Spotted Hyena
Overview
The spotted hyena also known as laughing hyena, is a carnivorous
Carnivore
A carnivore meaning 'meat eater' is an organism that derives its energy and nutrient requirements from a diet consisting mainly or exclusively of animal tissue, whether through predation or scavenging...

 mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

 of the family Hyaenidae, of which it is the largest extant member. Though the species' prehistoric range included Eurasia
Eurasia
Eurasia is a continent or supercontinent comprising the traditional continents of Europe and Asia ; covering about 52,990,000 km2 or about 10.6% of the Earth's surface located primarily in the eastern and northern hemispheres...

 extending from Atlantic Europe
Atlantic Europe
Atlantic Europe is a geographical and anthropological term for the western portion of Europe which borders the Atlantic Ocean. The term may refer to the idea of Atlantic Europe as a cultural unit and/or as an biogeographical region....

 to China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

, it now only occurs in all of Africa
Africa
Africa is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...

 south of the Sahara
Sahara
The Sahara is the world's second largest desert, after Antarctica. At over , it covers most of Northern Africa, making it almost as large as Europe or the United States. The Sahara stretches from the Red Sea, including parts of the Mediterranean coasts, to the outskirts of the Atlantic Ocean...

 save for the Congo Basin
Congo Basin
The Congo Basin is the sedimentary basin that is the drainage of the Congo River of west equatorial Africa. The basin begins in the highlands of the East African Rift system with input from the Chambeshi River, the Uele and Ubangi Rivers in the upper reaches and the Lualaba River draining wetlands...

. Spotted hyenas live in large matriarchal communities called clans, which can consist of up to 80 individuals.

Though often mislabeled as cowardly scavenger
Scavenger
Scavenging is both a carnivorous and herbivorous feeding behavior in which individual scavengers search out dead animal and dead plant biomass on which to feed. The eating of carrion from the same species is referred to as cannibalism. Scavengers play an important role in the ecosystem by...

s, spotted hyenas derive the majority of their nourishment by hunting medium sized ungulate
Ungulate
Ungulates are several groups of mammals, most of which use the tips of their toes, usually hoofed, to sustain their whole body weight while moving. They make up several orders of mammals, of which six to eight survive...

s, and frequently clash with lion
Lion
The lion is one of the four big cats in the genus Panthera, and a member of the family Felidae. With some males exceeding 250 kg in weight, it is the second-largest living cat after the tiger...

s over food and territory.
Encyclopedia
The spotted hyena also known as laughing hyena, is a carnivorous
Carnivore
A carnivore meaning 'meat eater' is an organism that derives its energy and nutrient requirements from a diet consisting mainly or exclusively of animal tissue, whether through predation or scavenging...

 mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

 of the family Hyaenidae, of which it is the largest extant member. Though the species' prehistoric range included Eurasia
Eurasia
Eurasia is a continent or supercontinent comprising the traditional continents of Europe and Asia ; covering about 52,990,000 km2 or about 10.6% of the Earth's surface located primarily in the eastern and northern hemispheres...

 extending from Atlantic Europe
Atlantic Europe
Atlantic Europe is a geographical and anthropological term for the western portion of Europe which borders the Atlantic Ocean. The term may refer to the idea of Atlantic Europe as a cultural unit and/or as an biogeographical region....

 to China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

, it now only occurs in all of Africa
Africa
Africa is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...

 south of the Sahara
Sahara
The Sahara is the world's second largest desert, after Antarctica. At over , it covers most of Northern Africa, making it almost as large as Europe or the United States. The Sahara stretches from the Red Sea, including parts of the Mediterranean coasts, to the outskirts of the Atlantic Ocean...

 save for the Congo Basin
Congo Basin
The Congo Basin is the sedimentary basin that is the drainage of the Congo River of west equatorial Africa. The basin begins in the highlands of the East African Rift system with input from the Chambeshi River, the Uele and Ubangi Rivers in the upper reaches and the Lualaba River draining wetlands...

. Spotted hyenas live in large matriarchal communities called clans, which can consist of up to 80 individuals.

Though often mislabeled as cowardly scavenger
Scavenger
Scavenging is both a carnivorous and herbivorous feeding behavior in which individual scavengers search out dead animal and dead plant biomass on which to feed. The eating of carrion from the same species is referred to as cannibalism. Scavengers play an important role in the ecosystem by...

s, spotted hyenas derive the majority of their nourishment by hunting medium sized ungulate
Ungulate
Ungulates are several groups of mammals, most of which use the tips of their toes, usually hoofed, to sustain their whole body weight while moving. They make up several orders of mammals, of which six to eight survive...

s, and frequently clash with lion
Lion
The lion is one of the four big cats in the genus Panthera, and a member of the family Felidae. With some males exceeding 250 kg in weight, it is the second-largest living cat after the tiger...

s over food and territory. Studies indicate that their social intelligence is on par with some primate
Primate
A primate is a mammal of the order Primates , which contains prosimians and simians. Primates arose from ancestors that lived in the trees of tropical forests; many primate characteristics represent adaptations to life in this challenging three-dimensional environment...

 species. The relative amount of frontal cortex in their brains not used for motor control is higher than in other carnivorans examined, which has been attributed to their complex social lives.

The spotted hyena features prominently in African mythology and folklore
African Traditional Religion
The traditional religions indigenous to Africa have, for most of their existence, been orally rather than scripturally transmitted. They are generally associated with animism. Most have ethno-based creations stories...

, where its portrayal varies from being a bringer of light, to a symbol of immorality and depravity.

Evolution

It is thought that the ancestors of the spotted hyena branched off from the true hyenas (striped
Striped Hyena
The Striped Hyena is a species of true hyena native to North and East Africa, the Caucasus, the Middle East, Middle and Central Asia and the Indian Subcontinent...

 and brown hyena
Brown Hyena
The brown hyena is a species of hyena which occurs in Namibia, Botswana, western and southern Zimbabwe, southern Mozambique and South Africa. It is currently the rarest hyena.-Description:...

s) during the Pliocene
Pliocene
The Pliocene Epoch is the period in the geologic timescale that extends from 5.332 million to 2.588 million years before present. It is the second and youngest epoch of the Neogene Period in the Cenozoic Era. The Pliocene follows the Miocene Epoch and is followed by the Pleistocene Epoch...

 era, 5.332 million to 1.806 million years ago. Ancestral spotted hyenas probably developed social behaviours in response to increased pressure from rivals on carcasses, thus forcing them to operate in teams. Spotted hyenas evolved sharp carnassials behind their crushing premolars, therefore they did not need to wait for their prey to die, as is the case for brown and striped hyenas, and thus became pack hunters as well as scavengers. They began forming increasingly larger territories, necessitated by the fact that their prey was often migratory, and long chases in a small territory would have caused them to encroach into another clan's turf. The evolution of pack behaviour in hyenas likely influenced the ancestors of lion
Lion
The lion is one of the four big cats in the genus Panthera, and a member of the family Felidae. With some males exceeding 250 kg in weight, it is the second-largest living cat after the tiger...

s into first forming prides, in order to better defend their kills. According to the fossil record, the species first evolved
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

 in the Indian Subcontinent
Indian subcontinent
The Indian subcontinent, also Indian Subcontinent, Indo-Pak Subcontinent or South Asian Subcontinent is a region of the Asian continent on the Indian tectonic plate from the Hindu Kush or Hindu Koh, Himalayas and including the Kuen Lun and Karakoram ranges, forming a land mass which extends...

. Spotted hyenas colonized the Middle East
Middle East
The Middle East is a region that encompasses Western Asia and Northern Africa. It is often used as a synonym for Near East, in opposition to Far East...

, Africa and the Ice Age
Ice age
An ice age or, more precisely, glacial age, is a generic geological period of long-term reduction in the temperature of the Earth's surface and atmosphere, resulting in the presence or expansion of continental ice sheets, polar ice sheets and alpine glaciers...

 plains of Eurasia
Eurasia
Eurasia is a continent or supercontinent comprising the traditional continents of Europe and Asia ; covering about 52,990,000 km2 or about 10.6% of the Earth's surface located primarily in the eastern and northern hemispheres...

 extending from Atlantic Europe
Atlantic Europe
Atlantic Europe is a geographical and anthropological term for the western portion of Europe which borders the Atlantic Ocean. The term may refer to the idea of Atlantic Europe as a cultural unit and/or as an biogeographical region....

 to China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

 where a large subspecies
Subspecies
Subspecies in biological classification, is either a taxonomic rank subordinate to species, ora taxonomic unit in that rank . A subspecies cannot be recognized in isolation: a species will either be recognized as having no subspecies at all or two or more, never just one...

 known as C. c. spelaea or "cave hyena
Cave Hyena
The Cave Hyena is an extinct subspecies of spotted hyena native to Eurasia, ranging from Northern China to Spain and into the British Isles...

" developed as a response to the cold climate. Naturalists and paleontologists originally assumed that the cave hyena was a separate species from the spotted hyena, due to large differences in fore and hind extremities. This was first put into question by Björn Kurtén, who stated “[...] there is evidence that this European population was continuous with southern, typical representatives of the nominate subspecies”. This was corroborated by genetic analysis' in 2004, showing no differences in DNA
DNA
Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...

 between the two populations. With the decline of grasslands 12,500 years ago, Europe experienced a massive loss of lowland habitats favoured by cave hyenas, and a corresponding increase in mixed woodlands. Cave hyenas, under these circumstances, would have been outcompeted by wolves and human
Human
Humans are the only living species in the Homo genus...

s which were as much at home in forests as in open lands, and in highlands as in lowlands. Cave hyena populations began to shrink after roughly 20,000 years ago, completely disappearing from Western Europe
Western Europe
Western Europe is a loose term for the collection of countries in the western most region of the European continents, though this definition is context-dependent and carries cultural and political connotations. One definition describes Western Europe as a geographic entity—the region lying in the...

 between 14–11,000 years ago, and earlier in some areas. The spotted hyena only vanished from the Middle East in the early Holocene
Holocene
The Holocene is a geological epoch which began at the end of the Pleistocene and continues to the present. The Holocene is part of the Quaternary period. Its name comes from the Greek words and , meaning "entirely recent"...

 around 8000 years ago, and was replaced in this region by the striped hyena. Since then, it has been confined to Sub-Saharian Africa.

History, systematics and naming

It is thought that the spotted hyena conforms to the chaus described by Pliny the Elder
Pliny the Elder
Gaius Plinius Secundus , better known as Pliny the Elder, was a Roman author, naturalist, and natural philosopher, as well as naval and army commander of the early Roman Empire, and personal friend of the emperor Vespasian...

, which was later described by Linnaeus as being part of the cat tribe. It is also thought to be the Crocotta
Crocotta
The crocotta , is a mythical dog-wolf of India or Ethiopia, linked to the hyena and said to be a deadly enemy of men and dogs.-Ancient Accounts:...

of Strabo
Strabo
Strabo, also written Strabon was a Greek historian, geographer and philosopher.-Life:Strabo was born to an affluent family from Amaseia in Pontus , a city which he said was situated the approximate equivalent of 75 km from the Black Sea...

