a pride, preventing another male from taking over, the cubs he has sired are at lower risk of infanticide.
Parental Investment: no parental involvement; altricial ; male parental care ; female parental care ; pre-fertilization (Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Male, Female); pre-independence (Provisioning: Male, Female, Protecting: Male, Female); post-independence association with parents; inherits maternal/paternal territory
- Alden, P., R. Estes, D. Schlitter, B. McBride. 1998. National Audubon Society Field Guide to African Wildlife. New York, United States of America: Alfred A. Knopf, Inc..
- Estes, R. 1993. The Safari Companion: A Guide to Watching African Mammals. Vermont, United States of America: Chelsea Green Publishing Company.
- Packer, C., A. Pusey. 2001. Egalitarianism in Female African Lions. Science, 293/5530: 690-693. Accessed February 09, 2004 at http://www.sciencemag.org/cgi/content/full/.
- Schaller, G. 1972. The Serengeti Lion. Chicago, United States of America: The University of Chicago Press.
- Urban, M., P. West. 2002. "Lion Research Center" (On-line). Accessed February 10, 2004 at http://www.lionresearch.org/.
Evolution and Systematics
The claws cats provide versatility for a variety of functions, such as gripping or running, because they are retractable.
"The paws of a lion resemble those of most of the cat family. Cats and dogs walk in what is called the digitigrade position: the heel and instep are raised off the ground, making locomotion quieter and more versatile. The large pads on the ball of the foot and on the toes provide a cushion when walking and also help silence the feet. The lion has retractile claws -- it can retract them while at rest or when walking, so that they do not catch in the ground and reduce his speed." (Foy and Oxford Scientific Films 1982:177)
Learn more about this functional adaptation.
- Foy, Sally; Oxford Scientific Films. 1982. The Grand Design: Form and Colour in Animals. Lingfield, Surrey, U.K.: BLA Publishing Limited for J.M.Dent & Sons Ltd, Aldine House, London. 238 p.
Molecular Biology and Genetics
Statistics of barcoding coverage: Panthera leo
Public Records: 0
Specimens with Barcodes: 16
Species With Barcodes: 1
Panthera leo leo (Barbary lions) and P. l. melanochaita (Cape lions) are two extinct subspecies of African lion.
African lion populations have greatly declined in West Africa and in many African countries they are restricted to protected areas. If there are no connecting corridors between wildlife reserves genetic viability will likely become a problem.
Asiatic lions, P. l. persica, are confined to one population in the Gir Forest reserve of India, consisting of less than 200 mature individuals. This subspecies is listed as critically endangered on the IUCN red list and is on Appendix I of CITES. Another population of Asiatic lions is desperately needed in order to safeguard the survival of this subspecies. The Palpur-Kuno Wildlife Sanctuary in northern Madhya Pradesh had been identified as a potential reintroduction site in India. Threats to the Gir Forest population include the close proximity of humans and their cattle and habitat degradation.
Some very small lion populations require genetic management in order to survive and maintain genetic diversity. Hluhluwe-Umfolozi Park (HUP) in Natal, for example, has a population of 120 lions produced from only three lions that were introduced into the park in the 1960s. In 2001, researchers tried artificial insemination techniques to rejuvenate the genetic pool of these South African lions. This process is very difficult and energy intensive. Inbred populations could also potentially be rejuvenated by introducing adult females and whole prides into an area (minimizing conflict between existing lions and introduced lions).
US Federal List: no special status
CITES: appendix i; appendix ii
IUCN Red List of Threatened Species: vulnerable
- Cat Specialist Group 2001, 2003. "Panthera leo" (On-line). 2003 IUCN Red List of Threatened Species. Accessed February 09, 2004 at http://www.redlist.org/search/details.php?species=15951.
- Hanby, J., J. Bygott, C. Packer. 1995. Ecology, Demography, and Behavior of Lions in Two Contrasting Habitats: Ngorongoro Crater and the Serengeti Plains. Pp. 315-331 in A Sinclair, P Arcese, eds. Serengeti II. Chicago, United States of America: The University of Chicago Press.
- Trivedi, B. 2004. "To Boost Gene Pool, Lions Artificially Inseminated" (On-line). National Geographic News. Accessed February 08, 2004 at http://news.nationalgeographic.com/news/2002/06/0612_020612_TVlion.html.
IUCN Red List Assessment
Red List Category
Red List Criteria
A species population reduction of approximately 30% is suspected over the past two decades (= approximately three Lion generations). The causes of this reduction (primarily indiscriminate killing in defense of life and livestock, coupled with prey base depletion: Bauer 2008), are unlikely to have ceased. This suspected reduction is based on direct observation; appropriate indices of abundance; a decline in area of occupation, extent of occupation and habitat quality; and actual and potential levels of exploitation.
Estimating the size of the African Lion population is an ambitious exercise involving many uncertainties. The five main efforts (Ferreras and Cousins 1996; Chardonnet 2002; Bauer and Van Der Merwe 2004; IUCN 2006a&b; Riggio et al. in press) all use different methods. The African Lion Working Group compiled individual population estimates primarily from protected areas (23,000 Lions: Bauer and Van Der Merwe 2004). In 1980, Ferreras and Cousins (1996) predicted 18,600 Lions to occur in protected areas. This was probably an underestimate as not all protected areas inhabited by Lions at that time were included. Still, the comparison suggests that the number of Lions in African protected areas has remained stable or possibly increased over time. But Ferreras and Cousins (1996) predicted that most Lions in 1980 were found outside protected areas. Chardonnet (2002) finds that unprotected areas still comprise a significant portion (half) of the Lion's current African range. Comparison of Ferreras and Cousin's (1996) prediction of 75,800 Lions in 1980 (roughly three Lion generations ago) to Chardonnet's (2002) estimate of 39,000 Lions yields a suspected decline of 48.5%. This calculation suggests a substantial decline in Lions outside protected areas over the past two decades. Ferreras and Cousins (1996) may have over-estimated the African Lion population in 1980, as their number was derived from a model rather than actual Lion counts, and so it is possible that the rate of decline of the African Lion population may be lower. A group exercise led by WCS and the IUCN SSC Cat Specialist Group estimated that 42% of major Lion populations were declining (IUCN 2006a&b, Bauer 2008). The rate of decline is most unlikely to have been as high as 90%, as reported in a series of news reports in 2003 (Kirby 2003, Frank and Parker 2003). All available data were reviewed and combined with satellite imagery leading to the most recent estimate of 32.000 lions in 67 lion areas comprising 3.4 million km2, 17% of historical range or about 25% of savannah Africa (Riggio et al. in press).
In Sub-Saharan Africa, the Lion conservation community works in the context of four regions: West, Central, East and Southern. The Lion population is classified as regionally Endangered in West Africa (Bauer and Nowell 2004). It is isolated from Lion populations of Central Africa, with little or no exchange of breeding individuals (Chardonnet 2002, Bauer and Van Der Merwe 2004). The number of mature individuals in West Africa is estimated by two separate recent surveys at 850 (Bauer and Van Der Merwe 2004) and 1,163 (Chardonnet 2002). Both estimates are well below the Endangered criterion level of 2,500. Lions in West Africa are grouped into three isolated subpopulations by Chardonnet (2002) and approximately seven by the African Lion Working Group (Bauer and Van Der Merwe 2004). Chardonnet's (2002) three subpopulations consist of 18 different individual populations, between which there may be some interchange of individuals, although this is unknown. There is disagreement over the size of the largest individual population in West Africa: the African Lion Working Group (Bauer and Van Der Merwe 2004) estimates 100 Lions in Burkina Faso's Arly-Singou ecosystem, while Chardonnet (2002) estimates 404 for the same area (mean 250).
IUCN Red List Status: VULNERABLE
There have been few efforts in the past to estimate the number of Lions in Africa. Myers (1975) wrote, "Since 1950, their [Lion] numbers may well have been cut in half, perhaps to as low as 200,000 in all or even less". Later, Myers (1986) wrote, "In light of evidence from all the main countries of its range, the Lion has been undergoing decline in both range and numbers, often an accelerating decline, during the past two decades". In the early 1990s, IUCN SSC Cat Specialist Group members made educated "guesstimates" of 30,000 to 100,000 for the African Lion population (Nowell and Jackson 1996).
The most quantitative historical estimate of the African Lion population in the recent past was made by Ferreras and Cousins (1996), who developed a GIS-based model to predict African Lion range and numbers, calibrated by surveying experts about the factors affecting Lion populations. First they correlated vegetation (Leaf Area Index) with Lion densities, using known values from 37 studies in 19 African protected areas, and mapped potential Lion range. Then the reduction effect of human activities on Lion range and numbers were estimated. Lion experts were surveyed in order to develop and rank a set of factors which would lead to lower Lion densities as well as Lion absence. These included agriculture, human population density, cattle grazing, and distance from a protected area, and were derived from GIS databases of varying age. For example, in areas identified as main cattle grazing areas Lion density was reduced by 90%, and in areas identified as having widespread agricultural cultivation or high human population density (> 2.5 people/km²) Lions were considered absent. Lion density was reduced by 50% in areas with low human population density (1-2.5 people/km²). Because of the age of their data sources on extent of agriculture and pastoralism, Ferreras and Cousins (1996) selected 1980 as the base year for their predicted African Lion population of 75,800. They emphasized the need for ground-truthing their estimate by censusing Lions, particularly outside protected areas.
Two recent surveys have provided current estimates of the African Lion population, with some ground-truthing. The African Lion Working Group, a network of specialists affiliated with the IUCN SSC Cat Specialist Group, conducted a mail survey and compiled estimates of 100 known African Lion populations. Not included were populations of known existence, but unknown or unestimated size. The ALWG African Lion population estimate is 23,000, with a range of 16,500-30,000 (Bauer and Van Der Merwe 2004). The second survey was carried out by Philippe Chardonnet and sponsored by the International Foundation for the Conservation of Wildlife and Conservation Force (Chardonnet 2002). He also compiled estimates for 144 individual African Lion populations, grouped into 36 largely isolated subpopulations. His methodology included extrapolation of estimates of known populations into areas where Lion status was unknown, and his total figure is not surprisingly larger: 39,000 Lions in Africa, with a range of 29,000-47,000.
Approximately 30% of the individual population estimates compiled by the African Lion Working Group were based on scientific surveys. Techniques for these surveys included total count based on individually identified body features, sampling by use of calling stations playing recordings of hyaena and/or Lion prey, and mark-recapture methods including radio telemetry, photo databases, and spoor counts (Bauer and Van Der Merwe 2004). Seventy percent of their population figures were derived from expert opinion or guesstimate. In the other survey, 63% of Chardonnet's (2002) individual population estimates were based on expert opinions or guesstimates. Twelve percent of Chardonnet's (2002) estimates were based on scientific surveys, and a further 25% were derived from extrapolation of variables from nearby populations and catch-per-unit effort-estimates based on Lion trophy hunting.
