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The spotted hyena (Crocuta crocuta), also known as the laughing hyena,[3] is a species of hyena, currently classed as the sole member of the genus Crocuta, native to Sub-Saharan Africa. It is listed as being of least concern by the IUCN
IUCN
on account of its widespread range and large numbers estimated between 27,000 and 47,000 individuals.[1] The species is, however, experiencing declines outside of protected areas due to habitat loss and poaching.[1] The species may have originated in Asia,[4] and once ranged throughout Europe
Europe
for at least one million years until the end of the Late Pleistocene.[5] The spotted hyena is the largest known member of the Hyaenidae, and is further physically distinguished from other species by its vaguely bear-like build,[6] its rounded ears,[7] its less prominent mane, its spotted pelt,[8] its more dual purposed dentition,[9] its fewer nipples[10] and the presence of a pseudo-penis in the female. It is the only mammalian species to lack an external vaginal opening.[11] The spotted hyena is the most social of the Carnivora
Carnivora
in that it has the largest group sizes and most complex social behaviours.[12] Its social organisation is unlike that of any other carnivore, bearing closer resemblance to that of cercopithecine primates (baboons and macaques) with respect to group-size, hierarchical structure, and frequency of social interaction among both kin and unrelated group-mates.[13] However, the social system of the spotted hyena is openly competitive rather than cooperative, with access to kills, mating opportunities and the time of dispersal for males depending on the ability to dominate other clan-members. Females provide only for their own cubs rather than assist each other, and males display no paternal care. Spotted hyena
Spotted hyena
society is matriarchal; females are larger than males, and dominate them.[14] The spotted hyena is a highly successful animal, being the most common large carnivore in Africa. Its success is due in part to its adaptability and opportunism; it is primarily a hunter but may also scavenge, with the capacity to eat and digest skin, bone and other animal waste. In functional terms, the spotted hyena makes the most efficient use of animal matter of all African carnivores.[15] The spotted hyena displays greater plasticity in its hunting and foraging behaviour than other African carnivores;[16] it hunts alone, in small parties of 2–5 individuals or in large groups. During a hunt, spotted hyenas often run through ungulate herds in order to select an individual to attack. Once selected, their prey is chased over a long distance, often several kilometres, at speeds of up to 60 km/h.[17] The spotted hyena has a long history of interaction with humanity; depictions of the species exist from the Upper Paleolithic
Upper Paleolithic
period, with carvings and paintings from the Lascaux
Lascaux
and Chauvet Caves.[18] The species has a largely negative reputation in both Western culture and African folklore. In the former, the species is mostly regarded as ugly and cowardly, while in the latter, it is viewed as greedy, gluttonous, stupid, and foolish, yet powerful and potentially dangerous. The majority of Western perceptions on the species can be found in the writings of Aristotle
Aristotle
and Pliny the Elder, though in relatively unjudgemental form. Explicit, negative judgements occur in the Physiologus, where the animal is depicted as a hermaphrodite and grave robber.[19] The IUCN's hyena specialist group identifies the spotted hyena's negative reputation as detrimental to the species' continued survival, both in captivity and the wild.[19][20]

Contents

1 Etymology, discovery and naming

1.1 Local and indigenous names

2 Taxonomy, origins and evolution 3 Description

3.1 Anatomy 3.2 Dimensions 3.3 Fur 3.4 Female genitalia

4 Behaviour

4.1 Social behaviour 4.2 Mating, reproduction, and development 4.3 Denning behaviour 4.4 Intelligence 4.5 Hunting behaviour

5 Ecology

5.1 Diet 5.2 Enemies and competitors

6 Communication

6.1 Body language 6.2 Vocalisations

7 Diseases and parasites 8 Range, habitat and population 9 Relationships with humans

9.1 Cultural depictions and perceptions

9.1.1 In prehistory 9.1.2 In Africa 9.1.3 In Western culture

9.2 Livestock predation 9.3 Attacks on humans and grave desecration 9.4 Urban hyenas 9.5 Hunting and use in traditional medicine 9.6 Spotted hyenas in captivity and as pets

10 See also 11 References

11.1 Notes 11.2 Bibliography

12 Further reading 13 External links

Etymology, discovery and naming[edit] The spotted hyena's scientific name Crocuta, was once widely thought to be derived from the Latin
Latin
loanword crocutus, which translates as "saffron-coloured one", in reference to the animal's fur colour. This was proven to be incorrect, as the correct spelling of the loanword would have been Crocāta, and the word was never used in that sense by Graeco-Roman sources. Crocuta actually comes from the Ancient Greek word Κροκόττας (Krokottas), which is derived from the Sanskrit
Sanskrit
koṭṭhâraka, which in turn originates from kroshṭuka (both of which were originally meant to signify the golden jackal). The earliest recorded mention of Κροκόττας is from Strabo's Geographica, where the animal is described as a mix of wolf and dog native to Ethiopia.[21]

Engraving of a spotted hyena from Thomas Pennant's History of Quadrupeds, one of the first authentic depictions of the species[22]

From Classical antiquity
Classical antiquity
until the Renaissance, the spotted and striped hyena were either assumed to be the same species, or distinguished purely on geographical, rather than physical grounds. Hiob Ludolf, in his Historia aethiopica, was the first to clearly distinguish the Crocuta from Hyaena
Hyaena
on account of physical, as well as geographical grounds, though he never had any first hand experience of the species, having gotten his accounts from an Ethiopian intermediary.[3] Confusion still persisted over the exact taxonomic nature of the hyena family in general, with most European travelers in Ethiopia
Ethiopia
referring to hyenas as "wolves". This partly stems from the Amharic
Amharic
word for hyena, ጅብ (jɨbb), which is linked to the Arabic word ذئب (dhiʾb) "wolf".[23] The first detailed first-hand descriptions of the spotted hyena by Europeans come from Willem Bosman and Peter Kolbe. Bosman, a Dutch tradesman who worked for the Dutch West India Company
Dutch West India Company
at the Gold Coast
Coast
(modern day Ghana) from 1688–1701, wrote of "Jakhals, of Boshond" (jackals or woodland dogs) whose physical descriptions match the spotted hyena. Kolben, a German mathematician and astronomer who worked for the Dutch East India Company
Dutch East India Company
in the Cape of Good Hope
Cape of Good Hope
from 1705–1713, described the spotted hyena in great detail, but referred to it as a "tigerwolf", because the settlers in southern Africa did not know of hyenas, and thus labelled them as "wolves".[24] Bosman and Kolben's descriptions went largely unnoticed until 1771, when the Welsh naturalist Thomas Pennant, in his Synopsis of Quadrupeds, used the descriptions, as well as his personal experience with a captive specimen, as a basis for consistently differentiating the spotted hyena from the striped. The description given by Pennant was precise enough to be included by Johann Erxleben in his Systema regni animalis by simply translating Pennant's text into Latin. Crocuta was finally recognised as a separate genus from Hyaena
Hyaena
in 1828.[25] Local and indigenous names[edit] Several languages of Africa lack species specific names for hyenas: for example, the spotted and striped species have identical names in Dyula, Swahili, Mandinka, Mossi, Ngambaye, Wolof and Fulani. In other languages, other species may simply be termed "small spotted hyena", such as in Swahili, where the spotted hyena is termed fisi and the aardwolf fisi ndogo.[26]

Indigenous names for Crocuta crocuta[27][28]

Linguistic group or area Indigenous name

Afrendille Walaba

Afrikaans Gevlekte hiëna

Amharic ጅብ (Djibb)

Arabic Marfaïn (CAR and Chad) D'ba (North Africa) Dibb (Ethiopia) Guervave (Hassānīya)

Ateso Ibuin

Avukaia Labagu

Babouté Mangou

Baguirmien Niougo

Baka Libagu

Bakola Massobé

Bambara Namakoro Suruku

Banda Bongo

Bechuana Piri Phiri

Bemba (Zambia) Chimbwi

Bornouan Boultou

Creole Lobo

Danakil Jangóula

Dioula Suruku Namakoro

Elkoni Makatiet nyenegea

English Spotted hyena

French Hyène tachetée

Fula (Fulfulde/Pulaar/Pular) Bonooru Fowru Deppuru

Galaorabéjsa Wårabéssa Orabéjsa

Gambe Mangili

Ganda and Runyoro Empisi

German Tüpfelhyäne Fleckenhyäne

Gourage Woraba

Gourmatche Namlino

Harari ዎራባ (Worábba)

Hausa Kura

Herero Mbúngu-mbidíwa

Ila (Zambia) Kabwenga

Jita Imembe

Kalenjin Kimatet

Kaonde Mungolwe

Karamojong Ebu Etutui

Kichagga Ingurunju Ifulu

Kigogo and kikongo Misi

Kikondo Mbulu

Kiliangulu Warabes

Kiluba Kimburi

Kimeru Mbitingaau

Kinyarwanda Impysi

Kinyaturo Mpiti

Kinyiha Ipatama

Kipare
Kipare
and Kizigua Ibau

Kirangi Mbichi

Kisukuma, Kikamba and Kimaragoli Mbiti

Kisungwa Fifi

Kota Massoba

Kotoko Machi

Kunda Tika

Kitaita Mbisi

Lugbara Rara

Luhya Namuyu

Luo Otoyo

Lwo Lagwara

Madi Ebowu

Malinke Namakoro Souroukou

Mambakushu Dimbugurumba

Manding Tourouma

Mangbetu Neunga

Masai Ondilili Oln'gojine

Maure Chertat Gaboune Gougouh

Mboko Assoba

M'boum Baglak

Mondo Lepagu

Mongom Massobe

Moore Katre Swasa

Ngambaye Riguen ndah

Nkoya (Zambia) Muntambwi

Nsenga (Zambia) Chimbwe

Njanja (Zambia) Fisi

Ovambo kafukambungu Mbungu-omanini

Ovacuangari and Ovadirico Divúndu

Portuguese Hiena-malhada

Runyoro Empisi

Sara Nyéyi

Sebei Mangatiet

Serer Omone

seTswana Phiri LeHolo

Shona Bere

Somali Warabe

Songhai Koro

Soninke Tourouguė

Sotho (northern and southern) Phiri

Swahili Fisi Nyangao

Tigrinya ዝብእ (Zibb-i)

Tonga (Zambia) Suntwe

Tsonga Mhisi

Tumbuka (Zambia) Chimbwi

Twi Pataku

Wolof Bouki

Xhosa Impisi Mpisi IsAndawane

Yoruba Kòriko Ikookò

Zande Ngini Nzege

Taxonomy, origins and evolution[edit]

Pair of spotted hyenas at White River, Mpumalanga. Note the great degree of individual variation in fur colour, which was once used as a basis for separating the species into various subspecies.

Unlike the striped hyena, for which a number of subspecies were proposed in light of its extensive modern range, the spotted hyena is a genuinely variable species, both temporally and spatially. Its range once encompassed almost all of Africa and Eurasia, and displayed a large degree of morphological geographic variation, which led to an equally extensive set of specific and subspecific epithets. It was gradually realised that all of this variation could be applied to individual differences in a single subspecies. In 1939, biologist L. Harrison Matthews demonstrated through comparisons between a large selection of spotted hyena skulls from Tanzania
Tanzania
that all the variation seen in the then recognised subspecies could also be found in a single population, with the only set of characters standing out being pelage (which is subject to a high degree of individual variation) and size (which is subject to Bergmann's Rule). When fossils are taken into consideration, the species displayed even greater variation than it does in modern times, and a number of these named fossil species have since been classed as synonymous with Crocuta crocuta, with firm evidence of there being more than one species within the genus Crocuta still lacking.[28]

Skull of Crocuta sivalensis, an extinct Indian hyena proposed by Björn Kurtén as being the ancestor of the modern spotted hyena

Both Björn Kurtén and Camille Arambourg promoted an Asiatic origin for the species; Kurtén focussed his arguments on the Plio-Pleistocene taxon Crocuta sivalensis from the Siwaliks,[4] a view defended by Arambourg, who nonetheless allowed the possibility of an Indo-Ethiopian origin.[29] This stance was contested by Ficarelli and Torre, who referred to evidence of the spotted hyena's presence from African deposists dating from the early Pleistocene, a similar age to the Asian C. sivalensis.[30] Studies on the phylogeographic distribution of mtDNA haplotypes indicates three migration events from Africa to Eurasia, though neither the topology of the phylogenetic tree or the fossil record exclude the possibility of an Asian origin. The earliest migration of spotted hyenas from Africa to Eurasia
Eurasia
began less than 3.5 million years ago, most probably from the area where the first spotted hyena fossils were discovered, reaching East Asia
Asia
and most likely also Pakistan. The second migration of spotted hyenas occurred less than 1.3–1.5 million years ago and resulted in the first arrival of hyenas in Europe
Europe
and a separation of African spotted hyenas into a southern and a northern population. The third spotted hyena migration took place 0.36 million years ago, starting from the northern African population and reaching both Europe
Europe
and Asia. Unlike other African carnivores, with the exception of the leopard, there is no evidence to suggest that spotted hyenas underwent a genetic bottleneck during the Pleistocene.[31] The ancestors of the genus Crocuta diverged from Hyaena
Hyaena
(the genus of striped and brown hyenas) 10 million years ago.[31][32] The ancestors of the spotted hyena probably developed social behaviours in response to increased pressure from other predators on carcasses, which forced them to operate in teams. At one point in their evolution, spotted hyenas developed sharp carnassials behind their crushing premolars; this rendered waiting for their prey to die no longer a necessity, as is the case for brown and striped hyenas, and thus became pack hunters as well as scavengers. They began forming increasingly larger territories, necessitated by the fact that their prey was often migratory, and long chases in a small territory would have caused them to encroach into another clan's land.[9] It has been theorised that female dominance in spotted hyena clans could be an adaptation in order to successfully compete with males on kills, and thus ensure that enough milk is produced for their cubs.[14] Another theory is that it is an adaptation to the length of time it takes for cubs to develop their massive skulls and jaws, thus necessitating greater attention and dominating behaviours from females.[33] Its appearance in Europe
Europe
and China during the Cromerian
Cromerian
period coincided with the decline and eventual extinction of Pachycrocuta
Pachycrocuta
brevirostris, the giant short-faced hyena. As there is no evidence of environmental change being responsible, it is likely that the giant short-faced hyena became extinct due to competition with the spotted hyena.[34] Description[edit]

Skeleton

Spotted hyena
Spotted hyena
walking in profile

Skull, as illustrated by Frédéric Cuvier. Note the disproportionately large carnassials and premolars adapted for bone consumption.