, which he thought to be a wolf-dog hybrid. Sculptured representations indicate that the species was rarely encountered by the Ancient Egypt
Ancient Egypt
Ancient Egypt was an ancient civilization of Northeastern Africa, concentrated along the lower reaches of the Nile River in what is now the modern country of Egypt. Egyptian civilization coalesced around 3150 BC with the political unification of Upper and Lower Egypt under the first pharaoh...

ians, who considered them exotic enough to include them in their menageries of foreign animals and to exclude them from their sacred animals. Certain scholars interpret Aristotle
Aristotle
Aristotle was a Greek philosopher and polymath, a student of Plato and teacher of Alexander the Great. His writings cover many subjects, including physics, metaphysics, poetry, theater, music, logic, rhetoric, linguistics, politics, government, ethics, biology, and zoology...

's inaccurate description of striped hyenas as being hermaphroditic animals as being a confusion between the striped and spotted species.

Systematics

In his 12th edition
12th edition of Systema Naturae
The 12th edition of was the last edition of to be overseen by its author, Carl Linnaeus. It was published in three volumes, with parts appearing from 1766 to 1768...

 of Systema Naturae
Systema Naturae
The book was one of the major works of the Swedish botanist, zoologist and physician Carolus Linnaeus. The first edition was published in 1735...

, Linnaeus placed hyenas into the genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 Canis
Canis
Canis is a genus containing 7 to 10 extant species, including dogs, wolves, coyotes, and jackals, and many extinct species.-Wolves, dogs and dingos:Wolves, dogs and dingos are subspecies of Canis lupus...

, between wolves and fox
Fox
Fox is a common name for many species of omnivorous mammals belonging to the Canidae family. Foxes are small to medium-sized canids , characterized by possessing a long narrow snout, and a bushy tail .Members of about 37 species are referred to as foxes, of which only 12 species actually belong to...

es. Brisson
Mathurin Jacques Brisson
Mathurin Jacques Brisson was a French zoologist and natural philosopher.Brisson was born at Fontenay-le-Comte. The earlier part of his life was spent in the pursuit of natural history, his published works in this department including Le Règne animal and Ornithologie...

 had already given the form a generic distinction under the name Hyæna. In his own edition of Linnaeus' Systema Naturæ, Johann Friedrich Gmelin
Johann Friedrich Gmelin
Johann Friedrich Gmelin was a German naturalist, botanist, entomologist, herpetologist and malacologist.- Education :Johann Friedrich Gmelin was born as the eldest son of Philipp Friedrich Gmelin in 1748 in Tübingen...

 gave the spotted species the binomial name Canis crocuta, though Thomas Pennant
Thomas Pennant
Thomas Pennant was a Welsh naturalist and antiquary.The Pennants were a Welsh gentry family from the parish of Whitford, Flintshire, who had built up a modest estate at Bychton by the seventeenth century...

 had previously described it under the title of Hyæna, and placed it under the category of "Spotted Hyænas". Georges Cuvier
Georges Cuvier
Georges Chrétien Léopold Dagobert Cuvier or Jean Léopold Nicolas Frédéric Cuvier , known as Georges Cuvier, was a French naturalist and zoologist...

 made Hyænas into the last subdivision of digitigrade
Digitigrade
A digitigrade is an animal that stands or walks on its digits, or toes. Digitigrades include walking birds , cats, dogs, and many other mammals, but not plantigrades or unguligrades...

s, following viverrids and preceding felids. Cuvier was convinced that there were at least two different species of spotted hyena, based on regional differences in coat colours. However, subsequent naturalists did not accept this, for although they noted coat variations, there were no other differences to fully warrant classing them as different species. John Edward Gray
John Edward Gray
John Edward Gray, FRS was a British zoologist. He was the elder brother of George Robert Gray and son of the pharmacologist and botanist Samuel Frederick Gray ....

 later brought the spotted hyena under the Felidae
Felidae
Felidae is the biological family of the cats; a member of this family is called a felid. Felids are the strictest carnivores of the thirteen terrestrial families in the order Carnivora, although the three families of marine mammals comprising the superfamily pinnipedia are as carnivorous as the...

, placing it within a category including other hyenas and the aardwolf
Aardwolf
The aardwolf is a small, insectivorous mammal, native to Eastern and Southern Africa. The name means "earth wolf" in Afrikaans/Dutch. It is also called "maanhaar jackal". It is related to hyenas, but unlike its relatives, it does not hunt large prey. This unusual animal preys on insects...

. M. Lesson arranged the hyænids under his third section of digitigrades, a section consisting of animals lacking a small tooth behind the lower molar. The spotted hyena was placed between aardwolves and cats, and was termed Hyæna capensis.

Local and indigenous names

Several languages of Africa
Languages of Africa
There are over 2100 and by some counts over 3000 languages spoken natively in Africa in several major language families:*Afro-Asiatic spread throughout the Middle East, North Africa, the Horn of Africa, and parts of the Sahel...

 lack species specific names for hyenas: for example, the spotted and striped species have identical names in Dioula
Dioula language
Jula is a Mande language spoken in Burkina Faso, Côte d'Ivoire and Mali. It is one of the Manding languages, and is most closely related to Bambara, being mutually intelligible with Bambara as well as Malinke. It is a trade language in West Africa and is spoken by millions of people, either as a...

, Swahili
Swahili language
Swahili or Kiswahili is a Bantu language spoken by various ethnic groups that inhabit several large stretches of the Mozambique Channel coastline from northern Kenya to northern Mozambique, including the Comoro Islands. It is also spoken by ethnic minority groups in Somalia...

, Malinké
Mandinka people
The Mandinka, Malinke are one of the largest ethnic groups in West Africa with an estimated population of eleven million ....

, Mòoré, Ngambaye
Ngambay language
Ngambay is one of the major languages spoken by Sara people in southwestern Chad, northeastern Cameroon and eastern Nigeria, with about a million native speakers. Ngambay is the most widely-spoken of the Sara languages, and is used as a trade language between speakers of other dialects...

, Oulof
Wolof language
Wolof is a language spoken in Senegal, The Gambia, and Mauritania, and is the native language of the Wolof people. Like the neighbouring languages Serer and Fula, it belongs to the Atlantic branch of the Niger–Congo language family...

 and Fula
Fula language
The Fula or Fulani language is a language of West Africa. It is spoken as a first language by the and related groups from Senegambia and Guinea to Cameroon and Sudan...

. 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
Aardwolf
The aardwolf is a small, insectivorous mammal, native to Eastern and Southern Africa. The name means "earth wolf" in Afrikaans/Dutch. It is also called "maanhaar jackal". It is related to hyenas, but unlike its relatives, it does not hunt large prey. This unusual animal preys on insects...

 fisi ndogo.

Afrendille: Walaba Afrikaans
Afrikaans
Afrikaans is a West Germanic language, spoken natively in South Africa and Namibia. It is a daughter language of Dutch, originating in its 17th century dialects, collectively referred to as Cape Dutch .Afrikaans is a daughter language of Dutch; see , , , , , .Afrikaans was historically called Cape...

:
Gevlekte hiëna Amharinja: Djibb Arabic (CAR
Central African Republic
The Central African Republic , is a landlocked country in Central Africa. It borders Chad in the north, Sudan in the north east, South Sudan in the east, the Democratic Republic of the Congo and the Republic of the Congo in the south, and Cameroon in the west. The CAR covers a land area of about ,...

)
and Arabic (Chad
Chad
Chad , officially known as the Republic of Chad, is a landlocked country in Central Africa. It is bordered by Libya to the north, Sudan to the east, the Central African Republic to the south, Cameroon and Nigeria to the southwest, and Niger to the west...

)
: Marfaïn Arabic (North Africa):D'ba Arabi (Ethiopia): Dibb Ateso
Ateso language
Ateso is a Nilo-Saharan language, spoken by the Iteso people of Uganda and Kenya. It is one of the Teso–Turkana cluster of languages....

:
Ibuin Avukaia: Labagu Babouté: Mangou Baguirmien: Niougo Baka
Bongo-Baka languages
The Bongo languages, or Bongo–Baka, comprise half a dozen languages spoken in South Sudan. They are members of the Central Sudanic language family.The most populous Bongo language is Jur Modo, spoken by a hundred thousand people...

:
Libagu Bakola: Mazzobé Bambara
Bambara language
Bambara, more correctly known as Bamanankan , its designation in the language itself , is a language spoken in Mali by as many as six million people...

:
Namakoro, suruku Banda
Banda languages
Banda is a family of Ubangian languages spoken by the Banda people of Central Africa.-Languages:Olson classfies the Banda family as follows :*Central**Central Banda...

:
Bongo Bechuana: Piri, phiri Bemba (Zambia)
Bemba language
The Bemba language, ChiBemba , is a major Bantu language spoken primarily in north-eastern Zambia by the Bemba people and as a lingua franca by about 18 related ethnic groups, including the Bisa people of Mpika and Lake Bangweulu, and to a lesser extent in Katanga in the Democratic Republic of the...

:
Chimbwi Bornouan: Boultou Creole: Lobo Danakil
Danakil
Danakil may refer to the following:*Danakil, an old ethnonym referring to:**Afar people**Afar language*Afar Depression *Danakil Desert...

:
Jangóula Dioula: Suruku, namakoro Elkoni: Makatiet nyenegea English
English language
English is a West Germanic language that arose in the Anglo-Saxon kingdoms of England and spread into what was to become south-east Scotland under the influence of the Anglian medieval kingdom of Northumbria...

:
Spotted hyena, Tiger wolf, Wolf of the Cape colonists French
French language
French is a Romance language spoken as a first language in France, the Romandy region in Switzerland, Wallonia and Brussels in Belgium, Monaco, the regions of Quebec and Acadia in Canada, and by various communities elsewhere. Second-language speakers of French are distributed throughout many parts...

:
Hyène tachetée Fula (Fulfulde/Pulaar/Pular)
Fula language
The Fula or Fulani language is a language of West Africa. It is spoken as a first language by the and related groups from Senegambia and Guinea to Cameroon and Sudan...

: Bonooru, fowru, deppuru Galaorabéjsa: Wårabéssa, orabéjsa Gambe: Mangili German
German language
German is a West Germanic language, related to and classified alongside English and Dutch. With an estimated 90 – 98 million native speakers, German is one of the world's major languages and is the most widely-spoken first language in the European Union....

:
Tüpfelhyäne, Fleckenhyäne Gouragi: Woraba Gourmatche
Gurma
Gourma redirects here. For the Burkina Faso province, see Gourma Province.Gurma is an ethnic group living mainly in Burkina Faso, around Fada N'Gourma, and also in northern areas of Togo and Benin, as well as southwestern Niger...

:
Namlino Harari: Worábba Hassānīya Arabic
Hassānīya Arabic
Hassānīya is the variety of Arabic originally spoken by the Beni Hassān Bedouin tribes, who extended their authority over most of Mauritania and the Western Sahara between the fifteenth and seventeenth centuries. It has almost completely replaced the Berber languages spoken in this region...

:
Guervave Haussa
Hausa people
The Hausa are one of the largest ethnic groups in West Africa. They are a Sahelian people chiefly located in northern Nigeria and southeastern Niger, but having significant numbers living in regions of Cameroon, Ghana, Cote d'Ivoire, Chad and Sudan...

