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Hyenas or hyaenas are feliform carnivoran mammals belonging to the family Hyaenidae. With just four extant species, it is the fifth-smallest family in the order Carnivora and one of the smallest in the class Mammalia. Despite their low diversity, hyenas are unique and vital components of most African ecosystems.

Evolution
Origins Hyenas originated in the jungles of Miocene Eurasia 22 million years ago, when most early feliform species were still largely arboreal. The first ancestral hyenas were likely similar to the modern African civet; one of the earliest hyena species described, Plioviverrops, was a lithe, civet-like animal that inhabited Eurasia 20–22 million years ago, and is identifiable as a hyaenid by the structure of the middle ear and dentition. The lineage of Plioviverrops prospered and gave rise to descendants with longer legs and more pointed jaws, a direction similar to that taken by canids in North America. Hyenas then diversified into two distinct types: lightly built dog-like hyenas and robust bone-crushing hyenas. Although the dog-like hyenas thrived 15 million years ago (with one taxon having colonised North America), most became extinct after a change in climate, along with the arrival of canids into Eurasia. Of the dog-like hyena lineage, only the insectivorous aardwolf survived, while the bone-crushing hyenas (including the extant spotted, brown, and striped hyenas) became the undisputed top scavengers of Eurasia and Africa. Rise of modern hyenas (left) and a spotted hyena (right), two species of the "bone-crushing" hyenas The four extant species are the striped hyena (Hyaena hyaena), the brown hyena (Parahyaena brunnea), the spotted hyena (Crocuta crocuta), and the aardwolf (Proteles cristata). The aardwolf can trace its lineage directly back to Plioviverrops 15 million years ago, and is the only survivor of the dog-like hyena lineage. Its success is partly attributed to its insectivorous diet, for which it faced no competition from canids crossing from North America. It is likely that its unrivaled ability to digest the terpene excretions from soldier termites is a modification of the strong digestive system its ancestors used to consume fetid carrion. The spotted hyena (Crocuta crocuta) diverged from the striped and brown hyena 10 million years ago. Its direct ancestor was the Indian Crocuta sivalensis, which lived during the Villafranchian. The eventual disappearance of the spotted hyena from Europe has traditionally been attributed to the end of the last glacial period and a subsequent displacement of open grassland by closed forests, which favoured wolves and humans instead. However, analyses have shown that climate change alone is insufficient to explain the spotted hyena's disappearance from Europe, suggesting that other factors – such as human pressure – must have played a role. This suggests that the events must be seen within the broader context of late-Quaternary extinctions, as the late Pleistocene and early Holocene saw the disappearance of many primarily large mammals from Europe and the world. Expansion or duplication of the olfatory receptor gene family has been found in all 4 extant species, which would have led to the evolution of the more specialised feeding habits of hyenas. Expansion in immune-related gene families was also found in the spotted hyena, striped hyena and brown hyena, which would have led to the evolution of the scavenging in these species. Mutations and variants were also found in digestion-related genes (ASH1L, PTPN5, PKP3, AQP10). One of these digestion-related genes has variants also related to enhanced bone mineralisation (PTPN5), while other have also a role in inflammatory skin responses (PKP3). In aardwolves, expansion of genes related to toxin response were found (Lipocalin and UDP Glucuronosyltransferase gene families), which would have led to the evolution of the feeding of termites Trinervitermes in this species. Mutations and variants in genes related to craniofacial shape were also found (GARS, GMPR, STIP1, SMO and PAPSS2). Another gene is related to protective epidermis function (DSC1). ==Genera of the Hyaenidae (extinct and recent)==
Genera of the Hyaenidae (extinct and recent)