Anatomy[edit] The spotted hyena has a strong and well developed neck and forequarters, but relatively underdeveloped hindquarters. The rump is rounded rather than angular, which prevents attackers coming from behind from getting a firm grip on it.[35] The head is wide and flat with a blunt muzzle and broad rhinarium. In contrast to the striped hyena, the ears of the spotted hyena are rounded rather than pointed. Each foot has four digits, which are webbed and armed with short, stout and blunt claws. The paw-pads are broad and very flat, with the whole undersurface of the foot around them being naked. The tail is relatively short, being 300–350 mm (12–14 in) long,[7] and resembles a pompom in appearance.[6] Unusually among hyaenids, and mammals in general, the female spotted hyena is considerably larger than the male.[36] Both sexes have a pair of anal glands which open into the rectum just inside the anal opening. These glands produce a white, creamy secretion which is pasted onto grass stalks by everting the rectum. The odour of this secretion is very strong, smelling of boiling cheap soap or burning, and can be detected by humans several metres downwind.[37] The spotted hyena has a proportionately large heart, constituting close to 1% of its body weight, thus giving it great endurance in long chases. In contrast, a lion's heart makes up only 0.45–0.57 percent of its body weight.[38] The now extinct Eurasian populations were distinguished from the modern African populations by their shorter distal extremities and longer humerus and femur.[39] The skull of the spotted hyena differs from that of the striped hyena by its much greater size and narrower sagittal crest. For its size, the spotted hyena has one of the most powerfully built skulls among the Carnivora.[40] The dentition is more dual purposed than that of other modern hyena species, which are mostly scavengers; the upper and lower third premolars are conical bone-crushers, with a third bone-holding cone jutting from the lower fourth premolar. The spotted hyena also has its carnassials situated behind its bone-crushing premolars, the position of which allows it to crush bone with its premolars without blunting the carnassials.[9] Combined with large jaw muscles and a special vaulting to protect the skull against large forces, these characteristics give the spotted hyena a powerful bite which can exert a pressure of 80 kgf/cm2 (1140 lbf/in²),[41] which is 40% more force than a leopard can generate.[42] The jaws of the spotted hyena outmatch those of the brown bear in bonecrushing ability,[43] and free ranging hyenas have been observed to crack open the long bones of giraffes measuring 7 cm in diameter.[44] A 63.1 kg (139 lb) spotted hyena is estimated to have a bite force of 565.7 newtons at the canine tip and 985.5 newtons at the carnassial eocone.[45] One individual in a study was found to exert a bite force of 4,500 newtons on the measuring instruments.[46] Dimensions[edit] The spotted hyena is the largest extant member of the Hyaenidae.[47] Adults measure 95–165.8 cm (37–65 in) in body length, and have a shoulder height of 70–91.5 cm (28–36 in).[27] Adult male spotted hyenas in the Serengeti
Serengeti
weigh 40.5–55.0 kg (89–121 lb), while females weigh 44.5–63.9 kg (98–141 lb). Spotted hyenas in Zambia
Zambia
tend to be heavier, with males weighing on average 67.6 kg (149 lb), and females 69.2 kg (153 lb).[36] Exceptionally large weights of 81.7 kg (180 lb)[9] and 90 kg (198 lb)[27] are known. It has been estimated that adult members of the now extinct Eurasian populations weighed 102 kg (225 lb).[48] Fur[edit] Fur colour varies greatly and changes with age.[35] Unlike the fur of the striped and brown hyena, that of the spotted hyena consists of spots rather than stripes and is much shorter, lacking the well defined spinal mane of the former two species.[8] The base colour generally is a pale greyish-brown or yellowish-grey on which an irregular pattern of roundish spots is superimposed on the back and hind quarters. The spots, which are of variable distinction, may be reddish, deep brown or almost blackish. The spots vary in size, even on single individuals, but are commonly 20 mm (0.79 in) in diameter. A less distinct spot pattern is present on the legs and belly but not on the throat and chest. A set of five, pale and barely distinct bands replace the spots on the back and sides of the neck. A broad, medial band is present on the back of the neck, and is lengthened into a forward facing crest. The crest is mostly reddish-brown in colour. The crown and upper part of the face is brownish, save for a white band above both eyes, though the front of the eyes, the area around the rhinarium, the lips and the back portion of the chin are all blackish. The limbs are spotted, though the feet vary in colour, from light brown to blackish. The fur is relatively sparse and consists of two hair types; moderately fine underfur (measuring 15–20 mm (0.59–0.79 in)) and long, stout bristle hairs (30–40 mm (1.2–1.6 in)).[7] European Paleolithic rock art depicting the species indicates that the Eurasian populations retained the spots of their modern-day African counterparts.[18] Female genitalia[edit] Further information: Clitoris
Clitoris
§ Spotted hyenas

Male and female reproductive systems of the spotted hyena, from Schmotzer & Zimmerman, Anatomischer Anzeiger (1922). Abb. 1 (Fig. 1.) Male reproductive anatomy. Abb. 2 (Fig. 2.) Female reproductive anatomy.[49] Principal abbreviations (from von Eggeling) are: T, testis; Vd, vas deferens; BU, urethral bulb; Ur, urethra; R, rectum; P, penis; S, scrotum; O, ovary; FT, tuba Fallopii; RL, ligament uteri; Ut, uterus; CC, Corpus clitoris. Remaining abbreviations, in alphabetical order, are: AG, parotid analis; B, vesica urinaria; CG, parotid Cowperi; CP, Corpus penis; CS, corpus spongiosum; GC, glans; GP, glans penis; LA, levator ani muscle; Pr, prepuce; RC, musculus retractor clitoris; RP, Musculus retractor penis; UCG, Canalis urogenital.

The genitalia of the female closely resembles that of the male; the clitoris is shaped and positioned like a penis, and is capable of erection. The female also possesses no external vagina (vaginal opening), as the labia are fused to form a pseudo-scrotum. The pseudo-penis is traversed to its tip by a central urogenital canal, through which the female urinates, copulates and gives birth.[50][51] The pseudo-penis can be distinguished from the males' genitalia by its greater thickness and more rounded glans.[11][52][53] In both males and females, the base of the glans is covered with penile spines.[54][55] The formation of the pseudo-penis appears largely androgen independent, as the pseudo-penis appears in the female fetus before differentiation of the fetal ovary and adrenal gland.[11] After parturition, the pseudo-penis is stretched, and loses many of its original aspects; it becomes a slack-walled and reduced prepuce with an enlarged orifice with split lips.[40] Behaviour[edit] Social behaviour[edit] Spotted hyenas are social animals which live in large communities (referred to as "clans") which can consist of at most 80 individuals.[56] Group-size varies geographically; in the Serengeti, where prey is migratory, clans are smaller than those in the Ngorongoro Crater, where prey is sedentary.[57] Spotted hyena
Spotted hyena
clans are more compact and unified than wolf packs, but are not as closely knit as those of African wild dogs.[58] Females dominate males, with even the lowest ranking females being dominant over the highest ranking males. It is typical for females to remain with their natal clan, thus large clans usually contain several matrilines, whereas males typically disperse from their natal clan at the age of 2½ years. The clan is a fission-fusion society, in which clan-members do not often remain together, but may forage alone or in small groups.[59] High-ranking hyenas maintain their position through aggression directed against lower-ranking clan-members.[10] Spotted hyena hierarchy is nepotistic; the offspring of dominant females automatically outrank adult females subordinate to their mother.[60] However, rank in spotted hyena cubs is greatly dependent on the presence of the mother; low-ranking adults may act aggressively toward higher-ranking cubs when the mother is absent. Although individual spotted hyenas only care for their own young, and males take no part in raising their young, cubs are able to identify relatives as distantly related as great-aunts. Also, males associate more closely with their own daughters rather than unrelated cubs, and the latter favour their fathers by acting less aggressively toward them.[13] Spotted hyena
Spotted hyena
societies are more complex than those of other carnivorous mammals, and are remarkably similar to those of cercopithecine primates in respect to group size, structure, competition and cooperation. Like cercopithecine primates, spotted hyenas use multiple sensory modalities, recognise individual conspecifics, are conscious that some clan-mates may be more reliable than others, recognise third-party kin and rank relationships among clan-mates, and adaptively use this knowledge during social decision making. Also, like cercopithecine primates, dominance ranks in hyena societies are not correlated with size or aggression, but with ally networks.[10][13] In this latter trait, the spotted hyena further show parallels with primates by acquiring rank through coalition. However, rank reversals and overthrows in spotted hyena clans are very rare.[10] The social network dynamics of spotted hyenas are determined by multiple factors.[61] Environmental factors include rainfall and prey abundance; individual factors include preference to bond with females and with kin; and topological effects include the tendency to close triads in the network. Female hyenas are more flexible than males in their social bonding preferences.[61] Higher ranking adult spotted hyenas tend to have higher telomere length, and therefore are healthier, naturally live longer, and reproduce more.[62] Territory size is highly variable, ranging from less than 40 km2 in the Ngorongoro Crater
Ngorongoro Crater
to over 1,000 km2 in the Kalahari. Home ranges are defended through vocal displays, scent marking and boundary patrols.[59] Clans mark their territories by either pasting or pawing in special latrines located on clan range boundaries. Clan boundaries are usually respected; hyenas chasing prey have been observed to stop dead in their tracks once their prey crosses into another clan's range. Hyenas will however ignore clan boundaries in times of food shortage. Males are more likely to enter another clan's territory than females are, as they are less attached to their natal group and will leave it when in search of a mate. Hyenas travelling in another clan's home range typically exhibit bodily postures associated with fear, particularly when meeting other hyenas. An intruder can be accepted into another clan after a long period of time if it persists in wandering into the clan's territory, dens or kills.[63]

Mating, reproduction, and development[edit]

Female nursing cub, Amboseli National Park, Kenya

The spotted hyena is a non-seasonal breeder, though a birth peak does occur during the wet season. Females are polyestrus, with an estrus period lasting two weeks.[64] Like many felid species, the spotted hyena is promiscuous, and no enduring pair bonds are formed. Members of both sexes may copulate with several mates over the course of several years.[51] Males will show submissive behaviour when approaching females in heat, even if the male outweighs his partner.[65] Females usually favour younger males born or joined into the clan after they were born. Older females show a similar preference, with the addition of preferring males with whom they have had long and friendly prior relationships.[66] Passive males tend to have greater success in courting females than aggressive ones.[67] Copulation in spotted hyenas is a relatively short affair,[65] lasting 4–12 minutes,[55] and typically only occurs at night with no other hyenas present.[65] The mating process is complicated, as the male's penis enters and exits the female's reproductive tract through her pseudo-penis rather than directly through the vagina, which is blocked by the false scrotum and testes. These unusual traits make mating more laborious for the male than in other mammals, while also ensuring that rape is physically impossible.[50][51] Once the female retracts her clitoris, the male enters the female by sliding beneath her, an operation facilitated by the penis's upward angle. Once this is accomplished, a typical mammalian mating posture is adopted.[51][68] Copulation may be repeated multiple times during a period of several hours.[51] Both partners usually lick their genitals for several minutes after mating.[69]

Spotted hyena
Spotted hyena
cub in the Serengeti, Tanzania. Note the well defined spots, which will fade with age.

The length of the gestation period tends to vary greatly, though 110 days is the average length of time.[64] In the final stages of pregnancy, dominant females provide their developing offspring with higher androgen levels than lower-ranking mothers do. The higher androgen levels – the result of high concentrations of ovarian androstenedione – are thought to be responsible for the extreme masculinization of female behavior and morphology.[70] This has the effect of rendering the cubs of dominant females more aggressive and sexually active than those of lower ranking hyenas; high ranking male cubs will attempt to mount females earlier than lower ranking males.[71] The average litter consists of two cubs, with three occasionally being reported.[64] Males take no part in the raising of young.[72] Giving birth is difficult for female hyenas, as the females give birth through their narrow clitoris, and spotted hyena cubs are the largest carnivoran young relative to their mothers' weight.[73] During parturition, the clitoris ruptures in order to facilitate the passage of the young, and may take weeks to heal.[59] Cubs are born with soft, brownish black hair, and weigh 1.5 kg on average.[74] Unique among carnivorous mammals, spotted hyenas are also born with their eyes open and with 6–7 mm long canine teeth and 4 mm long incisors. Also, cubs will attack each other shortly after birth. This is particularly apparent in same sexed litters, and can result in the death of the weaker cub.[73] This neonatal siblicide kills an estimated 25% of all hyenas in their first month. Male cubs which survive grow faster and are likelier to achieve reproductive dominance, while female survivors eliminate rivals for dominance in their natal clan.[68] Lactating females can carry 3–4 kg (6.6–8.8 lb) of milk in their udders.[60] Spotted hyena
Spotted hyena
milk has the highest protein and fat content of any terrestrial carnivore.[59][75] Cubs will nurse from their mother for 12–16 months, though they can process solid food as early as three months.[76] Mothers do not regurgitate food for their young.[77] Females are very protective of their cubs, and will not tolerate other adults, particularly males, approaching them. Spotted hyenas exhibit adult behaviours very early in life; cubs have been observed to ritually sniff each other and mark their living space before the age of one month. Within ten days of birth, they are able to move at considerable speed. Cubs begin to lose the black coat and develop the spotted, lighter coloured pelage of the adults at 2–3 months. They begin to exhibit hunting behaviours at the age of eight months, and will begin fully participating in group hunts after their first year.[76] Spotted hyenas reach sexual maturity at the age of three years. The average lifespan in zoos is 12 years, with a maximum of 25 years.[78] Denning behaviour[edit]

Spotted hyena
Spotted hyena
and two cubs in their den, Ngorongoro Crater, Tanzania

