Ictitheriinae
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Hyenas or hyaenas ( ; from
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
, ) are
feliform Feliformia is a suborder within the order Carnivora consisting of "cat-like" carnivorans, including cats (large and small), hyenas, mongooses, viverrids, and related taxa. Feliformia stands in contrast to the other suborder of Carnivora, Cani ...
carnivora Carnivora ( ) is an order of placental mammals specialized primarily in eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the sixth largest order of mammals, comprising at least 279 species. Carnivor ...
n mammals belonging to the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Hyaenidae (). With just four extant
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
(each in its own
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
), it is the fifth-smallest family in the order Carnivora and one of the smallest in the class
Mammalia A mammal () is a vertebrate animal of the class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three middle ear bon ...
. Despite their low diversity, hyenas are unique and vital components of most
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
n ecosystems. Although
phylogenetically In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
closer to felines and
viverrid Viverridae is a family (biology), family of small to medium-sized feliform mammals, comprising 14 genera with 33 species. This family was named and first described by John Edward Gray in 1821. Viverrids occur all over Africa, in southern Europe, ...
s, hyenas are behaviourally and morphologically similar to canids in several elements due to
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
: both hyenas and canines are non-
arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally (scansorial), but others are exclusively arboreal. The hab ...
,
cursorial A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often ...
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 In ethology, territory is the sociographical area that an animal consistently defends against conspecific competition (or, occasionally, against animals of other species) using agonistic behaviors or (less commonly) real physical aggression. ...
, defecation habits, mating, and parental behavior are consistent with the behavior of other feliforms. 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. Other cultures associate them with witchcraft, using their body parts in
traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) refers to the knowledge, skills, and practices rooted in the cultural beliefs of various societies, especially Indigenous groups, used for maintaining health and treatin ...
.


Evolution


Origins

Hyenas originated in the jungles of
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
Eurasia 22 million years ago, when most early feliform species were still largely
arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally (scansorial), but others are exclusively arboreal. The hab ...
. The first ancestral hyenas were likely similar to the modern
African civet The African civet (''Civettictis civetta'') is a large viverrid native to sub-Saharan Africa, where it is considered common and widely distributed in woodlands and secondary forests. It is listed as Least Concern on the IUCN Red List since 2008 ...
; 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 The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear). The mammalian middle ear contains three ossicles (malleus, incus, and stapes), which transfer the vibrations ...
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 North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
. 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 In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
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 Extant or Least-concern species, least concern is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Exta ...
spotted,
brown Brown is a color. It can be considered a composite color, but it is mainly a darker shade of orange. In the CMYK color model used in printing and painting, brown is usually made by combining the colors Orange (colour), orange and black. In the ...
, and striped hyenas) became the undisputed top scavengers of Eurasia and Africa.


Rise and fall of the dog-like hyenas

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 ''Ictitherium'' (meaning "weasel beast") is an extinct genus belonging to the family Hyaenidae and the subfamily Ictitheriinae erected by Trouessart in 1897. ''Ictitherium'' species were endemic to Africa and Eurasia during the Middle Miocene t ...
viverrinum'', which was similar to a
jackal Jackals are Canidae, canids native to Africa and Eurasia. While the word has historically been used for many canines of the subtribe Canina (subtribe), canina, in modern use it most commonly refers to three species: the closely related black-b ...
. 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
canid Canidae (; from Latin, ''canis'', "dog") is a family (biology), biological family of caniform carnivorans, constituting a clade. A member of this family is also called a canid (). The family includes three subfamily, subfamilies: the Caninae, a ...
s crossing the
Bering land bridge Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 72° north latitude in the Chukchi Sea; and on the south by the tip of the ...
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 The cheetah (''Acinonyx jubatus'') is a large Felidae, cat and the Fastest animals, fastest land animal. It has a tawny to creamy white or pale buff fur that is marked with evenly spaced, solid black spots. The head is small and rounded, wit ...
-like sprinter. Most of the dog-like hyenas had died off by 1.5 million years ago.