:
Koura Herero
Herero language
The Herero language is a language of the Bantu family . It is spoken by the Herero people in Namibia and Botswana...

:
Mbúngu-mbidíwa Ila (Zambia): Kabwenga Jita
Jita
The Jita are an ethnic and linguistic group based in Mara Region in northern Tanzania, on the southeastern shore of Lake Victoria. In 1987 the Jita population was estimated to number 217,000 .-References:...

:
Imembe Kalenjin
Kalenjin language
The Nandi languages, or Kalenjin proper, are a dialect cluster of the Kalenjin branch of the Nilotic language family.In Kenya, where speakers make up 18% of the population, the name Kalenjin, a Nandi expression meaning "I say ", gained prominence in the late 1940s and the early 1950s, when several...

:
Kimatet Kaonde
Kaonde language
Kaonde, also known as Chikaonde and Kawonde, is a Bantu language that is spoken primarily in Zambia but also in the Democratic Republic of the Congo. Kaonde and its dialects are spoken and understood by perhaps 350,000 people or more. It is estimated that approximately 3% of Zambians are native...

:
Mungolwe Karamojong
Karamojong
The Karamojong or Karimojong, are an ethnic group of agro-pastoral herders living mainly in the north-east of Uganda. Their language is also known as Karamojong or Karimojong, and is part of the Nilo-Saharan language group.-History:...

:
Ebu, Etutui Kichagga: Ingurunju, ifulu Kigogo
Kigogo
Kigogo is an administrative ward in the Kinondoni district of the Dar es Salaam Region of Tanzania. According to the 2002 census, the ward has a total population of 37,964.-References:...

and kikongo: Misi Kikamba and Kisukuma
Sukuma
The Sukuma is the largest ethnic group in Tanzania, with an estimated 5.5 million members representing about 16 percent of the country's total population. Sukuma means "north" and refers to "people of the north"...

:
Mbiti Kikondo: Mbulu Kiliangulu: Warabes Kiluba
Luba-Katanga language
Luba-Katanga, also known as Luba-Shaba and Kiluba, is one of two major Bantu language spoken in the Democratic Republic of the Congo called "Luba". It is spoken mostly in the south-east area of the country by the Luba people....

:
Kimburi Kimeru
Meru language
Meru is the language spoken by the Meru people who live on the Eastern and Northern slopes of Mount Kenya, Kenya, Africa and on the Nyambene ranges. They settled in this area after centuries of migration from the north.-Tradition:...

:
Mbitingaau Kinyarwanda: Impysi Kinyaturo: Mpiti Kinyiha: Ipatama Kipare
Kiparé
Kiparé is a village in the commune of Parakou in the Borgou Department of central-eastern Benin. It is located south of Parakou city centre.-External links:*...

 and Kizigua:
Ibau Kirangi: Mbichi Kisungwa: Fifi Koniagui: Iriguni Kota
Bakota
The Bakota are a Bantu ethnic group from the northeastern region of Gabon. The language they speak is called iKota, but is sometimes referred to as Bakota, ikuta, Kota, and among the Fang, they are known as Mekora. The language has several dialects, which include: Ndambomo, Mahongwe,...

:
Massoba Kotoko: Machi Kunda
Kunda people
The Kunda people are an ethnic group of Zambia, Zimbabwe, and Mozambique. They number approximately at 193,913 people. They speak the Chikunda/Chinyanja language, a Bantu language in the Bantu subfamily....

:
Tika Jita
Jita
The Jita are an ethnic and linguistic group based in Mara Region in northern Tanzania, on the southeastern shore of Lake Victoria. In 1987 the Jita population was estimated to number 217,000 .-References:...

:
Imembe Kikuyu
Gikuyu language
Gikuyu or Kikuyu is a language of the Bantu family spoken primarily by the Kikuyu people of Kenya. Numbering about 6 million , they are the largest ethnic group in Kenya. Gikuyu is spoken in the area between Nyeri and Nairobi. Gikuyu is one of the five languages of the Thagichu subgroup of the...

:
Hiti Kimeru
Meru language
Meru is the language spoken by the Meru people who live on the Eastern and Northern slopes of Mount Kenya, Kenya, Africa and on the Nyambene ranges. They settled in this area after centuries of migration from the north.-Tradition:...

:
Mbitingaau Kinyiha: Impatama Kirangi: Mbichi Kisukuma
Sukuma language
Sukuma is a Bantu language spoken by over five million people in central Tanzania. It forms a dialect continuum with Nyamwezi, but is more distinct from other neighboring languages....

, Kikamba and Kimaragoli: Mbiti Kisungwa: Fifi Swahili
Swahili language
Swahili or Kiswahili is a Bantu language spoken by various ethnic groups that inhabit several large stretches of the Mozambique Channel coastline from northern Kenya to northern Mozambique, including the Comoro Islands. It is also spoken by ethnic minority groups in Somalia...

:
Fisi, Nyangao Kitaita: Mbisi Ganda and Runyoro: Empisi Lugbara
Lugbara language
Lugbara is the language of the Lugbara people. It is spoken in the West Nile region in northwestern Uganda, as well as the Democratic Republic of the Congo's Orientale Province. Lugbara was first written by Christian missionaries in 1918, based on the Ayivu dialect...

:
Rara Luhya: Namunyu Luo
Luo languages
The Nilotic Luo languages, or Lwoian, are a dozen languages spoken by the Luo peoples in an area ranging from southern Sudan via Uganda to southern Kenya, with Dholuo extending into northern Tanzania and Alur into the Democratic Republic of the Congo. They form one of the two branches of Western...

:
Otoyo Lwo
Dholuo language
The Luo language, Dholuo or Luo proper, is the eponymous language of the Luo group of Nilotic languages, spoken by about 4.4 million Luo people of Kenya and Tanzania, who occupy parts of the eastern shore of Lake Victoria and areas to the south...

:
Lagwara Madi
Madi language (Sudan and Uganda)
The Ma'di language is a found in Uganda and South Sudan. . The Madi people refer to their language as Ma'di ti literally: Ma'di mouth....

:
Ebowu Maasai
Maasai language
The Maasai language is an Eastern Nilotic language spoken in Southern Kenya and Northern Tanzania by the Maasai people, numbering about 800,000...

:
Ondilili, Oln'gojine Sebei
Sebei
Sebei is an ethnic group of Uganda and Sudan. They speak Sebei, a Nilotic language. Many members of this ethnic group live in Kapchorwa District of Uganda....

:
Mangatiet Malay
Malay language
Malay is a major language of the Austronesian family. It is the official language of Malaysia , Indonesia , Brunei and Singapore...

:
Dubuk

Physical description

Spotted hyenas are the largest of extant hyenas. Their hair is shorter than those of striped hyena
Striped Hyena
The Striped Hyena is a species of true hyena native to North and East Africa, the Caucasus, the Middle East, Middle and Central Asia and the Indian Subcontinent...

s, and their mane
Mane
- Biology :* mane of a horse, the line of hair along the spine of the neck* mane of a lion, found around the male mammal's neck- Places :* Mane Department, a department in the Sanmatenga Province of Burkina Faso* French communes:...

s less full. Spotted hyenas have powerful forequarters and necks which rival those of leopard
Leopard
The leopard , Panthera pardus, is a member of the Felidae family and the smallest of the four "big cats" in the genus Panthera, the other three being the tiger, lion, and jaguar. The leopard was once distributed across eastern and southern Asia and Africa, from Siberia to South Africa, but its...

s, though comparatively small hindquarters. The rump
Rump (animal)
The rump or croup, in the external morphology of an animal, is the portion of the posterior dorsum that is posterior to the loins and anterior to the tail. Anatomically, the rump corresponds to the sacrum....

 is rounded rather than angular, which prevents attackers chasing from behind getting a firm grip on it. Female spotted hyenas are considerably larger than males, weighing 12% more. Adults measure 95–165.8 cm (37.4–65.3 in) in body length, plus a tail of 25–35 cm (9.8–13.8 in), and have a shoulder height of 70 –. Adult male spotted hyenas in the Serengeti
Serengeti
The Serengeti ecosystem is a geographical region in Africa. It is located in north Tanzania and extends to south-western Kenya between latitudes 1 and 3 S and longitudes 34 and 36 E. It spans some ....

 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
Zambia , officially the Republic of Zambia, is a landlocked country in Southern Africa. The neighbouring countries are the Democratic Republic of the Congo to the north, Tanzania to the north-east, Malawi to the east, Mozambique, Zimbabwe, Botswana and Namibia to the south, and Angola to the west....

 tend to be heavier, with males weighing on average 67.6 kg (149 lb), and females 69.2 kg (153 lb). Macdonald
David W. Macdonald
David Whyte Macdonald CBE FRSE is a Scottish zoologist and conservationist. He is the Director of the Wildlife Conservation Research Unit , Oxford University, which he founded in 1986...

 (1992) gives a maximum weight of 81.7 kg (180 lb), while Kingdon (1977) gives one of 86 kg (190 lb). The skulls of Zambian hyenas are also 7% longer and wider than those of Serengeti populations. It has been estimated that adult members of the now extinct Eurasian populations weighed 102 kg (225 lbs).

Their dentition is more dual purposed than that of other modern hyena species, which are mostly scavengers: the upper and lower third premolar
Premolar
The premolar teeth or bicuspids are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant, making eight premolars total in the mouth. They have at least two cusps. Premolars can be considered as a 'transitional tooth' during chewing, or...

s are conical bone-crushers, with a third bone-holding cone jutting from the lower fourth premolar. Spotted hyenas also have carnassial
Carnassial
Carnassials are large teeth found in many carnivorous mammals, used for shearing flesh and bone in a scissor- or shear-like way. In the Carnivora, the carnassials are the modified last upper premolar and the first molar, but in the prehistoric creodonts, the carnassials were further back in the...

s behind their bone-crushing premolars, the position of which allows hyenas to crush bone with their premolars without blunting their carnassials. The carnassials themselves are proportionately larger than those of other carnivorous mammals. Although they possess disproportionately large teeth to counteract wear, three year old spotted hyenas have teeth as worn as those of six year old lions. Combined with large jaw muscles and a special vaulting to protect the skull against large forces, these characteristics give spotted hyenas a powerful bite which can exert a pressure of 453 kgf/cm2 (6443 lbf/in²), which is 40% more force than a leopard can generate. An experiment conducted by Savage (1955) demonstrated how the jaws of spotted hyenas outmatch those of brown bear
Brown Bear
The brown bear is a large bear distributed across much of northern Eurasia and North America. It can weigh from and its largest subspecies, the Kodiak Bear, rivals the polar bear as the largest member of the bear family and as the largest land-based predator.There are several recognized...

s in bonecrushing ability, and free ranging hyenas have been observed to crack open the long bones of giraffes measuring 7 cm in diameter. Although once thought to have the most powerful jaws among extant carnivorous mammals, other animals such as the Tasmanian Devil
Tasmanian Devil
The Tasmanian devil is a carnivorous marsupial of the family Dasyuridae, now found in the wild only on the Australian island state of Tasmania. The size of a small dog, it became the largest carnivorous marsupial in the world following the extinction of the thylacine in 1936...

 have been proven to have even stronger bites.

