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113-407: The lion ( Panthera leo ) is a large cat of the genus Panthera , native to Africa and India . It has a muscular, broad-chested body ; a short, rounded head; round ears; and a dark, hairy tuft at the tip of its tail. It is sexually dimorphic ; adult male lions are larger than females and have a prominent mane. It is a social species , forming groups called prides. A lion's pride consists of

226-536: A Mitochondrial Eve that lived in Africa 150,000 years ago provided early support for the Out-of-Africa model. While this study had its shortcomings, it received significant attention both within scientific circles and a wider audience. A more thorough phylogeographic analysis that used ten different genes instead of a single mitochondrial marker indicates that at least two major expansions out of Africa after

339-894: A generalist hypercarnivore and is considered to be both an apex and keystone predator due to its wide prey spectrum. Its prey consists mainly of medium-sized to large ungulates , particularly blue wildebeest , plains zebra , African buffalo , gemsbok and giraffe . It also frequently takes common warthog despite it being much smaller. In India, chital and sambar deer are the most common wild prey, while livestock contributes significantly to lion kills outside protected areas. It usually avoids fully grown adult elephants , rhinoceros and hippopotamus and small prey like dik-dik , hyraxes , hares and monkeys . Unusual prey include porcupines and small reptiles. Lions kill other predators but seldom consume them. Young lions first display stalking behaviour at around three months of age, although they do not participate in hunting until they are almost

452-415: A buildup of scent attracting the attention of predators, 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. Usually, the mother does not integrate herself and her cubs back into the pride until the cubs are six to eight weeks old. Sometimes the introduction to pride life occurs earlier, particularly if other lionesses have given birth at about

565-547: A closer relationship with P. leo melanochaita . Lion samples from some parts of the Ethiopian Highlands cluster genetically with those from Cameroon and Chad, while lions from other areas of Ethiopia cluster with samples from East Africa. Researchers, therefore, assume Ethiopia is a contact zone between the two subspecies. Genome -wide data of a wild-born historical lion sample from Sudan showed that it clustered with P. l. leo in mtDNA-based phylogenies, but with

678-467: A constant lookout for circling vultures, which indicate the death or distress of an animal. Most carrion on which both hyenas and lions feed upon are killed by hyenas rather than lions. Carrion is thought to provide a large part of lion diet. Lions and spotted hyenas occupy a similar ecological niche and compete for prey and carrion; a review of data across several studies indicates a dietary overlap of 58.6%. Lions typically ignore hyenas unless they are on

791-423: A favoured 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 fleeing from other lionesses. Males attached to prides do not usually participate in group hunting. Some evidence suggests, however, that males are just as successful as females; they are typically solo hunters who ambush prey in small bushland. They may join in

904-413: A few adult males, related females, and cubs. Groups of female lions usually hunt together, preying mostly on medium-sized and large ungulates . The lion is an apex and keystone predator ; although some lions scavenge when opportunities occur and have been known to hunt humans , lions typically do not actively seek out and prey on humans. The lion inhabits grasslands , savannahs , and shrublands . It

1017-464: A few reported cases of old and wounded lions falling prey to wild dogs. Most lionesses reproduce by the time they are four years of age. Lions do not mate at a specific time of year and the females are polyestrous . Like those of other cats, 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

1130-502: A half of the body length, although with some exceptions, like the Lynx species and margay ( Leopardus wiedii ). Cat species vary greatly in body and skull sizes, and weights: Most cat species have a haploid number of 18 or 19. Central and South American cats have a haploid number of 18, possibly due to the combination of two smaller chromosomes into a larger one. Felidae have type IIx muscle fibers three times more powerful than

1243-544: A heavier mane. On average, Asiatic lions have sparser manes than African lions. This feature likely evolved to signal the fitness of males to females. Males with darker manes appear to have greater reproductive success and are more likely to remain in a pride for longer. They have longer and thicker hair and higher testosterone levels, but they are also more vulnerable to heat stress. The core body temperature does apparently not increase regardless of sex, season, feeding time, length and colour of mane, but only surface temperature

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1356-597: A high affinity to P. l. melanochaita . This result suggested that the taxonomic position of lions in Central Africa may require revision. Other lion subspecies or sister species to the modern lion existed in prehistoric times: The Panthera lineage is estimated to have genetically diverged from the common ancestor of the Felidae around 9.32 to 4.47 million years ago to 11.75 to 0.97 million years ago . Results of analyses differ in

1469-499: A kill or are being harassed, while the latter tend to visibly react to the presence of lions with or without the presence of food. In the Ngorongoro crater , lions subsist largely on kills stolen from hyenas, causing them to increase their kill rate. In Botswana's Chobe National Park, the situation is reversed as hyenas there frequently challenge lions and steal their kills, obtaining food from 63% of all lion kills. When confronted on

1582-511: A kill, hyenas may either leave or wait patiently at a distance of 30–100 m (98–328 ft) until the lions have finished. Hyenas may feed alongside lions and force them off a kill. The two species attack one another even when there is no food involved for no apparent reason. Lions can account for up to 71% of hyena deaths in Etosha National Park . Hyenas have adapted by frequently mobbing lions that enter their home ranges. When

