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Hyenas or hyaenas ( ; from Ancient Greek ὕαινα, ) ὕαινα, Henry George Liddell, Robert Scott, A Greek-English Lexicon, on Perseus. Etymologically, this is a feminine of ὕς "swine". are feliformia mammals belonging to the family Hyaenidae (). With just four extant species (each in its own genus), it is the fifth-smallest family in the order Carnivora and one of the smallest in the class Mammalia. (2025). 9788496553491, Lynx Edicions. ISBN 9788496553491 Despite their low diversity, hyenas are unique and vital components of most ecosystems.
Although phylogenetics closer to Felidae and Viverridae, hyenas are Ethology and morphologically similar to canidae in several elements due to convergent evolution: both hyenas and canines are non-arboreal, cursorial hunters that catch prey with their teeth rather than claws. Both eat food quickly and may store it, and their calloused feet with large, blunt, nonretractable claws are adapted for running and making sharp turns. However, hyenas' grooming, scent marking, defecation habits, mating, and parental behavior are consistent with the behavior of other feliforms.
Hyenas feature prominently in the folklore and mythology of human cultures that live alongside them. Hyenas are commonly viewed as frightening and worthy of contempt. In some cultures, hyenas are thought to influence people's spirits, rob graves, and steal livestock and children. Other cultures associate them with witchcraft, using their body parts in traditional medicine.
Hyenas then diversified into two distinct types: lightly built dog-like hyenas and robust bone-crushing hyenas. Although the dog-like hyenas thrived 15 million years ago (with one taxon having colonised North America), most became extinct after a change in climate, along with the arrival of canids into Eurasia. Of the dog-like hyena lineage, only the insectivorous aardwolf survived, while the bone-crushing hyenas (including the Extant taxon spotted, brown hyena, and striped hyenas) became the undisputed top scavengers of Eurasia and Africa.
The aardwolf can trace its lineage directly back to Plioviverrops 15 million years ago, and is the only survivor of the dog-like hyena lineage. Its success is partly attributed to its insectivorous diet, for which it faced no competition from canids crossing from North America. It is likely that its unrivaled ability to digest the terpene excretions from soldier is a modification of the strong digestive system its ancestors used to consume fetid carrion.
The striped hyena may have evolved from Hyaenictitherium namaquensis of Pliocene Africa. Striped hyena fossils are common in Africa, with records going back as far as the Villafranchian. As fossil striped hyenas are absent from the Mediterranean region, it is likely that the species is a relatively late invader to Eurasia, having likely spread outside Africa only after the extinction of in Asia at the end of the Ice Age. The striped hyena occurred for some time in Europe during the Pleistocene, having been particularly widespread in France and Germany. It also occurred in Montmaurin, Hollabrunn in Austria, the Furninha in Portugal and the Genista Caves in Gibraltar. The European form was similar in appearance to modern populations, but was larger, being comparable in size to the brown hyena.
The spotted hyena ( Crocuta crocuta) diverged from the striped and brown hyena 10 million years ago. Its direct ancestor was the Indian Crocuta sivalensis, which lived during the Villafranchian. Ancestral spotted hyenas probably developed social behaviours in response to increased pressure from rivals on carcasses, thus forcing them to operate in teams. Spotted hyenas evolved sharp behind their crushing premolars, therefore they did not need to wait for their prey to die, and thus became pack hunters as well as scavengers. They began forming increasingly larger territories, necessitated by the fact that their prey was often migratory, and long chases in a small territory would have caused them to encroach into another Hyena clan turf. Spotted hyenas spread from their original homeland during the Middle Pleistocene, and quickly colonised a very wide area from Europe, to southern Africa and China.
The eventual disappearance of the spotted hyena from Europe has traditionally been attributed to the end of the last glacial period and a subsequent displacement of open grassland by closed forests, which favoured wolves and humans instead. However, analyses have shown that climate change alone is insufficient to explain the spotted hyena's disappearance from Europe, suggesting that other factors – such as human pressure – must have played a role. This suggests that the events must be seen within the broader context of late-Quaternary extinctions, as the late Pleistocene and early Holocene saw the disappearance of many primarily large mammals from Europe and the world.