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
Percrocutidae Percrocutidae is an extinct family of hyena-like feliform carnivorans endemic to Asia, Africa, and Southern Europe from the Middle Miocene through the Pliocene, existing for about . The first percrocutids are known from the middle Miocene of E ...
. 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 The extinct Borophaginae form one of three subfamilies found within the canid family. The other two canid subfamilies are the extinct Hesperocyoninae and extant Caninae. Borophaginae, called "bone-crushing dogs", were endemic to North America d ...
. 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 ''Pachycrocuta'' is an extinct genus of hyena. The largest and most well-researched species is ''Pachycrocuta brevirostris'', colloquially known as the giant short-faced hyena as it stood about at the shoulder and it is estimated to have average ...
'', was a mega-scavenger that could splinter the bones of
elephant Elephants are the largest living land animals. Three living species are currently recognised: the African bush elephant ('' Loxodonta africana''), the African forest elephant (''L. cyclotis''), and the Asian elephant ('' Elephas maximus ...
s. Starting in the early
Middle Pleistocene The Chibanian, more widely known as the Middle Pleistocene (its previous informal name), is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocen ...
, ''Pachycrocuta'' was replaced by the smaller ''Crocuta'' and ''Hyena'', which corresponds to a general faunal change, perhaps in connection to the
Mid-Pleistocene transition The Mid-Pleistocene Transition (MPT), also known as the Mid-Pleistocene Revolution (MPR), is a fundamental change in the behaviour of glacial cycles during the Quaternary glaciations. The transition lasted around 550,000 years, from 1.25 million ...
.


Rise of modern 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 Terpenes () are a class of natural products consisting of compounds with the formula (C5H8)n for n ≥ 2. Terpenes are major biosynthetic building blocks. Comprising more than 30,000 compounds, these unsaturated hydrocarbons are produced predomi ...
excretions from soldier
termite Termites are a group of detritivore, detritophagous Eusociality, eusocial cockroaches which consume a variety of Detritus, decaying plant material, generally in the form of wood, Plant litter, leaf litter, and Humus, soil humus. They are dist ...
s is a modification of the strong digestive system its ancestors used to consume fetid carrion. The
striped hyena The striped hyena (''Hyaena hyaena'') is a species of hyena native to North and East Africa, the Middle East, the Caucasus, Central Asia, and the Indian subcontinent. It is the only extant species in the genus ''Hyaena''. It is listed by the IU ...
may have evolved from ''Hyaenictitherium namaquensis'' of
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
. 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 The Mediterranean Sea ( ) is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean basin and almost completely enclosed by land: on the east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
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 hyena The spotted hyena (''Crocuta crocuta''), also known as the laughing hyena, is a hyena species, currently classed as the sole extant member of the genus ''Crocuta'', native to sub-Saharan Africa. It is listed as being of least concern by the IUC ...
s in Asia at the end of the
Ice Age An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages, and g ...
. The striped hyena occurred for some time in
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
during the Pleistocene, having been particularly widespread in
France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
and
Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
. It also occurred in Montmaurin,
Hollabrunn Hollabrunn () is a district capital town in the Austrian state of Lower Austria, on the Göllersbach river. It is situated in the heart of the biggest wine region of Austria, the Weinviertel. History The surroundings of Hollabrunn were firs ...
in
Austria Austria, formally the Republic of Austria, is a landlocked country in Central Europe, lying in the Eastern Alps. It is a federation of nine Federal states of Austria, states, of which the capital Vienna is the List of largest cities in Aust ...
, the Furninha Cave in
Portugal Portugal, officially the Portuguese Republic, is a country on the Iberian Peninsula in Southwestern Europe. Featuring Cabo da Roca, the westernmost point in continental Europe, Portugal borders Spain to its north and east, with which it share ...
and the Genista Caves in
Gibraltar Gibraltar ( , ) is a British Overseas Territories, British Overseas Territory and British overseas cities, city located at the southern tip of the Iberian Peninsula, on the Bay of Gibraltar, near the exit of the Mediterranean Sea into the A ...
. The European form was similar in appearance to modern populations, but was larger, being comparable in size to the
brown hyena The brown hyena (''Parahyaena brunnea''), also called the strandwolf, is a species of hyena found in Namibia, Botswana, western and southern Zimbabwe, southern Mozambique and South Africa. It is the only extant species in the genus ''Parahyaena' ...
. The
spotted hyena The spotted hyena (''Crocuta crocuta''), also known as the laughing hyena, is a hyena species, currently classed as the sole extant member of the genus ''Crocuta'', native to sub-Saharan Africa. It is listed as being of least concern by the IUC ...
(''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. 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
carnassial Carnassials are paired upper and lower teeth modified in such a way as to allow enlarged and often self-sharpening edges to pass by each other in a shearing manner. This adaptation is found in carnivorans, where the carnassials are the modified f ...
s 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 A territory is an area of land, sea, or space, belonging or connected to a particular country, person, or animal. In international politics, a territory is usually a geographic area which has not been granted the powers of self-government, ...
, 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. Spotted hyenas spread from their original homeland during the
Middle Pleistocene The Chibanian, more widely known as the Middle Pleistocene (its previous informal name), is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocen ...
, and quickly colonised a very wide area from Europe, to southern Africa and
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
. 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 Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...
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 Duplication, duplicate, and duplicator may refer to: Biology and genetics * Gene duplication, a process which can result in free mutation * Chromosomal duplication, which can cause Bloom and Rett syndrome * Polyploidy, a phenomenon also known ...
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 Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding be ...
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)