1695-513: A kill. Crippled lions and cubs may fall victim to hyenas and leopards or be trampled by buffalo or elephants. Careless lions may be maimed when hunting prey. Nile crocodiles may also kill and eat lions, evidenced by the occasional lion claw found in crocodile stomachs. Ticks commonly infest the ears, neck and groin regions of the lions. Adult forms of several tapeworm species of the genus Taenia have been isolated from lion intestines, having been ingested as larvae in antelope meat. Lions in

1808-472: A longer evolutionary history than genetically less diverse lion samples from Asia and West and Central Africa. A whole genome-wide sequence of lion samples showed that samples from West Africa shared alleles with samples from Southern Africa, and samples from Central Africa shared alleles with samples from Asia. This phenomenon indicates that Central Africa was a melting pot of lion populations after they had become isolated, possibly migrating through corridors in

1921-400: A loose pride while females associate with up to 12 other females, forming a stronger pride together with their cubs. Female and male lions associate only when mating. Coalitions of males hold territory for a longer time than single lions. Males in coalitions of three or four individuals exhibit a pronounced hierarchy, in which one male dominates the others and mates more frequently. The lion is

2034-430: A nomad is a "range". Males associated with a pride patrol the fringes. Both males and females 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 defense against intruders, while others lag behind. Lions tend to assume specific roles in the pride; slower-moving individuals may provide other valuable services to

2147-552: A nomadic phase before gaining residence in a pride. A study undertaken in the Serengeti National Park revealed that nomadic coalitions gain residency at between 3.5 and 7.3 years of age. In Kruger National Park, dispersing male lions move more than 25 km (16 mi) away from their natal pride in search of their own territory. Female lions stay closer to their natal pride. Therefore, female lions in an area are more closely related to each other than male lions in

2260-475: A phylogeographic approach will likely play a key role in understanding the vectors and spread of avian influenza ( HPAI H5N1 ), demonstrating the relevance of phylogeography to the general public. Phylogeographic analysis of ancient and modern languages has been used to test whether Indo-European languages originated in Anatolia or in the steppes of Central Asia. Language evolution was modeled in terms of

2373-423: A reed-bed, a cave, or some other sheltered area, usually away from the pride. She will often hunt alone while the cubs are still helpless, staying relatively close to the den. Lion cubs are born blind, their eyes opening around seven days 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. To avoid

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2486-565: A reversal in climate change allows for rapid migration out of refugial areas, these species spread rapidly into newly available habitat. A number of empirical studies find genetic signatures of both animal and plant species that support this scenario of refugia and postglacial expansion. This has occurred both in the tropics (where the main effect of glaciation is increasing aridity , i.e. the expansion of savanna and retraction of tropical rainforest ) as well as temperate regions that were directly influenced by glaciers. Phylogeography can help in

2599-562: A short, rounded head, a reduced neck, and round ears; males have broader heads. The fur varies in colour from light buff to silvery grey, yellowish red, and dark brown. The colours of the underparts are generally lighter. A new-born lion has dark spots , which fade as the cub reaches adulthood, although faint spots may still be seen on the legs and underparts. The tail of all lions ends in a dark, hairy tuft that, in some lions, conceals an approximately 5 mm (0.20 in)-long, hard "spine" or "spur" composed of dermal papillae. The functions of

2712-426: A species' biogeography /biogeographical past sets phylogeography apart from classical population genetics and phylogenetics . Past events that can be inferred include population expansion, population bottlenecks , vicariance , dispersal, and migration . Recently developed approaches integrating coalescent theory or the genealogical history of alleles and distributional information can more accurately address

2825-553: A very characteristic manner starting with a few deep, long roars that subside into grunts. The lion is listed as Vulnerable on the IUCN Red List . The Indian population is listed on CITES Appendix I and the African population on CITES Appendix II . Felidae Felidae ( / ˈ f ɛ l ɪ d iː / ) is the family of mammals in the order Carnivora colloquially referred to as cats . A member of this family

2938-472: A vicariance event affected multiple taxa during the Pleistocene or Pliocene . Phylogeography also gives an important historical perspective on community composition. History is relevant to regional and local diversity in two ways. One, the size and makeup of the regional species pool results from the balance of speciation and extinction . Two, at a local level community composition is influenced by

3051-425: A year old and begin to hunt effectively when nearing the age of two. Single lions are capable of bringing down zebra and wildebeest, while larger prey like buffalo and giraffe are riskier. In Chobe National Park , large prides have been observed hunting African bush elephants up to around 15 years old in exceptional cases, with the victims being calves, juveniles, and even subadults. In typical hunts, each lioness has

3164-399: Is in heat . Lions of both sexes may be involved in group homosexual and courtship activities. Males will also head-rub and roll around with each other before mounting each other. Generation length of the lion is about seven years. The average gestation period is around 110   days; the female gives birth to a litter of between one and four cubs in a secluded den, which may be a thicket,

3277-548: Is a mixture of dry savannah forest and very dry, deciduous scrub forest . In Africa, the range of the lion originally spanned most of the central African rainforest zone and the Sahara desert. In the 1960s, it became extinct in North Africa, except in the southern part of Sudan. During the mid- Holocene , around 8,000-6,000 years ago, the range of lions expanded into Southeastern and Eastern Europe, partially re-occupying