The clan's social life revolves around a communal den. While some clans may use particular den sites for years, others may use several different dens within a year or several den sites simultaneously.[59] Spotted hyena
Spotted hyena
dens can have more than a dozen entrances, and are mostly located on flat ground. The tunnels are usually oval in section, being wider than they are high, and narrow down from an entrance width of ½–1 metre (1.6–7.7 ft) to as small as 25 cm (9.8 in). In the rocky areas of East Africa and Congo, spotted hyenas use caves as dens, while those in the Serengeti
Serengeti
use kopjes as resting areas in daylight hours. Dens have large bare patches around their entrances, where hyenas move or lie down on. Because of their size, adult hyenas are incapable of using the full extent of their burrows, as most tunnels are dug by cubs or smaller animals. The structure of the den, consisting of small underground channels, is likely an effective anti-predator device which protects cubs from predation during the absence of the mother. Spotted hyenas rarely dig their own dens, having been observed for the most part to use the abandoned burrows of warthogs, springhares and jackals. Faeces are usually deposited 20 metres (66 feet) away from the den, though they urinate wherever they happen to be. Dens are used mostly by several females at once, and it is not uncommon to see up to 20 cubs at a single site.[79] The general form of a spotted hyena den is tunnel-shaped, with a spacious end chamber used for sleeping or breeding. This chamber measures up to 2 metres (6.6 feet) in width, the height being rather less.[80] Females generally give birth at the communal den or a private birth den. The latter is primarily used by low status females in order to maintain continual access to their cubs, as well as ensure that they become acquainted with their cubs before transferral to the communal den.[59] Intelligence[edit] Compared to other hyenas, the spotted hyena shows a greater relative amount of frontal cortex which is involved in the mediation of social behavior. Studies strongly suggest convergent evolution in spotted hyena and primate intelligence.[13] A study done by evolutionary anthropologists demonstrated that spotted hyenas outperform chimpanzees on cooperative problem-solving tests; captive pairs of spotted hyenas were challenged to tug two ropes in unison to earn a food reward, successfully cooperating and learning the maneuvers quickly without prior training. Experienced hyenas even helped inexperienced clan-mates to solve the problem. In contrast, chimps and other primates often require extensive training, and cooperation between individuals is not always as easy for them.[81] The intelligence of the spotted hyena was attested to by Dutch colonists in 19th-century South Africa, who noted that hyenas were exceedingly cunning and suspicious, particularly after successfully escaping from traps.[82] Spotted hyenas seem to plan on hunting specific species in advance; hyenas have been observed to indulge in activities such as scent marking before setting off to hunt zebras, a behaviour which does not occur when they target other prey species.[83] Also, spotted hyenas have been recorded to utilise deceptive behaviour, including giving alarm calls during feeding when no enemies are present, thus frightening off other hyenas and allowing them to temporarily eat in peace. Similarly, mothers will emit alarm calls in attempting to interrupt attacks on their cubs by other hyenas.[13] Hunting behaviour[edit] Unlike other large African carnivores, spotted hyenas do not preferentially prey on any species, and only African buffalo
African buffalo
and giraffe are significantly avoided. Spotted hyenas prefer prey with a body mass range of 56–182 kg (123–401 lb), with a mode of 102 kg (225 lb).[84] When hunting medium to large sized prey, spotted hyenas tend to select certain categories of animal; young animals are frequently targeted, as are old ones, though the latter category is not so significant when hunting zebras, due to their aggressive anti-predator behaviours.[85] The spotted hyena tracks live prey by sight, hearing and smell. Carrion is detected by smell and the sound of other predators feeding. During daylight hours, they watch vultures descending upon carcasses. Their auditory perception is powerful enough to detect sounds of predators killing prey or feeding on carcasses over distances of up to 10 km (6.2 mi).[17] Unlike the grey wolf, the spotted hyena relies more on sight than smell when hunting, and does not follow its prey's prints or travel in single file.[58] Small prey is killed by being shaken in the mouth, while large prey is eaten alive.[86] Spotted hyenas usually hunt wildebeest either singly, or in groups of two or three. They catch adult wildebeest usually after 5 km (3.1 mi) chases at speeds of up to 60 km/h (37 mi/h). Chases are usually initiated by one hyena and, with the exception of cows with calves, there is little active defence from the wildebeest herd. Wildebeest
Wildebeest
will sometimes attempt to escape hyenas by taking to water although, in such cases, the hyenas almost invariably catch them.[87] Zebras require different hunting methods to those used for wildebeest, due to their habit of running in tight groups and aggressive defence from stallions. Typical zebra hunting groups consist of 10–25 hyenas,[83] though there is one record of a hyena killing an adult zebra unaided.[88] During a chase, zebras typically move in tight bunches, with the hyenas pursuing behind in a crescent formation. Chases are usually relatively slow, with an average speed of 15–30 km/h. A stallion will attempt to place himself between the hyenas and the herd, though once a zebra falls behind the protective formation it is immediately set upon, usually after a chase of 3 km (1.9 mi). Though hyenas may harass the stallion, they usually only concentrate on the herd and attempt to dodge the stallion's assaults. Unlike stallions, mares typically only react aggressively to hyenas when their foals are threatened. Unlike wildebeest, zebras rarely take to water when escaping hyenas.[83] When hunting Thomson's gazelles, spotted hyenas usually operate alone, and prey primarily on young fawns. Chases against both adult and young gazelles can cover distances of 5 km (3.1 mi) with speeds of 60 km/h (37 mi/h). Female gazelles do not defend their fawns, though they may attempt to distract hyenas by feigning weakness.[89] Ecology[edit] Diet[edit]

Spotted hyena
Spotted hyena
with a wildebeest skeleton in Karatu, Arusha, Tanzania

The spotted hyena is the most carnivorous member of the Hyaenidae.[14] Unlike its brown and striped cousins, the spotted hyena is a predator, not a scavenger; this has been shown since the 1960s.[90] One of the earliest studies to demonstrate their hunting abilities was done by Hans Kruuk, an African wildlife ecologist, and he showed through a 7-year study of hyena populations in Africa that spotted hyenas hunt as much as lions, and with later studies this has been shown to be the average in all areas of Africa.[91] However spotted hyenas remain being mislabeled as scavengers, often even by ecologists and wildlife documentary channels. The spotted hyena is very efficient at eating its prey; not only is it able to splinter and eat the largest ungulate bones, it is also able to digest them completely. Spotted hyenas can digest all organic components in bones, not just the marrow. Any inorganic material is excreted with the faeces, which consist almost entirely of a white powder with few hairs. They react to alighting vultures more readily than other African carnivores, and are more likely to stay in the vicinity of lion kills or human settlements.[92] Wildebeest
Wildebeest
are the most commonly taken medium-sized ungulate prey item in both Ngorongoro and the Serengeti, with zebra and Thomson's gazelles coming close behind.[93] Cape buffalo
Cape buffalo
are rarely attacked due to differences in habitat preference, though adult bulls have been recorded to be taken on occasion.[94] In Kruger National Park, blue wildebeest, cape buffalo, Burchell's zebra, greater kudu and impala are the spotted hyena's most important prey, while giraffe, impala, wildebeest and zebra are its major food sources in the nearby Timbavati
Timbavati
area. Springbok
Springbok
and kudu are the main prey in Namibia's Etosha
Etosha
National Park, and springbok in the Namib. In the southern Kalahari, gemsbok, wildebeest and springbok are the principal prey. In Chobe, the spotted hyena's primary prey consists of migratory zebra and resident impala. In Kenya's Masai Mara, 80% of the spotted hyena's prey consists of topi and Thomson's gazelle, save for during the four-month period when zebra and wildebeest herds migrate to the area. Bushbuck, suni and buffalo are the dominant prey items in the Aberdare Mountains, while Grant's gazelle, gerenuk, sheep, goats and cattle are likely preyed upon in northern Kenya. In west Africa, it is thought that the spotted hyena is primarily a scavenger, but will occasionally attack domestic stock and medium-size antelopes in some areas. In Cameroon, it is common for spotted hyenas to feed on small antelopes like kob, but may also scavenge on reedbuck, kongoni, buffalo, giraffe, African elephant, topi and roan antelope carcasses. Records indicate that spotted hyenas in Malawi
Malawi
feed on medium to large-sized ungulates such as waterbuck and impala. In Tanzania's Selous Game Reserve, spotted hyenas primarily prey on wildebeest, followed by buffalo, zebra, impala, giraffe, reedbuck and kongoni. In Uganda, it is thought that the species primarily preys on birds and reptiles, while in Zambia
Zambia
it is primarily a scavenger.[95] Spotted hyenas have also been found to catch fish, tortoises, humans, black rhino, hippo calves, young African elephants, pangolins and pythons.[96] There is at least one record of four hyenas killing an adult or subadult hippopotamus in Kruger National Park.[97] The fossil record indicates that the now extinct European spotted hyenas primarily fed on Przewalski's horses, Irish elk, reindeer, red deer, roe deer, fallow deer, wild boar, ibex, steppe wisent, aurochs, and woolly rhinoceros. Spotted hyenas are thought to be responsible for the dis-articulation and destruction of some cave bear skeletons. Such large carcasses were an optimal food resource for hyenas, especially at the end of winter, when food was scarce.[98] Spotted hyenas may consume leather articles such as boots and belts around campsites. Jane Goodall
Jane Goodall
recorded spotted hyenas attacking or savagely playing with the exterior and interior fittings of cars, and the species is thought to be responsible for eating car tyres.[99] A single spotted hyena can eat at least 14.5 kg of meat per meal.[100] Although spotted hyenas act aggressively toward each other when feeding, they compete with each other mostly through speed of eating, rather than by fighting as lions do.[101] When feeding on an intact carcass, spotted hyenas will first consume the meat around the loins and anal region, then open the abdominal cavity and pull out the soft organs. Once the stomach, its wall and contents are consumed, the hyenas will eat the lungs and abdominal and leg muscles. Once the muscles have been eaten, the carcass is disassembled and the hyenas carry off pieces to eat in peace.[102] Spotted hyenas are adept at eating their prey in water: they have been observed to dive under floating carcasses to take bites, then resurface to swallow.[103] A single hyena can take less than two minutes in eating a gazelle fawn,[103] while a group of 35 hyenas can completely consume an adult zebra in 36 minutes.[104] Spotted hyenas do not require much water, and typically only spend 30 seconds drinking.[105] Enemies and competitors[edit]

Spotted hyenas mobbing a lion, Sabi Sand Game Reserve

Spotted hyena
Spotted hyena
confronting African wild dogs, Sabi Sand Game Reserve

Where spotted hyenas and lions occupy the same geographic area, the two species occupy the same ecological niche, and are thus in direct competition with one another. In some cases, the extent of dietary overlap can be as high as 68.8%.[84] Lions typically ignore spotted hyenas, unless they are on a kill or are being harassed by them. There exists a common misconception that hyenas steal kills from lions, but most often it is the other way around.[106] Lions will readily steal the kills of spotted hyenas; in the Ngorongoro Crater, it is common for lions to subsist largely on kills stolen from hyenas. Lions are quick to follow the calls of hyenas feeding, a fact demonstrated by field experiments, during which lions repeatedly approached whenever the tape-recorded calls of hyenas feeding were played.[107] When confronted on a kill by lions, spotted hyenas will either leave or wait patiently at a distance of 30–100 metres until the lions have finished eating.[108] In some cases, spotted hyenas are bold enough to feed alongside lions, and may occasionally force lions off a kill.[101] This mostly occurs during the nighttime, when hyenas are bolder.[109] Spotted hyenas usually prevail against groups of lionesses unaccompanied by males if they outnumber them 4:1.[110] In some instances they were seen to have taken on and routed two pride males while outnumbering them 5:1.[111] The two species may act aggressively toward one another even when there is no food at stake.[109] Lions may charge at hyenas and maul them for no apparent reason; one male lion was filmed killing two hyenas on separate occasions without eating them,[112] and lion predation can account for up to 71% of hyena deaths in Etosha. Spotted hyenas have adapted to this pressure by frequently mobbing lions which enter their territories.[113] Experiments on captive spotted hyenas revealed that specimens with no prior experience with lions act indifferently to the sight of them, but will react fearfully to the scent.[114] Although cheetahs and leopards preferentially prey on smaller animals than those hunted by spotted hyenas, hyenas will steal their kills when the opportunity presents itself. Cheetahs are usually easily intimidated by hyenas, and put up little resistance,[115] while leopards, particularly males, may stand up to hyenas. There are records of some male leopards preying on hyenas.[116] Hyenas are nonetheless dangerous opponents for leopards; there is at least one record of a young adult male leopard dying from a septicemia infection caused by wounds inflicted by a spotted hyena.[117] There is also a case of two spotted hyenas killing and eating a young leopard in Timbavati
Timbavati
Game Reserve, after a young hyena was killed by the leopard.[118] Spotted hyenas will follow packs of African wild dogs in order to appropriate their kills. They will typically inspect areas where wild dogs have rested and eat any food remains they find. When approaching wild dogs at a kill, solitary hyenas will approach cautiously and attempt to take off with a piece of meat unnoticed, though they may be mobbed by the dogs in the attempt. When operating in groups, spotted hyenas are more successful in pirating dog kills, though the dog's greater tendency to assist each other puts them at an advantage against spotted hyenas, who rarely work in unison. Cases of dogs scavenging from spotted hyenas are rare. Although wild dog packs can easily repel solitary hyenas, on the whole, the relationship between the two species is a one sided benefit for the hyenas,[119] with wild dog densities being negatively correlated with high hyena populations.[120] Spotted hyenas dominate other hyena species wherever their ranges overlap. Brown hyenas encounter spotted hyenas in the Kalahari, where the brown species outnumbers the spotted. The two species typically encounter each other on carcasses, which the larger spotted species usually appropriate. Sometimes, brown hyenas will stand their ground and raise their manes while emitting growls. This usually has the effect of seemingly confusing spotted hyenas, which will act bewildered, though they will occasionally attack and maul their smaller cousins. Similar interactions have been recorded between spotted and striped hyenas in the Serengeti.[121] Black-backed and side-striped jackals, and African golden wolves will feed alongside hyenas, though they will be chased if they approach too closely. Spotted hyenas will sometimes follow jackals and wolves during the gazelle fawning season, as jackals and wolves are effective at tracking and catching young animals. Hyenas do not take to eating wolf flesh readily; four hyenas were reported to take half an hour in eating a golden wolf. Overall, the two animals typically ignore each other when there is no food or young at stake.[122] Spotted hyenas usually keep a safe distance from Nile crocodiles. Though they readily take to water to catch and store prey, hyenas will avoid crocodile-infested waters.[123] Recent observations shows that African rock pythons can hunt adult spotted hyenas.[124] Communication[edit] Body language[edit]