With the exception of size, there is little sexual dimorphism
Sexual dimorphism
Sexual dimorphism is a phenotypic difference between males and females of the same species. Examples of such differences include differences in morphology, ornamentation, and behavior.-Examples:-Ornamentation / coloration:...

 in spotted hyenas. The external genitalia of females closely resemble those of males: the 15 cm (6 inch) clitoris
Clitoris
The clitoris is a sexual organ that is present only in female mammals. In humans, the visible button-like portion is located near the anterior junction of the labia minora, above the opening of the urethra and vagina. Unlike the penis, which is homologous to the clitoris, the clitoris does not...

 is similar in shape and position to a penis
Penis
The penis is a biological feature of male animals including both vertebrates and invertebrates...

, and is capable of erection
Erection
Penile erection is a physiological phenomenon where the penis becomes enlarged and firm. Penile erection is the result of a complex interaction of psychological, neural, vascular and endocrine factors, and is usually, though not exclusively, associated with sexual arousal...

. The only visible difference between the penis of male spotted hyenas and the clitoris of females is that the latter's organ has a blunter tip. The labia are fused together into a pair of fibrous sacs resembling a scrotum
Scrotum
In some male mammals the scrotum is a dual-chambered protuberance of skin and muscle containing the testicles and divided by a septum. It is an extension of the perineum, and is located between the penis and anus. In humans and some other mammals, the base of the scrotum becomes covered with curly...

. Typically, when observing sexually mature animals, naturalists use the presence of nipple
Nipple
In its most general form, a nipple is a structure from which a fluid emanates. More specifically, it is the projection on the breasts or udder of a mammal by which breast milk is delivered to a mother's young. In this sense, it is often called a teat, especially when referring to non-humans, and...

s as an indicator of gender when observing spotted hyenas at a distance. Females have two nipples and rarely four. The color and spotting of the coat varies with age and individual. The number of spots tends to decrease with age.

Although there are no different extant subspecies, spotted hyenas do display a degree of regional variation, particularly in their southern range, where they tend to be darker and browner in color, particularly on the back and legs. Due to this darker hue, the spots of southern spotted hyenas are less defined and angular than their cousins on the West Coast
West Coast, Western Cape
The West Coast is a region of the Western Cape province in South Africa and is bordered by the Atlantic Ocean on the west and the Swartland region on the east....

. Also, the fur is longer in the South African form, particularly around the ears. Specimens from the former cameroons
Cameroons
British Cameroons was a British Mandate territory in West Africa, now divided between Nigeria and Cameroon.The area of present-day Cameroon was claimed by Germany as a protectorate during the "Scramble for Africa" at the end of the 19th century...

, the Epukiro district of former German West Africa and northern and western Togo
Togo
Togo, officially the Togolese Republic , is a country in West Africa bordered by Ghana to the west, Benin to the east and Burkina Faso to the north. It extends south to the Gulf of Guinea, on which the capital Lomé is located. Togo covers an area of approximately with a population of approximately...

 have proportionately longer tails than average. The spots of Cameroon spotted hyenas are greatly elongated in the hind region, with the main spots being almost streaks.

Spotted hyenas have powerful night vision
Night vision
Night vision is the ability to see in low light conditions. Whether by biological or technological means, night vision is made possible by a combination of two approaches: sufficient spectral range, and sufficient intensity range...

, which allows them to recognize each other in complete darkness, even if they are downwind.

Behavior

Spotted hyenas will rest and give birth in dens, which they rarely dig themselves: they will often use the abandoned lairs of warthog
Warthog
The Warthog or Common Warthog is a wild member of the pig family that lives in grassland, savanna, and woodland in Sub-Saharan Africa. In the past it was commonly treated as a subspecies of P...

s, springhare
Springhare
The springhare , or springhaas, is not actually a hare, but a member of the order Rodentia. It is one of a number of species in the genus Pedetes, and is native to southern Africa. Synonyms are P. caffer or P...

s and jackal
Jackal
Although the word jackal has been historically used to refer to many small- to medium-sized species of the wolf genus of mammals, Canis, today it most properly and commonly refers to three species: the black-backed jackal and the side-striped jackal of sub-Saharan Africa, and the golden jackal of...

s. A single den can house several females and dozens of cubs at once. Unlike grey wolves, it is not uncommon for spotted hyenas to accommodate cubs of different litters in one den. Spotted hyenas will sometimes live in close proximity to warthog
Warthog
The Warthog or Common Warthog is a wild member of the pig family that lives in grassland, savanna, and woodland in Sub-Saharan Africa. In the past it was commonly treated as a subspecies of P...

s, sharing mud holes and sleeping within a few metres of each other. Spotted hyenas may sleep in the open if the weather is not too hot, but otherwise they will rest near lakes, streams or in mud or dense shrubs. Unlike most social carnivores, spotted hyenas still display some atavistic behaviours of their solitary ancestors: spotted hyenas still head out for food alone, but later return to their community. Like other hyenas, spotted hyenas have two anal scent glands, which open into the rectum
Rectum
The rectum is the final straight portion of the large intestine in some mammals, and the gut in others, terminating in the anus. The human rectum is about 12 cm long...

 just inside the anal opening, though these glands are less elaborate than those of other hyena species. The white paste produced by these glands is deposited on grass stalks, and produces a powerful soapy odour which even humans can detect. Pasting is performed on a number of different occasions, such as when walking alone, when around a kill, when lions are present, by males and cubs near dens, and most frequently by parties of hyenas at territorial boundaries. Pasting is often followed by scratching the ground with their forepaws, which adds further scents from their interdigital glands.

Social behavior

Spotted hyenas are more social than grey wolves, but their groups are not as closely knit as African wild dog
African Wild Dog
Lycaon pictus is a large canid found only in Africa, especially in savannas and lightly wooded areas. It is variously called the African wild dog, African hunting dog, Cape hunting dog, painted dog, painted wolf, painted hunting dog, spotted dog, or ornate wolf...

s. Spotted hyena societies are more complex than those of other carnivorous mammals, and have been reported to be remarkably similar to those of cercopithecine primate
Primate
A primate is a mammal of the order Primates , which contains prosimians and simians. Primates arose from ancestors that lived in the trees of tropical forests; many primate characteristics represent adaptations to life in this challenging three-dimensional environment...

s in respect to group size, structure, competition and cooperation. Like primates, spotted hyenas use multiple sensory modalities, recognise individual conspecifics, are conscious that some clan-mates may be more reliable than others, recognise 3rd 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. Group size
Group size measures
Many animals, including humans, tend to live in groups, herds, flocks, bands, packs, shoals, or colonies of conspecific individuals. The size of these groups, as expressed by the number of participant individuals, is an important aspect of their social environment...

 is variable; a "clan" of spotted hyenas can include 5–90 members and is led by a single alpha female called the matriarch. Scientists theorise that female hyena dominance could be an adaptation to the length of time it takes for cubs to develop the massive skulls and jaws, and intense feeding competition within clans, thus necessitating greater attention and dominating behaviours from females. Female hyena dominance is sometimes explained by the unusually high concentration of androgens produced by the ovaries. However adult hyena males display a higher concentration of androgens than adult hyena females. This would suggest that adult concentrations of androgens probably do not account for the difference of social dominance.

Reproduction and development

Spotted hyenas are not seasonal breeders, and can reproduce at any time of the year, though a birth peak does occur during the wet season
Wet season
The the wet season, or rainy season, is the time of year, covering one or more months, when most of the average annual rainfall in a region occurs. The term green season is also sometimes used as a euphemism by tourist authorities. Areas with wet seasons are dispersed across portions of the...

. Females are polyestrous, with an estrous period lasting two weeks. The average litter consists of two cubs, with three occasionally being reported. Mating
Mating
In biology, mating is the pairing of opposite-sex or hermaphroditic organisms for copulation. In social animals, it also includes the raising of their offspring. Copulation is the union of the sex organs of two sexually reproducing animals for insemination and subsequent internal fertilization...

 in spotted hyenas is a relatively short affair which typically only occurs at night with no other hyenas present. Males will show submissive behaviour when approaching females in heat, even if the male outweighs its partner. There is no copulatory tie, as in canids. Females usually favour younger males born into, 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. Passive males tend to have greater success in courting females than aggressive ones. Males take no part in the raising of young. The length of the gestation period tends to vary greatly, though 110 days is the average length of time. In the final stages of pregnancy, dominant females provide their developing offspring with higher androgen
Androgen
Androgen, also called androgenic hormone or testoid, is the generic term for any natural or synthetic compound, usually a steroid hormone, that stimulates or controls the development and maintenance of male characteristics in vertebrates by binding to androgen receptors...

 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. 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.

Birth is difficult, as females have to give birth through their narrow clitoris
Clitoris
The clitoris is a sexual organ that is present only in female mammals. In humans, the visible button-like portion is located near the anterior junction of the labia minora, above the opening of the urethra and vagina. Unlike the penis, which is homologous to the clitoris, the clitoris does not...

. Also, spotted hyena cubs are the largest carnivoran cubs relative to their mother's weight. In captivity, many cubs of first time mothers are stillborn
Stillbirth
A stillbirth occurs when a fetus has died in the uterus. The Australian definition specifies that fetal death is termed a stillbirth after 20 weeks gestation or the fetus weighs more than . Once the fetus has died the mother still has contractions and remains undelivered. The term is often used in...

 because of the long labour times involved, and in the wild, it is estimated that 10% of first time mothers die during labour. The cubs are born with soft, brownish black hair, and weigh on average 1.5 kg. 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 from the moment they are born. This is particularly apparent in same sexed litters, and can result in the death of the weaker cub. This neonatal siblicide
Siblicide
Siblicide is the killing of an infant individual by its close relatives . It may occur directly between siblings or be mediated by the parents. The evolutionary drivers may be either indirect benefits for the genetic viability of a population or direct benefits for the perpetrators...

 can amount to 25% of overall spotted hyena cub mortality factors. Spotted hyena hierarchy
Hierarchy
A hierarchy is an arrangement of items in which the items are represented as being "above," "below," or "at the same level as" one another...

 is nepotistic: the offspring of dominant females automatically outrank adult females subordinate to their mother, though they can lose their privileges if the mother dies. 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 two-three 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.

Lactating females can carry 3–4 kg (6.5–9 lb) of milk in their udders. Spotted hyena milk is very rich, having the highest protein content (14.9%) of any terrestrial carnivore. The fat content (14.1%) is second only to the polar bear
Polar Bear
The polar bear is a bear native largely within the Arctic Circle encompassing the Arctic Ocean, its surrounding seas and surrounding land masses. It is the world's largest land carnivore and also the largest bear, together with the omnivorous Kodiak Bear, which is approximately the same size...

, so unlike lions and wild dogs, they can leave their cubs for about a week without feeding them. Cubs will nurse from their mother for 12 or 16 months, though they can process solid food as early as three months.

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.

Dietary habits

Spotted hyenas are better equipped for scavenging than other African predators: not only are they able to splinter and eat the largest ungulate bones, they are also able to digest them completely. Spotted hyenas can digest all organic components in bones, not just the marrow
Bone marrow
Bone marrow is the flexible tissue found in the interior of bones. In humans, bone marrow in large bones produces new blood cells. On average, bone marrow constitutes 4% of the total body mass of humans; in adults weighing 65 kg , bone marrow accounts for approximately 2.6 kg...