3390-946: Is affected. Unlike in other felid species, female lions consistently interact with multiple males at once. Another hypothesis suggests that the mane also serves to protect the neck in fights, but this is disputed. During fights, including those involving maneless females and adolescents, the neck is not targeted as much as the face, back, and hindquarters. Injured lions also begin to lose their manes. Almost all male lions in Pendjari National Park are either maneless or have very short manes. Maneless lions have also been reported in Senegal, in Sudan 's Dinder National Park and in Tsavo East National Park , Kenya. Castrated lions often have little to no mane because

3503-525: Is also called a felid ( / ˈ f iː l ɪ d / ). The 41 extant Felidae species exhibit the greatest diversity in fur patterns of all terrestrial carnivores. Cats have retractile claws , slender muscular bodies and strong flexible forelimbs. Their teeth and facial muscles allow for a powerful bite. They are all obligate carnivores , and most are solitary predators ambushing or stalking their prey. Wild cats occur in Africa , Europe , Asia and

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3616-414: Is distributed and is found (illustrated in the inset maps below, including Amazon basin, Andes, Guiana-Venezuela, Central America-Chocó). The combination of techniques used in this study exemplifies more generally how phylogeographic studies proceed and test for patterns of common influence. Paleogeographic data establishes geological time records for historical events that explain the branching patterns in

3729-656: Is generally mutual and the recipient appears to express pleasure. The head and neck are the most common parts of the body licked; this behaviour may have arisen out of utility because lions cannot lick these areas themselves. Lions have an array of facial expressions and body postures that serve as visual gestures. A common facial expression is the "grimace face" or flehmen response , which a lion makes when sniffing chemical signals and involves an open mouth with bared teeth, raised muzzle, wrinkled nose, closed eyes and relaxed ears. Lions also use chemical and visual marking; males spray urine and scrape plots of ground and objects within

3842-430: Is integrated into phylogeographic analyses. Phylogeography takes a population genetics and phylogenetic perspective on biogeography . In the mid-1970s, population genetic analyses turned to mitochondrial markers. The advent of the polymerase chain reaction (PCR), the process where millions of copies of a DNA segment can be replicated, was crucial in the development of phylogeography. Thanks to this breakthrough,

3955-476: Is no evidence for gene flow between the two lineages, indicating that they did not share the same geographic area. The Eurasian and American cave lions became extinct at the end of the last glacial period without mitochondrial descendants on other continents. The modern lion was probably widely distributed in Africa during the Middle Pleistocene and started to diverge in sub-Saharan Africa during

4068-429: Is reconstructed using phylogenetic trees derived from molecular data. The molecular trees are mapped in relation to paleogeographic history of the region for a complete phylogeographic study. The tree shown in the center of the figure has its branch lengths calibrated to a molecular clock , with the geological time bar shown at the bottom. The same phylogenetic tree is duplicated four more times to show where each lineage

4181-644: Is referred to as the Out-of-Africa with replacement model, which contends that the last expansion out of Africa around 100,000 years ago resulted in the modern humans displacing all previous Homo spp. populations in Eurasia that were the result of an earlier wave of emigration out of Africa. The multiregional scenario claims that individuals from the recent expansion out of Africa intermingled genetically with those human populations of more ancient African emigrations. A phylogeographic study that uncovered

4294-411: Is short and powerful; it attempts to catch prey with a fast rush and final leap, usually pulls it down by the rump, and kills with a clamping bite to the throat or muzzle . It can hold the prey's throat for up to 13 minutes, until the prey stops moving. It has a bite force from 1593.8 to 1768 Newtons at the canine tip and up 4167.6 Newtons at the carnassial notch. Lions typically consume prey at

4407-518: Is shown in panels A-D) the authors of this study proposed a null-hypothesis that assumes no spatial structure and two alternative hypothesis involving dispersal and other biogeographic constraints (hypothesis are shown in panels E-G, listed as SMO, SM1, and SM2). The phylogeographers visited the ranges of each frog species to obtain tissue samples for genetic analysis; researchers can also obtain tissue samples from museum collections. The evolutionary history and relations among different poison frog species

4520-585: Is the Tsavo lion pride that always has just one adult male. Prides act as fission–fusion societies , and members will split into subgroups that keep in contact with roars . Nomadic lions range widely and move around sporadically, either in pairs or alone. Pairs are more frequent among related males. A lion may switch lifestyles; nomads can become residents and vice versa. Interactions between prides and nomads tend to be hostile, although pride females in estrus allow nomadic males to approach them. Males spend years in

4633-611: Is the most social of all wild felid species, living in groups of related individuals with their offspring. Such a group is called a " pride ". Groups of male lions are called "coalitions". Females form the stable social unit in a pride and do not tolerate outside females. The majority of females remain in their birth prides while all males and some females will disperse . The average pride consists of around 15 lions, including several adult females and up to four males and their cubs of both sexes. Large prides, consisting of up to 30 individuals, have been observed. The sole exception to this pattern

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4746-785: Is the oldest known cat that occurred after the Eocene–Oligocene extinction event about 33.9  million years ago ; fossil remains were excavated in France and Mongolia's Hsanda Gol Formation . Fossil occurrences indicate that the Felidae arrived in North America around 18.5  million years ago . This is about 20   million years later than the Ursidae and the Nimravidae , and about 10 million years later than

4859-609: Is the oldest known pantherine cat dated to the late Messinian to early Zanclean ages about 5.95 to 4.1 million years ago . A fossil skull was excavated in 2010 in Zanda County on the Tibetan Plateau . Panthera palaeosinensis from North China probably dates to the Late Miocene or Early Pliocene . The skull of the holotype is similar to that of a lion or leopard. Panthera zdanskyi dates to