Spotted hyenas interacting aggressively in the Masai Mara

Spotted hyenas greeting one another in Kruger National Park

Spotted hyenas have a complex set of postures in communication. When afraid, the ears are folded flat, and are often combined with baring of the teeth and a flattening of the mane. When attacked by other hyenas or by wild dogs, the hyena lowers its hindquarters. Before and during an assertive attack, the head is held high with the ears cocked, mouth closed, mane erect and the hindquarters high. The tail usually hangs down when neutral, though it will change position according to the situation. When a high tendency to flee an attacker is apparent, the tail is curled below the belly. During an attack, or when excited, the tail is carried forward on the back. An erect tail does not always accompany a hostile encounter, as it has also been observed to occur when a harmless social interaction occurs. Although they do not wag their tails, spotted hyenas will flick their tails when approaching dominant animals or when there is a slight tendency to flee. When approaching a dominant animal, subordinate spotted hyenas will walk on the knees of their forelegs in submission.[125] Greeting ceremonies among clan-members consist of two individuals standing parallel to each other and facing opposite directions. Both individuals raise their hind legs and lick each other's anogenital area.[59] During these greeting ceremonies, the penis or pseudo-penis often becomes erect, in both males and females. Erection
Erection
is usually a sign of submission, rather than dominance, and is more common in males than in females.[126] Vocalisations[edit]

It is said that feasting Hyaenas engage in violent fights, and there is such a croaking, shrieking and laughing at such times that a superstitious person might really think all the inhabitants of the infernal regions had been let loose. —  Alfred Brehm
Alfred Brehm
(1895)[127]

The spotted hyena has an extensive vocal range, with sounds ranging from whoops, fast whoops, grunts, groans, lows, giggles, yells, growls, soft grunt-laughs, loud grunt-laughs, whines and soft squeals. The loud "who-oop" call, along with the maniacal laughter, are among the most recognisable sounds of Africa. Typically, very high-pitched calls indicate fear or submission, while loud, lower-pitched calls express aggression.[128] The pitch of the laugh indicates the hyena's age, while variations in the frequency of notes used when hyenas make noises convey information about the animal's social rank.[129] Dr. Hans Kruuk compiled the following table on spotted hyena calls in 1972;[130]

Name Sound description Posture Context

Whoop A series of 6–9 (sometimes 15) calls lasting 2–3 seconds each and spaced 2–10 seconds apart. The general tone is a "oo" tone which begins in a low pitch and ends with a high note. This sound can be heard more than 5 km away.

Usually done standing, with the mouth opened slightly and the head bent down Used by both sexes when alone or in a group, and appears to be done spontaneously without external cause

Fast whoop Similar to the whoop, but higher pitched and with shorter intervals Tail is either horizontal or high with the ears cocked. Often done while running, with the mouth bent down Used with other hyenas present just before the onset of an attack, often during a dispute over a kill with lions or other hyenas

Grunt A soft, very low pitched growling sound which lasts several seconds.

Example from the Florida Teaching Zoo

The mouth is closed, and the posture aggressive. Emitted on the approach of another, unwelcome hyena, and may be followed by chasing

Groan Similar to above, but more "ooo" sounding and higher in pitch

Before and during meeting ceremonies

Low "Ooo" sound with a usually low pitch and lasts several seconds The mouth is slightly open with the head horizontal. Like the fast whoop, but with less tendency to attack

Giggle A series of loud, very high pitched "hee-hee-hee" sounds usually lasting less than 5 seconds.

Giggling call of a spotted hyena.

Running in a fleeing posture with the mouth slightly open When attacked or chased, usually over a kill

Yell A loud, high pitched call lasting several seconds As with the giggle As with the giggle, but when actually being bitten

Growl A loud, rattling, low pitched sound lasting several seconds, with an "aa" and "oh" quality Defensive posture When under attack, preceding a retaliatory bite

Soft grunt-laugh A rapid succession of low pitched, soft sounding staccato grunts lasting several seconds The mouth is closed or slightly open with a fleeing posture and the tail horizontal or high and the ears cocked When fleeing in surprise from a lion, man or when attacking large prey

Loud grunt-laugh Louder than the soft grunt-laugh, though still not very loud, and often lasts more than 5 minutes The mouth is the same as in the soft grunt-laugh, but with the tail high and ears cocked In encounters with lions or other hyena clans

Whine Loud, high pitched, rapid, drawn out "eeee" sounding squeals The mouth is slightly open with the head and tail hanging low Mostly used by cubs when following a female before suckling, or when thwarted from getting food

Soft squeal Same as above, but softer and without the staccato quality The mouth is slightly open with the ears flattened and the head tilted to one side with the teeth bared Used by both cubs and adults encountering a clan-mate after a long separation

Diseases and parasites[edit] Spotted hyenas may contract brucellosis, rinderpest[citation needed] and anaplasmosis. They are vulnerable to Trypanosoma congolense, which is contracted by consuming already infected herbivores, rather than through direct infection from tsetse flies.[131] It is known that adult spotted hyenas in the Serengeti
Serengeti
have antibodies against rabies, canine herpes, canine brucella, canine parvovirus, feline calici, leptospirosis, bovine brucella, rinderpest and anaplasmosis. During the canine distemper outbreak of 1993–94, molecular studies indicated that the viruses isolated from hyenas and lions were more closely related to each other than to the closest canine distemper virus in dogs. Evidence of canine distemper in spotted hyenas has also been recorded in the Masai Mara. Exposure to rabies does not cause clinical symptoms or affect individual survival or longevity. Analyses of several hyena saliva samples showed that the species is unlikely to be a rabies vector, thus indicating that the species catches the disease from other animals rather than from intraspecifics. The microfilaria of Dipetalonema dracuneuloides have been recorded in spotted hyenas in northern Kenya. The species is known to carry at least three cestode species of the genus Taenia, none of which are harmful to humans. It also carries protozoan parasites of the genus Hepatozoon
Hepatozoon
in the Serengeti, Kenya
Kenya
and South Africa.[132] Spotted hyenas may act as hosts in the life-cycles of various parasites which start life in herbivores; Taenia hyaenae and T. olnogojinae occur in hyenas in their adult phase. Trichinella spiralis
Trichinella spiralis
are found as cysts in hyena muscles.[131] Range, habitat and population[edit] The spotted hyena's distribution once ranged in Europe
Europe
from the Iberian Peninsula
Iberian Peninsula
to the Urals, where it remained for at least one million years.[5] Remains have also been found in the Russian Far East, and it has been theorised that the presence of hyenas there may have delayed the colonisation of North America.[133] The causes of the species' extinction in Eurasia
Eurasia
are still largely unknown.[5] In Western Europe
Europe
at least, the spotted hyena's extinction coincided with a decline in grasslands 12,500 years ago. Europe
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 which were as much at home in forests as in open lands, and in highlands as in lowlands. Spotted hyena
Spotted hyena
populations began to shrink roughly 20,000 years ago, completely disappearing from Western Europe
Europe
between 14–11,000 years ago, and earlier in some areas.[134] Historically, the spotted hyena was widespread throughout Sub-Saharan Africa. It is present in all habitats save for the most extreme desert conditions, tropical rainforests and the top of alpine mountains. Its current distribution is patchy in many places, especially in West Africa. Populations are concentrated in protected areas and surrounding land. There is a continuous distribution over large areas of Ethiopia, Kenya, Tanzania, Botswana, Namibia
Namibia
and the Transvaal Lowveld areas of South Africa.[135] During the 1770s and 1780s the species was still widespread in southern and western South Africa, being recorded i.a. on the Cape Peninsula
Cape Peninsula
and Cape Flats, and near present-day Somerset West, Riviersonderend, Mossel Bay, George, Joubertina, Gamtoos River, Jansenville, Cannon Rocks, Alice, Onseepkans
Onseepkans
and Augrabies Falls.[136] The species dwells in semi-deserts, savannah, open woodland, dense dry woodland, and mountainous forests up to 4,000 m in altitude. It is scarce or absent in tropical rainforests and coastal areas. Its preferred habitats in west Africa include the Guinea and Sudan savannahs, and is absent in the belt of dense coastal forest. In the Namib
Namib
Desert, it occurs in riverine growth along seasonal rivers, the sub-desertic pro- Namib
Namib
and the adjoining inland plateau. In ideal habitats, the spotted hyena outnumbers other large carnivores, including other hyena species. However, the striped and brown hyena occur at greater densities than the spotted species in desert and semi-desert regions.[137] Population densities based on systematic censuses vary substantially, from 0.006 to 1.7 individuals per km2.[1]

Country Population Status Threats/protection

 Angola Unknown[138] Data deficient[139] Although present, its numbers are unknown due to the Angolan Civil War[139]

 Benin 50–100[138] Threatened[139] Although generally tolerated, spotted hyenas in Benin
Benin
are declining due to persecution and prey depletion.[139]

 Botswana 1,000–2,000[138] Lower risk[139] Botswana's spotted hyena population is stable and legally protected by the Fauna Conservation Acts of 1982 and 1987, where it is listed as a game animal. A single game licence is required to hunt one, and control measures have largely removed the species from settled areas.[139]

 Burkina Faso 100-1,000[138] Data deficient[139] Burkina Faso's hyenas are depleted in number because of hunting, poaching and declining prey populations. Though still poisoned and trapped in retaliation against livestock depredations, the species is otherwise tolerated.[139]

 Burundi Unknown[138] Threatened[139] Probably on the verge of extinction[139]

 Cameroon 100-1,000[138] Threatened[139] Although protected within national parks, Cameroon
Cameroon
hyenas suffer from habitat desertification and human expansion, and are sometimes killed by tourist hunters and herdsmen.[139]

 Central African Republic 100-1,000[138] Data deficient[139] Although spotted hyena populations in CAR are probably stable and are generally tolerated, their level of legal protection is unknown.[139]

 Chad Unknown[138] Data deficient[139]

 Republic of the Congo Unknown[138] Data deficient[139]

 Democratic Republic of Congo Unknown[138] Data deficient[139]

 Djibouti Unknown[138] Data deficient[139]

 Equatorial Guinea Unknown[138] Data deficient[139]

 Eritrea Unknown[138] Data deficient[140]

 Ethiopia 1,000–2,000[138] Lower risk[139] Ethiopia
Ethiopia
has a stable population of hyenas which are considered valuable in clearing urban and rural populated areas, and are occasionally provisioned by the hyena men of Harar. There is no active bounty on them, even though they are considered vermin. They may be hunted without licence outside of national parks for five Ethiopian Birr (US$2.50).[139]

 Gabon Unknown[138] No Record[139]

 Ghana Unknown[138] Data deficient[139]

 Guinea-Bissau Unknown[138] Lower risk[139] Although still relatively common in some protected areas, spotted hyenas are likely declining due to persecution.[139]

 Kenya 2,000–4,000[138] Lower risk[139] Kenya's spotted hyena population differs in status for protected and unprotected areas, being lower risk in protected areas and threatened elsewhere. They are thought to be declining due to persecution through poisoning, shooting, snaring and trapping.[139]

 Liberia 0[138] No Record[139]

 Malawi 100-1,000[138] Data deficient[139] Malawi
Malawi
hyena populations occur at reasonable densities, though a growth in human population, habitat destruction and reduction in prey have caused the species to disappear from the central highlands. They are protected inside national parks, but are not tolerated elsewhere.[139]

 Mali 10–50[138] Threatened[139]

 Mauritania 100–500[138] Threatened[139] Spotted hyenas are still present in the Adrar.[139]

 Mozambique Unknown[138] Data deficient[139] Although protected by law, spotted hyenas are still hunted for their body parts.[139]

 Namibia 2,000–3,000[138] Lower risk[139] Namibian hyena populations are classed as lower risk in protected areas and threatened elsewhere. They are occasionally killed in communal and commercial farming areas.[139]

 Niger 20–50[138] Threatened[139] Niger's hyena populations are declining due to drought, desertification, eradication and poisoning.[139]

 Nigeria 100[138] Threatened[139] Nigerian spotted hyenas are on the verge of extinction, due to lack of legal protection, declines in prey populations and persecution in retaliation to livestock predation.[139]

 Rwanda Unknown[138] Threatened[139] Although present in Rwanda, it is unlikely that many individuals have survived the destruction of conservation areas during the Rwandan Civil War.[139]

 Senegal 100-1,000[138] Data deficient[139] Although considered useful scavengers and are afforded legal protection in national parks, spotted hyenas are not tolerated elsewhere.[139]

 Sierra Leone Unknown[138] Threatened[139] Although no longer regarded as pests, the population size and dynamics of spotted hyenas in Sierra Leone
Sierra Leone
are unknown.[139]

 Somalia Unknown[138] Data deficient[139]

 South Africa Cape: 80–100[138] Transvaal: 50–100[138] Kruger: 1,300–3,900[138] Natal: 250-1,000[138] Cape: threatened[139] Transvaal: threatened[139] Kruger: data deficient[139] Natal: threatened[139] Cape spotted hyena populations are even smaller than those of local brown hyenas, due to past persecution. Transvaal's spotted hyenas are not protected, though they are not bountied. Natal's spotted hyenas have been increasing in number within all game preserves and nature reserves, where they are protected.[139]

 Sudan Unknown[138] Data deficient[139]

 Tanzania Serengeti: 7,200–7,700[138] Elsewhere: 3,000–4,500[138] Lower risk[139] Threatened[139]