of subfamily Hyaeninae The list follows McKenna and Bell's Classification of Mammals for prehistoric genera (1997) and Wozencraft (2005) in Wilson and Reeders Mammal Species of the World for extant genera. The percrocutids are, in contrast to McKenna and Bell's classification, not included as a subfamily into the Hyaenidae, but as the separate family Percrocutidae, although they are generally grouped as sister-taxa to hyenas. However, many experts consider them members of Hyaenidae. Furthermore, the living brown hyena and its closest extinct relatives are not included in the genus Pachycrocuta, but in the genus Parahyaena. However, some research has suggested Parahyaena may be synonymous with Pachycrocuta, making the brown hyena the only extant member of this genus. • Family Hyaenidae • 'Subfamily Incertae sedis''''' • †Tongxinictis (Middle Miocene of Asia) • †Subfamily Ictitheriinae • †Herpestides (Early Miocene of Africa and Eurasia) • †Plioviverrops (including Jordanictis, Protoviverrops, Mesoviverrops; Early Miocene to Early Pliocene of Europe, Late Miocene of Asia) • †Ictitherium (=Galeotherium; including Lepthyaena, Sinictitherium, Paraictitherium; Middle Miocene of Africa, Late Miocene to Early Pliocene of Eurasia) • †Thalassictis (including Palhyaena, Miohyaena, Hyaenictitherium, Hyaenalopex; Middle to Late Miocene of Asia, Late Miocene of Africa and Europe) • †Hyaenotherium (Late Miocene to Early Pliocene of Eurasia) • †Miohyaenotherium (Late Miocene of Europe) • †Lycyaena (Late Miocene of Eurasia) • †Tungurictis (Middle Miocene of Africa and Eurasia) • †Protictitherium (Middle Miocene of Africa and Asia, Middle to Late Miocene of Europe) • Subfamily Hyaeninae • †Palinhyaena (Late Miocene of Asia) • †Ikelohyaena (Early Pliocene of Africa) • Hyaena (=Euhyaena,=Parahyaena; including striped hyena, Pliohyaena, Pliocrocuta, Anomalopithecus) Early Pliocene (?Middle Miocene) to Recent of Africa, Late Pliocene (?Late Miocene) to Late Pleistocene of Europe, Late Pliocene to recent in Asia • Parahyaena (=Hyaena; brown hyena Pliocene to recent of Africa) • †Hyaenictis (Late Miocene of Asia?, Late Miocene of Europe, Early Pliocene (?Early Pleistocene) of Africa) • †LeecyaenaProteles (=Geocyon; aardwolf. Pleistocene to Recent of Africa) Phylogeny The following cladogram illustrates the phylogenetic relationships between extant and extinct hyaenids based on the morphological analysis by Werdelin & Solounias (1991), as updated by Turner et al. (2008). {{Clade |style=font-size:80%; line-height:85%;background-color:#eeeeff |caption=Evolution of hyaenas |captionstyle=background-color:#8899ff;padding:10px;font-weight:bold; |footer=Phylogenic relationships based on morphological characteristics, after Werdelin & Solounias (1991) and Turner et al (2008) |footerstyle=background-color:#eeeeee;text-align:center; |1={{clade |style=background-color:#ccffcc; |label1= |1=Protictitherium crassum |2="Protictitherium" cingulatum |3="Protictitherium" intermedium |4="Protictitherium" llopisi |5="Protictitherium" punicum |6="Protictitherium" gaillardi |state7=dotted |7="Protictitherium" sumegense |state8=dotted |8="Protictitherium" csakvarense |style9=background-color:#eeeeff; |9={{clade |label1= |1= |2={{clade |label1= |1=Tungurictis spocki |2={{clade |label1= |1=Thalassictis robusta |2="Thalassictis" certa |3="Thalassictis" montadai |4="Thalassictis" proava |5="Thalassictis" sarmatica |6="Thalassictis" spelaea |7={{clade |label1= |1=Tongxinictis primordialis |2={{clade |label1= |1= |2={{clade |label1= |1= |2={{clade |label1= |1=Miohyaenotherium bessarabicum |2={{clade |1= |style2=background-color:#ccccff; |2={{clade |1= |2={{clade |1= |2={{clade |1= |2={{clade |label1=Chasmaporthetes |sublabel1=(running hyaenas) |1= |style2=background-color:#ddccff |label2=Hyaeninae |sublabel2=(bone-cracking hyenas) |2={HYAENINAE} }} }} }} }} }} }} }} }} }} }} }} }} }} |caption=Bone-cracking hyaenas |captionstyle=font-size:12px;font-weight:bold;color:darkred;text-align:right;padding:5px 20px 0px 0px; |1=Metahyaena confector |2=Palinhyaena reperta |3= }} }} A more recent molecular analysis agrees on the phylogenetic relationship between the four extant hyaenidae species (Koepfli et al, 2006). }} ==Characteristics==
Characteristics
Build Hyenas have relatively short torsos and are fairly massive and wolf-like in build, but have lower hind quarters, high withers and their backs slope noticeably downward towards their rumps. The forelegs are high, while the hind legs are very short and their necks are thick and short. Their skulls superficially resemble those of large canids, but are much larger and heavier, with shorter facial portions. Hyenas are digitigrade, with the fore and hind paws having four digits each and sporting bulging pawpads. With the exception of the spotted hyena, hyaenids have striped coats, which they likely inherited from their viverrid ancestors. Males in most hyena species are larger than females, though the spotted hyena is an exception, as it is the female of the species that outweighs and dominates the male. Also, unlike other hyenas, the female spotted hyena's external genitalia closely resembles that of the male. Their dentition is similar to that of the canid, but is more specialised for consuming coarse food and crushing bones. The carnassials, especially the upper, are very powerful and are shifted far back to the point of exertion of peak pressure on the jaws. The other teeth, save for the underdeveloped upper molars, are powerful, with broad bases and cutting edges. The canines