4972-476: Is typically brownish and tinged with yellow, rust, and black hairs. Mutations in the genes microphthalmia-associated transcription factor and tyrosinase are possibly responsible for the colour of manes. It starts growing when lions enter adolescence, when testosterone levels increase, and reach their full size at around four years old. Cool ambient temperatures in European and North American zoos may result in

5085-424: Is unable to consume all of the kill, it rests for a few hours before continuing to eat. On hot days, the pride retreats to shade with one or two males standing guard. Lions defend their kills from scavengers such as vultures and hyenas. Lions scavenge on carrion when the opportunity arises, scavenging animals dead from natural causes such as disease or those that were killed by other predators. Scavenging lions keep

5198-546: Is usually more diurnal than other wild cats, but when persecuted, it adapts to being active at night and at twilight . During the Neolithic period, the lion ranged throughout Africa and Eurasia, from Southeast Europe to India, but it has been reduced to fragmented populations in sub-Saharan Africa and one population in western India . It has been listed as Vulnerable on the IUCN Red List since 1996 because populations in African countries have declined by about 43% since

5311-577: The Americas . Some wild cat species are adapted to forest and savanna habitats , some to arid environments, and a few also to wetlands and mountainous terrain. Their activity patterns range from nocturnal and crepuscular to diurnal , depending on their preferred prey species. Reginald Innes Pocock divided the extant Felidae into three subfamilies: the Pantherinae , the Felinae and

5424-591: The Canidae . In the Early Miocene about 20 to 16.6 million years ago , Pseudaelurus lived in Africa. Its fossil jaws were also excavated in geological formations of Europe's Vallesian , Asia's Middle Miocene and North America's late Hemingfordian to late Barstovian epochs. In the Early or Middle Miocene, the saber-toothed Machairodontinae evolved in Africa and migrated northwards in

5537-583: The Gelasian about 2.55 to 2.16 million years ago . Several fossil skulls and jawbones were excavated in northwestern China. Panthera gombaszoegensis is the earliest known pantherine cat that lived in Europe about 1.95 to 1.77 million years ago . Living felids fall into eight evolutionary lineages or species clades . Genotyping of the nuclear DNA of all 41 felid species revealed that hybridization between species occurred in

5650-772: The Late Miocene . With their large upper canines , they were adapted to prey on large-bodied megaherbivores . Miomachairodus is the oldest known member of this subfamily. Metailurus lived in Africa and Eurasia about 8 to 6 million years ago . Several Paramachaerodus skeletons were found in Spain. Homotherium appeared in Africa, Eurasia and North America around 3.5  million years ago , and Megantereon about 3  million years ago . Smilodon lived in North and South America from about 2.5  million years ago . This subfamily became extinct in

5763-607: The Late Pleistocene . Results of mitochondrial analysis indicate that the living Felidae species descended from a common ancestor , which originated in Asia in the Late Miocene epoch. They migrated to Africa, Europe and the Americas in the course of at least 10 migration waves during the past ~11 million years. Low sea levels and interglacial and glacial periods facilitated these migrations. Panthera blytheae

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5876-509: The Narmada River in central India. Lions spend much of their time resting; they are inactive for about twenty hours per day. Although lions can be active at any time, their activity generally peaks after dusk with a period of socialising, grooming, and defecating. Intermittent bursts of activity continue until dawn, when hunting most often takes place. They spend an average of two hours a day walking and fifty minutes eating. The lion

5989-478: The Nile Basin during the early Holocene . In zoos, lions have been bred with tigers to create hybrids for the curiosity of visitors or for scientific purpose. The liger is bigger than a lion and a tiger, whereas most tigons are relatively small compared to their parents because of reciprocal gene effects. The leopon is a hybrid between a lion and leopard. The lion is a muscular, broad-chested cat with

6102-458: The snow leopard ( Panthera uncia ) and the Pallas's cat ( Otocolobus manul ). Those living in tropical and hot climate zones have short fur. Several species exhibit melanism with all-black individuals, cougars are notable for lacking melanism but leucism and albinism are present in cougars along with many other felids. In the great majority of cat species, the tail is between a third and

6215-475: The 1000 genomes project, genomic-scale SNP databases sampling thousands of individuals globally and samples taken from two non-Homo sapiens hominins (Neanderthals and Denisovans), the picture of human evolutionary has become more resolved and complex involving possible Neanderthal and Denisovan admixture, admixture with archaic African hominins, and Eurasian expansion into the Australasian region that predates

6328-530: The Acinonychinae, differing from each other by the ossification of the hyoid apparatus and by the cutaneous sheaths which protect their claws. This concept has been revised following developments in molecular biology and techniques for the analysis of morphological data. Today, the living Felidae are divided into two subfamilies: the Pantherinae and Felinae, with the Acinonychinae subsumed into

6441-771: The Asiatic lion population. In 2017, the Cat Classification Task Force of the Cat Specialist Group revised lion taxonomy , and recognises two subspecies based on results of several phylogeographic studies on lion evolution , namely: However, there seems to be some degree of overlap between both groups in northern Central Africa. DNA analysis from a more recent study indicates that Central African lions are derived from both northern and southern lions, as they cluster with P. leo leo in mtDNA-based phylogenies whereas their genomic DNA indicates