. 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. Wildebeest
Wildebeest
The wildebeest , also called the gnu is an antelope of the genus Connochaetes. It is a hooved mammal...

 are the most commonly taken medium sized ungulate prey item in both Ngorongoro and the Serengeti, with zebra
Zebra
Zebras are several species of African equids united by their distinctive black and white stripes. Their stripes come in different patterns unique to each individual. They are generally social animals that live in small harems to large herds...

 and Thomson's gazelle
Thomson's Gazelle
The Thomson's gazelle is one of the best-known gazelles. It is named after explorer Joseph Thomson and, as a result, is sometimes referred to as a "tommie"...

s coming close behind. Cape buffalo are rarely attacked due to differences in habitat preference, though adult bulls have been recorded to be taken on occasion. Spotted hyenas have also been found to catch fish
Fish
Fish are a paraphyletic group of organisms that consist of all gill-bearing aquatic vertebrate animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish, as well as various extinct related groups...

, tortoise
Tortoise
Tortoises are a family of land-dwelling reptiles of the order of turtles . Like their marine cousins, the sea turtles, tortoises are shielded from predators by a shell. The top part of the shell is the carapace, the underside is the plastron, and the two are connected by the bridge. The tortoise...

s, human
Human
Humans are the only living species in the Homo genus...

s, black rhino, hippo
Hippo
A hippo or hippopotamus is either of two species of large African mammal which live mainly in and near water:* Hippopotamus* Pygmy HippopotamusHippo may also refer to:-Given names:...

 calves, young African elephants, pangolin
Pangolin
A pangolin , also scaly anteater or Trenggiling, is a mammal of the order Pholidota. There is only one extant family and one genus of pangolins, comprising eight species. There are also a number of extinct taxa. Pangolins have large keratin scales covering their skin and are the only mammals with...

s, pythons, and a large number of different ungulate species. The fossil record indicates that Eurasian Spotted hyenas in what is now the Czech Republic
Czech Republic
The Czech Republic is a landlocked country in Central Europe. The country is bordered by Poland to the northeast, Slovakia to the east, Austria to the south, and Germany to the west and northwest....

 primarily fed on Przewalski's Horse
Przewalski's Horse
Przewalski's Horse or Dzungarian Horse, is a rare and endangered subspecies of wild horse native to the steppes of central Asia, specifically China and Mongolia.At one time extinct in the wild, it has been reintroduced to its native habitat in Mongolia at the Khustain Nuruu...

s. Other prey included woolly rhinoceros
Woolly Rhinoceros
The woolly rhinoceros is an extinct species of rhinoceros that was common throughout Europe and Asia during the Pleistocene epoch and survived the last glacial period. The genus name Coelodonta means "cavity tooth"...

, Reindeer
Reindeer
The reindeer , also known as the caribou in North America, is a deer from the Arctic and Subarctic, including both resident and migratory populations. While overall widespread and numerous, some of its subspecies are rare and one has already gone extinct.Reindeer vary considerably in color and size...

, Steppe Wisent
Steppe Wisent
The Steppe Bison or steppe wisent was a bison found on steppes throughout Europe, Central Asia, Beringia, and North America during the Quaternary...

, Irish elk
Irish Elk
The Irish Elk or Giant Deer , was a species of Megaloceros and one of the largest deer that ever lived. Its range extended across Eurasia, from Ireland to east of Lake Baikal, during the Late Pleistocene. The latest known remains of the species have been carbon dated to about 7,700 years ago...

, chamois
Chamois
The chamois, Rupicapra rupicapra, is a goat-antelope species native to mountains in Europe, including the Carpathian Mountains of Romania, the European Alps, the Tatra Mountains, the Balkans, parts of Turkey, and the Caucasus. The chamois has also been introduced to the South Island of New Zealand...

 and ibex. In Italy
Italy
Italy , officially the Italian Republic languages]] under the European Charter for Regional or Minority Languages. In each of these, Italy's official name is as follows:;;;;;;;;), is a unitary parliamentary republic in South-Central Europe. To the north it borders France, Switzerland, Austria and...

, their prey consisted of red deer
Red Deer
The red deer is one of the largest deer species. Depending on taxonomy, the red deer inhabits most of Europe, the Caucasus Mountains region, Asia Minor, parts of western Asia, and central Asia. It also inhabits the Atlas Mountains region between Morocco and Tunisia in northwestern Africa, being...

, aurochs
Aurochs
The aurochs , the ancestor of domestic cattle, were a type of large wild cattle which inhabited Europe, Asia and North Africa, but is now extinct; it survived in Europe until 1627....

, horse
Horse
The horse is one of two extant subspecies of Equus ferus, or the wild horse. It is a single-hooved mammal belonging to the taxonomic family Equidae. The horse has evolved over the past 45 to 55 million years from a small multi-toed creature into the large, single-toed animal of today...

s, roe deer
Roe Deer
The European Roe Deer , also known as the Western Roe Deer, chevreuil or just Roe Deer, is a Eurasian species of deer. It is relatively small, reddish and grey-brown, and well-adapted to cold environments. Roe Deer are widespread in Western Europe, from the Mediterranean to Scandinavia, and from...

, fallow deer
Fallow Deer
The Fallow Deer is a ruminant mammal belonging to the family Cervidae. This common species is native to western Eurasia, but has been introduced widely elsewhere. It often includes the rarer Persian Fallow Deer as a subspecies , while others treat it as an entirely different species The Fallow...

, wild boar and ibex. Spotted hyenas are thought to be responsible for the dis-articulation and destruction of some cave bear
Cave Bear
The cave bear was a species of bear that lived in Europe during the Pleistocene and became extinct at the beginning of the Last Glacial Maximum about 27,500 years ago....

 skeletons. Such large carcasses were an optimal food resource for the hyenas, especially at the end of winter, when food was scarce. However, they were less successful than cave lion
Cave lion
Panthera leo spelaea also known as the European or Eurasian cave lion, is an extinct subspecies of lion known from fossils and many examples of prehistoric art.-Physical characteristics:This subspecies was one of the largest lions...

s in navigating through cave bear dens, due to their inferior climbing abilities.

A single spotted hyena can eat at least 14.5 kg of meat per meal. 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. 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. 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. A single hyena can take less than two minutes in eating a gazelle fawn, while a group of 35 hyenas can completely consume an adult zebra in 36 minutes. Spotted hyenas do not require much water, and typically only spend 30 seconds drinking.

Hunting behavior

Unlike other large African carnivores, spotted hyenas do not preferentially prey on any species, and only buffalo, giraffe
Giraffe
The giraffe is an African even-toed ungulate mammal, the tallest of all extant land-living animal species, and the largest ruminant...

 and plains zebra
Plains Zebra
The plains zebra , also known as the common zebra or Burchell's zebra, is the most common and geographically widespread species of zebra. It ranges from the south of Ethiopia through East Africa to as far south as Angola and eastern South Africa...

 are significantly avoided. Spotted hyenas prefer prey with a body mass range of 56 –, with a mode of 102 kg (224.9 lb). 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 antipredator behaviours. Unlike grey wolves, spotted hyenas rely more on sight than smell when hunting, and do not follow their prey's prints or travel in single file.

Spotted hyenas usually hunt wildebeest either singly, or in groups of two or three. They catch adult wildebeest usually after 5 km chases at speeds of up to 60 km/h. Chases are usually initiated by one hyena, and with the exception of cows with calves, there is little active defense by the wildebeest herd. Wildebeest will sometimes attempt to escape hyenas by taking to water, though in such cases, the hyenas almost invariably catch them. Zebras require different hunting methods to those used for wildebeest, due to their habit of running in tight groups and aggressive defence from stallion
Stallion
A Stallion is a male horse.Stallion may also refer to:* Stallion , an American pop rock group* Stallion , a figure in the Gobot toyline* Stallion , a character in the console role-playing game series...

s. Hyenas seem to plan on hunting zebras in advance, as they tend to indulge in activities such as scent marking before setting off, a behaviour which does not occur when they target other prey species. Typical zebra hunting groups consist of 10–25 hyenas. During a chase, zebras typically move in tight bunches, with the hyenas pursuing behind in a crescent
Crescent
In art and symbolism, a crescent is generally the shape produced when a circular disk has a segment of another circle removed from its edge, so that what remains is a shape enclosed by two circular arcs of different diameters which intersect at two points .In astronomy, a crescent...

 formation. Chases are usually relatively slow, with an average speed of 15–30 km/h. A stallion will attempt to place itself 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. 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. Once prey is caught, spotted hyenas will kill their prey by eating it alive.

Spotted hyenas will increase their kill rate during the calving seasons of their prey, or when they are frequently displaced from their kills by other predators.

Interspecific predatory relationships

In areas where spotted hyenas and lion
Lion
The lion is one of the four big cats in the genus Panthera, and a member of the family Felidae. With some males exceeding 250 kg in weight, it is the second-largest living cat after the tiger...

s are sympatric, 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%. Lions typically ignore spotted hyenas, unless they are on a kill or are being harassed by them. Spotted hyenas themselves tend to visibly react to the presence of lions, whether there is food or not. Lions will readily appropriate the kills of spotted hyenas: in the Ngorongoro crater, it is common for lions to subsist largely on kills stolen from hyenas, causing the hyenas to increase their kill rate. Lions are quick to follow the calls of hyenas feeding, a fact which was proven by Dr. Hans Kruuk, who found that lions repeatedly approached him whenever he played the tape-recorded calls of hyenas feeding. 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. In some cases, spotted hyenas are bold enough to feed alongside lions, and may occasionally force the lions off a kill. Spotted hyenas usually prevail against groups of lionesses unaccompanied by males if they outnumber them 4:1. The two species may act aggressively toward one another even when there is no food involved. Lions may charge at hyenas and maul them for no apparent reason: one male lion was filmed killing two matriarch hyenas on separate occasions without eating them, 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. Occasionally, lion prides and spotted hyena clans may engage in full warfare, as with a case in early April, 1999 in Ethiopia
Ethiopia
Ethiopia , officially known as the Federal Democratic Republic of Ethiopia, is a country located in the Horn of Africa. It is the second-most populous nation in Africa, with over 82 million inhabitants, and the tenth-largest by area, occupying 1,100,000 km2...

, in which 6 lions and 35 hyenas were killed over a two week period. 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.

Although cheetah
Cheetah
The cheetah is a large-sized feline inhabiting most of Africa and parts of the Middle East. The cheetah is the only extant member of the genus Acinonyx, most notable for modifications in the species' paws...

s and leopard
Leopard
The leopard , Panthera pardus, is a member of the Felidae family and the smallest of the four "big cats" in the genus Panthera, the other three being the tiger, lion, and jaguar. The leopard was once distributed across eastern and southern Asia and Africa, from Siberia to South Africa, but its...

s 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, while leopards, particularly males, may stand up to hyenas. There are records of some male leopards preying on hyenas.