Estimating the size of the African Lion population is an ambitious exercise involving many uncertainties. The three main efforts (Ferreras and Cousins 1996; Chardonnet 2002; Bauer and Van Der Merwe 2004) all use different methods. The African Lion Working Group compiled individual population estimates primarily from protected areas (23,000 Lions: Bauer and Van Der Merwe 2004). In 1980, Ferreras and Cousins (1996) predicted 18,600 Lions to occur in protected areas. This was probably an underestimate as not all protected areas inhabited by Lions at that time were included. Still, the comparison suggests that the number of Lions in African protected areas has remained stable or possibly increased over time. But Ferreras and Cousins (1996) predicted that most Lions in 1980 were found outside protected areas. Chardonnet (2002) finds that unprotected areas still comprise a significant portion (half) of the Lion's current African range. Comparison of Ferreras and Cousin's (1996) prediction of 75,800 Lions in 1980 (roughly three Lion generations ago) to Chardonnet's (2002) estimate of 39,000 Lions yields a suspected decline of 48.5%. This calculation suggests a substantial decline in Lions outside protected areas over the past two decades. Ferreras and Cousins (1996) may have over-estimated the African Lion population in 1980, as their number was derived from a model rather than actual Lion counts, and so it is possible that the rate of decline of the African Lion population may be lower. A group exercise led by WCS and the IUCN SSC Cat Specialist Group estimated that 42% of major Lion populations were declining (Bauer 2008). The rate of decline is most unlikely to have been as high as 90%, as reported in a series of news reports in 2003 (Kirby 2003, Frank and Parker 2003).
Genetic population models indicate that large populations (50-100 Lion prides) are necessary to conserve genetic diversity and avoid inbreeding, which increases significantly when populations fall below 10 prides. Male dispersal is also an important factor in conserving genetic variation (Bjorklund 2003). These conditions are met in few wild Lion populations, although there are at least 17 Lion "strongholds" >50,000 km² in extent (Bauer 2008).
Outside sub-Saharan Africa, the Asiatic Lion P. leo persica occurs as an isolated single wild population in India's Gir Forest. The Gir Lion population had been reduced to a very low number in the early years of the 20th century, possibly fewer than 20. In 1936, 234 Lions were estimated, falling to approximately 100 iin the 1970s, and estimates in the 1980s and 1990s around 200 adults (Nowell and Jackson 1996). In 2005, the population was estimated at 359, +/- 10, including cubs, based on, according to the Chief Minister of Forests for Gujarat state, a total count of Lions by direct sighting in a block-system methodology. This is an increase from 327 reported in 2001 (Anon. 2006). The Wildlife Protection Society of India reported 34 Lion deaths in 2007, due to poaching, electrocution, falling into open wells, and death by motor vehicle and unknown causes. Single populations are vulnerable to extinction from catastrophic events, and it has long been recommended by scientists to establish a second wild population in Madhya Pradesh's Kuno Wildlife Sanctuary, but this has not yet been done (Jackson 2008). The Asiatic Lion shows reduced genetic variation due to small population size (O'Brien et al. 1987). It is feared that the size of the population is larger than the estimated carrying capacity of the habitat and prey base (Nowell and Jackson 1996).
The main threats to Lions are indiscriminate killing (primarily as a result of retaliatory or pre-emptive killing to protect life and livestock) and prey base depletion. Habitat loss and conversion has led to a number of populations becoming small and isolated (Bauer 2008).
The economic impact of stock raiding can be significant: Patterson et al. (2004) estimated that each Lion cost ranchers in Kenya living alongside Tsavo East National Park US$290 per year in livestock losses. Likewise, annual losses of cattle to Lions in areas adjacent to Waza National Park in Cameroon comprised only about 3.1% of all livestock losses, but were estimated to represent more than 22% of financial losses amounting to about US$370 per owner (Bauer 2003). Consequently, Lions are persecuted intensely in livestock areas across Africa; their scavenging behaviour makes them particularly vulnerable to poisoned carcasses put out to eliminate predators. Little actual information exists on the number of Lions killed as problem animals by local people, even though this is considered the primary threat to their survival outside protected areas. Implementation of appropriate livestock management measures, coupled with problem animal control measures and mechanisms for compensating livestock losses, are some of the primary responses to resolving human-Lion conflict (Frank et al. 2006).
Trophy hunting is carried out in a number of sub-Saharan African countries and is considered an important management tool for providing financial resource for Lion conservation for both governments and local communities. However, there is concern that current management regimes can lead to unsustainable offtakes (Packer et al. 2006).
Disease has also been a threat to Lion populations (Ray et al. 2005).
In parts of southeastern Tanzania there have been alarmingly high incidences of people killed by Lions, with up to 400 human Lion-related fatalities recorded from 1997-2007 (Ikanda 2007).
Historically African Lions have been killed for sport. Today their habitat and natural prey is disappearingas more and more land is usurped for agriculture. As a result, lions are raiding domestic livestock, putting them in direct conflict with farmers and cattle ranchers.
P. leo is included in CITES Appendix II; the Endangered Asiatic Lion subspecies P. leo persica is included in CITES Appendix I.
In Africa, Lions are present in a number of large and well-managed protected areas, and remain one of the most popular animals on the must-see lists of tourists and visitors to Africa. Most range states in East and Southern Africa have an infrastructure which supports wildlife tourism, and in this way Lions generate significant cash revenue for park management and local communities and provide a strong incentive for wildland conservation.
Regional conservation strategies have been developed for Lions in west and central Africa (IUCN 2006a) and eastern and southern Africa (IUCN 2006b). The West and Central African Lion Conservation Strategy focuses on three primary objectives to address threats that directly impact Lions: to reduce Lion-human conflict, and to conserve and increase Lion habitat and wild prey base. The objectives of the Eastern and Southern African Lion Conservation Strategy are articulated around the root issues in Lion conservation, including policy and land use, socio-economics, trade, and conservation politics. For example, the policy and land use objective is "to develop and implement harmonious and comprehensive legal and institutional frameworks that provide for the expansion of wildife-integrated land use, Lion conservation and associated socio-economic benefits in current and potential Lion range." The trade objective is "to prevent illegal trade in Lions and Lion products while promoting and safeguarding sustainable legal trade." Both regional strategies share common priorities of conserving and restoring Lion populations, improving management capacity, and increasing the flow of benefits to communities living with Lions. These strategies should be used by governments to guide national Lion action plans, policies and programs, and by the conservation community to guide their project development. By setting out common priorities to guide action on both national, community and landscape levels, the regional conservation strategies have the potential for broad and significant improvement of Lion status and management (Nowell et al. 2006).
For the Asiatic Lion, resolving human Lion-conflict is a high priority, as well as establishing a second wild population (Jackson 2008).
Relevance to Humans and Ecosystems
Humans are fearful of being attacked by lions and of their livestock being killed by lions. In most cases, this has not been a large problem. Lions have historically coexisted with the Masai and their cows in east Africa. When the prey base is plentiful, lions do not typically attack cattle. Also, if a lion sees a human on foot, it will typically run in the opposite direction.
There have been well-publicized attacks on humans by lions, such as the man-eating lions of Tsavo (killed 135 construction workers) portrayed in the 1996 movie The Ghost and the Darkness. As lions lose habitat, they are more frequently entering human inhabited areas, thus creating more conflict and potential for "problem lions" that attack humans.
Lions are frequently infected with FIV (Feline Immunodeficiency Virus) which is similar to HIV. In Tanzania’s Serengeti National Park, Ngorongoro Crater, and Kruger National Park of South Africa, 92% of lions tested had FIV. FIV does not seem to adversely affect lions but it will kill domestic cats.
Negative Impacts: injures humans (bites or stings); causes or carries domestic animal disease
- Pickrell, J. 2004. "Man-Eating Lions Not Aberrant, Experts Say" (On-line). National Geographic News. Accessed February 12, 2004 at http://news.nationalgeographic.com/news/2003/02/0211_030211_tsavolions.html.
Lions are a glamorous species, well-known throughout the world. They are a cultural icon in England and are one of the highest valued eco-tourism species in Africa. They are also the subject of many documentaries and research efforts.
Positive Impacts: ecotourism ; research and education
- Dudley, J. 2002. Issues and Priorities for Mammal Conservation. Conservation Biology, 16/4: 1169.
The lion (Panthera leo) is one of the five big cats in the genus Panthera and a member of the family Felidae. The commonly used term African lion collectively denotes the several subspecies found in Africa. With some males exceeding 250 kg (550 lb) in weight, it is the second-largest living cat after the tiger. Wild lions currently exist in sub-Saharan Africa and in Asia (where an endangered remnant population resides in Gir Forest National Park in India) while other types of lions have disappeared from North Africa and Southwest Asia in historic times. Until the late Pleistocene, about 10,000 years ago, the lion was the most widespread large land mammal after humans. They were found in most of Africa, across Eurasia from western Europe to India, and in the Americas from the Yukon to Peru. The lion is a vulnerable species, having seen a major population decline in its African range of 30–50% per two decades during the second half of the 20th century. Lion populations are untenable outside 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. Within Africa, the West African lion population is particularly endangered.
Lions live for 10–14 years in the wild, although in captivity they can live more than 20 years. In the wild, males seldom live longer than 10 years, as injuries sustained from continual fighting with rival males greatly reduce their longevity. 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. Lions are apex and keystone predators, although they are also expert scavengers obtaining over 50 percent of their food by scavenging as opportunity allows. While lions do not typically hunt humans, some have been known to do so. Sleeping mainly during the day, lions are primarily nocturnal, although bordering on crepuscular in nature.
Highly distinctive, the male lion is easily recognised by its mane, and its face is one of the most widely recognised 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 once occurred. It has been extensively depicted in sculptures, in paintings, on national flags, and in contemporary films and literature. Lions have been kept in menageries since the time of the Roman Empire, and have been a key species sought for exhibition in zoos over the world since the late eighteenth century. Zoos are cooperating worldwide in breeding programs for the endangered Asiatic subspecies.
- 1 Etymology
- 2 Taxonomy and evolution
- 3 Characteristics
- 4 Behaviour
- 5 Distribution and habitat
- 6 Population and conservation status
- 7 Cultural depictions
- 8 Heraldic depictions
- 9 See also
- 10 References
- 11 External links
The lion's name, similar in many Romance languages, is derived from the Latin leo, and the Ancient Greek λέων (leon). The Hebrew word לָבִיא (lavi) may also be related. It was one of the species originally described by Linnaeus, who gave it the name Felis leo, in his eighteenth-century work, Systema Naturae.
Taxonomy and evolution
The lion's closest relatives are the other species of the genus Panthera: the tiger, the jaguar, and the leopard. P. leo evolved in Africa between 1 million and 800,000 years ago, before spreading throughout the Holarctic region. It appeared in the fossil record 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. Lions died out in northern Eurasia at the end of the last glaciation, about 10,000 years ago; this may have been secondary to the extinction of Pleistocene megafauna.