Expansion or Gene duplication of the olfatory receptor gene family has been found in all 4 extant species, which would have led to the evolution of the more specialised feeding habits of hyenas.
Expansion in immune-related gene families was also found in the spotted hyena, striped hyena and brown hyena, which would have led to the evolution of the scavenging in these species. Mutations and variants were also found in digestion-related genes ( ASH1L, PTPN5, PKP3, AQP10). One of these digestion-related genes has variants also related to enhanced bone mineralisation ( PTPN5), while other have also a role in inflammatory skin responses ( PKP3).
In aardwolves, expansion of genes related to toxin response were found (Lipocalin and UDP Glucuronosyltransferase gene families), which would have led to the evolution of the feeding of termites Trinervitermes in this species. Mutations and variants in genes related to craniofacial shape were also found ( GARS, GMPR, STIP1, SMO and PAPSS2). Another gene is related to protective epidermis function ( DSC1).
A more recent molecular analysis agrees on the phylogenetic relationship between the four extant hyaenidae species (Koepfli et al, 2006).
Their dentition is similar to that of the canid, but is more specialised for consuming coarse food and crushing bones. The , especially the upper, are very powerful and are shifted far back to the point of exertion of peak pressure on the jaws. The other teeth, save for the underdeveloped upper molars, are powerful, with broad bases and cutting edges. The canine teeth are short, but thick and robust. Labiolingually, their are much stronger at the canine teeth than in canids, reflecting the fact that hyenas crack bones with both their anterior dentition and premolars, unlike canids, which do so with their post-carnassial molars. The strength of their jaws is such that both striped and spotted hyenas have been recorded to kill dogs with a single bite to the neck without breaking the skin.Daniel Johnson (1827) Sketches of Indian Field Sports: With Observations on the Animals; Also an Account of Some of the Customs of the Inhabitants; with a Description of the Art of Catching Serpents, as Practiced by the Conjoors and Their Method of Curing Themselves when Bitten: with Remarks on Hydrophobia and Rabid Animals p. 45-46, R. Jennings, 1827Stevenson-Hamilton, James (1917) Animal life in Africa, Vol. 1, p.95, London : William Heinemann The spotted hyena is renowned for its strong bite proportional to its size, but a number of other animals (including the Tasmanian devil) are proportionately stronger. The aardwolf has greatly reduced cheek teeth, sometimes absent in the adult, but otherwise has the same dental formula as the other three species. The dentition for all hyena species is:
Although hyenas lack perineum scent glands, they have a large pouch of naked skin located at the anal opening. Large above the anus open into this pouch. Several are present between the openings of the anal glands and above them. These glands produce a white, creamy secretion that the hyenas paste onto grass stalks. The odor of hyena butter is very strong, smelling of boiling cheap soap or burning, and can be detected by humans several meters downwind. The secretions are primarily used for territorial marking, though both the aardwolf and the striped hyena will spray them when attacked.
Mating between hyenas involves a number of short copulations with brief intervals, unlike canids, who generally canine tying. Spotted hyena cubs are born almost fully developed, with their eyes open and erupting incisors and canines, though lacking adult markings. In contrast, striped hyena cubs are born with adult markings, closed eyes and small ears. Hyenas do not regurgitate food for their young and male spotted hyenas play no part in raising their cubs, though male striped hyenas do so.
The striped hyena is primarily a scavenger, though it will also attack and kill any animals it can overcome, and will supplement its diet with fruit. The spotted hyena, though it also scavenges occasionally, is an active pack hunter of medium to large sized ungulates, which it catches by wearing them down in long chases and dismembering them in a canid-like manner. Spotted hyenas may kill as many as 95% of the animals they eat. Hyaena Specialist Group - Spotted Hyena: Diet and Foraging . Hyaenidae.org. Retrieved on 2015-11-06.
The aardwolf is primarily an insectivore, specialised for feeding on of the genus Trinervitermes and Hodotermes, which it consumes by licking them up with its long, broad tongue. An aardwolf can eat 300,000 Trinervitermes on a single outing.