6554-627: The Late Pleistocene. Lion populations in East and Southern Africa became separated from populations in West and North Africa when the equatorial rainforest expanded 183,500 to 81,800 years ago. They shared a common ancestor probably between 98,000 and 52,000 years ago. Due to the expansion of the Sahara between 83,100 and 26,600 years ago, lion populations in West and North Africa became separated. As

6667-564: The Ngorongoro Crater were afflicted by an outbreak of stable fly ( Stomoxys calcitrans ) in 1962, resulting in lions becoming emaciated and covered in bloody, bare patches. Lions sought unsuccessfully to evade the biting flies by climbing trees or crawling into hyena burrows; many died or migrated and the local population dropped from 70 to 15 individuals. A more recent outbreak in 2001 killed six lions. Captive lions have been infected with canine distemper virus (CDV) since at least

6780-694: The adjacent Timbavati Private Game Reserve in eastern South Africa. They were removed from the wild in the 1970s, thus decreasing the white lion gene pool . Nevertheless, 17 births have been recorded in five prides between 2007 and 2015. White lions are selected for breeding in captivity. They have reportedly been bred in camps in South Africa for use as trophies to be killed during canned hunts . African lions live in scattered populations across sub-Saharan Africa. The lion prefers grassy plains and savannahs , scrub bordering rivers, and open woodlands with bushes. It rarely enters closed forests. On Mount Elgon ,

6893-420: The age of two. When a new male lion takes over a pride, adolescents both male and female may be evicted. Lions may live 12–17 years in the wild. Although adult lions have no natural predators, evidence suggests most die violently from attacks by humans or other lions. Lions often inflict serious injuries on members of other prides they encounter in territorial disputes or members of the home pride when fighting at

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7006-540: 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 the role of allogrooming among primates. Head rubbing, nuzzling the forehead, face and neck against another lion appears to be a form of greeting and 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

7119-495: The attack. They take advantage of factors that reduce visibility; many kills take place near some form of cover or at night. One study in 2018 recorded a lion running at a top speed of 74.1 km/h (46.0 mph). The lion accelerates at the start of the chase by 9.5 m/s² , whereas zebras, wildebeest and Thomson's gazelle accelerate by 5 m/s², 5.6 m/s² and 4.5 m/s², respectively; acceleration appears to be more important than steady displacement speed in lion hunts. The lion's attack

7232-480: The authors found that hidden within what was thought to be a single, widely distributed species, an ancient and previously undetected species was also present. Conservation decisions can now be made to ensure that both lineages received protection. Results like this are not an uncommon outcome from phylogeographic studies. An analysis of salamanders of the genus Eurycea , also in the Appalachians, found that

7345-579: The biogeographic distribution of organisms in this area, including the isolation and reconnection of South America , the uplift of the Andes, an extensive Amazonian floodbasin system during the Miocene, the formation of Orinoco and Amazon drainages , and dry−wet climate cycles throughout the Pliocene to Pleistocene epochs. Using this contextual paleogeographic information (paleogeographic time series

7458-433: The course of evolution within the majority of the eight lineages. Modelling of felid coat pattern transformations revealed that nearly all patterns evolved from small spots. Traditionally, five subfamilies had been distinguished within the Felidae based on phenotypical features: the Pantherinae , the Felinae , the Acinonychinae, and the extinct Machairodontinae and Proailurinae . Acinonychinae used to only contain

7571-410: The cubs varies—one male could patiently let the cubs play with his tail or his mane, while another may snarl and bat the cubs away. Pride lionesses often synchronise their reproductive cycles and communal rearing and suckling of the young, which suckle indiscriminately from any or all of the nursing females in the pride. The synchronisation of births is advantageous because the cubs grow to being roughly

7684-574: The current taxonomy of the group greatly underestimated species level diversity. The authors of this study also found that patterns of phylogeographic diversity were more associated with historical (rather than modern) drainage connections, indicating that major shifts in the drainage patterns of the region played an important role in the generation of diversity of these salamanders. A thorough understanding of phylogeographic structure will thus allow informed choices in prioritizing areas for conservation. The field of comparative phylogeography seeks to explain

7797-448: The early 1990s. Lion populations are untenable outside designated protected areas. Although the cause of the decline is not fully understood, habitat loss and conflicts with humans are the greatest causes for concern. One of the most widely recognised animal symbols in human culture, the lion has been extensively depicted in sculptures and paintings, on national flags, and in literature and films. Lions have been kept in menageries since

7910-508: The following cladogram: Sunda clouded leopard (N. diardi) Clouded leopard ( N. nebulosa ) Tiger (P. tigris) Snow leopard (P. uncia) Jaguar (P. onca) Lion ( P. leo ) Leopard ( P. pardus ) Serval (L. serval) African golden cat ( C. aurata ) Caracal ( C. caracal ) Andean mountain cat ( L. jacobita ) Phylogeographic This term was introduced to describe geographically structured genetic signals within and among species . An explicit focus on

8023-524: The gamut from white to black, and fur patterns from distinctive small spots, and stripes to small blotches and rosettes . Most cat species are born with spotted fur, except the jaguarundi ( Herpailurus yagouaroundi ), Asian golden cat ( Catopuma temminckii ) and caracal ( Caracal caracal ). The spotted fur of lion ( Panthera leo ), cheetah (Acinonyx jubatus) and cougar ( Puma concolor ) cubs change to uniform fur during their ontogeny . Those living in cold environments have thick fur with long hair, like

