Hyena

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

Hyenas

Temporal range: 22–0 Ma 

PreꞒ

Pg

Early Miocene – recent

The four living species of hyena,

clockwise from upper left:
spotted hyena (Crocuta
crocuta), brown hyena (Hyaena
brunnea), aardwolf (Proteles
cristata) and striped hyena
(Hyaena hyaena)

Scientific classification

Kingdom: Animalia

Phylum: Chordata

Class: Mammalia

Order: Carnivora

Suborder: Feliformia

Infraorder: Viverroidea

Family: Hyaenidae
Gray, 1821

Genera

Crocuta (olive overlay on map)

Hyaena (blue on map)

Proteles (magenta red on map)

Adcrocuta†

Allohyaena†

Belbus†

Chasmaporthetes†

Herpestides†

Hyaenictis†

Hyaenotherium†

Ictitherium†

Ikelohyaena†

Leecyaena†

Lycyaena†

Metahyaena†
 Miohyaenotherium†

Palinhyaena†

Pachycrocuta†

Pliocrocuta†

Plioviverrops†

Protictitherium†

Thalassictis†

Tongxinictis†

Tungurictis†

Werdelinus†

Synonyms

Protelidae Flower, 1869

Aardwolf, smallest member of the Hyena family, skeleton. (Museum of Osteology)

Hyenas, or hyaenas (from Ancient Greek ὕαινα, hýaina[1]), are feliform carnivoran mammals of the

family Hyaenidae /haɪˈɛnɪdiː/. With only four extant species (in three genera), 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 closer to felines and viverrids, as part of suborder Feliformia, hyenas are
behaviourally and morphologically similar to canids in several elements due to convergent evolution;
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
claws are adapted for running and making sharp turns. However, hyenas' grooming, scent marking,
defecation habits, mating and parental behaviour are consistent with the behaviour 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 African medicine.[6]

Contents

1Evolution

1.1Origins

1.2Rise and fall of the dog-like hyenas

1.3Bone-crushing hyenas

1.4Rise of modern hyenas

2Genera of the Hyaenidae (extinct and recent)

2.1Phylogeny

3Characteristics

3.1Build

3.2Behavior

4Relationships with humans

4.1Folklore, mythology and literature

4.2Attacks on humans

4.3Hyenas as food and medicine

5References

5.1Notes

5.2Bibliography

6Further reading

7External links
Evolution[edit]

Origins[edit]

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.[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 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.[7]

Rise and fall of the dog-like hyenas[edit]

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 Ictitherium viverrinum, which was similar to a jackal. The dog-like hyenas were numerous; in some
Miocene fossil sites, the remains of Ictitherium and other dog-like hyenas outnumber those of all other
carnivores combined. The decline of the dog-like hyenas began 5–7 million years ago during a period of
climate change, exacerbated by canids crossing 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. Chasmaporthetes managed to survive for some time in North America by deviating
from the endurance-running and bone-crushing niches monopolised 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[edit]

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 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 dog subfamily Borophaginae. 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.[7] With the decline of large
herbivores by the late ice age, Pachycrocuta was replaced by the smaller Crocuta.[7]

Rise of modern hyenas[edit]

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

The four extant species are the striped hyena (Hyaena hyaena), the brown hyena (Hyaena 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 striped hyena may have evolved from H. namaquensis of Pliocene Africa. Striped hyena fossils are
common in Africa, with records going back as far as 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.[8]

The spotted hyena (Crocuta crocuta) diverged from the striped and brown hyena 10 million years ago.
[9] Its direct ancestor was the Indian Crocuta sivalensis, which lived during the Villafranchian.
[10] 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, 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.[7] 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.[10] 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.[11]

Genera of the Hyaenidae (extinct and recent)[edit]

Reconstruction of Pachycrocuta brevirostris

A spotted hyena of subfamily Hyaeninae

The list follows McKenna and Bell's Classification of Mammals for prehistoric genera (1997)[12] and
Wozencraft (2005) in Wilson and Reeders Mammal Species of the World for extant genera.[13] 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[14]). 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

†Tongxinictis[15] (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[16](Late Miocene of Europe)

†Lycyaena (Late Miocene of Eurasia)

†Tungurictis[17] (Middle Miocene of Africa and Eurasia)

†Protictitherium (Middle Miocene of Africa and Asia, Middle to Late Miocene of Europe)

Subfamily Hyaeninae

†Palinhyaena[18] (Late Miocene of Asia)

†Ikelohyaena[19] (Early Pliocene of Africa)

Hyaena (=Euhyaena, =Parahyaena; 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[20] (Late Miocene of Asia?, Late Miocene of Europe, Early Pliocene (?Early Pleistocene) of
Africa)

†Leecyaena[18] (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

Proteles (=Geocyon; aardwolf. Pleistocene to Recent of Africa)