Except for the aardwolf, hyenas are known to drive off larger predators, like lions, from their kills, despite having a reputation in popular culture for being cowardly. Hyenas are primarily nocturnal animals, but sometimes venture from their lairs in the early-morning hours. With the exception of the highly social spotted hyena, hyenas are generally not gregarious animals, though the striped and brown hyenas may live in family groups and congregate at kills.
Spotted hyenas are one of the few mammals other than bats known to survive infection with rabies virus and have shown little or no disease-induced mortality during outbreaks in sympatric carnivores, in part due to the high concentration of Antibody present in their saliva. Despite this perceived unique disease resistance, little is known about the immune system of spotted hyenas, and even less is known about other Hyaenidae species.
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 use spotted hyenas as Riding animal. In the Mtwara Region of Tanzania, it is believed that a child born at night while a hyena is crying will likely grow up to be a thief. In the same area, hyena feces are believed to enable a child to walk at an early age, thus it is not uncommon in that area to see children with hyena dung wrapped in their clothes. The Kaguru of Tanzania and the Kujamaat of southern Senegal view hyenas as inedible and greedy hermaphrodites. A mythical African tribe called the Bouda is reputed to have members able to transform into hyenas. A similar myth occurs in Mansôa. These "" are killed when discovered, and do not revert to human form once dead.
Striped hyenas are often referred to in literature and folklore, typically as symbols of treachery and stupidity.Mounir R. Abi-Said (2006) Reviled as a grave robber: The ecology and conservation of striped hyaenas in the human dominated landscapes of Lebanon Ph.D. thesis, University of Kent (Biodiversity management) In the Near and Middle East, striped hyenas are generally regarded as physical incarnations of genie.Frembgen, Jürgen W. The Magicality of the Hyena: Beliefs and Practices in West and South Asia , Asian Folklore Studies, Volume 57, 1998: 331–344 Arab writer al-Qazwīnī (1204–1283) spoke of a tribe of people called al-Ḍabyūn meaning "hyena people". In his book
In a similar vein to al-Damīrī, the Greek people until the end of the 19th century believed that the bodies of werewolves, if not destroyed, would haunt battlefields as vampiric hyenas that drank the blood of dying soldiers. The image of striped hyenas in Afghanistan, India and Palestine is more varied. Though feared, striped hyenas were also symbolic of love and fertility, leading to numerous varieties of love medicine derived from hyena body parts. Among the Baluch people and in northern India, witches or magicians are said to ride striped hyenas at night.
The striped hyena is mentioned in the Bible. The Arabic word for the hyena, ḍab` or ḍabu` (plural ḍibā`), is alluded to in a valley in Israel known as Shaqq-ud-Diba` (meaning "cleft of the hyenas") and Wadi-Abu-Diba` (meaning "valley of the hyenas"). Both places have been interpreted by some scholars as being the Biblical Valley of Tsebo`im mentioned in 1 Samuel 13:18. In modern Hebrew language, the word for hyena and hypocrite are both the same: . Though the Authorized King James Version of the Bible interprets the term " `ayit tsavua`" (found in Jeremiah 12:9) as "speckled bird", Henry Baker Tristram argued that it was most likely a hyena being mentioned.
The vocalization of the spotted hyena resembling hysterical human laughter has been alluded to in numerous works of literature: "to laugh like a hyæna" or a "hyen" was a common simile, and is featured in The Cobbler's Prophecy (1594), Webster's Duchess of Malfi (1623) and Shakespeare's As You Like It, Act IV. Sc.1.
Die Strandjutwolf (The brown hyena) is an allegory poem by the renowned South African poet, N. P. van Wyk Louw, which evokes a sinister and ominous presence.
