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This article is about the family of animals. For other uses, see Hyena (disambiguation).

Hyenas
Temporal range: 26–0 Ma Early Miocene-recent
All extant species of hyenas in descending order of size: spotted hyena, brown hyena, striped hyena and aardwolf.
Scientific classification
Kingdom:
Animalia
Phylum:
Chordata
Class:
Mammalia
Order:
Carnivora
Suborder:
Feliformia
Family:
Hyaenidae

Gray, 1821
Genera
Synonyms

Hyenas or hyaenas (from Greek ὕαινα hýaina[1]) are any feliform carnivoran mammals of the family Hyaenidae /haɪˈɛn[invalid input: 'ɨ']diː/. With only four extant species, it is the fifth-smallest biological family in the Carnivora, and one of the smallest in the class Mammalia.[2] Despite their low diversity, hyenas are unique and vital components of most African ecosystems.[3]

Although phylogenetically they are closer to felines and viverrids, hyenas are behaviourally and morphologically similar to canines in several aspects; both hyenas and canines are non-arboreal, cursorial hunters that catch prey with their teeth rather than claws. Both eat food quickly and may store it, and their calloused feet with large, blunt, nonretractable nails are adapted for running and making sharp turns. However, the hyenas' grooming, scent marking, defecating habits, mating, and parental behaviour are consistent with the behaviour of other feliforms.[4]

Spotted hyenas may kill as many as 95% of the animals they eat,[5] while striped hyenas are largely scavengers.[6] Generally, hyenas are known to drive off larger predators, like lions, from their kills, despite having a reputation in popular culture for being cowardly.[6] Hyenas are primarily nocturnal animals, but sometimes venture from their lairs in the early-morning hours. With the exception of the highly social spotted hyena, hyenas are generally not gregarious animals, though they may live in family groups and congregate at kills.[7]

Hyenas first arose in Eurasia during the Miocene period from viverrid-like ancestors, and 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), they 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.[8]

Hyenas feature prominently in the folklore and mythology of human cultures with which they are sympatric. Hyenas are commonly viewed as frightening and worthy of contempt, and are associated with witchcraft, because their body parts are used as ingredients in traditional medicine. In some cultures, hyenas are thought to influence people’s spirits, rob graves, and steal livestock and children.[9]

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 banded palm 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.[8]

Rise and fall of the doglike hyenas

Skull of Ictitherium viverrinum, one of the "doglike" hyenas. American Museum of Natural History

The descendants of Plioviverrops reached their peak 15 million years ago, with more than 30 species having been identified. Unlike most modern hyena species, which are specialised bone-crushers, these doglike hyenas were nimble-bodied, wolfish animals; one species among them was Ictitherium viverrinum, which was similar to a jackal. The doglike hyenas were very numerous; in some Miocene fossil sites, the remains of Ictitherium and other doglike hyenas outnumber those of all other carnivores combined. The decline of the doglike hyenas began 5–7 million years ago during a period of climate change, which was exacerbated when canids crossed the Bering land bridge to Eurasia. One species, Chasmaporthetes ossifragus, managed to cross the land bridge into North America, being the only hyena to do so. Chasmopothertes managed to survive for some time in North America by deviating from the cursorial and bone-crushing niches monopolised by canids, and developing into a cheetah-like sprinter. Most of the doglike hyenas had died off by 1.5 million years ago.[8]

Bone-crushing hyenas

By 10–12 million years ago, the hyena family had split into two distinct groups: dog-like hyenas and bone-crushing hyenas. The arrival of the ancestral bone-crushing hyenas coincided with the decline of the similarly built but unrelated family Percrocutidae. The bone-crushing hyenas survived the changes in climate and the arrival of canids, which wiped out the dog-like hyenas, though they never crossed into North America, as their niche there had already been taken by the Borophaginae subfamily of dogs. By 5 million years ago, the bone-crushing hyenas had become the dominant scavengers of Eurasia, primarily feeding on large herbivore carcasses felled by sabre-toothed cats. One genus, Pachycrocuta, was a 200 kg (440 lb) mega-scavenger that could splinter the bones of elephants. With the decline of large herbivores by the late ice age, Pachycrocuta was replaced by the smaller Crocuta.[8]

Rise of modern hyenas

Skeletons of a striped hyena and a spotted hyena, two species of the "bone-crushing" hyenas.