Traditionally, 12 recent subspecies of lion were recognised, distinguished by mane appearance, size, and distribution. Because these characteristics are very insignificant and show a high individual variability, most of these forms were probably not true subspecies, especially as they were often based upon zoo material of unknown origin that may have had "striking, but abnormal" morphological characteristics. Today, only eight subspecies are usually accepted, although one of these, the Cape lion, formerly described as Panthera leo melanochaita, is probably invalid. Even the remaining seven subspecies might be too many. While the status of the Asiatic lion (P. l. persica) as a subspecies is generally accepted, the systematic relationships among African lions are still not completely resolved. Mitochondrial variation in living African lions seemed to be modest according to some newer studies, therefore all sub-Saharan lions sometimes have been considered a single subspecies, however, a recent study revealed lions from western and central Africa differ genetically from lions of southern or eastern Africa. According to this study, Western African lions are more closely related to Asian lions than to South or East African lions. These findings might be explained by a late Pleistocene extinction event of lions in western and central Africa and a subsequent recolonization of these parts from Asia. Previous studies, which were focused mainly on lions from eastern and southern parts of Africa, already showed these can be 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. Another study revealed there are three major types of lions, one North African–Asian, one southern African and one middle African. Conversely, Per Christiansen found that using skull morphology allowed him to identify the subspecies krugeri, nubica, persica, and senegalensis, while there was overlap between bleyenberghi with senegalensis and krugeri. The Asiatic lion persica was the most distinctive, and the Cape lion had characteristics allying it more with P. l. persica than the other sub-Saharan lions. He had analysed 58 lion skulls in three European museums.
The majority of lions kept in zoos are hybrids of different subspecies. Approximately 77% of the captive lions registered by the International Species Information System are of unknown origin. Nonetheless, they might carry genes that are extinct in the wild, and might be therefore important to maintain overall genetic variability of the lion. It is believed that those lions, imported to Europe before the middle of the nineteenth century, were mainly either Barbary lions from North Africa or lions from the Cape.
Eight recent (Holocene) subspecies are recognised today:
- P. l. persica, known as the Asiatic lion or South Asian, Persian, or Indian lion, once was widespread from Turkey, across Southwest Asia, to India and Pakistan, now around 400 exist in and near the Gir Forest of India. Genetic evidence suggests its ancestors split from the ancestors of sub-Saharan African lions between 203 and 74 thousand years ago.
- P. l. leo, known as the Barbary lion, originally ranged from Morocco to Egypt. It is extinct in the wild due to excessive hunting, as the last wild Barbary lion was killed in Morocco in 1942. This was one of the largest of the lion subspecies, with reported lengths of 3.0–3.3 m (9.8–10.8 ft) and weights of more than 200 kg (440 lb) for males. It appears to be more closely related to the Asiatic rather than sub-Saharan lions. A number of animals in captivity are likely to be Barbary lions, particularly the 90 animals descended from the Moroccan Royal collection at Rabat Zoo.
- P. l. senegalensis, known as the West African lion, is found in western Africa, from Senegal to the Central African Republic.
- 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 Masai lion is found in East Africa, from Ethiopia and Kenya to Tanzania and Mozambique; a local population is known as the Tsavo lion.
- P. l. bleyenberghi, known as the southwest African or Katanga lion, is found in southwestern Africa, Namibia, Botswana, Angola, Katanga (Democratic Republic of the Congo), Zambia, and Zimbabwe.
- P. l. krugeri, known as the southeast African 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 its status as a distinct subspecies. The Cape lion probably was only the southernmost population of the extant P. l. krugeri.
- A newly discerned lion subspecies could exist in captivity in Ethiopia's capital city of Addis Ababa. Researchers compared the microsatellite variations over ten loci of fifteen lions in captivity with those of six different wild lion populations. They determined that these lions are genetically unique and presumably that "their wild source population is similarly unique." These lions—with males that have a distinctly dark and luxuriant mane seam to define a new subspecies perhaps native only to Ethiopia. These lions were part of a collection of the late Haile Selassie I of Ethiopia.
Several additional subspecies of lion existed in prehistoric times:
- P. l. fossilis, known as the 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 sizes comparable to the American cave lion and slightly larger than the Upper Pleistocene European cave lion.
- 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. This species is known from Paleolithic cave paintings, ivory carvings, and clay busts, indicating it had protruding ears, tufted tails, perhaps faint tiger-like stripes, and at least some had a ruff or primitive mane around their necks, possibly indicating males.
- P. l. atrox, known as the American lion or American cave lion, was abundant in the Americas from Canada to Peru in the Pleistocene Epoch until about 10,000 years ago. This form is the sister clade of P. l. spelaea, and likely arose when an early P. l. spelaea population became isolated south of the North American continental ice sheet about 0.34 Mya. One of the largest purported lion subspecies to have existed, its body length is estimated to have been 1.6–2.5 m (5.2–8.2 ft).
- P. l. youngi or Panthera youngi, flourished 350,000 years ago. Its relationship to the extant lion subspecies is obscure, and it probably represents a distinct species.
- 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.
- P. l. vereshchagini, the Beringian cave lion of Yakutia (Russia), Alaska (United States), and the Yukon Territory (Canada), has been considered a subspecies separate from P. l. spelaea on morphological grounds. However, mitochondrial DNA sequences obtained from cave lion fossils from Europe and Alaska were indistinguishable.
- P. l. mesopotamica or Mesopotamian lion, flourished during the Neo-Assyrian Period (approximately 1000–600 BC). It inhabited the Mesopotamian Plain where it probably represents a distinct sub-species. Nearly all ancient Mesopotamian representations of male lions demonstrate full underbelly hair in which until recently was only identified in the Barbary lion (Panthera leo leo) from Northern Africa and in most Asiatic lions (P. l. persica) from captivity in colder climates. Ancient evidence from adjacent landmasses reveal no substantiation for lions with underbelly hair in this manner so that the distinct phenotype of depicted lions in ancient Mesopotamia (including Babylon, Elam and ancient Persia) represent an extinct sub-species. Many of the images of these lions are derived from lion hunting sculptures so that the extinction of this sub-species likely resulted from overhunting in the ancient world.
- P. l. europaea, known as the European lion, was probably identical with Panthera leo persica or Panthera leo spelea. 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 ancient Romans and Greeks.
- 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 coloured individuals, it has been extinct since 1931. A less likely identity is a natural leopard-lion hybrid commonly known as a leopon.
Lions have been known to breed with tigers (most often the Siberian and Bengal subspecies) to create hybrids called ligers and tiglons (or tigons). They also have been crossed with leopards to produce leopons, and jaguars to produce jaglions. The marozi is reputedly a spotted lion or a naturally occurring leopon, while the Congolese spotted lion is a complex lion-jaguar-leopard hybrid called a lijagulep. Such hybrids were once commonly bred in zoos, but this is now discouraged due to the emphasis on conserving species and subspecies. Hybrids are still bred in private menageries and in zoos in China.
The liger is a cross between a male lion and a tigress. Because the growth-inhibiting gene from the female tiger mother is absent, the growth-promoting gene passed on by the male lion father is unimpeded by a regulating gene and 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 often are fertile. Males have about a 50% chance of having a mane, but if they grow them, their manes will be modest: around 50% the size of a pure lion mane.
Ligers are much bigger than normal lions, typically 3.65 m (12.0 ft) in length, and can weigh up to 500 kg (1,100 lb).
The less common tiglon or tigon is a cross between a lioness and a male tiger. In contrast to ligers, tigons are often relatively small in comparison to their parents, because of reciprocal gene effects.
Behind only the tiger, the lion is the second largest living felid in length and weight. Its skull is very similar to that of the tiger, although 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. Lion colouration 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 specialised roles that each gender plays in the pride. For instance, the lioness, the hunter, lacks the male's thick mane. The colour of the male's mane varies from blond to black, generally becoming darker as the lion grows older. 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.
The size of adult lions varies across their range with those from the southern African populations in Rhodesia, Kalahari and Kruger Park averaging around 189.6 kg (418 lb) and 126.9 kg (280 lb) in males and females respectively compared to 174.9 kg (386 lb) and 119.5 kg (263 lb) of male and female lions from East Africa. Reported body measurements in males are head-body lengths ranging from 170 to 250 cm (5 ft 7 in to 8 ft 2 in), tail lengths of 90–105 cm (2 ft 11 in–3 ft 5 in). In females reported head-body lengths range from 140 to 175 cm (4 ft 7 in to 5 ft 9 in), tail lengths of 70–100 cm (2 ft 4 in–3 ft 3 in), however, the frequently cited maximum head and body length of 250 cm (8 ft 2 in) fits rather to extinct Pleistocene forms, like the American lion, with even large modern lions measuring several centimetres less in length. Record measurements from hunting records are supposedly a total length of nearly 3.6 m (12 ft) for a male shot near Mucsso, southern Angola in October 1973 and a weight of 313 kg (690 lb) for a male shot outside Hectorspruit in eastern Transvaal, South Africa in 1936. Another notably outsized male lion, which was shot near Mount Kenya, weighed in at 272 kg (600 lb).
The mane of the adult male lion, unique among cats, is one of the most distinctive characteristics of the species. It may provide an excellent intimidation display; aiding the lion during confrontations with other lions. The presence, absence, colour, 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. 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.
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 colour and size of a lion's mane, such as the ambient temperature. 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. The males of the Asiatic subspecies, however, are characterised by sparser manes than average African lions.
In the Pendjari National Park area almost all males are maneless or have very weak manes. Maneless male lions have also been reported from Senegal, from Sudan (Dinder National Park), and from Tsavo East National Park in Kenya, and the original male white lion from Timbavati also was maneless. The testosterone hormone has been linked to mane growth, therefore castrated lions often have minimal to no mane, as the removal of the gonads inhibits testosterone production.
Cave paintings of extinct European cave lions almost exclusively show animals with no manes, suggesting that either they were maneless, or that the paintings depict lionesses as seen hunting in a group.
The white lion is not a distinct subspecies, but a special morph with a genetic condition, leucism, that causes paler colouration akin to that of the white tiger; the condition is similar to melanism, which causes black panthers. 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 colour of their coats is due to a recessive allele. Reportedly, they have been bred in camps in South Africa for use as trophies to be killed during canned hunts.
Lions spend much of their time resting and are inactive for about 20 hours per day. Although lions can be active at any time, their activity generally peaks after dusk with a period of socializing, grooming, and defecating. Intermittent bursts of activity follow through the night hours until dawn, when hunting most often takes place. They spend an average of two hours a day walking and 50 minutes eating.
Lions are the most socially inclined of all wild felids, most of which remain quite solitary in nature. The lion is a predatory carnivore with two types of social organization. Some lions are residents, living in groups centering around related lionesses, called prides. Females form the stable social unit in a pride and do not tolerate outside females. Membership only changes with the births and deaths of lionesses, although some females do leave and become nomadic. Although extremely large prides, consisting of up to 30 individuals, have been observed, the average pride consists of five or six 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. The number of adult males in a coalition is usually two but may increase to as many as four before decreasing again over time. The sole exception to this pattern is the Tsavo lion pride which always has just one adult male. Male cubs are excluded from their maternal pride when they reach maturity at around 2–3 years of age. The second organizational behaviour is labeled nomads, who range widely and move about sporadically, either singularly or in pairs. 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, as a rule, live at least some portion of their lives as nomads, and some are never 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. 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 calorific 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 and hone their skills. 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.