 Uganda 100-1,000[138] Data deficient[139] Although protected and tolerated, Ugandan hyenas rarely occur outside protected areas[139]

 Zambia 1,000–2,000[138] Data deficient[139] Zambia's hyenas are increasing in the Luangwa Valley, but declining elsewhere.[139]

 Zimbabwe 5,600[138] Lower risk[139] Although possibly declining in number in some areas,[141] spotted hyenas are not protected and are considered problem animals in the 8th Schedule of the Parks and Wildlife.[139]

Relationships with humans[edit] Cultural depictions and perceptions[edit] In prehistory[edit]

Trace of a 20,000-year-old spotted hyena painting from the Chauvet Cave, France

Atlatl
Atlatl
mammoth ivory "creeping hyena", found in La Madeleine rock shelter, dated back to circa 12,000 to 17,000 years ago

The spotted hyena (cave hyena subspecies) is depicted in a few examples of Upper Palaeolithic
Upper Palaeolithic
rock art in France. A painting from the Chauvet Cave
Chauvet Cave
depicts a hyena outlined and represented in profile, with two legs, with its head and front part with well distinguishable spotted coloration pattern. Because of the specimen's steeped profile, it is thought that the painting was originally meant to represent a cave bear, but was modified as a hyena. In Lascaux, a red and black rock painting of a hyena is present in the part of the cave known as the Diverticule axial, and is depicted in profile, with four limbs, showing an animal with a steep back. The body and the long neck have spots, including the flanks. An image on a cave in Ariège shows an incompletely outlined and deeply engraved figure, representing a part of an elongated neck, smoothly passing into part of the animal’s forelimb on the proximal side. Its head is in profile, with a possibly re-engraved muzzle. The ear is typical of the spotted hyena, as it is rounded. An image in the Le Gabillou Cave in Dordogne
Dordogne
shows a deeply engraved zoomorphic figure with a head in frontal view and an elongated neck with part of the forelimb in profile. It has large round eyes and short, rounded ears which are set far from each other. It has a broad, line-like mouth that evokes a smile. Though originally thought to represent a composite or zoomorphic hybrid, it is probable it is a spotted hyena based on its broad muzzle and long neck.[18] The relative scarcity of hyena depictions in Paleolithic rock art has been theorised to be due to the animal's lower rank in the animal worship hierarchy; the spotted hyena's appearance was likely unappealing to Ice Age hunters, and it was not sought after as prey. Also, it was not a serious rival like the cave lion or cave bear, and it lacked the impressiveness of the mammoth or woolly rhino.[18] In Africa[edit]

Spotted hyena
Spotted hyena
mask from Burkina Faso, Musée barrois

Spotted hyena
Spotted hyena
being fed in Harar, Ethiopia

In Africa, the spotted hyena is usually portrayed as an abnormal and ambivalent animal, considered to be sly, brutish, necrophagous and dangerous. It further embodies physical power, excessivity, ugliness, stupidity, as well as sacredness. Spotted hyenas vary in their folkloric and mythological depictions, depending on the ethnic group from which the tales originate. It is often difficult to know whether or not spotted hyenas are the specific hyena species featured in such stories, particularly in West Africa, as both spotted and striped hyenas are often given the same names.[142] In west African tales, spotted hyenas symbolise immorality, dirty habits, the reversal of normal activities, and other negative traits, and are sometimes depicted as bad Muslims who challenge the local animism that exists among the Beng in Côte d’Ivoire. In East Africa, Tabwa mythology portrays the spotted hyena as a solar animal that first brought the sun to warm the cold earth.[142] In the culture of the Mbugwe in Tanzania, the spotted hyena is linked to witchcraft. According to Mbugwe folklore, every witch possesses one or more hyenas, which are referred to as "night cattle" and are branded with an invisible mark. It is said that all hyenas are owned by witches, and that truly wild hyenas are non-existent. Lactating female spotted hyenas are said to be milked by their owners every night to make hyena butter, and are further used as mounts. When a witch acquires a hyena mount, he rides it to distant lands in order to bewitch victims and return safely home before morning. The Mbugwe consider killing hyenas to be dangerous, as the bond between the hyena and its owner is very strong, and will likely result in the witch seeking retribution. In order to obviate this danger, a killed hyena usually has its ears, tail and front legs cut off and buried, as these are the parts which are supposed to be marked by the witches' brand.[143] In the Mtwara Region
Mtwara Region
of Tanzania, it is believed that a child born at night while a hyena is crying will likely grow up to be a thief. In the same area, hyena faeces are believed to enable a child to walk at an early age, thus it is not uncommon in that area to see children with hyena dung wrapped in their clothes.[144] The Kaguru of Tanzania
Tanzania
and the Kujamaat of Southern Senegal
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.[19] A similar myth occurs in Mansôa, Guinea-Bissau. These "werehyenas" are executed when discovered, but do not revert to their human form when killed.[144] In Khoikhoi mythology, the spotted hyena is often the butt of the jackal's tricks. Gogo folklore links the spotted hyena to the origin of death; in one tale, the hyena prevents humanity from achieving immortality, thus ensuring it can continue to eat corpses. A similar tale is present among the Meru. In their narrative, the supreme god Murungu sent a mole to inform humanity that they would be reborn after death. Fearing this would deprive it of corpses to eat, the hyena prevents the mole from ever delivering the message. Madi and Nuer mythology links the spotted hyena to the separation between heaven and earth; at one time, humanity kept in contact with the Creator in the sky via a cowhide rope, which was subsequently severed by a hungry hyena.[145] The spotted hyena is a sacred totem animal for some Pedi tribes, with the skin often being used as robes by chieftains and their bones as divining instruments.[146] According to the doctrine of the Ethiopian Orthodox Tewahedo Church, hyenas are unclean animals which represent sexual deviancy and lawlessness. The Egyptian Saint abba (Father) Matewos of Asfoni was associated with hyenas; one fable tells of how he rescued a cub trapped in a pit, and had his feet licked in gratitude by its mother. In Ethiopian folklore, an albino hyena called the "King of Hyenas" is ascribed great power. Some ethnic groups in Ethiopia associate themselves with hyenas; the Gurage traditionally believe that their ancestors migrated from Arabia
Arabia
to Ethiopia
Ethiopia
using hyenas as mounts. In Dorze tradition, the highest Demuṣa-priests have the ability to control hyenas, and will send them to punish defaulting debtors.[147] Spotted hyenas feature prominently in the rituals of certain African cultures. In the Gelede cult of the Yoruba people
Yoruba people
of Benin
Benin
and Southwest Nigeria, a spotted hyena mask is used at dawn to signal the end of the èfè ceremony. As the spotted hyena usually finishes the meals of other carnivores, the animal is associated with the conclusion of all things. Among the Korè cult of the Bambara people in Mali, the belief that spotted hyenas are hermaphrodites appears as an ideal in-between in the ritual domain. The role of the spotted hyena mask in their rituals is often to turn the neophyte into a complete moral being by integrating his male principles with femininity. The Beng people believe that upon finding a freshly killed hyena with its anus inverted, one must plug it back in, for fear of being struck down with perpetual laughter. They also view spotted hyena faeces as contaminating, and will evacuate a village if a hyena relieves itself within village boundaries.[142] In Harar, Ethiopia, spotted hyenas are regularly fed by the city's inhabitants, who believe the hyenas' presence keeps devils at bay, and associate mystical properties such as fortune telling to them.[148] In Western culture[edit]

As several distinguished authors of the present age have undertaken to reconcile the world to the Great Man-Killer of Modern times; as Aaron Burr has found an apologist, and almost a eulogist; and as learned commentators have recently discovered that even Judas Iscariot
Judas Iscariot
was a true disciple, we are rather surprised to find that someone has not undertaken to render the family of Hyenas popular and amiable in the eyes of mankind. Certain it is, that few marked characters in history have suffered more from the malign inventions of prejudice[149]

Traditional Western beliefs about the spotted hyena can be traced back to Aristotle's Historia Animalium, which described the species as a necrophagous, cowardly and potentially dangerous animal. He further described how the hyena uses retching noises to attract dogs. In On the Generation of Animals, Aristotle
Aristotle
criticised the erroneous belief that the spotted hyena is a hermaphrodite (which likely originated from the confusion caused by the masculinised genitalia of the female), though his physical descriptions are more consistent with the striped hyena. Pliny the Elder
Pliny the Elder
supported Aristotle's depiction, though he further elaborated that the hyena can imitate human voices. Additionally, he wrote how the hyena was held in high regard among the Magi, and that hyena body parts could cure different diseases, give protection and stimulate sexual desire in people.[19] The author of the Physiologus, who infused pagan tales with the spirit of Christian moral and mystical teaching, reactivated the myth that the hyena is a hermaphrodite. The author compared the species to "double-minded men" who are neither "man nor woman, that is, neither faithful nor unfaithful". He further states that "The sons of Israel are like this animal since in the beginning they served the living God but later, given over to pleasure and lust, they adored idols." The bestiaries of the Middle Ages
Middle Ages
embraced the Physiologus's descriptions, but further elaborated on the animal's necrophagous habits. These bestiaries almost invariably depict hyenas feeding on human corpses. These illustrations were largely based on the descriptions given by Aristotle
Aristotle
and Pliny, though the animals have no spots or other bodily markings, thus making it unlikely that the authors had ever seen hyenas first-hand.[19] During the 15th and 16th centuries, travellers to Africa provided further descriptions of the species. Leo Africanus
Leo Africanus
repeated some of the old concepts on the hyena, with the addition of describing its legs and feet as similar to those of men. In 1551, Swiss naturalist Conrad Gesner
Conrad Gesner
rejected the belief of the hyena's hermaphroditism, and theorised that it originated from confusion over an androgynous fish bearing the same name. He adds three other animals within the category of hyenas, including an Ethiopian quadruped named "Crocotta", which was thought to be a hybrid between a hyena and a lioness. Sir Thomas Browne also argued against the hyena's supposed hermaphroditism, stating that all animals follow their own "Law of Coition", and that a hermaphrodite would transgress this. Sir Walter Raleigh, in an attempt to rationalise how Noah's Ark
Noah's Ark
could have fitted all extant species of animal, wrote that hyenas were hybrids between foxes and wolves which originated after the Great Flood. References to the spotted hyena's vocalisations are referenced in numerous contemporary examples of English literature, including Shakespeare's As You Like It
As You Like It
and George Chapman's Eastward Ho. John Milton, in his Samson Agonistes, compares the species to Delila.[19] Natural historians of the 18th and 19th centuries rejected stories of hermaphroditism in hyenas, and recognised the differences between the spotted and striped hyena. However, they continued to focus on the species' scavenging habits, their potential to rob graves and their perceived cowardice. During the 20th century, Western and African stereotypes of the spotted hyena converged; in both Ernest Hemingway's Green Hills of Africa
Green Hills of Africa
and Disney's The Lion
Lion
King, the traits of gluttony and comical stupidity, common in African depictions of hyenas, are added to the Western perception of hyenas being cowardly and ugly.[19] After the release of The Lion
Lion
King, hyena biologists protested against the animal's portrayal: one hyena researcher sued Disney
Disney
studios for defamation of character,[150] and another – who had organized the animators' visit to the University of California's Field Station for Behavioural Research, where they would observe and sketch captive hyenas[19] – suggested boycotting the film.[151] Livestock predation[edit] When targeting livestock, the spotted hyena primarily preys upon cattle, sheep and goats,[12] though hyenas in the southern parts of Ethiopia's Tigray region preferentially target donkeys.[152] Reports of livestock damage are often not substantiated, and hyenas observed scavenging on a carcass may be mistaken for having killed the animal. The rate at which the species targets livestock may depend on a number of factors, including stock keeping practices, the availability of wild prey and human-associated sources of organic material, such as rubbish. Surplus killing
Surplus killing
has been recorded in South Africa's eastern Cape Province. Attacks on stock tend to be fewer in areas where livestock is coralled by thorn fences and where domestic dogs are present. One study in northern Kenya
Kenya
revealed that 90% of all cases of livestock predation by hyenas occurred in areas outside the protection of thorn fences.[12] Attacks on humans and grave desecration[edit] Like most mammalian predators, the spotted hyena is typically shy in the presence of humans, and has the highest flight distance (up to 300 metres) among African carnivores. However, this distance is reduced during the night, when hyenas are known to follow people closely.[153] Although spotted hyenas do prey on humans in modern times, such incidents are rare. However, attacks on humans by spotted hyenas are likely to be underreported.[154] Man-eating spotted hyenas tend to be very large specimens; a pair of man-eating hyenas, responsible for killing 27 people in Mlanje, Malawi, in 1962, were weighed at 72 and 77 kg (159 and 170 lb) after being shot.[155] Victims of spotted hyenas tend to be women, children and sick or infirm men,[156] and there are numerous cases of biologists in Africa being forced up trees in order to escape them.[155] Attacks occur most commonly in September, when many people sleep outdoors, and bush fires make the hunting of wild game difficult for hyenas.[154][155] In 1903, Hector Duff wrote of how spotted hyenas in the Mzimba district of Angoniland would wait at dawn outside people's huts and attack them when they opened their doors.[157] In 1908–09 in Uganda, spotted hyenas regularly killed sufferers of African sleeping sickness as they slept outside in camps.[156] Spotted hyenas are widely feared in Malawi, where they have been known to occasionally attack people at night, particularly during the hot season when people sleep outside. Hyena
Hyena
attacks were widely reported in Malawi's Phalombe
Phalombe
plain, to the north of Michesi Mountain. Five deaths were recorded in 1956, five in 1957 and six in 1958. This pattern continued until 1961 when eight people were killed.[157] During the 1960s, Flying Doctors received over two dozen cases of hyena attacks on humans in Kenya.[144] An anecdotal 2004 news report from the World Wide Fund for Nature indicates that 35 people were killed by spotted hyenas during a 12-month period in Mozambique
Mozambique
along a 20 km stretch of road near the Tanzanian border.[154] Although attacks against living humans are rare, the spotted hyena readily feeds on human corpses. In the tradition of the Maasai[144] and the Hadza,[158] corpses are left in the open for spotted hyenas to eat. A corpse rejected by hyenas is seen as having something wrong with it, and liable to cause social disgrace, therefore it is not uncommon for bodies to be covered in fat and blood from a slaughtered ox.[144] In Ethiopia, hyenas were reported to feed extensively on the corpses of victims of the 1960 attempted coup[159] and the Red Terror.[160] Hyenas habituated to scavenging on human corpses may develop bold behaviours towards living people; hyena attacks on people in southern Sudan
Sudan
increased during the Second Sudanese Civil War, when human corpses were readily available to them.[161] Urban hyenas[edit] In some parts of Africa, spotted hyenas have begun to frequent metropolitan areas, where groups or "clans" of the animals have become a menace. The Ethiopian capital Addis Ababa
Addis Ababa
is estimated to have up to a thousand resident hyenas which survive by scavenging rubbish tips and preying on feral dogs and cats. There have also been attacks on humans who are homeless. In 2013, a baby boy was killed by hyenas after being snatched from his mother as she camped near the Hilton Hotel. Some 40 of the animals were reportedly seen alongside a fence bordering the British Embassy compound. In December 2013, a cull was organised and marksmen killed ten hyenas which had occupied wasteland near the city centre.[162] Hunting and use in traditional medicine[edit]