The aardwolf (Proteles cristata) 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. Its unrivaled ability to digest the terpene excretions from soldier termites is likely a modification of the strong digestive system its ancestors used to digest fetid carrion.[8]

The striped hyena (Hyaena hyaena) may have evolved from H. namaquensis of Pliocene Africa. Striped hyena fossils are common in Africa, with records going back as far as the Middle Pleistocene and even to the Villafranchian. As fossil striped hyenas are absent from the Mediterranean region, it is likely that the species is a relatively late invader to Eurasia, having likely spread outside Africa only after the extinction of spotted hyenas in Asia at the end of the Ice Age. The striped hyena occurred for some time in Europe during the Pleistocene, having been particularly widespread in France and Germany. It also occurred in Montmaurin, Hollabrunn in Austria, the Furninha Cave in Portugal and the Genista Caves in Gibraltar. The European form was similar in appearance to modern populations, but was larger, being comparable in size to the brown hyena (Hyaena brunnea).[10]

The spotted hyena (Crocuta crocuta) diverged from the striped and brown hyena 10 million years ago.[11] Its direct ancestor was the Indian Crocuta sivalensis, which lived during the Villafranchian.[12] Ancestral spotted hyenas probably developed social behaviours in response to increased pressure from rivals on carcasses, thus forcing them to operate in teams. Spotted hyenas evolved sharp carnassials behind their crushing premolars, therefore they did not need to wait for their prey to die, as is the case for brown and striped hyenas, and thus became pack hunters as well as scavengers. They began forming increasingly larger territories, necessitated by the fact that their prey was often migratory, and long chases in a small territory would have caused them to encroach into another clan's turf.[8] Spotted hyenas spread from their original homeland during the Middle Pleistocene, and quickly colonised a very wide area from Europe, to southern Africa and China.[12] With the decline of grasslands 12,500 years ago, Europe experienced a massive loss of lowland habitats favoured by spotted hyenas, and a corresponding increase in mixed woodlands. Spotted hyenas, under these circumstances, would have been outcompeted by wolves and humans, who were as much at home in forests as in open lands—and in highlands as in lowlands. Spotted hyena populations began to shrink after roughly 20,000 years ago, completely disappearing from Western Europe between 11 and 14 thousand years ago, and earlier in some areas.[13]

Genera of the Hyaenidae (extinct and recent)

File:Short-faced hyena1.JPG
Reconstruction of Pachycrocuta brevirostris
A spotted hyena of subfamily Hyaeninae

The list follows McKenna and Bells Classification of Mammals for prehistoric genera (1997)[14] and Wozencraft (2005) in Wilson and Reeders Mammal Species of the World for extant genera.[15] 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 (though they are generally grouped as sister-taxa to hyenas[16]). Furthermore, the living brown hyena and its closest extinct relatives are not included in the genus Pachycrocuta, but in the genus Hyaena. The Protelinae (aardwolves) are not treated as a separate subfamily, but included in the Hyaeninae.