The list follows McKenna and Bell's ''Classification of Mammals'' for prehistoric genera (1997)Malcolm C. McKenna, Susan K. Bell: ''Classification of Mammals: Above the Species Level'', Columbia University Press, New York 1997, 631 Seiten, and Wozencraft (2005) in Wilson and Reeders ''
Mammal Species of the World ''Mammal Species of the World: A Taxonomic and Geographic Reference'' is a standard reference work in mammalogy giving descriptions and Bibliographic database, bibliographic data for the known species of mammals. It is now in its third edition, ...
'' 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 Percrocutidae is an extinct family of hyena-like feliform carnivorans endemic to Asia, Africa, and Southern Europe from the Middle Miocene through the Pliocene, existing for about . The first percrocutids are known from the middle Miocene of E ...
, 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 ''Ictitherium'' (meaning "weasel beast") is an extinct genus belonging to the family Hyaenidae and the subfamily Ictitheriinae erected by Trouessart in 1897. ''Ictitherium'' species were endemic to Africa and Eurasia during the Middle Miocene t ...
'' (=''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 ''Ikelohyaena'' (from Greek language, Greek: ἴκελος ''íkelos'', 'like' and Latin: ''hyaena'', 'hyena') is an extinct genus of hyaenids that lived in Southern and Eastern Africa during the Pliocene. ''I. abronia'', the type and only known ...
'' (Early Pliocene of Africa) *** ''
Hyaena The striped hyena (''Hyaena hyaena'') is a species of hyena native to North and East Africa, the Middle East, the Caucasus, Central Asia, and the Indian subcontinent. It is the only extant species in the genus ''Hyaena''. It is listed by the IUC ...
'' (=''Euhyaena'',=''Parahyaena''; including
striped hyena The striped hyena (''Hyaena hyaena'') is a species of hyena native to North and East Africa, the Middle East, the Caucasus, Central Asia, and the Indian subcontinent. It is the only extant species in the genus ''Hyaena''. It is listed by the IU ...
, ''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 The brown hyena (''Parahyaena brunnea''), also called the strandwolf, is a species of hyena found in Namibia, Botswana, western and southern Zimbabwe, southern Mozambique and South Africa. It is the only extant species in the genus ''Parahyaena' ...
Pliocene to recent of Africa) *** †'' 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 ''Chasmaporthetes'', also known as hunting or running hyena, is an extinct genus of hyenas distributed in Eurasia, North America, and Africa during the Pliocene-Pleistocene epochs, living from 4.9 million to 780,000 years ago, existing for about ...
'' (=''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 ''Pachycrocuta'' is an extinct genus of hyena. The largest and most well-researched species is ''Pachycrocuta brevirostris'', colloquially known as the giant short-faced hyena as it stood about at the shoulder and it is estimated to have average ...
'' (Pliocene and Pleistocene of Eurasia and Africa) *** †'' Adcrocuta'' (Late Miocene of Eurasia) *** '' Crocuta'' (=''Crocotta''; including ''Eucrocuta'';
spotted hyena The spotted hyena (''Crocuta crocuta''), also known as the laughing hyena, is a hyena species, currently classed as the sole extant member of the genus ''Crocuta'', native to sub-Saharan Africa. It is listed as being of least concern by the IUC ...
and cave hyena. Late Pliocene to recent of Africa, Late Pliocene to Late Pleistocene of Eurasia) ** Subfamily Protelinae *** †'' Gansuyaena'' *** '' Proteles'' (=''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). A more recent molecular analysis agrees on the phylogenetic relationship between the four extant hyaenidae species (Koepfli ''et al'', 2006).