Spotted hyena
Spotted hyena
attacked by Maasai warriors

Spotted hyena
Spotted hyena
shot by Abel Chapman
Abel Chapman
at the Lukenia Heights, 23 January 1906

The spotted hyena has been hunted for its body parts for use in traditional medicine,[138] for amusement,[19] and for sport, though this is rare, as the species is generally not considered attractive.[138][152] There is fossil evidence of humans in Middle Pleistocene Europe
Europe
butchering and presumably consuming spotted hyenas.[163] Such incidences are rare in modern Africa, where most tribes, even those known to eat unusual kinds of meat, generally despise hyena flesh.[153] Several authors during the Scramble for Africa
Scramble for Africa
attested that, despite its physical strength, the spotted hyena poses no danger to hunters when captured or cornered. It was often the case that native skinners refused to even touch hyena carcasses, though this was not usually a problem, as hyena skins were not considered attractive.[156][164] In Burkina Faso, the hyena's tail is used for medicinal and magical purposes. In Cameroon, Côte d'Ivoire and Senegal, the animals' whole body is harvested for bushmeat and medicine. In Malawi
Malawi
and Tanzania, the genitalia, nose tips and tails are used for traditional medicine. In Mozambique, traditional healers use various spotted hyena body parts, particularly the paws.[138] Oromo hunters typically go through ritual purification after killing hyenas.[147] Kujamaat hunters traditionally treat the spotted hyenas they kill with the same respect due to deceased tribal elders, in order to avoid retribution from hyena spirits acting on behalf of the dead animal.[142] During the early years of Dutch colonisation in southern Africa, hyenas (referred to as "wolves" by the colonists) were especially susceptible to trapping, as their prediliction for carrion and lack of caution about enclosed spaces worked against them. A feature of many frontier farms was the wolwehok (hyena trap), which was roughly constructed from stone or wood and baited with meat. The trap featured a trap-door, which was designed to shut once the bait was disturbed.[165] In the Cape Colony, spotted hyenas were often hunted by tracking them to their dens and shooting them as they escaped. Another hunting method was to trap them in their dens and dazzle them with torchlight, before stabbing them in the heart with a long knife.[82] When chased by hunting dogs, spotted hyenas often attack back, unless the dogs are of exceptionally large, powerful breeds. James Stevenson-Hamilton wrote that wounded spotted hyenas could be dangerous adversaries for hunting dogs, recording an incident in which a hyena managed to kill a dog with a single bite to the neck without breaking the skin.[166] Further difficulties in killing spotted hyenas with dogs include the species' thick skin, which prevents dogs from inflicting serious damage to the animal's muscles.[167] Spotted hyenas in captivity and as pets[edit]

South African zoologist Kevin Richardson with captive spotted hyenas

From a husbandry point of view, hyenas are easily kept, as they have few disease problems and it is not uncommon for captive hyenas to reach 15–20 years of age. Nevertheless, the spotted hyena was historically scantily represented in zoos, and was typically obtained in order to fill empty cages until a more prestigious species could be obtained. In subsequent years, animals considered to be more charismatic were allocated larger and better quality facilities, while hyenas were often relegated to inferior exhibits. In modern times, the species faces spatial competition from more popular animals, especially large canids. Also, many captive individuals have not been closely examined to confirm their sexes, thus resulting in non-breeding pairs often turning out to be same-sexed individuals. As a result, many captive hyena populations are facing extinction.[168] During the 19th century, the species was frequently displayed in travelling circuses as oddities. Alfred Brehm
Alfred Brehm
wrote that the spotted hyena is harder to tame than the striped hyena, and that performing specimens in circuses were not up to standard.[169] Sir John Barrow described how spotted hyenas in Sneeuberge
Sneeuberge
were trained to hunt game, writing that they were "as faithful and diligent as any of the common domestic dogs".[170] In Tanzania, spotted hyena cubs may be taken from a communal den by witchdoctors, in order to increase their social status.[144] An April 2004 BBC
BBC
article described how a shepherd living in the small town of Qabri Bayah about 50 kilometres from Jigjiga, Ethiopia
Ethiopia
managed to use a male spotted hyena as a livestock guardian dog, suppressing its urge to leave and find a mate by feeding it special herbs.[171] If not raised with adult members of their kind, captive spotted hyenas will exhibit scent marking behaviours much later in life than wild specimens.[74] Although easily tamed, spotted hyenas are exceedingly difficult to house train,[172] and can be very destructive; a captive, otherwise perfectly tame, specimen in the Tower of London
Tower of London
managed to tear an 8-foot (2.4 m) long plank nailed to its recently repaired enclosure floor with no apparent effort.[173] During the research leading to the composition of his monograph The Spotted Hyena: A Study of Predation
Predation
and Social Behavior, Hans Kruuk kept a tame hyena he named Solomon.[174] Kruuk found Solomon's company so congenial, he would have kept him, but Solomon had an insatiable taste for "cheese in the bar of the tourist lounge and bacon off the Chief Park Warden's breakfast table", and no door could hold him back, so Solomon was obliged to live out his days in the Edinburgh Zoo.[175] See also[edit]

Spotted hyenas in Harar Pachycrocuta

References[edit] Notes[edit]

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Retrieved 22 March 2015.  ^ Kruuk 1972, pp. 251–265 ^ a b c Kruuk 1972, pp. 27–31 ^ a b c Kruuk 1972, pp. 230–233 ^ How Hyenas Avoid Incest by LiveScience Staff, posted: 15 August 2007 01:00 pm ET. ^ It’s a dog’s life – aggressive male hyenas fail to impress the girls from innovations report, May 14, 2003. ^ a b Estes 1998, p. 293 ^ East, Marion L.; et al. (2003). "Sexual conflicts in spotted hyenas: male and female mating tactics and their reproductive outcome with respect to age, social status and tenure" (PDF). Proceedings of the Royal Society of London B: Biological Sciences. 270 (1521): 1247–1254. doi:10.1098/rspb.2003.2363. PMC 1691369 . PMID 12816637. CS1 maint: Explicit use of et al. (link) ^ Nelson RJ. 2005. Introduction to Behavioral Endocrinology. Sinauer Associates: Massachusetts. p. 115. ^ The Painful Realities of Hyena
Hyena
Sex, by Bjorn Carey, LiveScience Staff Writer, posted: 26 April 2006 01:00 pm ET. ^ Kruuk 1972, p. 273 ^ a b Macdonald 1992, p. 140 ^ a b Kruuk 1972, p. 247 ^ Mills & Mills 2011, p. 127 ^ a b Kruuk 1972, pp. 248–249 ^ Kruuk 1972, p. 242 ^ Kruuk 1972, p. 34 ^ Kruuk 1972, pp. 242–247 ^ Rosevear 1974, p. 362 ^ Hyenas Surprisingly Good at Cooperative Tasks By LiveScience Staff, posted: 28 September 2009 12:56 pm ET ^ a b Moodie, John Wedderburn Dunbar (1835) Ten Years in South Africa, Vol. 1, Richard Bentley ^ a b c Kruuk 1972, pp. 176–185 ^ a b Hayward, M. W. (2006). "Prey preferences of the spotted hyaena (Crocuta crocuta)" (PDF). Journal of Zoology. 270 (4): 606–614. doi:10.1111/j.1469-7998.2006.00183.x.  ^ Kruuk 1972, p. 101 ^ Rosevear 1974, pp. 365–366 ^ Kruuk 1972, pp. 148–176 ^ Brockmann, H. Jane; Snowdon, Charles T.; Macedo, Regina (2010) Behavioral Ecology of Tropical Animals: Volume 42 of Advances in the Study of Behavior, Academic Press, p. 206, ISBN 0-12-380894-4 ^ Kruuk 1972, pp. 196–197 ^ Kruuk 1972 ^ Tyrants of the Cenozoic: Evolution
Evolution
of Bone-Crushing Hyenas and Dogs - YouTube ^ Kruuk 1972, pp. 107–108 ^ Kruuk 1972, pp. 63–64 ^ Kruuk 1972, p. 199 ^ Mills & Hofer 1998, p. 32 ^ Kruuk 1972, p. 65 ^ Mills & Mills 2010, p. 104 ^ "Prey deposits and den sites of the Upper Pleistocene hyena Crocuta crocuta spelaea (Goldfuss, 1823)in horizontal and vertical caves of the Bohemian Karst". Cajusg. Diedrich & Karelzak. Retrieved 2008-01-20.  ^ Rosevear 1974, pp. 363–364 ^ Kruuk 1972, p. 124 ^ a b Kruuk 1972, p. 128 ^ Kruuk 1972, pp. 125–126 ^ a b Kruuk 1972, p. 119 ^ Kruuk 1972, p. 126 ^ Kruuk 1972, p. 127 ^ Carroll, Chris. Behind the snarl lies a cagey opportunist, proficient hunter, and dutiful parent. . National Geographic ^ Kruuk 1972, pp. 128–137 ^ Schaller 1976, p. 272 ^ a b Schaller 1976, p. 273 ^ Cooper, S. (1991). "Optimal hunting group size: the need for lions to defend their kills against loss to spotted hyaenas". African Journal of Ecology. 29 (2): 130–136. doi:10.1111/j.1365-2028.1991.tb00993.x.  ^ AfricaHunting.com - Hyena: Scavenger or Predator? The Human Influence on Hyena
Hyena
& Lion ^ Dereck and Beverley Joubert (1992). Eternal Enemies: Lions and Hyenas (DVD). National Geographic.  ^ Trinkel, Martina & Katsberger, Gerald (2005). "Competitive interactions between spotted hyenas and lions in the Etosha
Etosha
National Park, Namibia". African Journal of Ecology. 43 (3): 220–224. doi:10.1111/j.1365-2028.2005.00574.x.  ^ Kruuk 1972, p. 248 ^ Kruuk 1972, pp. 138–139 ^ Jonathan Scott & Angela Scott (2006). Big Cat
Cat
Diary: Leopard. p. 108. ISBN 0-00-721181-3.  ^ Balme, Guy; Hunter, Luke (2004). "Mortality in a protected Leopard population, Phinda Private Game Reserve, South Africa: A population in decline?" (PDF). Ecological Journal. 6. Retrieved 31 July 2014.  ^ http://africageographic.com/blog/hyenas-kill-a-leopard/ ^ Kruuk 1972, pp. 139–141 ^ Creel, Scott; Creel, Nancy Marusha (2002), The African wild dog: behavior, ecology, and conservation, Princeton University Press, pp. 253–254, ISBN 0-691-01654-2 ^ Spotted Hyaena
Hyaena
versus Brown Hyaena, Skirmishes in the Desert
Desert
from Martin Harvey and M. G. L. Mills' African Predators, Smithsonian Books (October 2001) ^ Kruuk 1972, pp. 141–143 ^ Kruuk 1972, p. 146 ^ https://news.nationalgeographic.com/2017/03/snakes-pythons-africa-hyena-kill/ ^ Kruuk 1972, pp. 216–220 ^ Richard C. Francis (2004). Why Men Won't Ask for Directions: The Seductions of Sociobiology. Princeton University Press. p. 178. ISBN 978-0-691-05757-6.  ^ Brehm 1895, p. 184 ^ Kruuk 1972, p. 220 ^ Walker, Matt Hyena
Hyena
laughs and giggles decoded, Earth
Earth
News, 01:55 GMT, Tuesday, 30 March 2010 02:55 UK ^ Kruuk 1972, pp. 310–311 ^ a b Kruuk 1972, p. 38 ^ Mills & Hofer 1998, p. 38 ^ Summerill, Lynette; Gnawed Bones tell Tales, Summer 2003, ASU Research ^ C. Stiner, Mary (2004) Comparative ecology and taphonomy of spotted hyenas, humans, and wolves in Pleistocene Italy, Revue de Paléobiologie, Genève ^ Mills & Hofer 1998, pp. 54–56 ^ Rookmaaker, L. C. (1989). The zoological exploration of southern Africa, 1650-1790. Rotterdam: A. Balkema. pp. 112, 114, 132, 173, 252, 254. ISBN 9789061918677.  ^ Mills & Hofer 1998, p. 31 ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq Mills & Hofer 1998, p. 74 ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq ar as at au av aw ax ay az ba bb bc bd be bf bg bh bi bj Mills & Hofer 1998, pp. 75–78 ^ Mills & Hofer 1998, p. 68 ^ Williams, Samual; Williams, Kathryn; Joubert, Christoffel; Hill, Russell (2016-01-14). "The impact of land reform on the status of large carnivores in Zimbabwe". PeerJ. 4: e1537. doi:10.7717/peerj.1537. PMC 4728035 . PMID 26819838. Retrieved 3 May 2017.  ^ a b c d Frembgen, Jürgen W. (1998). "The Magicality of the Hyena: Beliefs and Practices in West and South Asia". Asian Folklore Studies. 57 (2): 331–344. doi:10.2307/1178757. JSTOR 1178757.  ^ Middleton, John; Winter, E. H. (2004). Witchcraft
Witchcraft
and Sorcery in East Africa, Routledge, pp. 165–167, ISBN 0-415-33073-4 ^ a b c d e f Mills & Hofer 1998, p. 97 ^ Lynch, Patricia Ann (2004) African mythology A to Z, Infobase Publishing, pp. 47–48, ISBN 0-8160-4892-4 ^ Quin, P.J. 1959. Food and feeding habits of the Pedi. Johannesburg: Witwatersrand University Press, p. 125. ^ a b Uhlig, Siegbert (2007) Encyclopaedia Aethiopica: He-N, Volume 3 of Encyclopaedia Aethiopica, Otto Harrassowitz Verlag, ISBN 3-447-05607-X. p. 97. ^ Bierbaum, Bernd (2011) In Ethiopia, BoD – Books on Demand, ISBN 3844858849. p. 45. ^ Goodrich, S.G. and Winchell, A., Johnson's Natural History (New York: A.J. Johnson & Company, 1885), p. 248. ^ Mcpherson, James. The good, the bad and the hyena Archived 2014-10-27 at the Wayback Machine.. suzieszterhas.com ^ Frank, Laurence D. (May 2006). Girl Power, African Geographic ^ a b Yirga, Giday; Bauer, Hans (2010). "Livestock Depredation of the Spotted Hyena
Hyena
(Crocuta crocuta) in Southern Tigray, Northern Ethiopia" (PDF). International Journal of Ecology and Environmental Sciences. 36 (1): 67–73. Archived from the original (PDF) on December 16, 2012.  ^ a b Kruuk 1972, pp. 144–145 ^ a b c Begg, Colleen; Begg, Kieth; 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) ^ a b c Kruuk, Hans (2002) Hunter and hunted: relationships between carnivores and people Cambridge University Press, pp. 64–65, ISBN 0-521-89109-4 ^ a b c Roosevelt, Theodore (1910) African Game Trails: An Account of the African Wanderings of an American Hunter, Naturalist, New York, C. Scribner's sons ^ a b Knight, John (2000). Natural Enemies: People-Wildlife conflicts in Anthropological Perspective. ASIN 0415224411. ISBN 0-415-22441-1. CS1 maint: ASIN uses ISBN (link) ^ Marlowe, Frank (2010), The Hadza: hunter-gatherers of Tanzania, University of California Press, ISBN 0-520-25342-6 ^ Kapuściński, Ryszard, The Emperor: Downfall of an Autocrat. 1978. ISBN 0-679-72203-3 ^ Donham, Donald Lewis (1999) Marxist modern: an ethnographic history of the Ethiopian revolution, University of California Press, p. 135, ISBN 0-520-21329-7 ^ Copson, Raymond W. (1994) Africa's wars and prospects for peace, M.E. Sharpe, p. 6, ISBN 1-56324-300-8 ^ Fletcher, Martin (23 February 2014). "The urban hyenas that attack rough sleepers". BBC
BBC
website. Retrieved 23 February 2014.  ^ "The scavenger or the scavenged?" A. Rodríguez-Hidalgo. Journal of Taphonomy, 1(2010): 75–76 ^ Tjader, Richard (1910) The big game of Africa: with many illustrations from photographs by the author, D. Appleton and company in New York and, London ^ Beinart, William (2008) The Rise of Conservation in South Africa: Settlers, Livestock, and the Environment 1770–1950, Oxford University Press US, ISBN 0-19-954122-1 ^ Stevenson-Hamilton 1917, p. 95 ^ Daniell, Samuele; Barrow, Sir John; Somerville, William (1820), Sketches representing the native tribes, animals, and scenery of southern Africa: from drawings made by the late Mr. Samuel Daniell, W. Daniell, p. 22 ^ Mills & Hofer 1998, p. 92 ^ Brehm 1895, p. 185 ^ Barrow, Sir John (1801) An Account of Travels Into the Interior of Southern Africa, T. Cadell, jun and W. Davies ^ Adow, Mohammed. (2004-04-12) Africa Taming Ethiopia's hyenas. BBC News. Retrieved on 2011-02-15. ^ Rosevear 1974, p. 363 ^ Bingley, William (1829), Animal
Animal
biography, or, Popular zoology ^ op. cit in Kruuk (1972) ^ Kruuk, Hans (1975) Hyaena. Oxford University Press, London