are short, but thick and robust. Labiolingually, their mandibles are much stronger at the canine teeth than in canids, reflecting the fact that hyenas crack bones with both their anterior dentition and premolars, unlike canids, which do so with their post-carnassial molars. The strength of their jaws is such that both striped and spotted hyenas have been recorded to kill dogs with a single bite to the neck without breaking the skin. The spotted hyena is renowned for its strong bite proportional to its size, but a number of other animals (including the Tasmanian devil) are proportionately stronger. The aardwolf has greatly reduced cheek teeth, sometimes absent in the adult, but otherwise has the same dental formula as the other three species. The dental formula for all hyena species is: Although hyenas lack perineal scent glands, they have a large pouch of naked skin located at the anal opening. Large anal glands above the anus open into this pouch. Several sebaceous glands are present between the openings of the anal glands and above them. These glands produce a white, creamy secretion that the hyenas paste onto grass stalks. The odor of this secretion is very strong, smelling of boiling cheap soap or burning, and can be detected by humans several meters downwind. The secretions are primarily used for territorial marking, though both the aardwolf will spray them when attacked. Behavior Hyenas groom themselves often like felids and viverrids, and their way of licking their genitals is very cat-like (sitting on the lower back, legs spread with one leg pointing vertically upward). They defecate in the same manner as other Carnivora, though they never raise their legs as canids do when urinating, as urination serves no territorial function for them. Instead, hyenas mark their territories using their anal glands, a trait found also in viverrids and mustelids, but not canids and felids. When attacked by lions or dogs, striped will feign death, though the spotted hyena will defend itself ferociously. The striped hyena is comparatively silent, its vocalizations being limited to a chattering laugh and howling. Mating between hyenas involves a number of short copulations with brief intervals, unlike canids, who generally engage in a single, drawn out copulation. In contrast, striped hyena cubs are born with adult markings, closed eyes and small ears. Hyenas do not regurgitate food for their young and male spotted hyenas play no part in raising their cubs, The striped hyena is primarily a scavenger, though it will also attack and kill any animals it can overcome, and will supplement its diet with fruit. The spotted hyena, though it also scavenges occasionally, is an active pack hunter of medium to large sized ungulates, which it catches by wearing them down in long chases and dismembering them in a canid-like manner. Spotted hyenas may kill as many as 95% of the animals they eat. The aardwolf is primarily an insectivore, specialised for feeding on termites of the genus Trinervitermes and Hodotermes, which it consumes by licking them up with its long, broad tongue. An aardwolf can eat 300,000 Trinervitermes on a single outing. Spotted hyenas are one of the few mammals other than bats known to survive infection with rabies virus and have shown little or no disease-induced mortality during outbreaks in sympatric carnivores, in part due to the high concentration of antibodies present in their saliva. Despite this perceived unique disease resistance, little is known about the immune system of spotted hyenas, and even less is known about other Hyaenidae species. ==Relationships with humans==
Relationships with humans
Folklore, mythology and literature (Crocuta crocuta spelaea) painting found in the Chauvet Cave in 1994 Spotted hyenas vary in their folkloric and mythological depictions, depending on the ethnic group from which the tales originate. It is often difficult to know whether spotted hyenas are the specific hyena species featured in such stories, particularly in West Africa, as both spotted and striped hyenas are often given the same names. In West African tales, spotted hyenas are sometimes depicted as bad Muslims who challenge the local animism that exists among the Beng in Côte d'Ivoire. The Kaguru of Tanzania and the Kujamaat of southern Senegal view hyenas as inedible and greedy hermaphrodites. A mythical African tribe called the Bouda is reputed to have members able to transform into hyenas. A similar myth occurs in Mansôa. These "werehyenas" are killed when discovered, and do not revert to human form once dead. In the Near and Middle East, striped hyenas are generally regarded as physical incarnations of jinns. Arab writer al-Qazwīnī (1204–1283) spoke of a tribe of people called al-Ḍabyūn meaning "hyena people". In his book '''Ajā'ib Al-Makhlūqāt'' he wrote that should one of this tribe be in a group of 1,000 people, a hyena could pick him