8136-866: The genus Acinonyx but this genus is now within the Felinae subfamily. The following cladogram based on Piras et al. (2013) depicts the phylogeny of basal living and extinct groups. † Proailurus bourbonnensis † Proailurus lemanensis † Proailurus major † Pseudaelurus quadridentatus † Pseudaelurus cuspidatus † Pseudaelurus guangheesis † Machairodontinae [REDACTED] † Hyperailurictis intrepidus † Hyperailurictis marshi † Hyperailurictis stouti † Hyperailurictis validus † Hyperailurictis skinneri † Sivaelurus chinjiensis † Styriofelis turnauensis † Styriofelis romieviensis Felinae [REDACTED] † Miopanthera lorteti † Miopanthera pamiri Pantherinae [REDACTED] The phylogenetic relationships of living felids are shown in

8249-530: The group. Alternatively, there may be rewards associated with being a leader that fends off intruders; the rank of lionesses in the pride is reflected in these responses. The male or males associated with the pride must defend their relationship with the pride from outside males who may attempt to usurp them. Dominance hierarchies do not appear to exist among individuals of either sex in a pride. Asiatic lion prides differ in group composition. Male Asiatic lions are solitary or associate with up to three males, forming

8362-568: The hunting of large, slower-moving prey like buffalo; and even hunt them on their own. Moderately-sized hunting groups generally have higher success rates than lone females and larger groups. Lions are not particularly known for their stamina. For instance, a lioness's heart comprises only 0.57% of her body weight and a male's is about 0.45% of his body weight, whereas a hyena's heart comprises almost 1% of its body weight. Thus, lions run quickly only in short bursts at about 48–59 km/h (30–37 mph) and need to be close to their prey before starting

8475-454: The information contained in mitochondrial DNA sequences was much more accessible. Advances in both laboratory methods (e.g. capillary DNA sequencing technology) that allowed easier sequencing of DNA and computational methods that make better use of the data (e.g. employing coalescent theory ) have helped improve phylogeographic inference. By 2000, Avise generated a seminal review of the topic in book form, in which he defined phylogeography as

8588-471: The initial range extension of Homo erectus played an important role shaping the modern human gene pool and that recurrent genetic exchange is pervasive. These findings strongly demonstrated Africa's central role in the evolution of modern humans, but also indicated that the multiregional model had some validity. These studies have largely been supplanted by population genomic studies that use orders of magnitude more data. In light of these recent data from

8701-487: The interaction between local extinction of species’ populations and recolonization. A comparative phylogenetic approach in the Australian Wet Tropics indicates that regional patterns of species distribution and diversity are largely determined by local extinctions and subsequent recolonizations corresponding to climatic cycles. Phylogeography integrates biogeography and genetics to study in greater detail

8814-495: The latter. Pantherinae includes five Panthera and two Neofelis species , while Felinae includes the other 34 species in 12 genera . The first cats emerged during the Oligocene about 25 million years ago , with the appearance of Proailurus and Pseudaelurus . The latter species complex was ancestral to two main lines of felids: the cats in the extant subfamilies and a group of extinct "saber-tooth" felids of

8927-467: The lineal history of a species in context of the geoclimatic history of the planet. An example study of poison frogs living in the South American neotropics (illustrated to the left) is used to demonstrate how phylogeographers combine genetics and paleogeography to piece together the ecological history of organisms in their environments. Several major geoclimatic events have greatly influenced

9040-719: The lion has been recorded up to an elevation of 3,600 m (11,800 ft) and close to the snow line on Mount Kenya . Savannahs with an annual rainfall of 300 to 1,500 mm (12 to 59 in) make up the majority of lion habitat in Africa, estimated at 3,390,821 km (1,309,203 sq mi) at most, but remnant populations are also present in tropical moist forests in West Africa and montane forests in East Africa. The Asiatic lion now survives only in and around Gir National Park in Gujarat, western India. Its habitat

9153-805: The lion population in Kenya's Masai Mara National Reserve declined, the spotted hyena population increased rapidly. Lions tend to dominate cheetahs and leopards, steal their kills and kill their cubs and even adults when given the chance. Cheetahs often lose their kills to lions or other predators. A study in the Serengeti ecosystem revealed that lions killed at least 17 of 125 cheetah cubs born between 1987 and 1990. Cheetahs avoid their competitors by hunting at different times and habitats. Leopards, by contrast, do not appear to be motivated by an avoidance of lions, as they use heavy vegetation regardless of whether lions are present in an area and both cats are active around

9266-546: The location of the hunt but sometimes drag large prey into cover. They tend to squabble over kills, particularly the males. Cubs suffer most when food is scarce but otherwise all pride members eat their fill, including old and crippled lions, which can live on leftovers. Large kills are shared more widely among pride members. An adult lioness requires an average of about 5 kg (11 lb) of meat per day while males require about 7 kg (15 lb). Lions gorge themselves and eat up to 30 kg (66 lb) in one session. If it

9379-532: The mechanisms responsible for the phylogenetic relationships and distribution of different species. For example, comparisons across multiple taxa can clarify the histories of biogeographical regions. For example, phylogeographic analyses of terrestrial vertebrates on the Baja California peninsula and marine fish on both the Pacific and gulf sides of the peninsula display genetic signatures that suggest