Characteristics


Build

Hyenas have relatively short torsos and are fairly massive and
wolf The wolf (''Canis lupus''; : wolves), also known as the grey wolf or gray wolf, is a Canis, canine native to Eurasia and North America. More than thirty subspecies of Canis lupus, subspecies of ''Canis lupus'' have been recognized, includin ...
-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 In terrestrial vertebrates, digitigrade ( ) locomotion is walking or running on the toes (from the Latin ''digitus'', 'finger', and ''gradior'', 'walk'). A digitigrade animal is one that stands or walks with its toes (phalanges) on the ground, and ...
, with the fore and hind paws having four digits each and sporting bulging pawpads. Like canids, hyenas have short, blunt, non-retractable claws. Their
pelage A fur is a soft, thick growth of hair that covers the skin of almost all mammals. It consists of a combination of oily guard hair on top and thick underfur beneath. The guard hair keeps moisture from reaching the skin; the underfur acts as an ...
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. With the exception of the spotted hyena, hyaenids have striped coats, which they likely inherited from their
viverrid Viverridae is a family (biology), family of small to medium-sized feliform mammals, comprising 14 genera with 33 species. This family was named and first described by John Edward Gray in 1821. Viverrids occur all over Africa, in southern Europe, ...
ancestors. Their ears are large and have simple basal ridges and no marginal bursa. Their
vertebral column The spinal column, also known as the vertebral column, spine or backbone, is the core part of the axial skeleton in vertebrates. The vertebral column is the defining and eponymous characteristic of the vertebrate. The spinal column is a segmente ...
, including the cervical region are of limited mobility. Hyenas have no
baculum The baculum (: bacula), also known as the penis bone, penile bone, ''os penis'', ''os genitale'', or ''os priapi'', is a bone in the penis of many placental mammals. It is not present in humans, but is present in the penises of some primates, ...
. Hyenas have one more pair of ribs than canids do, and their tongues are rough like those of felids and viverrids. 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 Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiology ...
is similar to that of the
canid Canidae (; from Latin, ''canis'', "dog") is a family (biology), biological family of caniform carnivorans, constituting a clade. A member of this family is also called a canid (). The family includes three subfamily, subfamilies: the Caninae, a ...
, but is more specialised for consuming coarse food and crushing bones. The
carnassial Carnassials are paired upper and lower teeth modified in such a way as to allow enlarged and often self-sharpening edges to pass by each other in a shearing manner. This adaptation is found in carnivorans, where the carnassials are the modified f ...
s, 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 The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
, are powerful, with broad bases and cutting edges. The canines are short, but thick and robust. Labiolingually, their
mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
s are much stronger at the canine teeth than in canids, reflecting the fact that hyenas crack bones with both their anterior dentition and
premolars The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mout ...
, 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. Daniel Johnson (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 Practiced by the Conjoors and Their Method of Curing Themselves when Bitten: with Remarks on Hydrophobia and Rabid Animals
' p. 45-46, R. Jennings, 1827
Stevenson-Hamilton, James (1917)
Animal life in Africa
Vol. 1'', p.95, London : William Heinemann
The spotted hyena is renowned for its strong bite proportional to its size, but a number of other animals (including the
Tasmanian devil The Tasmanian devil (''Sarcophilus harrisii''; palawa kani: ''purinina'') is a carnivorous marsupial of the family Dasyuridae. It was formerly present across mainland Australia, but became extinct there around 3,500 years ago; it is now con ...
) 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 Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiology ...
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 gland The anal glands or anal sacs are small glands near the anus in many mammals. They are situated in between the external anal sphincter muscle and internal anal sphincter muscle. In non-human mammals, the secretions of the anal glands contain most ...
s above the anus open into this pouch. Several
sebaceous gland A sebaceous gland or oil gland is a microscopic exocrine gland in the skin that opens into a hair follicle to secrete an oily or waxy matter, called sebum, which lubricates the hair and skin of mammals. In humans, sebaceous glands occur in ...
s 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 Soap is a salt (chemistry), salt of a fatty acid (sometimes other carboxylic acids) used for cleaning and lubricating products as well as other applications. In a domestic setting, soaps, specifically "toilet soaps", are surfactants usually u ...
or burning, and can be detected by humans several meters downwind. The secretions are primarily used for
territorial marking In ethology, territory is the sociographical area that an animal consistently defends against conspecific competition (or, occasionally, against animals of other species) using agonistic behaviors or (less commonly) real physical aggression. ...
, though both the aardwolf and the striped hyena 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
mustelid The Mustelidae (; from Latin , weasel) are a diverse family of carnivoran mammals, including weasels, badgers, otters, polecats, martens, grisons, and wolverines. Otherwise known as mustelids (), they form the largest family in the suborde ...
s, but not canids and
felid Felidae ( ) is the Family (biology), family of mammals in the Order (biology), order Carnivora colloquially referred to as cats. A member of this family is also called a felid ( ). The 41 extant taxon, extant Felidae species exhibit the gre ...
s. When attacked by lions or dogs, striped and brown hyenas will feign death, though the spotted hyena will defend itself ferociously. The spotted hyena is very vocal, producing a number of different sounds consisting of whoops, grunts, groans, lows, giggles, yells, growls, laughs and whines. The striped hyena is comparatively silent, its vocalizations being limited to a chattering laugh and howling. left, Whoop of a spotted hyena in Umfolosi Game Park, South Africa.
Mating In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. '' Copulation'' is the union of the sex organs of two sexually repr ...
between hyenas involves a number of short copulations with brief intervals, unlike canids, who generally engage in a single, drawn out copulation. Spotted hyena cubs are born almost fully developed, with their eyes open and erupting incisors and canines, though lacking adult markings. 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, though male striped hyenas do so. 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 Ungulates ( ) are members of the diverse clade Euungulata ("true ungulates"), which primarily consists of large mammals with hooves. Once part of the clade "Ungulata" along with the clade Paenungulata, "Ungulata" has since been determined to b ...
, 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
termite Termites are a group of detritivore, detritophagous Eusociality, eusocial cockroaches which consume a variety of Detritus, decaying plant material, generally in the form of wood, Plant litter, leaf litter, and Humus, soil humus. They are dist ...
s 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. 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. 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. 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 An antibody (Ab) or immunoglobulin (Ig) is a large, Y-shaped protein belonging to the immunoglobulin superfamily which is used by the immune system to identify and neutralize antigens such as bacteria and viruses, including those that caus ...
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