Lionesses do most of the hunting for their pride. They are more effective hunters, as they are 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 coordinated group with members who perform the same role consistently in order to stalk and bring down the prey successfully. 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, although pride members often behave aggressively toward each other as each tries to consume as much food as possible. Near the conclusion of the hunt, males have a tendency to dominate the kill once the lionesses have succeeded. They are more likely to share this with the cubs than with the lionesses, but males rarely share food they have killed by themselves.
Both males and females can defend the pride against intruders, but the male lion is better-suited for this purpose due to its stockier, more powerful build. Some individuals consistently lead the defence against intruders, while others lag behind. Lions tend to assume specific roles in the pride. Those lagging behind may provide other valuable services to the group. 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. 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.
Hunting and diet
Lions prefer to scavenge when the opportunity presents itself with carrion providing more than 50% of their diet. They scavenge animals either dead from natural causes (disease) 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. In fact, most dead prey on which both hyenas and lions feed upon are killed by the hyenas instead of the lions.
The lionesses do most of the hunting for the pride. The male lion associated with the pride usually stays and watches over young cubs until the lionesses return from the hunt. Typically, several work together and encircle the herd from different points. Once they have closed in on the herd, they usually target the animal closest to them. 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, 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 (which would also result in asphyxia).
Lions 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% of her body weight (a male's is about 0.45% of his body weight), whereas a hyena's heart is close to 1% of its body weight. Thus, they only run fast in short bursts, and need 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. They sneak up to the victim until they reach a distance of approximately 30 metres (98 feet) or less.
The prey consists mainly of medium-sized mammals, with a preference for wildebeest, 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 ungulates weighing between 50 and 300 kg (110 and 660 lb) such as kudu, hartebeest, gemsbok, and eland. Occasionally, they 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 in most parts of their range they rarely attack very large prey such as fully grown male giraffes due to the danger of injury. Giraffes and buffaloes are almost invulnerable to a solitary lion as well.
Extensive studies show that lionesses normally prey on mammals with an average weight of 126 kg (278 lb), while kills made by male lions average 399 kg (880 lb). In Africa, wildebeest rank at the top of preferred prey (making nearly half of the lion prey in the Serengeti) followed by zebra. Lions do not prey on fully grown adult elephants; most adult hippopotamuses, rhinoceroses, and smaller gazelles, impala, and other agile antelopes are generally excluded. However, giraffes and buffaloes are often taken in certain regions. For instance, in Kruger National Park, giraffes are regularly hunted. In Manyara Park, Cape buffaloes constitute as much as 62% of the lion's diet, due to the high number density of buffaloes. Occasionally hippopotamus is also taken, but adult rhinoceroses are generally avoided. Warthogs are often taken depending on availability. The lions of Savuti, Botswana, have adapted to hunting young elephants during the dry season, and a pride of 30 lions has been recorded killing individuals between the ages of four and eleven years. In the Kalahari desert in South Africa, black-maned lions may chase baboons up a tree, wait patiently, then attack them when they try to escape:
Lions also attack domestic livestock and in India cattle contribute significantly to their diet. Lions 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. A lion may gorge itself and eat up to 30 kg (66 lb) in one sitting; if it is unable to consume all the kill it will rest for a few hours before consuming more. On a hot day, the pride may retreat to shade leaving a male or two to stand guard. An adult lioness requires an average of about 5 kg (11 lb) of meat per day, a male about 7 kg (15 lb).
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 kilometres 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. There is evidence that male lions are just as successful at hunting as females; they are solo hunters who ambush prey in small bush. Young lions first display stalking behaviour 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.
Lions and spotted hyenas occupy the same ecological niche, meaning they compete for prey and carrion in the areas where they coexist. A review of data across several studies indicates a dietary overlap of 58.6%. Lions typically ignore spotted hyenas unless the lions are on a kill or are being harassed by the hyenas, while the latter tend to visibly react to the presence of lions whether there is food or not. Lions seize the kills of spotted hyenas: in the Ngorongoro crater, it is common for lions to subsist largely on kills stolen from hyenas, causing the hyenas to increase their kill rate. On the other hand, in Northern Botswana's Chobe National Park, the situation is reversed: hyenas frequently challenge lions and steal their kills: they obtain food from 63% of all lion kills. When confronted on a kill by lions, spotted hyenas may either leave or wait patiently at a distance of 30–100 m (98–328 ft) until the lions have finished, but they are also bold enough to feed alongside lions, and even force the lions off a kill. The two species may attack one another even when there is no food involved for no apparent reason. Lion predation can account for up to 71% of hyena deaths in Etosha. Spotted hyenas have adapted by frequently mobbing lions that enter their territories. Experiments on captive spotted hyenas revealed that specimens with no prior experience with lions act indifferently to the sight of them, but will react fearfully to the scent. The size of male lions allows them occasionally to confront hyenas in otherwise evenly matched brawls and so to tip the balance in favour of the lions.
Lions tend to dominate smaller felines such as cheetahs and leopards where they co-occur, stealing their kills and killing their cubs and even adults when given the chance. The cheetah has a 50% chance of losing its kill to lions or other predators. Lions are major killers of cheetah cubs, up to 90% 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. Similarly, lions dominate African wild dogs, not only taking their kills but also preying on young and (rarely) adult dogs. Population densities of wild dogs are low in areas where lions are more abundant. However, there are a few reported cases of old and wounded lions falling prey to wild dogs.
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, while the reverse is true for lions entering waterways, as evidenced by the occasional lion claw found in crocodile stomachs.
While lions do not usually hunt people, some (usually males) seem to seek out human prey; one well-publicised case includes the Tsavo maneaters, where 28 officially recorded 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. The hunter who killed the lions wrote a book detailing the animals' predatory behaviour. The lions were larger than normal, lacked manes, and one 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.
In their analysis of Tsavo and general man-eating, Kerbis Peterhans and Gnoske acknowledge that sick or injured animals may be more prone to man-eating, but that the behaviour 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 among other pantherines and primates in the paleontological record.
The lion's proclivity for man-eating has been systematically examined. American and Tanzanian scientists report that man-eating behaviour 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. Another study of 1,000 people attacked by lions in southern Tanzania between 1988 and 2009 found that the weeks following the full moon (when there was less moonlight) were a strong indicator of increased night attacks on people.
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.
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 has documented that between 1990 and 2004, lions attacked 815 people in Tanzania, killing 563. Packer and Ikanda are among the few conservationists who believe western conservation efforts must take account of these matters not just because of ethical concerns about human life, but also for the long term success of conservation efforts and lion preservation.
A man-eating lion was killed by game scouts in Southern Tanzania in April 2004. It is believed to have killed and eaten at least 35 people in a series of incidents covering several villages in the Rufiji Delta coastal region. Dr Rolf D. Baldus, the GTZ wildlife programme coordinator, commented that it was likely that the lion preyed on humans because it had a large abscess underneath a molar that was cracked in several places. He further commented that "This lion probably experienced a lot of pain, particularly when it was chewing." GTZ is the German development cooperation agency and has been working with the Tanzanian government on wildlife conservation for nearly two decades. 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 incidents in the early 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 people in what is now Njombe district.
Reproduction and life cycle
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Most lionesses will have reproduced by the time they are four years of age. Lions do not mate at any specific time of year, and the females are polyestrous. As with other cats' penises, the male lion's penis has spines that point backward. During withdrawal of the penis, the spines rake the walls of the female's vagina, which may cause ovulation. A lioness may mate with more than one male when she is in heat; during a mating bout, which could last several days, the couple copulates twenty to forty times a day, often forgoing eating. Lions reproduce very well in captivity.
The average gestation period is around 110 days, the female giving birth to a litter of one to four cubs 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 while the cubs are still helpless, staying relatively close to the thicket or den where the cubs are kept. 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. 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.
Usually, the mother does not integrate herself and her cubs back into the pride until the cubs are six to eight weeks old. Sometimes this introduction to pride life occurs earlier, however, particularly if other lionesses have given birth at about the same time. For instance, lionesses in a pride often synchronise 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, usually are dominated by larger cubs at mealtimes – consequently, death by starvation is more common among the younger cubs.
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 toward the thicket or den where they are being kept, doing their best to trample the cubs to death while 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, perhaps because females do not become fertile and receptive until their cubs mature or die. All in all, as many as 80% of the cubs will die before the age of two.
When first introduced to the rest of the pride, the cubs initially lack confidence when confronted with adult lions other than their mother. They soon begin to immerse themselves in the pride life, however, playing among 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 toward 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.
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, if they have not already been critically injured while 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.
Contrary to popular belief, it is not only males that are ousted from their pride to become nomads, although most 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. 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. Lions are shown to involved in group homosexual and courtship activities. Male Lions will also head rub and roll around with each other before having sex together.
Although adult lions have no natural predators, evidence suggests that the majority die violently from humans or other lions. Lions often inflict serious injuries on each other, either members of different prides encountering each other in territorial disputes, or members of the same pride fighting at a kill. Crippled lions and lion cubs may fall victim to hyenas, leopards, or be trampled by buffalo or elephants, and careless lions may be maimed when hunting prey.
Various species of tick commonly infest the ears, neck and groin regions of most lions. Adult forms of several species of the tapeworm genus Taenia have been isolated from intestines, the lions having ingested larval forms from antelope meat. 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. A more recent outbreak in 2001 killed six lions. Lions, especially in captivity, are vulnerable to the canine distemper virus (CDV), feline immunodeficiency virus (FIV), and feline infectious peritonitis (FIP). 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. 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.
When resting, lion socialization occurs through a number of behaviours, and the animal's expressive movements are highly developed. The most common peaceful tactile gestures are head rubbing and social licking, which have been compared with grooming in primates. Head rubbing – nuzzling one's forehead, face and neck against another lion – appears to be a form of greeting, as it is seen often after an animal has been apart from others, or after a fight or confrontation. Males tend to rub other males, while cubs and females rub females. Social licking often occurs in tandem with head rubbing; it is generally mutual and the recipient appears to express pleasure. The head and neck are the most common parts of the body licked, which may have arisen out of utility, as a lion cannot lick these areas individually.
A lion in captivity roaring
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Lions have an array of facial expressions and body postures that serve as visual gestures. Their repertoire of vocalizations is also large; variations in intensity and pitch, rather than discrete signals, appear central to communication. Lion sounds include snarling, 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. Lions have the loudest roar of any big cat.
Distribution and habitat
In Africa, lions can be found in savanna grasslands with scattered Acacia trees, which serve as shade; their habitat in India is a mixture of dry savanna forest and very dry deciduous scrub forest. The habitat of lions originally 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 in 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. By 100 AD they were extirpated. A population of Asiatic lions survived until the tenth century in the Caucasus, their last European outpost.