9492-544: The mid-18th and mid-20th centuries, 26 lion specimens were described and proposed as subspecies, of which 11 were recognised as valid in 2005. They were distinguished mostly by the size and colour of their manes and skins. In the 19th and 20th centuries, several lion type specimens were described and proposed as subspecies , with about a dozen recognised as valid taxa until 2017. Between 2008 and 2016, IUCN Red List assessors used only two subspecific names: P. l. leo for African lion populations, and P. l. persica for

9605-487: The mid-1970s. CDV is spread by 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 . Feline immunodeficiency virus and lentivirus also affect captive lions. When resting, lion socialisation occurs through a number of behaviours;

9718-466: The molecular trees. This study rejected the null model and found that the origin for all extant Amazonian poison frog species primarily stem from fourteen lineages that dispersed into their respective areas after the Miocene floodbasin receded. Regionally based phylogeographic studies of this type are repeated for different species as a means of independent testing. Phylogeographers find broadly concordant and repeated patterns among species in most regions of

9831-544: The muscle fibers of human athletes. The family Felidae is part of the Feliformia , a suborder that diverged probably about 50.6 to 35 million years ago into several families. The Felidae and the Asiatic linsangs are considered a sister group , which split about 35.2 to 31.9 million years ago . The earliest cats probably appeared about 35 to 28.5 million years ago . Proailurus

9944-460: The past and current distribution of species. As part of historical biogeography, researchers had been evaluating the geographical and evolutionary relationships of organisms years before. Two developments during the 1960s and 1970s were particularly important in laying the groundwork for modern phylogeography; the first was the spread of cladistic thought, and the second was the development of plate tectonics theory . The resulting school of thought

10057-430: The phylogenetic relationship of the lion; it was thought to form a sister group with the jaguar that diverged 3.46 to 1.22 million years ago , but also with the leopard that diverged 3.1 to 1.95 million years ago to 4.32 to 0.02 million years ago . Hybridisation between lion and snow leopard ancestors possibly continued until about 2.1 million years ago. The lion-leopard clade

10170-409: The planet that is due to a common influence of paleoclimatic history. Phylogeography has also proven to be useful in understanding the origin and dispersal patterns of our own species, Homo sapiens . Based primarily on observations of skeletal remains of ancient human remains and estimations of their age, anthropologists proposed two competing hypotheses about human origins. The first hypothesis

10283-610: The previous males associated with a pride, the victors often kill any existing young cubs , perhaps because females do not become fertile and receptive until their cubs mature or die. Females often fiercely defend their cubs from a usurping male but are rarely successful unless a group of three or four mothers within a pride join forces against the male. Cubs also die from starvation and abandonment, and predation by leopards, hyenas and wild dogs. Male cubs are excluded from their maternal pride when they reach maturity at around two or three years of age, while some females may leave when they reach

10396-644: The prioritization of areas of high value for conservation. Phylogeographic analyses have also played an important role in defining evolutionary significant units (ESU), a unit of conservation below the species level that is often defined on unique geographic distribution and mitochondrial genetic patterns. A recent study on imperiled cave crayfish in the Appalachian Mountains of eastern North America demonstrates how phylogenetic analyses along with geographic distribution can aid in recognizing conservation priorities. Using phylogeographical approaches,

10509-537: The rainforest decreased and thus gave rise to more open habitats, lions moved from West to Central Africa. Lions from North Africa dispersed to southern Europe and Asia between 38,800 and 8,300 years ago. Extinction of lions in southern Europe, North Africa and the Middle East interrupted gene flow between lion populations in Asia and Africa. Genetic evidence revealed numerous mutations in lion samples from East and Southern Africa, which indicates that this group has

10622-452: The range of the now extinct cave lion . In Hungary , the modern lion was present from about 4,500 to 3,200 years Before Present . In Ukraine , the modern lion was present from about 6,400 to 2,000 years Before Present. In Greece, it was common, as reported by Herodotus in 480 BC; it was considered rare by 300 BC and extirpated by AD 100. In Asia the lion once ranged in regions where climatic conditions supported an abundance of prey. It

10735-405: The relative roles of these different historical forces in shaping current patterns. The term phylogeography was first used by John Avise in his 1987 work Intraspecific Phylogeography: The Mitochondrial DNA Bridge Between Population Genetics and Systematics . Historical biogeography is a synthetic discipline that addresses how historical, geological, climatic and ecological conditions influenced

10848-484: The removal of the gonads inhibits testosterone production. Rarely, both wild and captive lionesses have manes. Increased testosterone may be the cause of maned lionesses reported in northern Botswana. The white lion is a rare morph with a genetic condition called leucism , which is caused by a double recessive allele . It is not albino; it has normal pigmentation in the eyes and skin. White lions have occasionally been encountered in and around Kruger National Park and

10961-426: The same area. The evolution of sociability in lions was likely driven both by high population density and the clumped resources of savannah habitats. The larger the pride, the more high-quality territory they can defend; "hotspots" being near river confluences , where the cats have better access to water, prey and shelter (via vegetation). The area occupied by a pride is called a "pride area" whereas that occupied by