Folklore, mythology and literature

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
Muslim Muslims () are people who adhere to Islam, a Monotheism, monotheistic religion belonging to the Abrahamic religions, Abrahamic tradition. They consider the Quran, the foundational religious text of Islam, to be the verbatim word of the God ...
s who challenge the local
animism Animism (from meaning 'breath, spirit, life') is the belief that objects, places, and creatures all possess a distinct spiritual essence. Animism perceives all things—animals, plants, rocks, rivers, weather systems, human handiwork, and in ...
that exists among the Beng in Côte d’Ivoire. In
East Africa East Africa, also known as Eastern Africa or the East of Africa, is a region at the eastern edge of the Africa, African continent, distinguished by its unique geographical, historical, and cultural landscape. Defined in varying scopes, the regi ...
, 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 Tanzania, officially the United Republic of Tanzania, is a country in East Africa within the African Great Lakes region. It is bordered by Uganda to the northwest; Kenya to the northeast; the Indian Ocean to the east; Mozambique and Malawi to t ...
, there is a belief that
witch Witchcraft is the use of magic by a person called a witch. Traditionally, "witchcraft" means the use of magic to inflict supernatural harm or misfortune on others, and this remains the most common and widespread meaning. According to ''Enc ...
es use spotted hyenas as mounts. 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 feces 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. The Kaguru of Tanzania and the Kujamaat of southern
Senegal Senegal, officially the Republic of Senegal, is the westernmost country in West Africa, situated on the Atlantic Ocean coastline. It borders Mauritania to Mauritania–Senegal border, the north, Mali to Mali–Senegal border, the east, Guinea t ...
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 Mansôa is a town located in the Oio Region of Guinea-Bissau. Population 8,313 (2009 est). The Sua language is spoken in Mansôa. Geography Mansôa is located 60 km east of the capital Bissau Bissau () is the Capital (political), capital ...
. These " werehyenas" are killed when discovered, and do not revert to human form once dead. Striped hyenas are often referred to in
Middle East The Middle East (term originally coined in English language) is a geopolitical region encompassing the Arabian Peninsula, the Levant, Turkey, Egypt, Iran, and Iraq. The term came into widespread usage by the United Kingdom and western Eur ...
ern literature and folklore, typically as symbols of treachery and stupidity.Mounir R. Abi-Said (2006) ''Reviled as a grave robber: The ecology and conservation of striped hyaenas in the human dominated landscapes of Lebanon'' Ph.D. thesis, University of Kent (Biodiversity management) In the Near and Middle East, striped hyenas are generally regarded as physical incarnations of
jinn Jinn or djinn (), alternatively genies, are supernatural beings in pre-Islamic Arabian religion and Islam. Their existence is generally defined as parallel to humans, as they have free will, are accountable for their deeds, and can be either ...
s.Frembgen, Jürgen W.
The Magicality of the Hyena: Beliefs and Practices in West and South Asia
'', Asian Folklore Studies, Volume 57, 1998: 331–344
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. A
Persia Iran, officially the Islamic Republic of Iran (IRI) and also known as Persia, is a country in West Asia. It borders Iraq to the west, Turkey, Azerbaijan, and Armenia to the northwest, the Caspian Sea to the north, Turkmenistan to the nort ...
n medical treatise written in 1376 tells how to cure cannibalistic people known as ''kaftar'', who are said to be "half-man, half-hyena". al-Damīrī in his writings in ''Ḥawayān al-Kubrā'' (1406) wrote that striped hyenas were
vampiric A vampire is a mythical creature that subsists by feeding on the Vitalism, vital essence (generally in the form of blood) of the living. In European folklore, vampires are undead, undead humanoid creatures that often visited loved ones and c ...
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. In a similar vein to al-Damīrī, the
Greeks Greeks or Hellenes (; , ) are an ethnic group and nation native to Greece, Greek Cypriots, Cyprus, Greeks in Albania, southern Albania, Greeks in Turkey#History, Anatolia, parts of Greeks in Italy, Italy and Egyptian Greeks, Egypt, and to a l ...
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. The image of striped hyenas in
Afghanistan Afghanistan, officially the Islamic Emirate of Afghanistan, is a landlocked country located at the crossroads of Central Asia and South Asia. It is bordered by Pakistan to the Durand Line, east and south, Iran to the Afghanistan–Iran borde ...
, India and
Palestine Palestine, officially the State of Palestine, is a country in West Asia. Recognized by International recognition of Palestine, 147 of the UN's 193 member states, it encompasses the Israeli-occupied West Bank, including East Jerusalem, and th ...
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 North India is a geographical region, loosely defined as a cultural region comprising the northern part of India (or historically, the Indian subcontinent) wherein Indo-Aryans (speaking Indo-Aryan languages) form the prominent majority populati ...
, witches or magicians are said to ride striped hyenas at night. The striped hyena is mentioned in the
Bible The Bible is a collection of religious texts that are central to Christianity and Judaism, and esteemed in other Abrahamic religions such as Islam. The Bible is an anthology (a compilation of texts of a variety of forms) originally writt ...
. The
Arabic Arabic (, , or , ) is a Central Semitic languages, Central Semitic language of the Afroasiatic languages, Afroasiatic language family spoken primarily in the Arab world. The International Organization for Standardization (ISO) assigns lang ...
word for the hyena, ''ḍab`'' or ''ḍabu`'' (plural ''ḍibā`''), is alluded to in a valley in