Spotted hyenas will follow packs of African wild dog
African Wild Dog
Lycaon pictus is a large canid found only in Africa, especially in savannas and lightly wooded areas. It is variously called the African wild dog, African hunting dog, Cape hunting dog, painted dog, painted wolf, painted hunting dog, spotted dog, or ornate wolf...

s in order to appropriate their kills. They will typically inspect areas where wild dogs have rested and eat any faeces 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.

Spotted hyenas dominate other hyena species wherever their ranges overlap. Brown hyena
Brown Hyena
The brown hyena is a species of hyena which occurs in Namibia, Botswana, western and southern Zimbabwe, southern Mozambique and South Africa. It is currently the rarest hyena.-Description:...

s 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 hyena
Striped Hyena
The Striped Hyena is a species of true hyena native to North and East Africa, the Caucasus, the Middle East, Middle and Central Asia and the Indian Subcontinent...

s in the Serengeti.

Jackal
Jackal
Although the word jackal has been historically used to refer to many small- to medium-sized species of the wolf genus of mammals, Canis, today it most properly and commonly refers to three species: the black-backed jackal and the side-striped jackal of sub-Saharan Africa, and the golden jackal of...

s will feed alongside hyenas, though they will be chased if they approach too closely. Spotted hyenas will sometimes follow jackals during the gazelle fawning season, as jackals are effective at tracking and catching young animals. Hyenas do not take to eating jackal flesh readily: four hyenas were reported to take half an hour in eating a golden jackal
Golden Jackal
The golden jackal , also known as the common jackal, Asiatic jackal, thos or gold-wolf is a Canid of the genus Canis indigenous to north and northeastern Africa, southeastern and central Europe , Asia Minor, the Middle East and southeast Asia...

. Overall, the two animals typically ignore each other when there is no food or young at stake.

Spotted hyenas usually keep a safe distance from Nile crocodile
Nile crocodile
The Nile crocodile or Common crocodile is an African crocodile which is common in Somalia, Ethiopia, Uganda, Kenya, Egypt, Tanzania, Zambia, Zimbabwe, Gabon, South Africa, Malawi, Sudan, Botswana, and Cameroon...

s. Though they readily take to water to catch and store prey, hyenas will avoid crocodile infested waters.

Now extinct spotted hyena populations living in Italy
Italy
Italy , officially the Italian Republic languages]] under the European Charter for Regional or Minority Languages. In each of these, Italy's official name is as follows:;;;;;;;;), is a unitary parliamentary republic in South-Central Europe. To the north it borders France, Switzerland, Austria and...

 shared their range with wolves, but managed to avoid competition by inhabiting lowlands, rather than the slopes favoured by wolves. Also, spotted hyenas primarily fed on horses, while wolves targeted ibex and roe deer. However, wolves and spotted hyenas seem to display negative abundance relations over time, with wolf populations expanding in the regions where hyenas disappeared.

Intelligence

Compared to other hyenas, spotted hyenas show a greater relative amount of frontal cortex exclusive to motor control functions. Studies strongly suggest convergent evolution
Convergent evolution
Convergent evolution describes the acquisition of the same biological trait in unrelated lineages.The wing is a classic example of convergent evolution in action. Although their last common ancestor did not have wings, both birds and bats do, and are capable of powered flight. The wings are...

 in spotted hyena and primate
Primate
A primate is a mammal of the order Primates , which contains prosimians and simians. Primates arose from ancestors that lived in the trees of tropical forests; many primate characteristics represent adaptations to life in this challenging three-dimensional environment...

 intelligence. A study done by evolutionary anthropologists demonstrated that spotted hyenas outperform chimpanzee
Chimpanzee
Chimpanzee, sometimes colloquially chimp, is the common name for the two extant species of ape in the genus Pan. The Congo River forms the boundary between the native habitat of the two species:...

s 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.

Body language

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.

Vocalisations

Spotted hyenas are very vocal animals, and produce a number of different calls. Generally, high pitched calls signify fear or submission, while low pitched calls accompany a high tendency to attack.

The loud "whoop" is a characteristic sound of the African night and is audible for over 5 km (3.1 mi) or more. It is a rallying cry, which varies in speed and pitch according to the urgency of the situation. Spotted hyenas also whoop to show off as individuals, the rate and style being an indicator of social status. Because of this, spotted hyenas whoop singly rather than in chorus as wolf packs do to display their collective strength. Although males tend to whoop more than females of similar rank, dominant females will engage in the longest bouts of whooping.
Giggles and grunt-laughter tend to be emitted in situations of great excitement, and perhaps indicate a conflicting tendency to flee or stay. The giggles, yells and grunts which accompany mass feeding tend to be directed at competing individuals at a carcass, and have the secondary, disadvantageous effect of attracting lions and other spotted hyenas. 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.
Soft grunts are made by females calling their cubs.
When attacked, spotted hyenas will emit loud growls and whimpers.

Current range and distribution

The largest known spotted hyena populations occur in the Serengeti ecosystem in Tanzania (where they number 7,200–7,700), Kruger National Park
Kruger National Park
Kruger National Park is one of the largest game reserves in Africa. It covers and extends from north to south and from east to west.To the west and south of the Kruger National Park are the two South African provinces of Limpopo and Mpumalanga. In the north is Zimbabwe, and to the east is...

 in South Africa
South Africa
The Republic of South Africa is a country in southern Africa. Located at the southern tip of Africa, it is divided into nine provinces, with of coastline on the Atlantic and Indian oceans...

 (1,300–3,900) and the Masai Mara
Masai Mara
The Maasai Mara National Reserve is a large game reserve in south-western Kenya, which is effectively the northern continuation of the Serengeti National Park in Tanzania...

 Game Reserve in Kenya
Kenya
Kenya , officially known as the Republic of Kenya, is a country in East Africa that lies on the equator, with the Indian Ocean to its south-east...

 (ca 500-1,000). Several hundred, unsurveyed individuals occur in Zimbabwe
Zimbabwe
Zimbabwe is a landlocked country located in the southern part of the African continent, between the Zambezi and Limpopo rivers. It is bordered by South Africa to the south, Botswana to the southwest, Zambia and a tip of Namibia to the northwest and Mozambique to the east. Zimbabwe has three...

an conservation areas, the Selous Game Reserve
Selous Game Reserve
The Selous Game Reserve is one of the largest faunal reserves of the world, located in the south of Tanzania. It was named after Englishman Sir Frederick Selous, a famous big game hunter and early conservationist, who died at Beho Beho in this territory in 1917 while fighting against the Germans...

 in Tanzania and the Okavango in Botswana
Botswana
Botswana, officially the Republic of Botswana , is a landlocked country located in Southern Africa. The citizens are referred to as "Batswana" . Formerly the British protectorate of Bechuanaland, Botswana adopted its new name after becoming independent within the Commonwealth on 30 September 1966...

. Spotted hyenas are considered by the IUCN to be of lower risk
Lower Risk
An animal with the conservation status of Lower Risk is one with populations sufficiently high so as to ensure its survival in the long run. Animals with this status do not qualify as being Vulnerable, Endangered or Critically Endangered....

 of extinction in Botswana, Ethiopia
Ethiopia
Ethiopia , officially known as the Federal Democratic Republic of Ethiopia, is a country located in the Horn of Africa. It is the second-most populous nation in Africa, with over 82 million inhabitants, and the tenth-largest by area, occupying 1,100,000 km2...

, Guinea Bissau, Kenya
Kenya
Kenya , officially known as the Republic of Kenya, is a country in East Africa that lies on the equator, with the Indian Ocean to its south-east...

, Namibia
Namibia
Namibia, officially the Republic of Namibia , is a country in southern Africa whose western border is the Atlantic Ocean. It shares land borders with Angola and Zambia to the north, Botswana to the east and South Africa to the south and east. It gained independence from South Africa on 21 March...

, South Africa, Tanzania and Zimbabwe. They are a threatened species
Threatened species
Threatened species are any speciesg animals, plants, fungi, etc.) which are vulnerable to endangerment in the near future.The World Conservation Union is the foremost authority on threatened species, and treats threatened species not as a single category, but as a group of three categories,...

 in Benin
Benin
Benin , officially the Republic of Benin, is a country in West Africa. It borders Togo to the west, Nigeria to the east and Burkina Faso and Niger to the north. Its small southern coastline on the Bight of Benin is where a majority of the population is located...

, Burundi
Burundi
Burundi , officially the Republic of Burundi , is a landlocked country in the Great Lakes region of Eastern Africa bordered by Rwanda to the north, Tanzania to the east and south, and the Democratic Republic of the Congo to the west. Its capital is Bujumbura...

 (where they are thought to be on the verge of extinction), Cameroon
Cameroon
Cameroon, officially the Republic of Cameroon , is a country in west Central Africa. It is bordered by Nigeria to the west; Chad to the northeast; the Central African Republic to the east; and Equatorial Guinea, Gabon, and the Republic of the Congo to the south. Cameroon's coastline lies on the...

, Mali
Mali
Mali , officially the Republic of Mali , is a landlocked country in Western Africa. Mali borders Algeria on the north, Niger on the east, Burkina Faso and the Côte d'Ivoire on the south, Guinea on the south-west, and Senegal and Mauritania on the west. Its size is just over 1,240,000 km² with...

, Mauritania
Mauritania
Mauritania is a country in the Maghreb and West Africa. It is bordered by the Atlantic Ocean in the west, by Western Sahara in the north, by Algeria in the northeast, by Mali in the east and southeast, and by Senegal in the southwest...

, Niger
Niger
Niger , officially named the Republic of Niger, is a landlocked country in Western Africa, named after the Niger River. It borders Nigeria and Benin to the south, Burkina Faso and Mali to the west, Algeria and Libya to the north and Chad to the east...

, Nigeria
Nigeria
Nigeria , officially the Federal Republic of Nigeria, is a federal constitutional republic comprising 36 states and its Federal Capital Territory, Abuja. The country is located in West Africa and shares land borders with the Republic of Benin in the west, Chad and Cameroon in the east, and Niger in...

, Rwanda
Rwanda
Rwanda or , officially the Republic of Rwanda , is a country in central and eastern Africa with a population of approximately 11.4 million . Rwanda is located a few degrees south of the Equator, and is bordered by Uganda, Tanzania, Burundi and the Democratic Republic of the Congo...

 and Sierra Leone
Sierra Leone
Sierra Leone , officially the Republic of Sierra Leone, is a country in West Africa. It is bordered by Guinea to the north and east, Liberia to the southeast, and the Atlantic Ocean to the west and southwest. Sierra Leone covers a total area of and has an estimated population between 5.4 and 6.4...

. They are extinct in Algeria
Algeria
Algeria , officially the People's Democratic Republic of Algeria , also formally referred to as the Democratic and Popular Republic of Algeria, is a country in the Maghreb region of Northwest Africa with Algiers as its capital.In terms of land area, it is the largest country in Africa and the Arab...

 and Lesotho
Lesotho
Lesotho , officially the Kingdom of Lesotho, is a landlocked country and enclave, surrounded by the Republic of South Africa. It is just over in size with a population of approximately 2,067,000. Its capital and largest city is Maseru. Lesotho is a member of the Commonwealth of Nations. The name...