Bibliography[edit]

Brehm, Alfred Edmund (1895). "Brehm's Life of Animals". Chicago: A. N. Marquis & Company.  Estes, Richard (1992). "The behavior guide to African mammals: including hoofed mammals, carnivores, primates". University of California Press. ISBN 0-520-08085-8.  Estes, Richard (1998). "The safari companion: a guide to watching African mammals, including hoofed mammals, carnivores, and primates". Chelsea Green Publishing. ISBN 1-890132-44-6.  Funk, Holger (2010). "Hyaena: On the Naming and Localisation of an Enigmatic Animal". GRIN Verlag. ISBN 3-640-69784-7.  Kingdon, Jonathan (1988). "East African mammals: an atlas of evolution in Africa, Volume 3, Part 1". University of Chicago Press. ISBN 0-226-43721-3.  Kruuk, Hans (1972). "The Spotted Hyena: A Study of Predation
Predation
and Social Behaviour". University of California Press. ISBN 0226455084.  Kurtén, Björn (1968). "Pleistocene mammals of Europe". Weidenfeld and Nicolson.  Lydekker, Richard (1908). "The Game Animals of Africa". London, R. Ward, limited.  Macdonald, David (1992). "The Velvet Claw: A Natural History of the Carnivores". New York: Parkwest. ISBN 0-563-20844-9.  Mills, Gus; Hofer, Heribert (1998). "Hyaenas: status survey and conservation action plan" (PDF). IUCN/SSC Hyena
Hyena
Specialist Group. ISBN 2-8317-0442-1.  Mills, Gus; Mills, Margie (2011). " Hyena
Hyena
Nights & Kalahari
Kalahari
Days". Jacana Media. ISBN 1-77009-811-9.  Roosevelt, Theodore (1910). "African Game Trails: An Account of the African Wanderings of an American Hunter, Naturalist". New York, C. Scribner's sons. ISBN 978-1-4510-0191-4.  Rosevear, Donovan Reginald (1974). "The carnivores of West Africa". London : Trustees of the British Museum (Natural History). ISBN 0-565-00723-8.  Schaller, George B. (1976). "The Serengeti
Serengeti
Lion: A Study of Predator-Prey Relations". University of Chicago Press. ISBN 0-226-73640-7.  Stevenson-Hamilton, James (1917). " Animal
Animal
life in Africa, Vol. 1". London : William Heinemann. 

Further reading[edit]

Hugo Van Lawick and Jane Goodall. Innocent Killers. Houghton Mifflin Company Boston, 1971 Mills, M.G.L. Kalahari
Kalahari
Hyenas: Comparative Behavioral Ecology of Two Species. The Blackburn Press, 2003

External links[edit]

Wikispecies
Wikispecies
has information related to Crocuta crocuta

Wikimedia Commons has media related to Crocuta crocuta.

The IUCN
IUCN
Hyaenidae Specialist Group page on spotted hyena Davis, Delbert Dwight & Story, Harriette Elizabeth (1949), The Female External Genitalia of the Spotted Hyena, Fieldiana: Zoology, vol. 31, no. 33, Chicago Natural History Museum McPherson, Jamie The Good, the Bad and the Hyena, BBC
BBC
Wildlife, Summer 2008. Spotted Hyena
Hyena
- Female Genitalia Video, Check123 Video Encyclopedia

v t e

Extant Carnivora
Carnivora
species

Kingdom: Animalia Phylum: Chordata Class: Mammalia Infraclass: Eutheria Superorder: Laurasiatheria

Suborder Feliformia

Nandiniidae

Nandinia

African palm civet
African palm civet
(N. binotata)

Herpestidae (Mongooses)

Atilax

Marsh mongoose
Marsh mongoose
(A. paludinosus)

Bdeogale

Bushy-tailed mongoose
Bushy-tailed mongoose
(B. crassicauda) Jackson's mongoose
Jackson's mongoose
(B. jacksoni) Black-footed mongoose
Black-footed mongoose
(B. nigripes)

Crossarchus

Alexander's kusimanse
Alexander's kusimanse
(C. alexandri) Angolan kusimanse
Angolan kusimanse
(C. ansorgei) Common kusimanse
Common kusimanse
(C. obscurus) Flat-headed kusimanse
Flat-headed kusimanse
(C. platycephalus)

Cynictis

Yellow mongoose
Yellow mongoose
(C. penicillata)

Dologale

Pousargues's mongoose
Pousargues's mongoose
(D. dybowskii)

Galerella

Angolan slender mongoose
Angolan slender mongoose
(G. flavescens) Black mongoose
Black mongoose
(G. nigrata) Somalian slender mongoose
Somalian slender mongoose
(G. ochracea) Cape gray mongoose
Cape gray mongoose
(G. pulverulenta) Slender mongoose
Slender mongoose
(G. sanguinea)

Helogale

Ethiopian dwarf mongoose
Ethiopian dwarf mongoose
(H. hirtula) Common dwarf mongoose
Common dwarf mongoose
(H. parvula)

Herpestes

Short-tailed mongoose
Short-tailed mongoose
(H. brachyurus) Indian gray mongoose
Indian gray mongoose
(H. edwardsii) Indian brown mongoose
Indian brown mongoose
(H. fuscus) Egyptian mongoose
Egyptian mongoose
(H. ichneumon) Small Asian mongoose
Small Asian mongoose
(H. javanicus) Long-nosed mongoose
Long-nosed mongoose
(H. naso) Collared mongoose
Collared mongoose
(H. semitorquatus) Ruddy mongoose
Ruddy mongoose
(H. smithii) Crab-eating mongoose
Crab-eating mongoose
(H. urva) Stripe-necked mongoose
Stripe-necked mongoose
(H. vitticollis)

Ichneumia

White-tailed mongoose
White-tailed mongoose
(I. albicauda)

Liberiictus

Liberian mongoose
Liberian mongoose
(L. kuhni)

Mungos

Gambian mongoose
Gambian mongoose
(M. gambianus) Banded mongoose
Banded mongoose
(M. mungo)

Paracynictis

Selous' mongoose
Selous' mongoose
(P. selousi)

Rhynchogale

Meller's mongoose
Meller's mongoose
(R. melleri)

Suricata

Meerkat
Meerkat
(S. suricatta)

Hyaenidae (Hyenas)

Crocuta

Spotted hyena
Spotted hyena
(C. crocuta)

Hyaena

Brown hyena
Brown hyena
(H. brunnea) Striped hyena
Striped hyena
(H. hyaena)

Proteles

Aardwolf
Aardwolf
(P. cristatus)

Felidae

Large family listed below

Viverridae

Large family listed below

Eupleridae

Small family listed below

Family Felidae

Felinae

Acinonyx

Cheetah
Cheetah
(A. jubatus)

Caracal

Caracal
Caracal
(C. caracal) African golden cat
African golden cat
(C. aurata)

Catopuma

Bay cat
Bay cat
(C. badia) Asian golden cat
Asian golden cat
(C. temminckii)

Felis

European wildcat
European wildcat
(F. silvestris) African wildcat
African wildcat
(F. lybica) Jungle cat
Jungle cat
(F. chaus) Black-footed cat
Black-footed cat
(F. nigripes) Sand cat
Sand cat
(F. margarita) Chinese mountain cat
Chinese mountain cat
(F. bieti) Domestic cat (F. catus)

Leopardus

Ocelot
Ocelot
(L. pardalis) Margay
Margay
(L. wiedii) Pampas cat
Pampas cat
(L. colocola) Geoffroy's cat
Geoffroy's cat
(L. geoffroyi) Kodkod
Kodkod
(L. guigna) Andean mountain cat
Andean mountain cat
(L. jacobita) Oncilla
Oncilla
(L. tigrinus) Southern tigrina
Southern tigrina
(L. guttulus)

Leptailurus

Serval
Serval
(L. serval)

Lynx

Canadian lynx (L. canadensis) Eurasian lynx
Eurasian lynx
(L. lynx) Iberian lynx
Iberian lynx
(L. pardinus) Bobcat
Bobcat
(L. rufus)

Otocolobus

Pallas's cat
Pallas's cat
(O. manul)

Pardofelis

Marbled cat
Marbled cat
(P. marmorata)

Prionailurus

Fishing cat
Fishing cat
(P. viverrinus) Leopard
Leopard
cat (P. bengalensis) Sundaland leopard cat (P. javanensis) Flat-headed cat
Flat-headed cat
(P. planiceps) Rusty-spotted cat
Rusty-spotted cat
(P. rubiginosus)

Puma

Cougar
Cougar
(P. concolor)

Herpailurus

Jaguarundi
Jaguarundi
(H. yagouaroundi)

Pantherinae

Panthera

Lion
Lion
(P. leo) Jaguar
Jaguar
(P. onca) Leopard
Leopard
(P. pardus) Tiger
Tiger
(P. tigris) Snow leopard
Snow leopard
(P. uncia)

Neofelis

Clouded leopard
Clouded leopard
(N. nebulosa) Sunda clouded leopard
Sunda clouded leopard
(N. diardi)

Family Viverridae
Viverridae
(includes Civets)

Paradoxurinae

Arctictis

Binturong
Binturong
(A. binturong)

Arctogalidia

Small-toothed palm civet
Small-toothed palm civet
(A. trivirgata)

Macrogalidia

Sulawesi palm civet
Sulawesi palm civet
(M. musschenbroekii)

Paguma

Masked palm civet
Masked palm civet
(P. larvata)

Paradoxurus

Golden wet-zone palm civet (P. aureus) Asian palm civet
Asian palm civet
(P. hermaphroditus) Jerdon's palm civet (P. jerdoni) Golden palm civet
Golden palm civet
(P. zeylonensis)

Hemigalinae

Chrotogale

Owston's palm civet
Owston's palm civet
(C. owstoni)

Cynogale

Otter civet
Otter civet
(C. bennettii)

Diplogale

Hose's palm civet
Hose's palm civet
(D. hosei)