Israel Israel, officially the State of Israel, is a country in West Asia. It Borders of Israel, shares borders with Lebanon to the north, Syria to the north-east, Jordan to the east, Egypt to the south-west, and the Mediterranean Sea to the west. Isr ...
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 Samuel is a figure who, in the narratives of the Hebrew Bible, plays a key role in the transition from the biblical judges to the United Kingdom of Israel under Saul, and again in the monarchy's transition from Saul to David. He is venera ...
13:18. In modern
Hebrew Hebrew (; ''ʿÎbrit'') is a Northwest Semitic languages, Northwest Semitic language within the Afroasiatic languages, Afroasiatic language family. A regional dialect of the Canaanite languages, it was natively spoken by the Israelites and ...
, the word for hyena and hypocrite are both the same: ''tsavua''. Though the
Authorized King James Version The King James Version (KJV), also the King James Bible (KJB) and the Authorized Version (AV), is an Early Modern English translation of the Christian Bible for the Church of England, which was commissioned in 1604 and published in 1611, by ...
of the Bible interprets the term "''`ayit tsavua`''" (found in
Jeremiah Jeremiah ( – ), also called Jeremias, was one of the major prophets of the Hebrew Bible. According to Jewish tradition, Jeremiah authored the Book of Jeremiah, book that bears his name, the Books of Kings, and the Book of Lamentations, with t ...
12:9) as "speckled bird",
Henry Baker Tristram Henry Baker Tristram FRS (11 May 1822 – 8 March 1906) was an English clergyman, Bible scholar, traveller and ornithologist. As a parson-naturalist he was an early, but short-lived, supporter of Darwinism, attempting to reconcile evolution an ...
argued that it was most likely a hyena being mentioned. The vocalization of the spotted hyena resembling hysterical human
laughter Laughter is a pleasant physical reaction and emotion consisting usually of rhythmical, usually audible contractions of the diaphragm and other parts of the respiratory system. It is a response to certain external or internal stimuli. Laug ...
has been alluded to in numerous works of literature: "to laugh like a hyæna" or a "hyen" was a common
simile A simile () is a type of figure of speech that directly ''compares'' two things. Similes are often contrasted with metaphors, where similes necessarily compare two things using words such as "like", "as", while metaphors often create an implicit c ...
, and is featured in ''The Cobbler's Prophecy'' (1594), Webster's ''
Duchess of Malfi ''The Duchess of Malfi'' (originally published as ''The Tragedy of the Dutchesse of Malfy'') is a Jacobean revenge tragedy written by English dramatist John Webster in 1612–1613. It was first performed privately at the Blackfriars Theatre, ...
'' (1623) and
Shakespeare William Shakespeare ( 23 April 1564 – 23 April 1616) was an English playwright, poet and actor. He is widely regarded as the greatest writer in the English language and the world's pre-eminent dramatist. He is often called England's natio ...
's ''
As You Like It ''As You Like It'' is a pastoral Shakespearean comedy, comedy by William Shakespeare believed to have been written in 1599 and first published in the First Folio in 1623. The play's first performance is uncertain, though a performance at Wil ...
'', Act IV. Sc.1. ''Die Strandjutwolf'' (The
brown hyena The brown hyena (''Parahyaena brunnea''), also called the strandwolf, is a species of hyena found in Namibia, Botswana, western and southern Zimbabwe, southern Mozambique and South Africa. It is the only extant species in the genus ''Parahyaena' ...
) is an
allegorical As a literary device or artistic form, an allegory is a narrative or visual representation in which a character, place, or event can be interpreted to represent a meaning with moral or political significance. Authors have used allegory throughou ...
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 Siberia ( ; , ) is an extensive geographical region comprising all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has formed a part of the sovereign territory of Russia and its predecessor states ...
was a significant factor in delaying human colonization of
Alaska Alaska ( ) is a non-contiguous U.S. state on the northwest extremity of North America. Part of the Western United States region, it is one of the two non-contiguous U.S. states, alongside Hawaii. Alaska is also considered to be the north ...
. 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 The Bering Strait ( , ; ) is a strait between the Pacific and Arctic oceans, separating the Chukchi Peninsula of the Russian Far East from the Seward Peninsula of Alaska. The present Russia–United States maritime boundary is at 168° 58' ...
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 The Red Terror () was a campaign of political repression and Mass killing, executions in Russian Soviet Federative Socialist Republic, Soviet Russia which was carried out by the Bolsheviks, chiefly through the Cheka, the Bolshevik secret police ...
. 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 The Second Sudanese Civil War was a conflict from 1983 to 2005 between the central Sudanese government and the Sudan People's Liberation Army/Movement, Sudan People's Liberation Army. It was largely a continuation of the First Sudanese Civil Wa ...
, 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.Begg, Colleen, Begg, Keith & Muemedi, Oscar (2007)
Preliminary data on human - carnivore conflict in Niassa National Reserve, Mozambique, particularly fatalities due to lion, spotted hyaena and crocodile
'', SGDRN (Sociedade para a Gestão e Desenvolvimento da Reserva do Niassa Moçambique)
Man-eating spotted hyenas tend to be very large specimens; a pair of man-eating hyenas, responsible for killing 27 people in
Mulanje Mulanje, formerly called Mlanje, is a town in the Southern Region, Malawi, Southern Region of Malawi, close to the border with Mozambique, to the east. It is near the Mulanje Massif. History A mission was established here and it was at first on ...
,
Malawi Malawi, officially the Republic of Malawi, is a landlocked country in Southeastern Africa. It is bordered by Zambia to the west, Tanzania to the north and northeast, and Mozambique to the east, south, and southwest. Malawi spans over and ...