  • Family Hyaenidae
    • †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)
      • †Lychyaena (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, =hyena; including striped hyena, brown 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
      • †Hyaenictis (Late Miocene of Asia?, Late Miocene of Europe, Early Pliocene (?Early Pleistocene) of Africa)
      • †Leecyaena (Late Miocene and/or Early Pliocene of Asia)
      • †Chasmaporthetes (=Ailuriaena; including Lycaenops, Euryboas; Late Miocene to Early Pleistocene of Eurasia, Early Pliocene to Late Pliocene or Early Pleistocene of Africa, Late Pliocene to Early Pleistocene of North America)
      • †Pachycrocuta (Pliocene and Pleistocene of Eurasia and Africa)
      • †Adcrocuta (Late Miocene of Eurasia)
      • Crocuta (=Crocotta; including Eucrocuta; spotted hyena and cave hyena. Late Pliocene to recent of Africa, Late Pliocene to Late Pleistocene of Eurasia)
    • Subfamily Protelinae

Characteristics

Build

Further information: Spotted hyena § Physical description, Striped hyena § Physical description, Aardwolf § Physical characteristics, and Brown hyena § Description
Striped hyena skull. Note the disproportionately large carnassials and premolars adapted for bone consumption
Aardwolf skull. Note the greatly reduced molars and carnassials, rendered redundant from insectivory

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 toward 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.[17] Like canids, hyenas have short, blunt, non-retractable claws.[18] Their pelage is sparse and coarse with poorly developed or absent underfur. Most species have a rich mane of long hair running from the withers or from the head.[17] With the exception of the spotted hyena, hyaenids have striped coats, which they likely inherited from their viverrid ancestors.[8] Their ears are large and have simple basal ridges and no marginal bursa.[18] Their vertebral column, including the cervical region are of limited mobility. Hyenas have no baculum.[19] Hyenas have an additional pair of ribs than canids, and their tongues are rough like those of felids and viverrids.[20] Males in most hyena species are larger than females,[21] though the spotted hyena is exceptional, 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.[22]

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.[19] 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.[23] 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.[24][25] 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.[26][27] The aardwolf has greatly reduced cheek teeth, sometimes absent in the adult, but otherwise has the same dentition as the other three species.[28] The dental formula for all hyena species is: 3.1.4.13.1.3.1

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.[18] These glands produce a white, creamy secretion that the hyenas paste onto grass stalks. The odour of this secretion is very strong, smelling of boiling cheap soap or burning, and can be detected by humans several metres downwind.[29] The secretions are primarily used for territorial marking, though both the aardwolf[8] and the striped hyena[30] will spray them when attacked.

Behaviour

See also: Spotted hyena § Behaviour, Brown hyena § Behavior, Striped hyena § Behavior, and Aardwolf § Behavior
Spotted hyena cubs at their den
Brown hyena cub standing on a path of stones

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). However, unlike other feliforms, they do not "wash" their faces. 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.[31] When attacked by lions or dogs, striped[6] and brown hyenas[32] will feign death, though the spotted hyena will defend itself ferociously.[25] The spotted hyena is very vocal, producing a number of different sounds consisting of whoops, grunts, groans, lows, giggles, yells, growls, laughs and whines.[33] The striped hyena is comparatively silent, its vocalisations being limited to a chattering laugh and howling.[34]

Mating between hyenas involves a number of short copulations with brief intervals, unlike canids, who generally engage in a single, drawn out copulation.[31] Spotted hyena cubs are born almost fully developed, with their eyes open and erupting incisors and canines, though lacking adult markings.[35] In contrast, striped hyena cubs are born with adult markings, closed eyes and small ears.[36] Hyenas do not regurgitate food for their young and male spotted hyenas play no part in raising their cubs,[31] though male striped hyenas do so.[37]

The striped hyena is primarily a scavenger, though it will occasionally attack and kill any defenseless animal it can overcome,[6] and will supplement its diet with fruits.[38] 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. 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.[8] Hyenas are also known for their characteristic calls, which sound like laughs.[39]

Relationships with humans

Folklore, mythology and literature

Cave hyena (Crocuta crocuta spelaea) painting found in the Chauvet Cave in 1994
A striped hyena, as depicted on the Nile mosaic of Palestrina