. There is a deficiency of data
Data Deficient
Data Deficient is a category applied by the IUCN, other agencies, and individuals to a species when the available information is not sufficient for a proper assessment of conservation status to be made...

 on the number of spotted hyenas in Angola
Angola
Angola, officially the Republic of Angola , is a country in south-central Africa bordered by Namibia on the south, the Democratic Republic of the Congo on the north, and Zambia on the east; its west coast is on the Atlantic Ocean with Luanda as its capital city...

, Burkina Faso
Burkina Faso
Burkina Faso – also known by its short-form name Burkina – is a landlocked country in west Africa. It is surrounded by six countries: Mali to the north, Niger to the east, Benin to the southeast, Togo and Ghana to the south, and Côte d'Ivoire to the southwest.Its size is with an estimated...

, the Central African Republic
Central African Republic
The Central African Republic , is a landlocked country in Central Africa. It borders Chad in the north, Sudan in the north east, South Sudan in the east, the Democratic Republic of the Congo and the Republic of the Congo in the south, and Cameroon in the west. The CAR covers a land area of about ,...

, Chad
Chad
Chad , officially known as the Republic of Chad, is a landlocked country in Central Africa. It is bordered by Libya to the north, Sudan to the east, the Central African Republic to the south, Cameroon and Nigeria to the southwest, and Niger to the west...

, the Republic of the Congo
Republic of the Congo
The Republic of the Congo , sometimes known locally as Congo-Brazzaville, is a state in Central Africa. It is bordered by Gabon, Cameroon, the Central African Republic, the Democratic Republic of the Congo , the Angolan exclave province of Cabinda, and the Gulf of Guinea.The region was dominated by...

, Côte d'Ivoire
Côte d'Ivoire
The Republic of Côte d'Ivoire or Ivory Coast is a country in West Africa. It has an area of , and borders the countries Liberia, Guinea, Mali, Burkina Faso and Ghana; its southern boundary is along the Gulf of Guinea. The country's population was 15,366,672 in 1998 and was estimated to be...

, the Democratic Republic of Congo, Djibouti
Djibouti
Djibouti , officially the Republic of Djibouti , is a country in the Horn of Africa. It is bordered by Eritrea in the north, Ethiopia in the west and south, and Somalia in the southeast. The remainder of the border is formed by the Red Sea and the Gulf of Aden at the east...

, Eritrea
Eritrea
Eritrea , officially the State of Eritrea, is a country in the Horn of Africa. Eritrea derives it's name from the Greek word Erethria, meaning 'red land'. The capital is Asmara. It is bordered by Sudan in the west, Ethiopia in the south, and Djibouti in the southeast...

, Equatorial Guinea
Equatorial Guinea
Equatorial Guinea, officially the Republic of Equatorial Guinea where the capital Malabo is situated.Annobón is the southernmost island of Equatorial Guinea and is situated just south of the equator. Bioko island is the northernmost point of Equatorial Guinea. Between the two islands and to the...

, Ghana
Ghana
Ghana , officially the Republic of Ghana, is a country located in West Africa. It is bordered by Côte d'Ivoire to the west, Burkina Faso to the north, Togo to the east, and the Gulf of Guinea to the south...

, Guinea
Guinea
Guinea , officially the Republic of Guinea , is a country in West Africa. Formerly known as French Guinea , it is today sometimes called Guinea-Conakry to distinguish it from its neighbour Guinea-Bissau. Guinea is divided into eight administrative regions and subdivided into thirty-three prefectures...

, Malawi
Malawi
The Republic of Malawi is a landlocked country in southeast Africa that was formerly known as Nyasaland. It is bordered by Zambia to the northwest, Tanzania to the northeast, and Mozambique on the east, south and west. The country is separated from Tanzania and Mozambique by Lake Malawi. Its size...

, Mozambique
Mozambique
Mozambique, officially the Republic of Mozambique , is a country in southeastern Africa bordered by the Indian Ocean to the east, Tanzania to the north, Malawi and Zambia to the northwest, Zimbabwe to the west and Swaziland and South Africa to the southwest...

, Senegal
Senegal
Senegal , officially the Republic of Senegal , is a country in western Africa. It owes its name to the Sénégal River that borders it to the east and north...

, Somalia
Somalia
Somalia , officially the Somali Republic and formerly known as the Somali Democratic Republic under Socialist rule, is a country located in the Horn of Africa. Since the outbreak of the Somali Civil War in 1991 there has been no central government control over most of the country's territory...

, Sudan
Sudan
Sudan , officially the Republic of the Sudan , is a country in North Africa, sometimes considered part of the Middle East politically. It is bordered by Egypt to the north, the Red Sea to the northeast, Eritrea and Ethiopia to the east, South Sudan to the south, the Central African Republic to the...

, Swaziland
Swaziland
Swaziland, officially the Kingdom of Swaziland , and sometimes called Ngwane or Swatini, is a landlocked country in Southern Africa, bordered to the north, south and west by South Africa, and to the east by Mozambique...

, Togo
Togo
Togo, officially the Togolese Republic , is a country in West Africa bordered by Ghana to the west, Benin to the east and Burkina Faso to the north. It extends south to the Gulf of Guinea, on which the capital Lomé is located. Togo covers an area of approximately with a population of approximately...

, Uganda
Uganda
Uganda , officially the Republic of Uganda, is a landlocked country in East Africa. Uganda is also known as the "Pearl of Africa". It is bordered on the east by Kenya, on the north by South Sudan, on the west by the Democratic Republic of the Congo, on the southwest by Rwanda, and on the south by...

 and Zambia
Zambia
Zambia , officially the Republic of Zambia, is a landlocked country in Southern Africa. The neighbouring countries are the Democratic Republic of the Congo to the north, Tanzania to the north-east, Malawi to the east, Mozambique, Zimbabwe, Botswana and Namibia to the south, and Angola to the west....

.

Relationships with humans

Prehistoric interactions

Kills partially processed by Neanderthal
Neanderthal
The Neanderthal is an extinct member of the Homo genus known from Pleistocene specimens found in Europe and parts of western and central Asia...

 and then by cave hyenas indicate that hyenas would occasionally steal Neanderthal kills, and cave hyenas and Neanderthal both competed for cave sites. Many caves show alternating occupations of hyenas and Neanderthals. The discovery of a cave in Siberia
Siberia
Siberia is an extensive region constituting almost all of Northern Asia. Comprising the central and eastern portion of the Russian Federation, it was part of the Soviet Union from its beginning, as its predecessor states, the Tsardom of Russia and the Russian Empire, conquered it during the 16th...

's Altai Mountains, which showed evidence of being inhabited by spotted hyenas for about 40,000 years, led to speculation that the presence of spotted hyenas there could have prevented humans from crossing the Bering land bridge
Bering land bridge
The Bering land bridge was a land bridge roughly 1,000 miles wide at its greatest extent, which joined present-day Alaska and eastern Siberia at various times during the Pleistocene ice ages. Like most of Siberia and all of Manchuria, Beringia was not glaciated because snowfall was extremely light...

 to the Americas
Americas
The Americas, or America , are lands in the Western hemisphere, also known as the New World. In English, the plural form the Americas is often used to refer to the landmasses of North America and South America with their associated islands and regions, while the singular form America is primarily...

, thus explaining why humans colonised the New World
New World
The New World is one of the names used for the Western Hemisphere, specifically America and sometimes Oceania . The term originated in the late 15th century, when America had been recently discovered by European explorers, expanding the geographical horizon of the people of the European middle...

 much later than when land bridge had formed. This scenario was brought up due to there being large numbers of fossil human remains dating back 50,000–60,000 years ago below the latitude of Mongolia
Mongolia
Mongolia is a landlocked country in East and Central Asia. It is bordered by Russia to the north and China to the south, east and west. Although Mongolia does not share a border with Kazakhstan, its western-most point is only from Kazakhstan's eastern tip. Ulan Bator, the capital and largest...

, and comparatively few remains, dated back less than 12,000 years ago, further north, where hyenas ranged. The discovery of a 14,000 year old dog
Dog
The domestic dog is a domesticated form of the gray wolf, a member of the Canidae family of the order Carnivora. The term is used for both feral and pet varieties. The dog may have been the first animal to be domesticated, and has been the most widely kept working, hunting, and companion animal in...

 skull further led to the theory that the domestication of the dog may have been a factor in aiding the eventual crossing, as dogs would have been valuable sentinels against hyena incursions into human encampments.

Impacts on human culture and thought

Spotted hyenas vary in their folkloric and mythological depictions, depending on the ethnic group from which the tales originate. It is often difficult to know whether or not spotted hyenas are the specific hyena species featured in such stories, particularly in West Africa, as both spotted and striped hyenas are often given the same names. In western African tales, spotted hyenas are sometimes depicted as bad Muslim
Muslim
A Muslim, also spelled Moslem, is an adherent of Islam, a monotheistic, Abrahamic religion based on the Quran, which Muslims consider the verbatim word of God as revealed to prophet Muhammad. "Muslim" is the Arabic term for "submitter" .Muslims believe that God is one and incomparable...

s who challenge the local animism
Animism
Animism refers to the belief that non-human entities are spiritual beings, or at least embody some kind of life-principle....

 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, while West African folklore generally shows the hyena as symbolizing immorality, dirty habits, the reversal of normal activities, and other negative traits. In Tanzania, there is a belief that witches use spotted hyenas as mounts. In the Mtwara Region
Mtwara Region
Mtwara is a southern region of Tanzania which has been underdeveloped for a long time: development is constrained by the lack of highway and energy infrastructures. The Dar es Salaam-Kibiti-Lindi-Mtwara road has been improved by the completion of the Mkapa bridge over the Rufiji River...

 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. The Kaguru
Kaguru
The Kaguru, or Kagulu, are an ethnic and linguistic group based in central Tanzania. In 1987 the Kaguru population was estimated to number 217,000.- Geography :...

 of Tanzania
Tanzania
The United Republic of Tanzania is a country in East Africa bordered by Kenya and Uganda to the north, Rwanda, Burundi, and the Democratic Republic of the Congo to the west, and Zambia, Malawi, and Mozambique to the south. The country's eastern borders lie on the Indian Ocean.Tanzania is a state...

 and the Kujamaat of Southern Senegal
Senegal
Senegal , officially the Republic of Senegal , is a country in western Africa. It owes its name to the Sénégal River that borders it to the east and north...

 view hyenas as inedible and greedy hermaphrodites. A mythical African tribe called the Bouda
Bouda
Werehyenas are mythological or folkloric creatures capable of assuming the shape of hyenas. They are present in the stories of several African and Eurasian cultures...

 is reputed to house members able to transform into hyenas. A similar myth occurs in Mansoa
Mansôa
Mansôa is a town located in the Oio Region of Guinea-Bissau. Population 7,376 .-References:...

. These "werehyenas" are executed when discovered, but do not revert back to their human form when killed.

Spotted hyenas feature prominently in the rituals of certain African tribes. In the Gelede cult of the Yoruba people
Yoruba people
The Yoruba people are one of the largest ethnic groups in West Africa. The majority of the Yoruba speak the Yoruba language...

 of Benin
Benin
Benin , officially the Republic of Benin, is a country in West Africa. It borders Togo to the west, Nigeria to the east and Burkina Faso and Niger to the north. Its small southern coastline on the Bight of Benin is where a majority of the population is located...

 and Southwest Nigeria
Nigeria
Nigeria , officially the Federal Republic of Nigeria, is a federal constitutional republic comprising 36 states and its Federal Capital Territory, Abuja. The country is located in West Africa and shares land borders with the Republic of Benin in the west, Chad and Cameroon in the east, and Niger in...