in 1962, weighed in at 72 kg (159 lb) and 77 kg (170 lb) after being shot.Kruuk, Hans (2002) ''Hunter and hunted: relationships between carnivores and people'' Cambridge University Press, A 1903 report describes spotted hyenas in the
Mzimba district Mzimba is a district in the Northern Region of Malawi. The capital is Mzimba and the biggest town is Mzuzu, which is also the administrative headquarters of the Northern Region. The district covers an area of 10,473 km2 and has a populatio ...
of Angoniland waiting at dawn outside people's huts to attack them when they opened their doors. Victims of spotted hyenas tend to be women, children and sick or infirm men;
Theodore Roosevelt Theodore Roosevelt Jr. (October 27, 1858 – January 6, 1919), also known as Teddy or T.R., was the 26th president of the United States, serving from 1901 to 1909. Roosevelt previously was involved in New York (state), New York politics, incl ...
wrote in 1908–1909 in
Uganda Uganda, officially the Republic of Uganda, is a landlocked country in East Africa. It is bordered to the east by Kenya, to the north by South Sudan, to the west by the Democratic Republic of the Congo, to the south-west by Rwanda, and to the ...
that spotted hyenas regularly killed sufferers of African sleeping sickness as they slept outside in camps. 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 Mozambique, officially the Republic of Mozambique, is a country located in Southeast Africa bordered by the Indian Ocean to the east, Tanzania to the north, Malawi and Zambia to the northwest, Zimbabwe to the west, and Eswatini and South Afr ...
along a 20-km stretch of road near the
Tanzania Tanzania, officially the United Republic of Tanzania, is a country in East Africa within the African Great Lakes region. It is bordered by Uganda to the northwest; Kenya to the northeast; the Indian Ocean to the east; Mozambique and Malawi to t ...
n border. In the 1880s, a hyena was reported to have attacked humans, especially sleeping children, over a three-year period in the
Iğdır Province Iğdır Province (, , , ) is a Provinces of Turkey, province in eastern Turkey, located along the borders with Armenia, Azerbaijan (the area of Nakhchivan Autonomous Republic), and Iran. Its adjacent provinces are Kars Province, Kars to the north ...
of Turkey, with 25 children and 3 adults being wounded in one year. The attacks provoked local authorities into announcing a reward of 100
ruble The ruble or rouble (; rus, рубль, p=rublʲ) is a currency unit. Currently, currencies named ''ruble'' in circulation include the Russian ruble (RUB, ₽) in Russia and the Belarusian ruble (BYN, Rbl) in Belarus. These currencies are s ...
s for every hyena killed. Further attacks were reported later in some parts of the
South Caucasus The South Caucasus, also known as Transcaucasia or the Transcaucasus, is a geographical region on the border of Eastern Europe and West Asia, straddling the southern Caucasus Mountains. The South Caucasus roughly corresponds to modern Armenia, ...
, particularly in 1908. Instances are known in
Azerbaijan Azerbaijan, officially the Republic of Azerbaijan, is a Boundaries between the continents, transcontinental and landlocked country at the boundary of West Asia and Eastern Europe. It is a part of the South Caucasus region and is bounded by ...
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 Turkmenistan's Bathyz Nature Reserve. A further attack on a child was reported around Serakhs in 1948. Several attacks have occurred in India; in 1962, 9 children were thought to have been taken by hyenas in the town of
Bhagalpur Bhagalpur, historically known as Champapuri, Champa Nagari, is a city in the Indian state of Bihar, situated on the southern bank of the Ganges river. It is the Bihar#Government and administration, third largest city of Bihar by population and ...
in the
Bihar Bihar ( ) is a states and union territories of India, state in Eastern India. It is the list of states and union territories of India by population, second largest state by population, the List of states and union territories of India by are ...
State in a six-week period, and 19 children up to the age of four were killed by hyenas in
Karnataka Karnataka ( ) is a States and union territories of India, state in the southwestern region of India. It was Unification of Karnataka, formed as Mysore State on 1 November 1956, with the passage of the States Reorganisation Act, 1956, States Re ...
in 1974. A survey of wild animal attacks during a five-year period in the Indian state of
Madhya Pradesh Madhya Pradesh (; ; ) is a state in central India. Its capital is Bhopal and the largest city is Indore, Indore. Other major cities includes Gwalior, Jabalpur, and Sagar, Madhya Pradesh, Sagar. Madhya Pradesh is the List of states and union te ...
reported that hyenas had attacked three people, causing fewer deaths than
wolves The wolf (''Canis lupus''; : wolves), also known as the grey wolf or gray wolf, is a canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, including the dog and dingo, though gr ...
,
gaur The gaur (''Bos gaurus''; ) is a large bovine native to the Indian Subcontinent and Southeast Asia, and has been listed as Vulnerable species, Vulnerable on the IUCN Red List since 1986. The global population was estimated at a maximum of 21,000 ...
,
boar The wild boar (''Sus scrofa''), also known as the wild swine, common wild pig, Eurasian wild pig, or simply wild pig, is a Suidae, suid native to much of Eurasia and North Africa, and has been introduced to the Americas and Oceania. The speci ...
,
elephants Elephants are the Largest and heaviest animals, largest living land animals. Three living species are currently recognised: the African bush elephant (''Loxodonta africana''), the African forest elephant (''L. cyclotis''), and the Asian ele ...
, tigers, leopards and
sloth bear The sloth bear (''Melursus ursinus''), also known as the Indian bear, is a myrmecophagous bear species native to the Indian subcontinent. It feeds on fruits, ants and termites. It is listed as vulnerable on the IUCN Red List, mainly because of ...
s.