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 or not 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 western African tales, spotted hyenas are sometimes depicted as bad Muslims who challenge the local animism that exists among the Beng in Côte d’Ivoire.[citation needed] In East Africa, Tabwa mythology portrays the spotted hyena as a solar animal that first brought the sun to warm the cold earth, while West African folklore generally shows the hyena as symbolizing immorality, dirty habits, the reversal of normal activities, and other negative traits. In Tanzania, there is a belief that witches use spotted hyenas as mounts.[40] In the Mtwara Region of Tanzania, it is believed that a child born at night while a hyena is crying will likely grow up to be a thief. In the same area, hyena faeces are believed to enable a child to walk at an early age, thus it is not uncommon in that area to see children with hyena dung wrapped in their clothes.[41] 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 house members able to transform into hyenas.[42] A similar myth occurs in Mansôa. These "werehyenas" are executed when discovered, but do not revert to their human form when killed.[41]

Striped hyenas are often referred to in Middle Eastern literature and folklore, typically as symbols of treachery and stupidity.[43] In the Near and Middle East, striped hyenas are generally regarded as physical incarnations of jinns.[40] Arab writer Al-Quazweeni (1204–1283) spoke of a tribe of people called Al-Dabeyoun meaning "hyena people". In his book Aajeb Al-Makhlouqat he wrote that should one of this tribe be in a group of 1000 people, a hyena could pick him out and eat him.[43] A Persian medical treatise written in 1376 tells how to cure cannibalistic people known as kaftar, who are said to be “half-man, half-hyena”.[40] Al-Doumairy in his writings in Hawayan Al-Koubra (1406) wrote that striped hyenas were vampiric creatures that attacked people at night and sucked the blood from their necks. He also wrote that hyenas only attacked brave people. Arab folklore tells of how hyenas can mesmerise victims with their eyes or sometimes with their pheromones.[43] In a similar vein to Al-Doumairy, the Greeks, until the end of the 19th century, believed that the bodies of werewolves, if not destroyed, would haunt battlefields as vampiric hyenas that drank the blood of dying soldiers.[44] The image of striped hyenas in Afghanistan, India and Palestine is more varied. Though feared, striped hyenas were also symbolic for 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.[40]

The striped hyena is mentioned in the Bible. The Arab word for the hyena, ḍab` or ḍabu` (plural ḍibā`), is alluded to in a valley in Israel known as Shaqq-ud-Diba` (meaning "cleft of the hyenas") and Wadi-Abu-Diba` (meaning "valley of the hyenas"). Both places have been interpreted by some scholars as being the Biblical Valley of Tsebo`im mentioned in 1 Samuel 13:18. The modern Hebrew word for hyena is tzavoa`, which is similar to the word "tsavua`" meaning "colored". Though the Authorized King James Version of the Bible interprets the term "`ayit tsavua`" (which appears in Jeremiah 12:9) as "speckled bird", Henry Baker Tristram argued that it was most likely a hyena being mentioned.[45]

The vocalisation of the spotted hyena resembling hysterical human laughter has been alluded to in numerous works of literature: "to laugh like a hyæna" was a common simile, and is featured in The Cobbler's Prophecy (1594), Webster's Duchess of Malfy (1623) and Shakespeares As You Like It, Act IV. Sc.1.

Hyenas are also featured in Disney's animated features, such as Dumbo, Lady and the Tramp, Noah's Ark, Bedknobs and Broomsticks, and The Lion King.

Attacks on humans

Illustration from Fraser's magazine showing an artist's impression of a "stag-hound" biting a spotted hyena attacking its master

Among hyenas, only the spotted and striped hyena have been known to become man-eaters. Hyenas are known to have preyed on humans in prehistory: Human hair has been found in fossilised hyena dung dating back 195,000 to 257,000 years.[46] 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 like modern spotted hyenas in Africa. The oldest Alaskan human remains coincide with roughly the same time cave hyenas became extinct, leading certain paleontologists to infer that hyena predation was what prevented humans crossing the Bering strait earlier.[47] Hyenas readily scavenge from human corpses; in Ethiopia, hyenas were reported to feed extensively on the corpses of victims of the 1960 attempted coup[48] and the Red Terror.[49] Hyenas habituated to scavenging on human corpses may develop bold behaviours 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.[50]