11074-417: The same size and have an equal chance of survival, and sucklings are not dominated by older cubs. Weaning occurs after six or seven months. Male lions reach maturity at about three years of age and at four to five years are capable of challenging and displacing adult males associated with another pride. They begin to age and weaken at between 10 and 15 years of age at the latest. When one or more new males oust

11187-403: The same time of day. In addition, there is no evidence that lions affect leopard abundance. Leopards take refuge in trees, though lionesses occasionally attempt to climb up and retrieve their kills. Lions similarly dominate African wild dogs , taking their kills and dispatching pups or adult dogs. Population densities of wild dogs are low in areas where lions are more abundant. However, there are

11300-417: The same time. When first introduced to the rest of the pride, lion cubs lack confidence when confronted with adults 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. Male tolerance of

11413-421: The spur are unknown. The tuft is absent at birth and develops at around 5 + 1 ⁄ 2 months of age. It is readily identifiable at the age of seven months. Its skull is very similar to that of the tiger, although the frontal region is usually more depressed and flattened and has a slightly shorter postorbital region and broader nasal openings than those of the tiger. Due to the amount of skull variation in

11526-443: The standard out of African expansion. Viruses are informative in understanding the dynamics of evolutionary change due to their rapid mutation rate and fast generation time. Phylogeography is a useful tool in understanding the origins and distributions of different viral strains. A phylogeographic approach has been taken for many diseases that threaten human health, including dengue fever , rabies , influenza and HIV . Similarly,

11639-444: The study of the "principles and processes governing the geographic distributions of genealogical lineages... within and among closely related species." Early phylogeographic work has recently been criticized for its narrative nature and lack of statistical rigor (i.e. it did not statistically test alternative hypotheses). The only real method was Alan Templeton 's Nested Clade Analysis, which made use of an inference key to determine

11752-638: The subfamily Machairodontinae , which range from the type genus Machairodus of the late Miocene to Smilodon of the Pleistocene. The "false saber-toothed cats", the Barbourofelidae and Nimravidae , are not true cats but are closely related. Together with the Felidae, Viverridae , hyenas and mongooses , they constitute the Feliformia . All members of the cat family have the following characteristics in common: The colour, length and density of their fur are very diverse. Fur colour covers

11865-453: The territory. The lion's repertoire of vocalisations is large; variations in intensity and pitch appear to be central to communication. Most lion vocalisations are variations of growling , snarling , meowing and roaring. Other sounds produced include puffing, bleating and humming. Roaring is used to advertise its presence. Lions most often roar at night, a sound that can be heard from a distance of 8 kilometres (5 mi). They tend to roar in

11978-402: The tiger in size. The size and weight of adult lions vary across its range and habitats. Accounts of a few individuals that were larger than average exist from Africa and India. The male lion's mane is the most recognisable feature of the species. It may have evolved around 320,000–190,000 years ago. It grows downwards and backwards, covering most of the head, neck, shoulders, and chest. The mane

12091-585: The time of the Roman Empire and have been a key species sought for exhibition in zoological gardens across the world since the late 18th century. Cultural depictions of lions were prominent in Ancient Egypt , and depictions have occurred in virtually all ancient and medieval cultures in the lion's historic and current range. The English word lion is derived via Anglo-Norman liun from Latin leōnem (nominative: leō ), which in turn

12204-401: The two species, usually only the structure of the lower jaw can be used as a reliable indicator of species. The skeletal muscles of the lion make up 58.8% of its body weight and represent the highest percentage of muscles among mammals. The lion has a high concentration of fast twitch muscle fibers, giving them quick bursts of speed but less stamina. Among felids, the lion is second only to

12317-475: The validity of a given process in explaining the concordance between geographic distance and genetic relatedness. Recent approaches have taken a stronger statistical approach to phylogeography than was done initially. Example Climate change, such as the glaciation cycles of the past 2.4 million years, has periodically restricted some species into disjunct refugia. These restricted ranges may result in population bottlenecks that reduce genetic variation. Once

12430-443: Was a borrowing from Ancient Greek λέων léōn . The Hebrew word לָבִיא lavi may also be related. The generic name Panthera is traceable to the classical Latin word 'panthēra' and the ancient Greek word πάνθηρ 'panther'. Felis leo was the scientific name used by Carl Linnaeus in 1758, who described the lion in his work Systema Naturae . The genus name Panthera was coined by Lorenz Oken in 1816. Between

12543-518: Was distributed in the Asian and African Palearctic since at least the early Pliocene . The earliest fossils recognisable as lions were found at Olduvai Gorge in Tanzania and are estimated to be up to 2 million years old. Estimates for the divergence time of the modern and cave lion lineages range from 529,000 to 392,000 years ago based on mutation rate per generation time of the modern lion. There

12656-947: Was present in the Caucasus until the 10th century. It lived in Palestine until the Middle Ages and in Southwest Asia until the late 19th century. By the late 19th century, it had been extirpated in most of Turkey. The last live lion in Iran was sighted in 1942, about 65 km (40 mi) northwest of Dezful , although the corpse of a lioness was found on the banks of the Karun river in Khuzestan province in 1944. It once ranged from Sind and Punjab in Pakistan to Bengal and

12769-426: Was vicariance biogeography, which explained the origin of new lineages through geological events like the drifting apart of continents or the formation of rivers. When a continuous population (or species) is divided by a new river or a new mountain range (i.e., a vicariance event), two populations (or species) are created. Paleogeography , geology and paleoecology are all important fields that supply information that

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