Hyenas as food and medicine

Hyenas have occasionally been used for food and medicinal purposes in
Somalia Somalia, officially the Federal Republic of Somalia, is the easternmost country in continental Africa. The country is located in the Horn of Africa and is bordered by Ethiopia to the west, Djibouti to the northwest, Kenya to the southwest, th ...
. Some Muslims consider it
halal ''Halal'' (; ) is an Arabic word that translates to in English. Although the term ''halal'' is often associated with Islamic dietary laws, particularly meat that is slaughtered according to Islamic guidelines, it also governs ethical practices ...
in
Islam Islam is an Abrahamic religions, Abrahamic monotheistic religion based on the Quran, and the teachings of Muhammad. Adherents of Islam are called Muslims, who are estimated to number Islam by country, 2 billion worldwide and are the world ...
although this is disputed by other Muslims.ilmhub.com
/ref> This practice dates back to the times of the
Ancient Greeks Ancient Greece () was a northeastern Mediterranean civilization, existing from the Greek Dark Ages of the 12th–9th centuries BC to the end of classical antiquity (), that comprised a loose collection of culturally and linguistically re ...
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


Citations


General and cited references

* * * * * * * *


Further reading

* Funk, Holdger (2010) ''Hyaena: On the Naming and Localisation of an Enigmatic Animal'', GRIN Verlag, . * 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.


External links


IUCN Conservation Union Hyaena Specialist Group
{{DEFAULTSORT:Hyena   Mammals of Asia Scavengers Extant Miocene first appearances Taxa named by John Edward Gray eo:Hieno