, 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
Mali
Mali , officially the Republic of Mali , is a landlocked country in Western Africa. Mali borders Algeria on the north, Niger on the east, Burkina Faso and the Côte d'Ivoire on the south, Guinea on the south-west, and Senegal and Mauritania on the west. Its size is just over 1,240,000 km² with...

, 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. Kujamaat hunters traditionally treat the spotted hyenas they kill with the respect due to human elders, in order to avoid retribution from malevolent hyena spirits acting on behalf of the dead animal. In Maasai tradition, and that of other tribes, 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
Ox
An ox , also known as a bullock in Australia, New Zealand and India, is a bovine trained as a draft animal. Oxen are commonly castrated adult male cattle; castration makes the animals more tractable...

.

The vocalisation of the spotted hyena resembling hysterical human laughter
Laughter
Laughing is a reaction to certain stimuli, fundamentally stress, which serves as an emotional balancing mechanism. Traditionally, it is considered a visual expression of happiness, or an inward feeling of joy. It may ensue from hearing a joke, being tickled, or other stimuli...

 has been alluded to in numerous works of literature: "to laugh like a hyæna" was a common proverb
Proverb
A proverb is a simple and concrete saying popularly known and repeated, which expresses a truth, based on common sense or the practical experience of humanity. They are often metaphorical. A proverb that describes a basic rule of conduct may also be known as a maxim...

, and is featured in The Cobbler's Prophecy (1594), Webster's Duchess of Malfy (1623) and Shakespeares As You Like It
As You Like It
As You Like It is a pastoral comedy by William Shakespeare believed to have been written in 1599 or early 1600 and first published in the folio of 1623. The play's first performance is uncertain, though a performance at Wilton House in 1603 has been suggested as a possibility...

, Act IV. Sc.1.

Attacks on humans

Spotted hyenas are usually timid around humans, and will typically flee over a distance of 300 metres when an approaching human is detected. Although spotted hyenas do prey on humans in modern times, such incidences are rare. However, according to the SGDRN (Sociedade para a Gestão e Desenvolvimento da Reserva do Niassa Moçambique), attacks on humans by spotted hyenas are likely to be underreported. Hyenas are known to have preyed on humans in prehistory: human hair has been found in fossilised hyena dung dating back 195,000 to 257,000 years. According to Dr. Hans Kruuk, man-eating
Man-eater
Man-eater is a colloquial term for an animal that preys upon humans. This does not include scavenging. Although human beings can be attacked by many kinds of animals, man-eaters are those that have incorporated human flesh into their usual diet...

 spotted hyenas tend to be very large specimens: a pair of man-eating hyenas, responsible for killing 27 people in Mlanje, Malawi
Malawi
The Republic of Malawi is a landlocked country in southeast Africa that was formerly known as Nyasaland. It is bordered by Zambia to the northwest, Tanzania to the northeast, and Mozambique on the east, south and west. The country is separated from Tanzania and Mozambique by Lake Malawi. Its size...

 in 1962 and were weighed at 72 kg (159 lb) and 77 kg (170 lb) after being shot. 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. According to R.G. Burton's A Book of Man-Eaters, spotted hyenas will enter human encampments without paying any notice of camp fire
Fire
Fire is the rapid oxidation of a material in the chemical process of combustion, releasing heat, light, and various reaction products. Slower oxidative processes like rusting or digestion are not included by this definition....

s. Victims of spotted hyenas tend to be women, children and sick or infirm men: Theodore Roosevelt
Theodore Roosevelt
Theodore "Teddy" Roosevelt was the 26th President of the United States . He is noted for his exuberant personality, range of interests and achievements, and his leadership of the Progressive Movement, as well as his "cowboy" persona and robust masculinity...

 wrote on how in 1908–09 in Uganda
Uganda
Uganda , officially the Republic of Uganda, is a landlocked country in East Africa. Uganda is also known as the "Pearl of Africa". It is bordered on the east by Kenya, on the north by South Sudan, on the west by the Democratic Republic of the Congo, on the southwest by Rwanda, and on the south by...

, spotted hyenas regularly killed sufferers of African sleeping sickness as they slept outside in camps. When attacking sleeping people, spotted hyenas usually bite the face
Face
The face is a central sense organ complex, for those animals that have one, normally on the ventral surface of the head, and can, depending on the definition in the human case, include the hair, forehead, eyebrow, eyelashes, eyes, nose, ears, cheeks, mouth, lips, philtrum, temple, teeth, skin, and...

, and attempt to drag their victims far from other humans. The Kikuyu of Kenya
Kenya
Kenya , officially known as the Republic of Kenya, is a country in East Africa that lies on the equator, with the Indian Ocean to its south-east...

 fear spotted hyenas more than the striped species. Spotted hyenas are widely feared in Malawi, where they have been known to occasionally attack people at night, particularly during the hot season when people sleep outside. Hyena attacks were widely reported in Malawi's Phalombe
Phalombe
Phalombe is a town approx. 40 km northeast of Mulanje in southern Malawi.It is situated below the Fort Lister Gap between the two larger parts of the Mulanje Massif. Due to this location it is vulnerable for flash floods, which can easily occur during the rainy season...

 plain, to the north of Michesi Mountain. Five deaths were recorded in 1956, five in 1957 and six in 1958. This pattern continued until 1961 when eight people were killed. Attacks occurred most commonly in September, when people slept outdoors, and bush fires made the hunting of wild game difficult for the hyenas. An anecdotal news report from the World Wide Fund for Nature 2004 indicates that 35 people were killed by spotted hyenas in a 12 month period in Mozambique
Mozambique
Mozambique, officially the Republic of Mozambique , is a country in southeastern Africa bordered by the Indian Ocean to the east, Tanzania to the north, Malawi and Zambia to the northwest, Zimbabwe to the west and Swaziland and South Africa to the southwest...

 along a 20 km stretch of road near the Tanzania
Tanzania
The United Republic of Tanzania is a country in East Africa bordered by Kenya and Uganda to the north, Rwanda, Burundi, and the Democratic Republic of the Congo to the west, and Zambia, Malawi, and Mozambique to the south. The country's eastern borders lie on the Indian Ocean.Tanzania is a state...

n border. Attitudes toward spotted hyena attacks tend to be muted when compared to the reactions evoked in areas where striped hyenas have attacked people.

Livestock predation

The degree with which spotted hyenas impact livestock varies from region to region: in the Laikipia district in Kenya, spotted hyenas have little impact on livestock compared to that perpetrated by lions, leopards and cheetahs. However, total reported losses in Tanzania during 2003 amounted to US $12,846 of which spotted hyena kills were reported to account for 98.2%. In a survey taken in seven different villages outside the Serengeti National Park in Tanzania in 2007, spotted hyenas accounted for 97.7% of livestock losses to predators. In the Maasai steppe landscape in Northern Tanzania, spotted hyenas frequently kill small stock (goat
Goat
The domestic goat is a subspecies of goat domesticated from the wild goat of southwest Asia and Eastern Europe. The goat is a member of the Bovidae family and is closely related to the sheep as both are in the goat-antelope subfamily Caprinae. There are over three hundred distinct breeds of...

, sheep and calves
Calf
Calves are the young of domestic cattle. Calves are reared to become adult cattle, or are slaughtered for their meat, called veal.-Terminology:...

) and dog
Dog
The domestic dog is a domesticated form of the gray wolf, a member of the Canidae family of the order Carnivora. The term is used for both feral and pet varieties. The dog may have been the first animal to be domesticated, and has been the most widely kept working, hunting, and companion animal in...

s, and usually commit their depredations at night, thus making them harder to retaliate against than lions, which mostly attack livestock in the daytime.

Spotted hyenas as pets

Spotted hyenas were occasionally present in the menagerie
Menagerie
A menagerie is/was a form of keeping common and exotic animals in captivity that preceded the modern zoological garden. The term was first used in seventeenth century France in reference to the management of household or domestic stock. Later, it came to be used primarily in reference to...

s of the Pharaoh
Pharaoh
Pharaoh is a title used in many modern discussions of the ancient Egyptian rulers of all periods. The title originates in the term "pr-aa" which means "great house" and describes the royal palace...

s. Sir John Barrow
Sir John Barrow, 1st Baronet
Sir John Barrow, 1st Baronet, FRS, FRGS was an English statesman.-Career:He was born the son of Roger Barrow in the village of Dragley Beck, in the parish of Ulverston then in Lancashire, now in Cumbria...

, in his An Account of Travels Into the Interior of Southern Africa, described how spotted hyenas in Sneeuberge
Sneeuberge
The Sneeuberge are located in the far western portions of the Eastern Cape province of South Africa. They are the highest mountain range in South Africa outside the Drakensberg and stretch between Murraysburg of the neighbouring Western Cape, north of Graaff Reinet and almost to Cradock.They are...

 were trained to hunt game, writing that they were "as faithful and diligent as any of the common domestic dogs". In Tanzania, spotted hyena cubs may be taken from a communal den by witchdoctor
Sangoma
A sangoma is a practitioner of herbal medicine, divination and counselling in traditional Nguni societies of Southern Africa .The philosophy is based on a belief in ancestral spirits...

s, in order to increase their status. During the research leading to the composition of his monograph The Spotted Hyena: A Study of Predation and Social Behavior, Hans Kruuk kept a tame hyena he named Solomon. 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. An April 2004 BBC
BBC
The British Broadcasting Corporation is a British public service broadcaster. Its headquarters is at Broadcasting House in the City of Westminster, London. It is the largest broadcaster in the world, with about 23,000 staff...

 article described how a shepherd living in the small town of Qabri Bayah about 50 kilometres from Jigjiga town in eastern Ethiopia
Ethiopia
Ethiopia , officially known as the Federal Democratic Republic of Ethiopia, is a country located in the Horn of Africa. It is the second-most populous nation in Africa, with over 82 million inhabitants, and the tenth-largest by area, occupying 1,100,000 km2...

 managed to use a male spotted hyena as a livestock guardian dog
Livestock guardian dog
A livestock guardian dog is a domesticated canine used to defend livestock against predators. LGDs are commonly referred to as "sheep dogs" since they most often have guarded flocks of sheep, but most are capable of guarding other species of livestock. They are classified as pastoral dogs...

, suppressing its urge to leave and find a mate by feeding it special herbs. If not raised with adult members of their kind, captive spotted hyenas will exhibit scent marking behaviours much later in life than wild specimens. Spotted hyenas can be very destructive: a captive, otherwise perfectly tame, specimen in the Tower of London
Tower of London
Her Majesty's Royal Palace and Fortress, more commonly known as the Tower of London, is a historic castle on the north bank of the River Thames in central London, England. It lies within the London Borough of Tower Hamlets, separated from the eastern edge of the City of London by the open space...

 managed to tear an 8 feet (2.4 m) long plank nailed to its recently repaired enclosure floor with no apparent effort. 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.

Further reading

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

External links

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