Although spotted hyenas do prey on humans in modern times, such incidents are rare. However, according to the SGDRN (Sociedade para a Gestão e Desenvolvimento da Reserva do Niassa Moçambique), attacks on humans by spotted hyenas are likely to be underreported.[51] According to hyena expert Dr. Hans Kruuk, man-eating spotted hyenas tend to be very large specimens: A pair of man-eating hyenas, responsible for killing 27 people in Mulanje (Mlanje), Malawi in 1962, were weighed at 72 kg (159 lb) and 77 kg (170 lb) after being shot.[52] In 1903, Hector Duff wrote of how spotted hyenas in the Mzimba district of Angoniland would wait at dawn outside people's huts and attack them when they opened their doors.[53] Victims of spotted hyenas tend to be women, children and sick or infirm men: Theodore Roosevelt wrote on how in 1908–1909 in Uganda, spotted hyenas regularly killed sufferers of African sleeping sickness as they slept outside in camps.[54] Spotted hyenas are widely feared in Malawi, where they have been known to occasionally attack people at night, particularly during the hot season when people sleep outside. Hyena attacks were widely reported in Malawi's Phalombe plain, to the north of Michesi Mountain. Five deaths were 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.[51][53] An anecdotal news report from the World Wide Fund for Nature 2004 indicates 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.[51]

In ordinary circumstances, striped hyenas are extremely timid around humans, though they may show bold behaviours toward people at night.[55] On rare occasions, striped hyenas have preyed on humans. In the 1880s, a hyena was reported to have attacked humans, especially sleeping children, over a three-year period in the Iğdır Province, with 25 children and 3 adults being wounded in one year. The attacks provoked local authorities into announcing a reward of 100 rubles for every hyena killed. Further attacks were reported later in some parts of Transcaucasia, particularly in 1908. Instances are known in Azerbaijan of striped hyenas killing children sleeping in courtyards during the 1930s and 1940s. In 1942, a sleeping guard was mauled in his hut by a hyena in Qalıncaq (Golyndzhakh). Cases of children being taken by hyenas by night are known in southeast Turkmenia's Bathyz Nature Reserve. A further attack on a child was reported around Serakhs in 1948.[56] Several attacks have occurred in India; in 1962, nine children were thought to have been taken by hyenas in the town of Bhagalpur in the Bihar State in a six-week period[45] and 19 children up to the age of four were killed by hyenas in Karnataka, Bihar in 1974.[57] A consensus on wild animal attacks during a five-year period in the Indian state of Madhya Pradesh showed that hyenas had only attacked three people, the lowest figure when compared to deaths caused by wolves, gaur, boar, elephants, tigers, leopards and sloth bears.[58]

Hyenas as food and medicine

File:Hyena-forcefeed.jpg
An Ancient Egyptian mural showing a striped hyena being forcefed

Hyenas are used for food and medicinal purposes in some areas, including Muslim nations under the Shafiite school, where hyenas are considered halal because of their omnivorous diet. 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.[40]