Among hyenas, only the spotted and striped hyenas have been known to become man-eaters. Hyenas are known to have preyed on humans in prehistory: human hair has been found in fossilized hyena dung dating back 195,000 to 257,000 years. Some paleontologists believe that competition and predation by ( Crocuta crocuta spelaea) in Siberia was a significant factor in delaying human colonization of Alaska. Hyenas may have occasionally stolen human kills, or entered campsites to drag off the young and weak, much as modern spotted hyenas do in Africa. The oldest Alaskan human remains coincide with roughly the same time cave hyenas became extinct, leading some paleontologists to infer that hyena predation prevented humans from crossing the Bering Strait earlier.
Hyenas readily scavenge from human corpses; in Ethiopia, hyenas were reported to feed extensively on the corpses of victims of the 1960 attempted coupKapuściński, Ryszard, . 1978. and the Red Terror.Donham, Donald Lewis (1999) Marxist modern: an ethnographic history of the Ethiopian revolution, University of California Press, page 135, Hyenas habituated to scavenging on human corpses may develop bold behaviors towards living people: hyena attacks on people in southern Sudan increased during the Second Sudanese Civil War, when human corpses were readily available to them.Copson, Raymond W. (1994) Africa's wars and prospects for peace, M.E. Sharpe, page 6,
Spotted hyenas have been known to prey on humans in modern times, but such incidents are rare. However, attacks on humans by spotted hyenas are likely to be underreported.Begg, Colleen, Begg, Keith & Muemedi, Oscar (2007) Preliminary data on human - carnivore conflict in Niassa National Reserve, Mozambique, particularly fatalities due to lion, spotted hyaena and crocodile , SGDRN (Sociedade para a Gestão e Desenvolvimento da Reserva do Niassa Moçambique) Man-eating spotted hyenas tend to be very large specimens; a pair of man-eating hyenas, responsible for killing 27 people in Mulanje, Malawi in 1962, weighed in at 72 kg (159 lb) and 77 kg (170 lb) after being shot.Kruuk, Hans (2002) Hunter and hunted: relationships between carnivores and people Cambridge University Press, A 1903 report describes spotted hyenas in the Mzimba District of Angoniland waiting at dawn outside people's huts to attack them when they opened their doors. Victims of spotted hyenas tend to be women, children and sick or infirm men; Theodore Roosevelt wrote in 1908–1909 in Uganda that spotted hyenas regularly killed sufferers of African sleeping sickness as they slept outside in camps.Roosevelt, Theodore (1910) African Game Trails: An Account of the African Wanderings of an American Hunter, Naturalist, New York, C. Scribner's sons
Spotted hyenas are widely feared in Malawi, where they have been known to attack people at night, particularly during the hot season when people sleep outside. A spate of hyena attacks was reported in Malawi's Phalombe plain, with five deaths recorded in 1956, five in 1957 and six in 1958. This pattern continued until 1961, when eight people were killed. Attacks occurred most commonly in September, when people slept outdoors and bush fires made the hunting of wild game difficult for the hyenas. A 2004 news report stated that 35 people were killed by spotted hyenas in a 12-month period in Mozambique along a 20-km stretch of road near the border.
In the 1880s, a hyena was reported to have attacked humans, especially sleeping children, over a three-year period in the Iğdır Province of Turkey, with 25 children and 3 adults being wounded in one year. The attacks provoked local authorities into announcing a reward of 100 for every hyena killed. Further attacks were reported later in some parts of the South Caucasus, particularly in 1908. Instances are known in Azerbaijan of striped hyenas killing children sleeping in courtyards during the 1930s and 1940s. In 1942, a sleeping guard was mauled in his hut by a hyena in Qalıncaq (Golyndzhakh). Cases of children being taken by hyenas by night are known in southeast Turkmenistan's Bathyz Nature Reserve. A further attack on a child was reported around Serakhs in 1948. Several attacks have occurred in India; in 1962, 9 children were thought to have been taken by hyenas in the town of Bhagalpur in the Bihar State in a six-week period, and 19 children up to the age of four were killed by hyenas in Karnataka in 1974. A survey of wild animal attacks during a five-year period in the Indian state of Madhya Pradesh reported that hyenas had attacked three people, causing fewer deaths than Indian wolf, gaur, boar, Asian elephant, Bengal tiger, Indian leopard and .
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