out and eat him. The image of striped hyenas in Afghanistan, India and Palestine is more varied. Though feared, striped hyenas were also symbolic of love and fertility, leading to numerous varieties of love medicine derived from hyena body parts. Among the Baluch and in northern India, witches or magicians are said to ride striped hyenas at night. The vocalization of the spotted hyena resembling hysterical human laughter has been alluded to in numerous works of literature: "to laugh like a hyæna" or a "hyen" was a common simile, and is featured in ''The Cobbler's Prophecy'' (1594), Webster's Duchess of Malfi (1623) and Shakespeare's As You Like It, Act IV. Sc.1. Die Strandjutwolf (The brown hyena) is an allegorical poem by the renowned South African poet, N. P. van Wyk Louw, which evokes a sinister and ominous presence. Attacks on humans Ordinarily, striped hyenas are extremely timid around humans, though they may show bold behaviors towards people at night. On rare occasions, striped hyenas have preyed on humans. Among hyenas, only the spotted and striped hyenas have been known to become man-eaters. Hyenas are known to have preyed on humans in prehistory: human hair has been found in fossilized hyena dung dating back 195,000 to 257,000 years. Some paleontologists believe that competition and predation by cave hyenas (Crocuta crocuta spelaea) in Siberia was a significant factor in delaying human colonization of Alaska. Hyenas may have occasionally stolen human kills, or entered campsites to drag off the young and weak, much as modern spotted hyenas do in Africa. The oldest Alaskan human remains coincide with roughly the same time cave hyenas became extinct, leading some paleontologists to infer that hyena predation prevented humans from crossing the Bering Strait earlier. Hyenas readily scavenge from human corpses; in Ethiopia, hyenas were reported to feed extensively on the corpses of victims of the 1960 attempted coup and the Red Terror. Hyenas habituated to scavenging on human corpses may develop bold behaviors towards living people: hyena attacks on people in southern Sudan increased during the Second Sudanese Civil War, when human corpses were readily available to them. Spotted hyenas have been known to prey on humans in modern times, but such incidents are rare. However, attacks on humans by spotted hyenas are likely to be underreported. Man-eating spotted hyenas tend to be very large specimens; a pair of man-eating hyenas, responsible for killing 27 people in Mulanje, Malawi in 1962, weighed in at 72 kg (159 lb) and 77 kg (170 lb) after being shot. A 1903 report describes spotted hyenas in the Mzimba district of Angoniland waiting at dawn outside people's huts to attack them when they opened their doors. Spotted hyenas are widely feared in Malawi, where they have been known to attack people at night, particularly during the hot season when people sleep outside. A spate of hyena attacks was reported in Malawi's Phalombe plain, with five deaths recorded in 1956, five in 1957 and six in 1958. This pattern continued until 1961, when eight people were killed. Attacks occurred most commonly in September, when people slept outdoors and bush fires made the hunting of wild game difficult for the hyenas. A 2004 news report stated that 35 people were killed by spotted hyenas in a 12-month period in Mozambique along a 20-km stretch of road near the Tanzanian border. Several attacks have occurred in India; in 1962, 9 children were thought to have been taken by hyenas in the town of Bhagalpur in the Bihar State in a six-week period, A survey of wild animal attacks during a five-year period in the Indian state of Madhya Pradesh reported that hyenas had attacked three people, causing fewer deaths than wolves, gaur, boar, elephants, tigers, leopards and sloth bears. File:Hyena attack (1842 woodcut).jpg|Illustration from ''Fraser's'' magazine showing an artist's impression of a "stag-hound" biting a spotted hyena attacking its master File:Bodleian Libraries, Handbill of Merchant's Hall, 1739, announcing A lion, lionesses, tigers, etc..jpg|A 1739 advertisement by Charles Benjamin Incledon featuring feliforms: the Mesopotamian lion from the vicinity of Bassorah, Cape lion, tiger from the East Indies, panther from Buenos Aires, Hyaena hyaena from West Africa, and leopard from Turkey, besides a "Man tyger" from Africa. The advertisement mentions that the 'hyaena' can mimic a human voice to lure humans. Hyenas as food and medicine Hyenas have occasionally been used for food and medicinal purposes in Somalia. Some Muslims consider it halal in Islam although this is disputed by other Muslims. This practice dates back to the times of the Ancient Greeks and Romans, who believed that different parts of the hyena's body were effective means to ward off evil and to ensure love and fertility. ==References==
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