The species was eradicated from Palestine by the Middle Ages and from most of the rest of Asia after the arrival of readily available firearms in the eighteenth century. Between the late nineteenth and early twentieth century, they became extinct in North Africa and Southwest Asia. By the late nineteenth century, the lion had disappeared from Turkey and most of northern India, while the last sighting of a live Asiatic lion in Iran was in 1941 (between Shiraz and Jahrom, Fars Province), although the corpse of a lioness was found on the banks of the Karun river, Khūzestān Province in 1944. There are no subsequent reliable reports from Iran. The subspecies now survives only in and around the Gir Forest of northwestern India. Approximately 400 lions live in the area of the 1,412 km2 (545 sq mi) sanctuary in the state of Gujarat, which covers most of the forest. Their numbers have increased from 180 to 411 animals mainly because the natural prey species have recovered.
Population and conservation status
Most lions now live in eastern and southern Africa, and their numbers there are rapidly decreasing, with an estimated 30–50% decline per 20 years in the late half of the 20th century. Estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004, down from early 1990s estimates that ranged as high as 100,000 and perhaps 400,000 in 1950. Primary causes of the decline include disease and human interference. Habitat loss and conflicts with humans are considered the most significant threats to the species. The remaining populations are often geographically isolated from one another, which can lead to inbreeding, and consequently, reduced genetic diversity. Therefore the lion is considered a vulnerable species by the International Union for Conservation of Nature, while the Asiatic subspecies is 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. Another population in northwestern Africa is found in Waza National Park, where approximately 14–21 animals persist.
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. The Ewaso Lions Project protects lions in the Samburu National Reserve, Buffalo Springs National Reserve and Shaba National Reserve of the Ewaso Ng'iro ecosystem in Northern Kenya. Outside these areas, the issues arising from lions' interaction with livestock and people usually results in the elimination of the former. In India, the last refuge of the Asiatic lion is the 1,412 km2 (545 sq mi) Gir Forest National Park in western India, which had approximately 180 lions in 1974 and about 400 in 2010. As in Africa, numerous human habitations are close by with the resultant problems between lions, livestock, locals and wildlife officials. The Asiatic Lion Reintroduction Project plans to establish a second independent population of Asiatic lions at the Kuno Wildlife Sanctuary in the Indian state of Madhya Pradesh. It is important to start a second population to serve as a 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 lions at Port Lympne Wild Animal Park in Kent, England that are descended from animals owned by the King of Morocco. Another eleven animals believed to be Barbary lions were found in Addis Ababa zoo, descendants of animals owned by Emperor Haile Selassie. WildLink International, in collaboration with Oxford University, launched their ambitious International Barbary Lion Project with the aim of identifying and breeding Barbary lions in captivity for eventual reintroduction into a national park in the Atlas Mountains of Morocco.
Following the discovery of the decline of lion population in Africa, several coordinated efforts involving lion conservation have been 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 hybridised 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.
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. Although many modern zoos are more selective about their exhibits, there are more than 1,000 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. 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, were still alive in August 2007. Breeding programs need to note origins to avoid breeding different subspecies and thus reducing conservation value. However, several Asiatic-African lion crosses have been bred.
At the ancient Egyptian cities of Taremu and Per-Bast were temples to the lioness goddesses of Egypt, Sekhmet and Bast and at Taremu there was a temple to the son of the deity, Maahes the lion prince, where live lions were kept and allowed to roam within his temple. The Greeks called the city Leontopolis, the "City of Lions" and documented that practice. Lions were kept and bred by Assyrian kings as early as 850 BC, and Alexander the Great was said to have been presented with tame lions by the Malhi of northern India. Later in Roman times, lions were kept by emperors to take part in the gladiator arenas or for executions (see bestiarii, damnatio ad bestias, and venatio). Roman notables, including Sulla, Pompey, and Julius Caesar, often ordered the mass slaughter of hundreds of lions at a time. In the East, lions were tamed by Indian princes, and Marco Polo reported that Kublai Khan kept lions inside. The first European "zoos" spread among 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. 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, probably stocked with animals from an earlier menagerie started in 1125 by Henry I at his hunting lodge in Woodstock, near Oxford; where lions had reportedly been stocked by William of Malmesbury.
Seraglios served as expressions of the nobility's power and wealth. Animals such as big cats and elephants, in particular, symbolised 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.
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. 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. A rival menagerie at the Exeter Exchange also exhibited lions until the early nineteenth century. The Tower menagerie was closed down by William IV, and animals transferred to the London Zoo, which opened its gates to the public on 27 April 1828.
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 giant pandas. Like other animals, lions were seen as little more than a natural, boundless commodity that was mercilessly exploited with terrible losses in capture and transportation. The widely reproduced imagery of the heroic hunter chasing lions would dominate a large part of the century. Explorers and hunters exploited a popular Manichean division of animals into "good" and "evil" to add thrilling value to their adventures, casting themselves as heroic figures. This resulted in big cats, always suspected of being man-eaters, representing "both the fear of nature and the satisfaction of having overcome it."
Lions were kept in cramped and squalid conditions at London Zoo until a larger lion house with roomier cages was built in the 1870s. Further changes took place in the early 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. 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. 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. 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 bond with them, particularly the latter. The lioness later achieved fame, her life being documented in a series of books and films.
Baiting and 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 seventeenth century. It was finally banned in Vienna by 1800 and England in 1835.
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. Van Amburgh performed before Queen Victoria 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. The ultimate proof of a tamer's dominance and control over a lion is demonstrated by placing his head in the lion's mouth. The now iconic lion tamer's chair was possibly first used by American Clyde Beatty (1903–1965).
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 and noble. A common depiction is their representation as "king of the jungle" or "king of beasts"; hence, the lion has been a popular symbol of royalty and stateliness, as well as a symbol of bravery; it is featured in several fables of the sixth century BC Greek storyteller Aesop.
Representations of lions date back to the early Upper Paleolithic. The lioness-headed ivory carving from Vogelherd cave in the Swabian Alb in southwestern Germany, dubbed Löwenmensch (lion-human) in German. The sculpture has been determined to be at least 32,000 years old from the Aurignacian culture, but it may date to as early as 40,000 years ago. The sculpture has been interpreted as anthropomorphic, giving human characteristics to an animal, however, it also may represent a deity.
Two lions were depicted mating in the Chamber of Felines in 15,000-year-old Paleolithic cave paintings in the Lascaux caves. Cave lions also are depicted in the Chauvet Cave, discovered in 1994; this has been dated at 32,000 years of age, though it may be of similar or younger age to Lascaux.
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; 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.
The lion was a prominent symbol in ancient Mesopotamia (from Sumer up to Assyrian and Babylonian times), where it was strongly associated with kingship. 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.
In the Talmud, Chullin 59b, Rabbi Joshua ben Hananiah tells Emperor Hadrian about the giant lion of the forest of Bei Ilai called the Tigris, a lion so huge that the space between its ears measures 9 cubits. The emperor asks the rabbi to call forth this lion. He reluctantly agrees. At a distance of 400 parasangs from Rome it roars, and all pregnant women miscarry and all the walls of Rome fall down. Then it comes to 300 parasangs and roars, and all the front teeth and molars of Roman men fall out, and even the emperor himself falls from his throne. He begs the rabbi to send it back. The rabbi prays and it returns to its place.
In the Puranic texts of Hinduism, Narasimha ("man-lion") a half-lion, half-man incarnation or (avatar) of Vishnu, is worshipped by his devotees and saved the child devotee Prahlada from his father, the evil demon king Hiranyakashipu; 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. 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. 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. Farther south on the Indian subcontinent, the Asiatic lion is symbolic for the Sinhalese, 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.
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 usually were wingless, with shorter, thicker bodies, and curly manes. 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.
The island nation of Singapore derives its name from the Malay words singa (lion) and pora (city/fortress), which in turn is from the Tamil-Sanskrit சிங்க singa सिंह siṃha and पुर புர pura, which is cognate to the Greek πόλις, pólis. According to the Malay Annals, this name was given by a fourteenth-century Sumatran Malay prince Sang Nila Utama, who, on alighting the island after a thunderstorm, spotted an auspicious beast on shore that appeared to be a lion.
The name of the nomadic Hadendoa people, inhabiting parts of Sudan, Egypt, and Eritrea, is made up of haɖa 'lion' and (n)ɖiwa 'clan'. Other variants are Haɖai ɖiwa, Hanɖiwa, and Haɖaatʼar (children of lioness).
"Lion" was the nickname of several medieval warrior rulers with a reputation for bravery, such as the English King Richard the Lionheart, Henry the Lion, (German: Heinrich der Löwe), Duke of Saxony, William the Lion, King of Scotland, 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, but the lioness 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. 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 of the NFL, Chelsea and Aston Villa of the English Premier League, (and the Premiership itself), Eintracht Braunschweig of the Bundesliga, and to a host of smaller clubs around the world.
Lions continue to be featured in modern literature, from the messianic Aslan in The Lion, the Witch and the Wardrobe and following books from The Chronicles of Narnia series written by C. S. Lewis, to the comedic Cowardly Lion in The Wonderful Wizard of Oz. 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, which has been the mascot for Metro-Goldwyn-Mayer (MGM) studios since the 1920s. The 1960s saw the appearance of what is possibly the most famous lioness, the Kenyan animal Elsa in the movie Born Free, based on the true-life international bestselling book of the same title. The lion's role as King of the Beasts has been used in cartoons, from the 1950s manga that 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, which also featured the popular song "The Lion Sleeps Tonight" in its soundtrack. A lion appears on the 50-rand South African banknote.
- Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. pp. 532–628. ISBN 978-0-8018-8221-0. OCLC 62265494.
- Bauer, H., Nowell, K.; Packer, C. (2008). "Panthera leo". IUCN Red List of Threatened Species. Version 2010.2. International Union for Conservation of Nature. Retrieved 28 August 2010.
- Linnaeus, Carolus (1758). Systema naturae per regna tria naturae :secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. (in Latin) 1 (10th ed.). Holmiae (Laurentii Salvii). p. 41. Retrieved 8 September 2008.
- Nowak, Ronald M. (1999). Walker's Mammals of the World. Baltimore: Johns Hopkins University Press. ISBN 0-8018-5789-9.
- Harington, C. R. "Dick" (1969). "Pleistocene remains of the lion-like cat (Panthera atrox) from the Yukon Territory and northern Alaska". Canadian Journal of Earth Sciences 6 (5): 1277–88. doi:10.1139/e69-127.
- Smuts, G. L. (1982). Lion. Johannesburg: Macmillan South Africa. p. 231. ISBN 0-86954-122-6.
- Lions' nocturnal chorus. Earth-touch.com. Retrieved on 31 July 2013.
- African Lion Panthera leo. Phoenix Zoo Fact Sheet.
- Simpson, D. P. (1979). Cassell's Latin Dictionary (5th ed.). London: Cassell Ltd. p. 342. ISBN 0-304-52257-0.