Hemigalus

Banded palm civet
Banded palm civet
(H. derbyanus)

Prionodontinae (Asiatic linsangs)

Prionodon

Banded linsang
Banded linsang
(P. linsang) Spotted linsang
Spotted linsang
(P. pardicolor)

Viverrinae

Civettictis

African civet
African civet
(C. civetta)

Genetta (Genets)

Abyssinian genet
Abyssinian genet
(G. abyssinica) Angolan genet
Angolan genet
(G. angolensis) Bourlon's genet
Bourlon's genet
(G. bourloni) Crested servaline genet
Crested servaline genet
(G. cristata) Common genet
Common genet
(G. genetta) Johnston's genet
Johnston's genet
(G. johnstoni) Rusty-spotted genet
Rusty-spotted genet
(G. maculata) Pardine genet
Pardine genet
(G. pardina) Aquatic genet
Aquatic genet
(G. piscivora) King genet
King genet
(G. poensis) Servaline genet
Servaline genet
(G. servalina) Haussa genet
Haussa genet
(G. thierryi) Cape genet
Cape genet
(G. tigrina) Giant forest genet
Giant forest genet
(G. victoriae)

Poiana

African linsang
African linsang
(P. richardsonii) Leighton's linsang
Leighton's linsang
(P. leightoni)

Viverra

Malabar large-spotted civet
Malabar large-spotted civet
(V. civettina) Large-spotted civet
Large-spotted civet
(V. megaspila) Malayan civet
Malayan civet
(V. tangalunga) Large Indian civet
Large Indian civet
(V. zibetha)

Viverricula

Small Indian civet
Small Indian civet
(V. indica)

Family Eupleridae

Euplerinae

Cryptoprocta

Fossa (C. ferox)

Eupleres

Eastern falanouc
Eastern falanouc
(E. goudotii) Western falanouc (E. major)

Fossa

Malagasy civet
Malagasy civet
(F. fossana)

Galidiinae

Galidia

Ring-tailed mongoose
Ring-tailed mongoose
(G. elegans)

Galidictis

Broad-striped Malagasy mongoose
Broad-striped Malagasy mongoose
(G. fasciata) Grandidier's mongoose
Grandidier's mongoose
(G. grandidieri)

Mungotictis

Narrow-striped mongoose
Narrow-striped mongoose
(M. decemlineata)

Salanoia

Brown-tailed mongoose
Brown-tailed mongoose
(S. concolor) Durrell's vontsira (S. durrelli)

Suborder Caniformia
Caniformia
(cont. below)

Ursidae (Bears)

Ailuropoda

Giant panda
Giant panda
(A. melanoleuca)

Helarctos

Sun bear
Sun bear
(H. malayanus)

Melursus

Sloth bear
Sloth bear
(M. ursinus)

Tremarctos

Spectacled bear
Spectacled bear
(T. ornatus)

Ursus

American black bear
American black bear
(U. americanus) Brown bear
Brown bear
(U. arctos) Polar bear
Polar bear
(U. maritimus) Asian black bear
Asian black bear
(U. thibetanus)

Mephitidae

Conepatus (Hog-nosed skunks)

Molina's hog-nosed skunk
Molina's hog-nosed skunk
(C. chinga) Humboldt's hog-nosed skunk
Humboldt's hog-nosed skunk
(C. humboldtii) American hog-nosed skunk
American hog-nosed skunk
(C. leuconotus) Striped hog-nosed skunk
Striped hog-nosed skunk
(C. semistriatus)

Mephitis

Hooded skunk
Hooded skunk
(M. macroura) Striped skunk
Striped skunk
(M. mephitis)

Mydaus

Sunda stink badger
Sunda stink badger
(M. javanensis) Palawan stink badger
Palawan stink badger
(M. marchei)

Spilogale (Spotted skunks)

Southern spotted skunk
Southern spotted skunk
(S. angustifrons) Western spotted skunk
Western spotted skunk
(S. gracilis) Eastern spotted skunk
Eastern spotted skunk
(S. putorius) Pygmy spotted skunk
Pygmy spotted skunk
(S. pygmaea)

Procyonidae

Bassaricyon (Olingos)

Eastern lowland olingo
Eastern lowland olingo
(B. alleni) Northern olingo
Northern olingo
(B. gabbii) Western lowland olingo
Western lowland olingo
(B. medius) Olinguito
Olinguito
(B. neblina)

Bassariscus

Ring-tailed cat
Ring-tailed cat
(B. astutus) Cacomistle
Cacomistle
(B. sumichrasti)

Nasua (Coatis inclusive)

White-nosed coati
White-nosed coati
(N. narica) South American coati
South American coati
(N. nasua)

Nasuella (Coatis inclusive)

Western mountain coati (N. olivacea) Eastern mountain coati (N. meridensis)

Potos

Kinkajou
Kinkajou
(P. flavus)

Procyon

Crab-eating raccoon
Crab-eating raccoon
(P. cancrivorus) Raccoon
Raccoon
(P. lotor) Cozumel raccoon
Cozumel raccoon
(P. pygmaeus)

Ailuridae

Ailurus

Red panda
Red panda
(A. fulgens)

Suborder Caniformia
Caniformia
(cont. above)

Otariidae (Eared seals) (includes fur seals and sea lions) ( Pinniped
Pinniped
inclusive)

Arctocephalus

South American fur seal
South American fur seal
(A. australis) Australasian fur seal (A. forsteri) Galápagos fur seal
Galápagos fur seal
(A. galapagoensis) Antarctic fur seal
Antarctic fur seal
(A. gazella) Juan Fernández fur seal
Juan Fernández fur seal
(A. philippii) Brown fur seal
Brown fur seal
(A. pusillus) Guadalupe fur seal
Guadalupe fur seal
(A. townsendi) Subantarctic fur seal
Subantarctic fur seal
(A. tropicalis)

Callorhinus

Northern fur seal
Northern fur seal
(C. ursinus)

Eumetopias

Steller sea lion
Steller sea lion
(E. jubatus)

Neophoca

Australian sea lion
Australian sea lion
(N. cinerea)

Otaria

South American sea lion
South American sea lion
(O. flavescens)

Phocarctos

New Zealand sea lion
New Zealand sea lion
(P. hookeri)

Zalophus

California sea lion
California sea lion
(Z. californianus) Galápagos sea lion
Galápagos sea lion
(Z. wollebaeki)

Odobenidae ( Pinniped
Pinniped
inclusive)

Odobenus

Walrus
Walrus
(O. rosmarus)

Phocidae (Earless seals) ( Pinniped
Pinniped
inclusive)

Cystophora

Hooded seal
Hooded seal
(C. cristata)

Erignathus

Bearded seal
Bearded seal
(E. barbatus)

Halichoerus

Gray seal (H. grypus)

Histriophoca

Ribbon seal
Ribbon seal
(H. fasciata)

Hydrurga

Leopard
Leopard
seal (H. leptonyx)

Leptonychotes

Weddell seal
Weddell seal
(L. weddellii)

Lobodon

Crabeater seal
Crabeater seal
(L. carcinophagus)

Mirounga (Elephant seals)

Northern elephant seal
Northern elephant seal
(M. angustirostris) Southern elephant seal
Southern elephant seal
(M. leonina)

Monachus

Mediterranean monk seal
Mediterranean monk seal
(M. monachus) Hawaiian monk seal
Hawaiian monk seal
(M. schauinslandi)

Ommatophoca

Ross seal
Ross seal
(O. rossi)

Pagophilus

Harp seal
Harp seal
(P. groenlandicus)

Phoca

Spotted seal
Spotted seal
(P. largha) Harbor seal
Harbor seal
(P. vitulina)

Pusa

Caspian seal
Caspian seal
(P. caspica) Ringed seal
Ringed seal
(P. hispida) Baikal seal
Baikal seal
(P. sibirica)

Canidae

Large family listed below

Mustelidae

Large family listed below

Family Canidae
Canidae
(includes dogs)

Atelocynus

Short-eared dog
Short-eared dog
(A. microtis)

Canis

Side-striped jackal
Side-striped jackal
(C. adustus) African golden wolf
African golden wolf
(C. anthus) Golden jackal
Golden jackal
(C. aureus) Coyote
Coyote
(C. latrans) Gray wolf
Gray wolf
(C. lupus) Black-backed jackal
Black-backed jackal
(C. mesomelas) Red wolf
Red wolf
(C. rufus) Ethiopian wolf
Ethiopian wolf
(C. simensis)

Cerdocyon

Crab-eating fox
Crab-eating fox
(C. thous)

Chrysocyon

Maned wolf
Maned wolf
(C. brachyurus)

Cuon

Dhole
Dhole
(C. alpinus)

Lycalopex

Culpeo
Culpeo
(L. culpaeus) Darwin's fox
Darwin's fox
(L. fulvipes) South American gray fox
South American gray fox
(L. griseus) Pampas fox
Pampas fox
(L. gymnocercus) Sechuran fox
Sechuran fox
(L. sechurae) Hoary fox
Hoary fox
(L. vetulus)

Lycaon

African wild dog
African wild dog
(L. pictus)

Nyctereutes

Raccoon
Raccoon
dog (N. procyonoides)

Otocyon

Bat-eared fox
Bat-eared fox
(O. megalotis)

Speothos

Bush dog
Bush dog
(S. venaticus)

Urocyon

Gray fox
Gray fox
(U. cinereoargenteus) Island fox
Island fox
(U. littoralis)

Vulpes (Foxes)

Bengal fox
Bengal fox
(V. bengalensis) Blanford's fox
Blanford's fox
(V. cana) Cape fox
Cape fox
(V. chama) Corsac fox
Corsac fox
(V. corsac) Tibetan sand fox
Tibetan sand fox
(V. ferrilata) Arctic fox
Arctic fox
(V. lagopus) Kit fox
Kit fox
(V. macrotis) Pale fox
Pale fox
(V. pallida) Rüppell's fox
Rüppell's fox
(V. rueppelli) Swift fox
Swift fox
(V. velox) Red fox
Red fox
(V. vulpes) Fennec fox
Fennec fox
(V. zerda)

Family Mustelidae

Lutrinae (Otters)

Aonyx

African clawless otter
African clawless otter
(A. capensis) Oriental small-clawed otter
Oriental small-clawed otter
(A. cinerea)

Enhydra

Sea otter
Sea otter
(E. lutris)

Hydrictis

Spotted-necked otter
Spotted-necked otter
(H. maculicollis)

Lontra

North American river otter
North American river otter
(L. canadensis) Marine otter
Marine otter
(L. felina) Neotropical otter
Neotropical otter
(L. longicaudis) Southern river otter
Southern river otter
(L. provocax)

Lutra

Eurasian otter
Eurasian otter
(L. lutra) Hairy-nosed otter
Hairy-nosed otter
(L. sumatrana)

Lutrogale

Smooth-coated otter
Smooth-coated otter
(L. perspicillata)

Pteronura

Giant otter
Giant otter
(P. brasiliensis)

Mustelinae (including badgers)

Arctonyx

Hog badger
Hog badger
(A. collaris)

Eira

Tayra
Tayra
(E. barbara)

Galictis

Lesser grison
Lesser grison
(G. cuja) Greater grison
Greater grison
(G. vittata)

Gulo

Wolverine
Wolverine
(G. gulo)

Ictonyx

Saharan striped polecat
Saharan striped polecat
(I. libyca) Striped polecat
Striped polecat
(I. striatus)

Lyncodon

Patagonian weasel
Patagonian weasel
(L. patagonicus)

Martes (Martens)

American marten
American marten
(M. americana) Yellow-throated marten
Yellow-throated marten
(M. flavigula) Beech marten
Beech marten
(M. foina) Nilgiri marten
Nilgiri marten
(M. gwatkinsii) European pine marten
European pine marten
(M. martes) Japanese marten
Japanese marten
(M. melampus) Sable
Sable
(M. zibellina)

Pekania

Fisher (P. pennanti)

Meles

Japanese badger
Japanese badger
(M. anakuma) Asian badger
Asian badger
(M. leucurus) European badger
European badger
(M. meles)

Mellivora

Honey badger
Honey badger
(M. capensis)

Melogale (Ferret-badgers)

Bornean ferret-badger
Bornean ferret-badger
(M. everetti) Chinese ferret-badger
Chinese ferret-badger
(M. moschata) Javan ferret-badger
Javan ferret-badger
(M. orientalis) Burmese ferret-badger
Burmese ferret-badger
(M. personata)

Mustela (Weasels and Ferrets)

Amazon weasel
Amazon weasel
(M. africana) Mountain weasel
Mountain weasel
(M. altaica) Stoat
Stoat
(M. erminea) Steppe polecat
Steppe polecat
(M. eversmannii) Colombian weasel
Colombian weasel
(M. felipei) Long-tailed weasel
Long-tailed weasel
(M. frenata) Japanese weasel
Japanese weasel
(M. itatsi) Yellow-bellied weasel
Yellow-bellied weasel
(M. kathiah) European mink
European mink
(M. lutreola) Indonesian mountain weasel
Indonesian mountain weasel
(M. lutreolina) Black-footed ferret
Black-footed ferret
(M. nigripes) Least weasel
Least weasel
(M. nivalis) Malayan weasel
Malayan weasel
(M. nudipes) European polecat
European polecat
(M. putorius) Siberian weasel
Siberian weasel
(M. sibirica) Back-striped weasel
Back-striped weasel
(M. strigidorsa) Egyptian weasel
Egyptian weasel
(M. subpalmata)

Neovison (Minks)

American mink
American mink
(N. vison)

Poecilogale

African striped weasel
African striped weasel
(P. albinucha)

Taxidea

American badger
American badger
(T. taxus)

Vormela

Marbled polecat
Marbled polecat
(V. peregusna)

Authority control

LCCN: sh85126973 GND: 4267568-6

Taxon
Taxon
identifiers

Wd: Q178089 ADW: Crocuta_crocuta ARKive: crocuta-crocuta EoL: 311569 Fossilworks: 53881 GBIF: 5218781 iNaturalist: 41886 ITIS: 621907 IUCN: 5674 MSW: 1

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