References

Notes

  1. ^ ὕαινα, Henry George Liddell, Robert Scott, A Greek-English Lexicon, on Perseus. Etymologically, this is a feminine of ὕς "swine".
  2. ^ Wilson, D.E.; Mittermeier, R.A., eds. (2009). Handbook of the Mammals of the World, Volume 1: Carnivora. Barcelona: Lynx Edicions. pp. 50–658. ISBN 978-84-96553-49-1.
  3. ^ Mills & Hofer 1998, p. iv
  4. ^ Kruuk 1972, p. 274
  5. ^ Hyaena Specialist Group - Spotted Hyena: Diet and Foraging. Hyaenidae.org. Retrieved on 2015-11-06.
  6. ^ a b c d Pocock 1941, p. 72
  7. ^ Rosevear 1974, pp. 343–344
  8. ^ a b c d e f g h i Macdonald 1992, pp. 119–144
  9. ^ Mills & Hofer 1998, p. 96
  10. ^ Kurtén 1968, pp. 66–68
  11. ^ Mills & Hofer 1998, p. 1
  12. ^ a b Kurtén 1968, pp. 69–72
  13. ^ "Comparative ecology and taphonomy of spotted hyenas, humans, and wolves in Pleistocene Italy" (PDF). C. Stiner, Mary. Revue de Paléobiologie, Genève. Retrieved 2008-09-16.
  14. ^ Malcolm C. McKenna, Susan K. Bell: Classification of Mammals: Above the Species Level, Columbia University Press, New York 1997, 631 Seiten, ISBN 0-231-11013-8
  15. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 532–548. ISBN 978-0-8018-8221-0. OCLC 62265494.
  16. ^ Borja Figueirido, Zhijie JACK Tseng, Alberto Martín-Serra, Skull Shape Evolution In Durophagous Carnivorans, Article in Evolution 67(7):1975-93 · July 2013 DOI: 10.1111/evo.12059 · Source: PubMed
  17. ^ a b Heptner & Sludskii 1992, p. 3
  18. ^ a b c Pocock 1941, pp. 62–63
  19. ^ a b Heptner & Sludskii 1992, pp. 4–5
  20. ^ Holl, William & Wood, Neville The Analyst: a quarterly journal of science, literature, natural history, and the fine arts, Volume 10, p. 59, Simpkin & Marshall, 1840
  21. ^ Mills & Hofer 1998, p. 21
  22. ^ Kruuk 1972, pp. 210–211
  23. ^ Therrien, François (2005). "Mandibular force profiles of extant carnivorans and implications for the feeding behaviour of extinct predators". Journal of Zoology. 267 (3): 249–270. doi:10.1017/S0952836905007430.
  24. ^ Johnson, Daniel (1827) Sketches of Indian Field Sports: With Observations on the Animals; Also an Account of Some of the Customs of the Inhabitants; with a Description of the Art of Catching Serpents, as Practised by the Conjoors and Their Method of Curing Themselves when Bitten: with Remarks on Hydrophobia and Rabid Animals p. 45-46, R. Jennings, 1827
  25. ^ a b Stevenson-Hamilton, James (1917) Animal life in Africa, Vol. 1, p.95, London : William Heinemann
  26. ^ Salleh, Anna (4 April 2005). "Marsupial has the deadliest bite". abc.net.au. Retrieved 24 January 2013.
  27. ^ Wroe, S, McHenry, C; Thomason, J. (2005). "Bite club: comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa". Proceedings of the Royal Society B: Biological Sciences. 272 (1563): 619–625. doi:10.1098/rspb.2004.2986. PMC 1564077. PMID 15817436.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  28. ^ Richardson, Philip K.R.; Bearder, Simon (1984). Macdonald, D. (ed.). The Encyclopedia of Mammals. New York: Facts on File. pp. 154–159. ISBN 0-87196-871-1.
  29. ^ Kruuk 1972, p. 222
  30. ^ Heptner & Sludskii 1992, p. 38
  31. ^ a b c Kruuk 1972, pp. 271–73
  32. ^ Mills & Mills 2010, pp. 60–61
  33. ^ Kruuk 1972, p. 220
  34. ^ Pocock 1941, p. 73
  35. ^ Kruuk 1972, pp. 247–249
  36. ^ Rosevear 1974, p. 350
  37. ^ Heptner & Sludskii 1992, pp. 40–42
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Bibliography

Further reading

  • Funk, Holdger (2010) Hyaena: On the Naming and Localisation of an Enigmatic Animal, GRIN Verlag, ISBN 3-640-69784-7
  • Lawick, Hugo & Goodall, Jane (1971) Innocent Killers, Houghton Mifflin Company Boston
  • Mills, M. G. L. (2003) Kalahari Hyenas: Comparative Behavioral Ecology of Two Species, The Blackburn Press
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