- Liddell, Henry George; Scott, Robert (1980). A Greek-English Lexicon (Abridged Edition). United Kingdom: Oxford University Press. p. 411. ISBN 0-19-910207-4.
- Simpson, John; Weiner, Edmund, ed. (1989). "Lion". Oxford English Dictionary (2nd ed.). Oxford: Clarendon Press. ISBN 0-19-861186-2.
- Yamaguchi, Nobuyuki; Cooper, Alan; Werdelin, Lars; MacDonald, David W. (August 2004). "Evolution of the mane and group-living in the lion (Panthera leo): a review". Journal of Zoology 263 (4): 329–42. doi:10.1017/S0952836904005242.
- Turner, Allen (1997). The big cats and their fossil relatives: an illustrated guide to their evolution and natural history. New York: Columbia University Press. ISBN 0-231-10229-1.
- Harington, C. R. (Dick) (1996). "American Lion". Yukon Beringia Interpretive Centre website. Yukon Beringia Interpretive Centre. Retrieved 25 January 2011.
- Burger, Joachim; Rosendahl, Wilfried; Loreille, Odile; Hemmer, Helmut; Eriksson, Torsten; Götherström, Anders; Hiller, Jennifer; Collins, Matthew J.; Wess, Timothy; Alt, Kurt W. (2004). "Molecular phylogeny of the extinct cave lion Panthera leo spelaea". Molecular Phylogenetics and Evolution (fulltext) 30 (3): 841–49. doi:10.1016/j.ympev.2003.07.020. PMID 15012963. Retrieved 20 September 2007.
- Grisham, Jack (2001). "Lion". In Bell, Catherine E. Encyclopedia of the World's Zoos. 2: G–P. Chofago: Fitzroy Dearborn. pp. 733–39. ISBN 1-57958-174-9.
- Barnett, Ross; Yamaguchi, Nobuyuki; Barnes, Ian; Cooper, Alan (August 2006). "Lost populations and preserving genetic diversity in the lion Panthera leo: Implications for its ex situ conservation". Conservation Genetics 7 (4): 507–14. doi:10.1007/s10592-005-9062-0.
- Bertola, L. D.; Van Hooft, W. F.; Vrieling, K.; Uit De Weerd, D. R.; York, D. S.; Bauer, H.; Prins, H. H. T.; Funston, P. J.; Udo De Haes, H. A.; Leirs, H.; Van Haeringen, W. A.; Sogbohossou, E.; Tumenta, P. N.; De Iongh, H. H. (2011). "Genetic diversity, evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa". Journal of Biogeography 38 (7): 1356–67. doi:10.1111/j.1365-2699.2011.02500.x.
- Dubach, Jean et al. (January 2005). "Molecular genetic variation across the southern and eastern geographic ranges of the African lion, Panthera leo". Conservation Genetics 6 (1): 15–24. doi:10.1007/s10592-004-7729-6.
- Barnett, Ross; Yamaguchi, Nobuyuki; Barnes, Ian; Cooper, Alan (2006). "The origin, current diversity and future conservation of the modern lion (Panthera leo)" (PDF). Proceedings of the Royal Society B: Biological Sciences 273 (1598): 2119–25. doi:10.1098/rspb.2006.3555. PMC 1635511. PMID 16901830. Retrieved 4 September 2007.
- Christiansen, Per (2008). "On the distinctiveness of the Cape lion (Panthera leo melanochaita Smith, 1842), and a possible new specimen from the Zoological Museum, Copenhagen". Mammalian Biology 73 (1): 58–65. doi:10.1016/j.mambio.2007.06.003.
- Barnett, Ross; Yamaguchi, Nobuyuki; Shapiro, Beth; Nijman, Vincent (2007). "Using ancient DNA techniques to identify the origin of unprovenanced museum specimens, as illustrated by the identification of a 19th century lion from Amsterdam". Contributions to Zoology 76 (2): 87–94.
- "Asiatic Lion—History". Asiatic Lion Information Centre. Wildlife Conservation Trust of India. 2006. Retrieved 15 September 2007.
- Singh, H. S.; Gibson, L. (2011). "A conservation success story in the otherwise dire megafauna extinction crisis: The Asiatic lion (Panthera leo persica) of Gir forest". Biological Conservation 144 (5): 1753–1757. doi:10.1016/j.biocon.2011.02.009.
- Nowell, Kristin; Jackson, Peter (1996). "Panthera Leo". Wild Cats: Status Survey and Conservation Action Plan (PDF). Gland, Switzerland: IUCN/SSC Cat Specialist Group. pp. 17–21. ISBN 2-8317-0045-0.
- Black, S. A.; Fellous, A.; Yamaguchi, N.; Roberts, D. L. (2013). "Examining the Extinction of the Barbary Lion and Its Implications for Felid Conservation". PLoS ONE 8 (4): e60174. doi:10.1371/journal.pone.0060174.
- Yadav, P. R. (2004). Vanishing And Endangered Species. Discovery Publishing House. pp. 176–78. ISBN 81-7141-776-0. Retrieved 14 December 2010.
- Burger, Joachim; Hemmer, Helmut (2006). "Urgent call for further breeding of the relic zoo population of the critically endangered Barbary lion (Panthera leo leo Linnaeus 1758)". European Journal of Wildlife Research 52: 54. doi:10.1007/s10344-005-0009-z.
- Black, Simon; Yamaguchi, Nobuyuki; Harland, Adrian; Groombridge, Jim (2010). "Maintaining the genetic health of putative Barbary lions in captivity: an analysis of Moroccan Royal Lions". European Journal of Wildlife Research 56: 21–31. doi:10.1007/s10344-009-0280-5.
- Sunquist, M. E. and Sunquist, F. C. (2009). Family Felidae (Cats). In: Don E. Wilson, Russell A. Mittermeier (eds.): Handbook of the Mammals of the World. Volume 1: Carnivores. Lynx Edicions, 2009, ISBN 978-84-96553-49-1, pp. 137 ff.
- Macdonald, David Wayne (2006). The Encyclopedia of Mammals. pp. 628–635. ISBN 0-87196-871-1.
- "A New, Genetically Distinct Lion Population is Found". News Watch. National Geographic Society. 30 November 2012. Retrieved 23 January 2013.
The Addis Ababa zoo lions have dark manes and small bodies, unlike other African lions. But life in captivity can sometimes influence appearance. A team of researchers, led by the Max Planck Institute for Evolutionary Anthropology in Germany and the University of York in the UK, checked to see if the lions really are different by comparing DNA samples of 15 lions from the zoo to six populations of wild lions. Their genetic analysis revealed that the gene sequence of all fifteen lions were unique and showed little sign of inbreeding. The study was recently published in the European Journal of Wildlife Research.
- Bruche, Susann; Gusset, Markus; Lippold, Sebastian; Barnett, Ross; Eulenberger, Klaus; Junhold, Jörg; Driscoll, Carlos A.; Hofreiter, Michael (2012). "A genetically distinct lion (Panthera leo) population from Ethiopia". European Journal of Wildlife Research 59 (2): 215–225. doi:10.1007/s10344-012-0668-5.
- Probst, Ernst (1999). Deutschland in der Urzeit (in German). Orbis. ISBN 3-572-01057-8.
- Packer, Craig; Jean Clottes (November 2000). "When Lions Ruled France". Natural History: 52–57.
- Koenigswald, Wighart von (2002). Lebendige Eiszeit: Klima und Tierwelt im Wandel (in German). Stuttgart: Theiss. ISBN 3-8062-1734-3.
- Barnett, Ross; Shapiro, Beth; Barnes, Ian; Ho, Simon Y.W.; Burger, Joachim; Yamaguchi, Nobuyuki; Higham, Thomas FG; Wheeler, H Todd; Rosendahl, Wilfried; Sher, Andrei V.; Sotnikova, Marina; Kuznetsova, Tatiana; Baryshnikov, Gennady F.; Martin, Larry D.; Harington, C. Richard; Burns, James A.; Cooper, Alan (April 2009). "Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity". Molecular Ecology 18 (8): 1668–77. doi:10.1111/j.1365-294X.2009.04134.x. PMID 19302360. Retrieved 23 November 2011.
- Martin, Paul S. (1984). Quaternary Extinctions. Tucson, AZ: University of Arizona Press. ISBN 0-8165-1100-4.
- Manamendra-Arachchi, Kelum; Pethiyagoda, Rohan; Dissanayake, Rajith; Meegaskumbura, Madhava (2005). "A second extinct big cat from the late Quaternary of Sri Lanka" (PDF). The Raffles Bulletin of Zoology (National University of Singapore). Supplement 12: 423–34. Retrieved 31 July 2007.
- Ashrafian, Hutan (2011). "An Extinct Mesopotamian Lion Subspecies". Veterinary Heritage 34 (2): 47–49.
- Shuker, Karl P. N. (1989). Mystery Cats of the World. Robert Hale. ISBN 0-7090-3706-6.
- Guggisberg, Charles Albert Walter (1975). Wild Cats of the World. New York: Taplinger Publishing. ISBN 0-8008-8324-1.
- Doi, H; Reynolds, B (1967). The Story of Leopons. New York: Putnam. OCLC 469041.
- Rafferty, John P. (2011). Carnivores. The Rosen Publishing Group. p. 120. ISBN 9781615303403. Retrieved 4 July 2014.
- Shi, Wei (2005). Growth and Behviour: Epigenetic and Genetic Factors Involved in Hybrid Dysgenesis. Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 11 (Uppsala: Acta Universitatis Upsaliensis). p. 9. ISBN 91-554-6147-6.
- "Tigon—Encyclopædia Britannica Article". Retrieved 25 January 2011.
- Heptner, V. G.; Sludskii, A. A. (1992). Mammals of the Soviet Union, Volume II, Part 2. Leiden u.a.: Brill. ISBN 90-04-08876-8.
- Schaller, p. 28.
- Smuts, G. L.; Robinson, G.A.; Whyte, I.J. (1980). "Comparative growth of wild male and female lions (Panthera leo)". Journal of Zoology 190 (3): 365–373. doi:10.1111/j.1469-7998.1980.tb01433.x.
- Mazák, Vratislav (April 2004) Der Tiger, Westarp Wissenschaften, 5th ed., pp. 178 ff., ISBN 3-89432-759-6.
- Wood, The Guinness Book of Animal Facts and Feats. Sterling Pub Co Inc (1983), ISBN 978-0-85112-235-9
- Trivedi, Bijal P. (2005). "Are Maneless Tsavo Lions Prone to Male Pattern Baldness?". National Geographic. Retrieved 7 July 2007.
- Trivedi, Bijal P. (22 August 2002). "Female Lions Prefer Dark-Maned Males, Study Finds". National Geographic News. National Geographic. Retrieved 1 September 2007.
- West, Peyton M.; Packer, Craig (August 2002). "Sexual Selection, Temperature, and the Lion's Mane". Science 297 (5585): 1339–43. doi:10.1126/science.1073257. PMID 12193785.
