Hyena

Source: Wikipedia, the free encyclopedia.
This article is about the family of animals. For other uses, see Hyena (disambiguation).

Hyenas
Temporal range: 22–0 
Ma
 Early Miocene – recent
The four extant species of hyena, clockwise from upper left: spotted hyena (
Hyaena hyaena
)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Superfamily: Herpestoidea
Family: Hyaenidae
Gray, 1821
Type genus
Hyaena
, 1762
Genera
Synonyms
Aardwolf, smallest member of the Hyena family, skeleton. (Museum of Osteology)

Hyenas or hyaenas (

Mammalia.[2] Despite their low diversity, hyenas are unique and vital components of most African ecosystems.[3]

Although

scent marking, defecation habits, mating and parental behavior are consistent with the behavior of other feliforms.[4]

Hyenas feature prominently in the folklore and mythology of human cultures that live alongside them. Hyenas are commonly viewed as frightening and worthy of contempt. In some cultures, hyenas are thought to influence people's spirits, rob graves, and steal livestock and children.[5] Other cultures associate them with witchcraft, using their body parts in traditional medicine.[6]

Evolution

Origins

Hyenas originated in the jungles of

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.[7]
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
extant spotted, brown and striped hyenas) became the undisputed top scavengers of Eurasia and Africa.[7]

Rise and fall of the dog-like hyenas

Skull of Ictitherium viverrinum, one of the "dog-like" 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 dog-like hyenas were nimble-bodied, wolfish animals; one species among them was

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. Chasmaporthetes managed to survive for some time in North America by deviating from the endurance-running and bone-crushing niches monopolized by canids, and developing into a cheetah-like sprinter. Most of the dog-like hyenas had died off by 1.5 million years ago.[7]

Bone-crushing hyenas

By 10–14 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 family

sabre-toothed cats. One genus, Pachycrocuta, was a 200 kg (440 lb) mega-scavenger that could splinter the bones of elephants.[7] Starting in the early Middle Pleistocene Pachycrocuta was replaced by the smaller Crocuta and Hyena, which corresponds to a general faunal change, perhaps in connection to the Mid-Pleistocene transition.[8]

Rise of modern hyenas

Skeletons of a striped hyena (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.[7]

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.[9]

The

Middle Pleistocene, and quickly colonised a very wide area from Europe, to southern Africa and China.[11] 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.[12] 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.[13] 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.

See also: Cave hyena

Genera of the Hyaenidae (extinct and recent)

A spotted hyena of subfamily Hyaeninae

The list follows McKenna and Bell's 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 Parahyaena. However, some research has suggested Parahyaena may be synonymous with Pachycrocuta, making the brown hyena the only extant member of this genus.[17]

Phylogeny

The following cladogram illustrates the phylogenetic relationships between extant and extinct hyaenids based on the morphological analysis by Werdelin & Solounias (1991),[25] as updated by Turner et al. (2008).[26]

Evolution of hyaenas
 

Protictitherium crassum

"Protictitherium" cingulatum

"Protictitherium" intermedium

"Protictitherium" llopisi

"Protictitherium" punicum

"Protictitherium" gaillardi

"Protictitherium" sumegense

"Protictitherium" csakvarense

 
 Plioviverrops 
 

Plioviverrops gervaisi

    Civet/mongoose-like insectivore-omnivores

Plioviverrops orbignyi

 

Plioviverrops guerini

Plioviverrops faventinus

Plioviverrops gaudryi

 

Tungurictis spocki

 

Thalassictis robusta

"Thalassictis" certa

"Thalassictis" montadai

"Thalassictis" proava

"Thalassictis" sarmatica

"Thalassictis" spelaea

 

Tongxinictis primordialis

 
 Proteles 
 

Proteles cristatus (aardwolf)

Proteles amplidentus

 
 Ictitherium 
 

Ictitherium viverrinum

Ictitherium ebu

    Jackal-like hyaenas
 
 

Ictitherium tauricum

Ictitherium ibericum

 

Ictitherium kurteni

Ictitherium intuberculatum

Ictitherium pannonicum

 

Miohyaenotherium bessarabicum

 Hyaenotherium 

Hyaenotherium wongii

Hyaenictitherium hyaenoides

"Hyaenictitherium" pilgrimi

"Hyaenictitherium" parvum

"Hyaenictitherium" namaquensis

"Hyaenictitherium" minimum

 Lycyaena 

Lycyaena chaeretis

Lycyaena dubia

    Cursorial hunting hyaenas

Lycyaena macrostoma

Lycyaena crusafonti

 Hyaenictis 

Hyaenictis graeca

Hyaenictis almerai

Hyaenictis hendeyi

 Lycyaenops 

Lycyaenops rhomboideae

Lycyaenops silberbergi

 Chasmaporthetes 

Chasmaporthetes exitelus

Chasmaporthetes bonisi

Chasmaporthetes borissiaki

Chasmaporthetes lunensis

Chasmaporthetes melei

Chasmaporthetes ossifragus

Chasmaporthetes sp. Florida

Chasmaporthetes nitidula

Chasmaporthetes australis

(running hyaenas)
 