- Yamaguchi, Nobuyuki; Haddane, B. (2002). "The North African Barbary Lion and the Atlas Lion Project". International Zoo News 49: 465–81.
- Menon, Vivek (2003). A Field Guide to Indian Mammals. Delhi: Dorling Kindersley India. ISBN 0-14-302998-3.
- Schoe, Marjolein; Sogbohossou, Etotépé A.; Kaandorp, Jacques and de Iongh, Hans (2010) Progress Report – collaring operation Pendjari Lion Project, Benin. Funded by the Dutch Zoo Conservation Fund.
- Munson, Linda (3 March 2006). "Contraception in felids". Theriogenology 66 (1): 126–34. doi:10.1016/j.theriogenology.2006.03.06. PMID 16626799.
- McBride, Chris (1977). The White Lions of Timbavati. Johannesburg: E. Stanton. ISBN 0-949997-32-3.
- Tucker, Linda (2003). Mystery of the White Lions—Children of the Sun God. Mapumulanga: Npenvu Press. ISBN 0-620-31409-5.
- Schaller, p. 122.
- Schaller, pp. 120–21.
- Schaller, p. 33.
- Schaller, p. 37.
- Schaller, p. 39.
- Schaller, p. 44.
- Milius, Susan (April 2002). "Biology: Maneless lions live one guy per pride". Society for Science & the Public 161 (16): 253. doi:10.1002/scin.5591611614.
- Schaller, p. 133.
- Join the Pride. Lion Conservation Fund. Retrieved on 31 July 2013.
- Heinsohn, R.; C. Packer (1995). "Complex cooperative strategies in group-territorial African lions". Science 269 (5228): 1260–62. doi:10.1126/science.7652573. PMID 7652573.
- Morell, V. (1995). "Cowardly lions confound cooperation theory". Science 269 (5228): 1216–17. doi:10.1126/science.7652566. PMID 7652566.
- Jahn, Gary C. (1996). "Lioness Leadership". Science 271 (5253): 1215. doi:10.1126/science.271.5253.1215a. PMID 17820922.
- Schaller, p. 213.
- >"Behavior and Diet". African Wildlife Foundation website. African Wildlife Foundation. 1996. Retrieved 6 June 2014.
- Mills, Gus. "About lions—Ecology and behaviour". African Lion Working Group. Retrieved 20 July 2007.
- Schaller, p. 248.
- Schaller, pp. 247–48.
- Schaller, p. 237.
- Schaller, p. 259.
- Radloff, Frans G. T.; Du Toit, Johan T. (2004). "Large predators and their prey in a southern African savanna: a predator's size determines its prey size range". Journal of Animal Ecology 73: 410–423. doi:10.1111/j.0021-8790.2004.00817.x.
- Denis-Huot, Christine; Denis-Huot, Michel (2002). The Art of Being a Lion. Friedman/Fairfax. p. 186. ISBN 88-8095-752-X.
- Pienaar, U. de V. (1969). "Predator-prey relationships among the larger mammals of the Kruger National Park". Koedoe 12 (1). doi:10.4102/koedoe.v12i1.753.
- Douglas-Hamilton, Iain; Douglas-Hamilton, Oria (1975). "Among the Elephants".
- Hayward, Matt W.; Kerley, Graham (2005). "Prey preferences of the lion (Panthera leo)". Journal of Zoology 267 (3): 309–22. doi:10.1017/S0952836905007508.
- Power, R. J.; Shem Compion, R. X. (2009). "Lion predation on elephants in the Savuti, Chobe National Park, Botswana". African Zoology 44 (1): 36–44. doi:10.3377/004.044.0104.
- Guggisberg, Charles Albert Walter (1961). Simba: the life of the lion. Cape Town: Howard Timmins.
- Schaller, pp. 270–76.
- Schaller, p. 276.
- Stander, PE (1992). "Cooperative hunting in lions: the role of the individual". Behavioral Ecology and Sociobiology 29 (6): 445–54. doi:10.1007/BF00170175.
- Scott R. Loariea; Craig J. Tamblingb; Gregory P. Asnera (2013). "Lion hunting behaviour and vegetation structure in an African savanna". Animal Behaviour 85 (5): 899–906. doi:10.1016/j.anbehav.2013.01.018.
- Schaller, p. 153.
- Hayward, Matt W. (2006). "Prey preferences of the spotted hyaena (Crocuta crocuta) and degree of dietary overlap with the lion (Panthera leo)" (PDF). Journal of Zoology 270: 606–14. doi:10.1111/j.1469-7998.2006.00183.
- Kruuk, Hans (1972). "Interactions between Hyenas and other Carnivorous Animals". The Spotted Hyena: A Study of Predation and Social Behaviour. Chicago: The University of Chicago Press. ISBN 0-226-45508-4.
- Gittleman, John L.; Funk, Stephan M.; Macdonald, David W.; Wayne, Robert K., eds. (2001). Carnivore Conservation (1 ed.). Cambridge University Press. ISBN 978-0-521-66232-1.
- Schaller, p. 272.
- Schaller, pp. 273–274.
- Joubert, Dereck and Joubert, Beverley (1992). Eternal Enemies: Lions and Hyenas (DVD). National Geographic.
- Trinkel, Martina; Kastberger, Gerald (2005). "Competitive interactions between spotted hyenas and lions in the Etosha National Park, Namibia". African Journal of Ecology 43 (3): 220–24. doi:10.1111/j.1365-2028.2005.00574.x.
- O'Brien, Stephen J; Wildt, David E; Bush, Mitchell (1986). "The Cheetah in Genetic Peril" (PDF). Scientific American (254): 68–76.
- Schaller, p. 293.
- Woodroffe, Rosie; Ginsberg, Joshua R (1999). "Conserving the African wild dog Lycaon pictus. I. Diagnosing and treating causes of decline". Oryx 33 (2): 132–42. doi:10.1046/j.1365-3008.1999.00052.x.
- Schaller, p. 188.
- "Crocodiles!". Nova (transcript). PBS. 28 April 1998. Retrieved 21 November 2010.
- Guggisberg, Charles Albert Walter (1972). Crocodiles: Their Natural History, Folklore, and Conservation. Newton Abbot: David & Charles. p. 195. ISBN 0-7153-5272-5.
- Patterson, Bruce D. (2004). The Lions of Tsavo: Exploring the Legacy of Africa's Notorious Man-Eaters. McGraw-Hill. ISBN 0-07-136333-5.
- Patterson, Bruce D.; Neiburger, Ellis J.; Kasiki, Samuel M. (February 2003). "Tooth Breakage and Dental Disease as Causes of Carnivore–Human Conflicts". Journal of Mammalogy 84 (1): 190–96. doi:10.1644/1545-1542(2003)084<0190:TBADDA>2.0.CO;2.
- Peterhans, Julian C. Kerbis; Thomas Patrick Gnoske (2001). "The Science of Man-eating". Journal of East African Natural History 90 (1&2): 1–40. doi:10.2982/0012-8317(2001)90[1:TSOMAL]2.0.CO;2. Archived from the original on 30 October 2007. Retrieved 7 July 2007.
- Packer, C.; Ikanda, D.; Kissui, B.; Kushnir, H. (August 2005). "Conservation biology: lion attacks on humans in Tanzania". Nature 436 (7053): 927–28. doi:10.1038/436927a. PMID 16107828.
- Packer, C; Swanson A; Ikanda D; Kushnir H (July 2011). Rands, Sean A., ed. "Fear of Darkness, the Full Moon and the Nocturnal Ecology of African Lions". PLoS ONE 6 (7): e22285. doi:10.1371/journal.pone.0022285.
- Frump, RR (2006). The Man-Eaters of Eden: Life and Death in Kruger National Park. The Lyons Press. ISBN 1-59228-892-8.
- Dickinson, Daniel (19 October 2004). "Toothache 'made lion eat humans'". BBC News. Retrieved 25 January 2011.
- Baldus, R (March 2006). "A man-eating lion (Panthera leo) from Tanzania with a toothache". European Journal of Wildlife Research 52 (1): 59–62. doi:10.1007/s10344-005-0008-0.
- Rushby, George G. (1965). No More the Tusker. London: W. H. Allen.
- Schaller, p. 29.
- Schaller, p. 174.
- Asdell, Sydney A. (1993) . Patterns of mammalian reproduction. Ithaca: Cornell University Press. ISBN 978-0-8014-1753-5.
- Schaller, p. 142.
- Scott, p. 45.
- Schaller, p. 143.
- Packer, C.; Pusey, A. E. (May 1983). "Adaptations of female lions to infanticide by incoming males". American Naturalist 121 (5): 716–28. doi:10.1086/284097.
- Macdonald, David (1984). The Encyclopedia of Mammals. New York: Facts on File. p. 31. ISBN 0-87196-871-1.
- Scott, p. 46.
- Crandall, Lee S. (1964). The management of wild animals in captivity. Chicago: University of Chicago Press. OCLC 557916.
- Scott, p. 68.
- Bagemihl, Bruce (1999). Biological Exuberance: Animal Homosexuality and Natural Diversity. New York: St. Martin's Press. pp. 302–05. ISBN 0-312-19239-8.
- Schaller, p. 183.
- Schaller, pp. 188–89.
- Schaller, pp. 189–90.
- Schaller, p. 184.
- Yeoman, Guy Henry; Walker, Jane Brotherton (1967). The ixodid ticks of Tanzania. London: Commonwealth Institute of Entomology. OCLC 955970.
- Sachs, R (1969). "Untersuchungen zur Artbestimmung und Differenzierung der Muskelfinnen ostafrikanischer Wildtiere [Differentiation and species determination of muscle-cysticerci in East African game animals]". Zeitschrift für tropenmedizin und Parasitologie (in German) 20 (1): 39–50. PMID 5393325.
- Fosbrooke, Henry (1963). "The stomoxys plague in Ngorongoro". East African Wildlife Journal 1 (6): 124–26. doi:10.1111/j.1365-2028.1963.tb00190.x.
- Nkwame, Valentine M (9 September 2006). "King of the jungle in jeopardy". The Arusha Times. Retrieved 4 September 2007.
- Roelke-Parker, M. E.; Munson, Linda; Packer, Craig; Kock, Richard; Cleaveland, Sarah; Carpenter, Margaret; O'Brien, Stephen J.; Pospischil, Andreas; Hofmann-Lehmann, Regina et al. (February 1996). "A canine distemper epidemic in Serengeti lions (Panthera leo)". Nature 379 (6564): 441–45. doi:10.1038/379441a0. PMID 8559247.
- Schaller, p. 85.
- Sparks, J (1967). "Allogrooming in primates:a review". In Desmond Morris. Primate Ethology. Chicago: Aldine. ISBN 0-297-74828-9. (2007 edition: 0-202-30826-X)
- Leyhausen, Paul (1960). Verhaltensstudien an Katzen (in German) (2nd ed.). Berlin: Paul Parey. ISBN 3-489-71836-4.
- Schaller, pp. 85–88.
- Schaller, pp. 88–91.
- Schaller, pp. 92–102.
- Ananthakrishnan, G.; Eklund, Robert; Peters, Gustav; Mabiza, Evans (2011). "An acoustic analysis of lion roars. II: Vocal tract characteristics". Speech, Music and Hearing Quarterly Progress and Status Report TMH-QPSR 51: 5.
- Eklund, Robert; Peters, Gustav;; Ananthakrishnan, G; Mabiza, Evans (2011). "An acoustic analysis of lion roars. I: Data collection and spectrogram and waveform analyses". Speech, Music and Hearing Quarterly Progress and Status Report TMH-QPSR 51: 1.
- Schaller, pp. 103–113.
- Miller, Brian (2000). Endangered animals: a reference guide to conflicting issues. Greenwood Publishing Group. ISBN 0-313-30816-0.
- Rudnai, Judith A. (1973). The social life of the lion. Wallingford: s.n. ISBN 0-85200-053-7.
- "The Gir—Floristic". Asiatic Lion Information Centre. Wildlife Conservation Trust of India. 2006. Retrieved 14 September 2007.
- Schaller, p. 5.
- Heptner, V. G.; A. A. Sludskii (1989). Mammals of the Soviet Union: Volume 1, Part 2: Carnivora (Hyaenas and Cats). New York: Amerind. ISBN 90-04-08876-8.
- "Past and present distribution of the lion in North Africa and Southwest Asia.". Asiatic Lion Information Centre. 2001. Archived from the original on 10 May 2008. Retrieved 1 June 2006.
- "Asiatic Lion—Population". Asiatic Lion Information Centre. Wildlife Conservation Trust of India. 2006. Retrieved 15 September 2007.
- Desai, Darshan (23 June 2003). "The Mane Don't Fit". Outlook India Magazine. Outlookindia.com. Retrieved 21 November 2010.
- Bauer, H.; Van Der Merwe, S. (2002). "The African lion database". Cat news 36: 41–53.
- Chardonnet, P. (2002). Conservation of African lion. Paris, France: International Foundation for the Conservation of Wildlife.
- "AWF Wildlife: Lion". African Wildlife Foundation. Retrieved 29 August 2007.
- "NATURE. The Vanishing Lions". PBS. Retrieved 25 January 2011.
- Breitenmoser, U., Mallon, D.P., Ahmad Khan, J. & Driscoll, C. (2008). "Panthera leo ssp. persica". IUCN Red List of Threatened Species. Version 2010.4. International Union for Conservation of Nature. Retrieved 13 December 2010.
- Tumenta, P. N.; Kok, J. S.; van Rijssel, J. C.; Buij, R.; Croes, B. M.; Funston, P. J.; de Iongh, H. H.; de Haes, H. A. Udo (2009). "Threat of rapid extermination of the lion (Panthera leo leo) in Waza National Park, Northern Cameroon". African Journal of Ecology 48 (4): 1–7. doi:10.1111/j.1365-2028.2009.01181.x.
- "Winning with lions". The East African Magazine.
- Roach, John (16 July 2003). "Lions Vs. Farmers: Peace Possible?". National Geographic News. National Geographic. Retrieved 1 September 2007.
- Saberwal, Vasant K.; Gibbs, James P.; Chellam, Ravi; Johnsingh, A. J. T. (1 June 1994). "Lion‐Human Conflict in the Gir Forest, India". Conservation Biology (Blackwell Science Inc) 8 (2): 501–7. doi:10.1046/j.1523-1739.1994.08020501.x. ISSN 1523-1739. Retrieved 1 May 2014.
- Johnsingh, A. J. T. (2004). "WII in the Field: Is Kuno Wildlife Sanctuary ready to play second home to Asiatic lions?". Wildlife Institute of India Newsletter 11 (4). Archived from the original on 27 September 2007. Retrieved 6 November 2010.
- "Barbary Lion News". Archived from the original on 17 December 2005. Retrieved 24 September 2007.
- de Courcy, p. 81
- de Courcy, p. 82
- Dollinger, P; Geser, S. "Lion: In the Zoo (subpage)". Visit the Zoo. WAZA (World Association of Zoos and Aquariums). Retrieved 5 April 2011.
- Aguiar, Eloise (August 2007). "Honolulu zoo's old lion roars no more". Honolulu Advertiser. Retrieved 4 September 2007.
- Frankham, Richard; Ballou, Jonathan; Briscoe, David (2009). Introduction to Conservation Genetics. Cambridge, England: Cambridge University Press. p. 437. ISBN 0-521-70271-2. Retrieved 12 September 2010.
- Avise, J. C.; Hamrick, J. L. (31 January 1996). Conservation Genetics. Springer Science & Business Media. p. 67. ISBN 9780412055812.
Furthermore, when Asiatic lions were inadvertently bred to African lion subspecies in North America, the fecundity, reproductive success, and spermatozoal development improved dramatically (Box 3.3; O'Brien et al., 1987b).
- Smith, Vincent Arthur (1924). The Early History of India. Oxford: Clarendon Press. p. 97.
- Wiedemann, Thomas (1995). Emperors and Gladiators. Routledge. p. 60. ISBN 0-415-12164-7.
- Baratay & Hardouin-Fugier, p. 17.
- Baratay & Hardouin-Fugier, pp. 19–21, 42.
- Baratay & Hardouin-Fugier, p. 20.
- Owen, James (3 November 2005). "Medieval Lion Skulls Reveal Secrets of Tower of London "Zoo"". National Geographic Magazine. National Geographic. Retrieved 5 September 2007.
- Blunt, p. 15.
- Baratay & Hardouin-Fugier, pp. 24–28.
- Blunt, p. 16.
- Blunt, p. 17.
- de Courcy, pp. 8–9.
- Blunt, p. 32.
- Baratay & Hardouin-Fugier, p. 122.
- Baratay & Hardouin-Fugier, pp. 114, 117.
- Baratay & Hardouin-Fugier, p. 113.
- Baratay & Hardouin-Fugier, pp. 173, 180–83.
- Blunt, p. 208.
- de Courcy, p. 69.
- Hone, William (2004) [1825–1826]. "July". In Kyle Grimes. The Every-Day Book. University of Alabama at Birmingham. p. 26. Archived from the original on 24 June 2008. Retrieved 25 January 2011.
- Blaisdell, Warren H. (November 1997). "How A Lion Fight Caused England To Stop The Breeding Of Both Ring And Pit Bulldogs". American Bulldog Review 3 (4). Archived from the original on 24 April 2008. Retrieved 5 September 2007.
- Baratay & Hardouin-Fugier, p. 187.
- Feldman, David (1993). How Does Aspirin Find a Headache?. HarperCollins. ISBN 0-06-016923-0.
- Garai, Jana (1973). The Book of Symbols. New York: Simon & Schuster. ISBN 0-671-21773-9.
- Aesop (2002). Gibbs, L, ed. Fables. Oxford World's Classics. Oxford: Oxford University Press. ISBN 0-19-284050-9.
- Martin Bailey Ice Age Lion Man is world’s earliest figurative sculpture The Art Newspaper, 31 January 2013, accessed 1 February 2013.
- Züchner, Christian (September 1998). Grotte Chauvet Archaeologically Dated. International Rock Art Congress IRAC ’98, Vila Real, Portugal. Retrieved 6 November 2010.
- Graves, R (1955). "The First Labour: The Nemean Lion". Greek Myths. London: Penguin. pp. 465–69. ISBN 0-14-001026-2.
- Cassin, Elena (1981). "Le roi et le lion" [The king & the lion] (PDF). Revue de l'histoire des religions (in French) 298 (198–4): 355–401. doi:10.3406/rhr.1981.4828. Retrieved 3 December 2009.
- Daniel 6
- "Bhag-P". Srimadbhagavatam. 1.3.18.
In the fourteenth incarnation, the Lord appeared as Nrisimha and bifurcated the strong body of the atheist Hiranyakasipu with His nails, just as a carpenter pierces cane.
- "Bhag-P". Srimadbhagavatam. 7.8.19–22. Retrieved 21 November 2010.
- Singh, Khushwant (2004). A History of the Sikhs: 1469–1838 I. Oxford University Press. ISBN 0-19-567308-5.
- "Know India: State Emblem". National Portal of India. National Informatics Centre. 2005. Archived from the original on 22 August 2007. Retrieved 27 August 2007.
- "National Flag". Government of Sri Lanka. Archived from the original on 27 March 2008. Retrieved 6 August 2007.
- "Article 6: The National Flag". Constitution. Government of Sri Lanka. 1978. Retrieved 6 August 2007.
- Ling, Li (May 2002). "The Two-Way Process in the Age of Globalization". Ex/Change (newsletter) (4). Egan, Ronald transl. (City University of Hong Kong). Archived from the original on 6 April 2005. Retrieved 26 September 2007.
- "Lion Dance Club". MIT.
- "Singapore". The American Heritage Dictionary of the English Language (4th ed.). Bartleby. 2000. Archived from the original on 29 June 2008. Retrieved 14 April 2006.
- "Early History". Singapore: Ministry of Information, Communications and the Arts. Archived from the original on 12 April 2010. Retrieved 6 November 2010.
- "Arms of Margaret Norrie McCain". The Public Register of Arms, Flags and Badges. CA. Retrieved 30 June 2010.
- "Heraldic Dictionary: Beasts". University of Notre Dame. Retrieved 20 July 2007.
- "Detroit Lions" (official Website). 2001. Retrieved 8 July 2007.
- "Chelsea centenary crest unveiled". BBC. 12 November 2004. Retrieved 2 January 2007.
- "The Aston Villa Crest: 2007 Onwards...". Aston Villa F.C. 2007. Archived from the original on 11 October 2007. Retrieved 6 August 2007.
- Lewis, Clive Staples (1950). The Lion, the Witch and the Wardrobe. HarperCollins. ISBN 0-06-023481-4.
- Baum, L. Frank; Hearn, Michael Patrick (1973). The Annotated Wizard of Oz. New York, NY: Clarkson N. Potter. p. 148. ISBN 0-517-50086-8.
- "Advertising Mascots—Animals—Leo the MGM Lion (MGM Studios)". TV Acres.
- Adamson, George (1969). Bwana Game: the life story of George Adamson. Fontana. ISBN 0-00-612145-4.
- Adamson, Joy (2000) . Born Free: A Lioness of Two Worlds. Pantheon. ISBN 0-375-71438-3.
- Schweizer, Peter (1998). Disney: The Mouse Betrayed. Washington D.C.: Regnery. pp. 164–69. ISBN 0-89526-387-4.
- "King Leonardo and His Short Subjects (1960– )". Internet Movie Database. 2007. Retrieved 14 September 2007.
- Baratay, Eric; Hardouin-Fugier, Elisabeth (2002). Zoo: a history of zoological gardens in the West. London: Reaktion Books. ISBN 1-86189-111-3.
- 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.
- Scott, Jonathon; Scott, Angela (2002). Big Cat Diary: Lion. Harper Collins. ISBN 0-00-714666-3.
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