Hyaeninae
 
Bone-cracking hyaenas    

Metahyaena confector

Palinhyaena reperta

Hyaenid sp. E Langebaar

Belbus beaumonti

Hyaena abronia

Hyaena hyaena (striped hyaena)

Parahyaena howelli

Parahyaena brunnea (brown hyaena)

Pliocrocuta perrieri

Pachycrocuta brevirostris (giant hyaena)

Adcrocuta eximia

Allohyaena kadici

Crocuta crocuta (spotted hyaena)
  

Crocuta eturono

(bone‑cracking hyenas)

Phylogenic relationships based on morphological characteristics, after Werdelin & Solounias (1991) and Turner et al (2008)

A more recent molecular analysis agrees on the phylogenetic relationship between the four extant hyaenidae species (Koepfli et al, 2006[27]).

 Hyaenidae 
 
   

Proteles cristatus (aardwolf)

 
 

Crocuta crocuta (spotted hyena)

Hyaena hyaena (striped hyena)

Parahyaena brunnea (brown hyena)

Characteristics

Build

Further information: Spotted hyena § Description, Striped hyena § Description, Aardwolf § Description, 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

female spotted hyena's external genitalia closely resembles that of the male.[33]

Their

premolars, unlike canids, which do so with their post-carnassial molars.[34] 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.[35][36] 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.[37][38] The aardwolf has greatly reduced cheek teeth, sometimes absent in the adult, but otherwise has the same dental formula as the other three species.[39] The dental formula
for all hyena species is: 3.1.4.13.1.3.1

Although hyenas lack

spray
them when attacked.

Behavior

Brown hyena marking its territory with its anal glands
Spotted hyena cubs at their den

Hyenas

mark their territories using their anal glands, a trait found also in viverrids and mustelids, but not canids and felids.[42] When attacked by lions or dogs, striped[43] and brown hyenas[44] will feign death, though the spotted hyena will defend itself ferociously.[36] The spotted hyena is very vocal, producing a number of different sounds consisting of whoops, grunts, groans, lows, giggles, yells, growls, laughs and whines.[45] The striped hyena is comparatively silent, its vocalizations being limited to a chattering laugh and howling.[46]

Whoop of a spotted hyena in Umfolosi Game Park, South Africa.

engage in a single, drawn out copulation.[42] Spotted hyena cubs are born almost fully developed, with their eyes open and erupting incisors and canines, though lacking adult markings.[47] In contrast, striped hyena cubs are born with adult markings, closed eyes and small ears.[48] Hyenas do not regurgitate food for their young and male spotted hyenas play no part in raising their cubs,[42] though male striped hyenas do so.[49]

The striped hyena is primarily a scavenger, though it will also attack and kill any animals it can overcome,

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.[51] 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.[7] Except for the aardwolf, hyenas are known to drive off larger predators, like lions, from their kills, despite having a reputation in popular culture for being cowardly.[43] 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 the striped and brown hyenas may live in family groups and congregate at kills.[52]
Spotted hyenas are one of the few mammals other than bats known to survive infection with rabies virus[53] 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.[54] Despite this perceived unique disease resistance, little is known about the immune system of spotted hyenas,[55][56][57] and even less is known about other Hyaenidae species.

Relationships with humans

Folklore, mythology and literature

Cave hyena (Crocuta crocuta spelaea) painting found in the Chauvet Cave in 1994
A depiction of the legendary striped hyena, Krokottas of Kytheros Island, from 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 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

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.[60] A similar myth occurs in Mansôa. These "werehyenas" are executed when discovered, but do not revert to their human form when killed.[59]

Striped hyenas are often referred to in Middle Eastern literature and folklore, typically as symbols of treachery and stupidity.

northern India, witches or magicians are said to ride striped hyenas at night.[58]

The striped hyena is mentioned in 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.[63]

The vocalization of the spotted hyena resembling hysterical human

Shakespeare's As You Like It, Act IV. Sc.1.[citation needed
]

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.[citation needed]

Attacks on humans

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

In ordinary circumstances, striped hyenas are extremely timid around humans, though they may show bold behaviors towards people at night.[64] On rare occasions, striped hyenas have preyed on humans.

Among hyenas, only the spotted and striped hyenas have been known to become

Bering strait earlier.[66] Hyenas readily scavenge from human corpses; in Ethiopia, hyenas were reported to feed extensively on the corpses of victims of the 1960 attempted coup[67] and the Red Terror.[68] 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.[69]

Although spotted hyenas have been known to prey on humans in modern times, such incidents are rare. However, attacks on humans by spotted hyenas are likely to be underreported.

African sleeping sickness as they slept outside in camps.[73] 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 were 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.[70][72] 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.[70]

In the 1880s, a hyena was reported to have attacked humans, especially sleeping children, over a three-year period in the

boar, elephants, tigers, leopards and sloth bears.[76]

Hyenas as food and medicine

Hyenas are used for food and medicinal purposes in

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.[58]

References

Citations

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General and cited references

Further reading

External links

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