Spotted hyena
The spotted hyena, Crocuta crocuta, is the animal behind a maniacal laughter that ranks among the most recognisable sounds of Africa. It is also called the laughing hyena. Its reputation is that of an ugly and cowardly creature, yet it is the most common large carnivore on the continent. It lives across sub-Saharan Africa, and the IUCN rates it as of least concern, with estimates between 27,000 and 47,000 individuals. Outside protected areas, though, its numbers are falling because of habitat loss and poaching. This documentary asks why an animal so successful has such a poor name. It asks how a society ruled by females really works. And it asks what that famous laugh is telling the animals that hear it. The first clues lie inside the clan.
In the Serengeti, where prey migrates, spotted hyena clans are smaller than in the Ngorongoro Crater, where prey stays put. The largest clans hold at most 80 animals. No other carnivore lives in groups so large or behaves so intricately. Females outweigh males and usually dominate them, which makes the society matriarchal. Even low-ranking females generally dominate high-ranking males. Now and then a female shares the top spot with a male, and a few clans have been led by a male. Such a male is a son of the matriarch who has taken the rank just below hers.
Cubs inherit their mother's rank and copy her friendships, and they keep those ties long after they stop depending on her. When a matriarch dies, her daughters already hold the standing and support to replace her. Daughters usually stay in the clan where they were born, so large clans contain several female family lines. Males typically leave at the age of two and a half years.
Researchers compare this world to that of baboons and macaques rather than to other carnivores. As in those monkeys, rank follows networks of allies instead of size or aggression. The hierarchy is nepotistic, so the offspring of a dominant female outrank adult females who are subordinate to their mother. Yet a cub's rank depends on its mother being present, and low-ranking adults may bully higher-ranking cubs when she is away.
Overthrows are rare, but a slip in status is costly. Hyenas that fall short of the rank they inherit during their formative months are about 1.5 times more likely to die at any given point than those that keep or improve it.
Cubs can recognise relatives as distant as great-aunts, and males stay closer to their own daughters than to unrelated cubs. Those cubs, in turn, treat their fathers more gently. Evolutionary anthropologists once asked captive pairs to pull two ropes in unison to earn food. The hyenas learned quickly without training and outperformed chimpanzees, and experienced animals even helped novices.
Rank also shows up in the body. Higher-ranking adults tend to have longer telomeres, which are structures at the ends of chromosomes, and higher levels of some immune defence proteins in their blood.
Copulation among spotted hyenas lasts only 4-12 minutes and typically happens at night with no other hyenas present. No lasting pair bonds form. Males act submissively toward females in heat, and passive males tend to have more success than aggressive ones. The male's penis enters the female's reproductive tract through her pseudo-penis, which makes forced copulation physically impossible.
That pseudo-penis is an enlarged clitoris, shaped and positioned like a penis and capable of erection. The female also has no external vaginal opening, and she is the only placental mammal known with this arrangement. She urinates, mates and gives birth through one central canal. Cubs are the largest carnivoran young relative to their mothers' weight, so the tissue ruptures during delivery and can take weeks to heal.
Late in pregnancy, dominant mothers give their developing cubs higher androgen levels than lower-ranking mothers do. This is thought to drive the strongly masculinised behaviour and anatomy of females. Cubs of dominant mothers are more aggressive and sexually active, and high-ranking males try to mount females earlier than low-ranking ones.
A newborn weighs 1.5 kg on average, with soft dark brown hair, after a pregnancy that averages 110 days. It is unique among carnivorous mammals in being born with its eyes open, with canine teeth 6-7 mm long and incisors of 4 mm. The average litter is two cubs, and three are occasionally reported.
Shortly after birth, cubs attack each other, especially in litters of the same sex, and the weaker one may die. This killing of a sibling accounts for an estimated 25% of all hyena deaths in the first month. Surviving males grow faster and are likelier to achieve reproductive dominance, while surviving females remove rivals for dominance in their natal clan.
Lactating mothers can carry 3-4 kg of milk, which has the highest protein and fat content of any land carnivore. Cubs nurse for 12-16 months, though they can process solid food at three months. Mothers do not regurgitate food for them. The black coat gives way to the lighter spotted pelage at 2-3 months, and hunting behaviours appear at eight months. Cubs join group hunts fully after their first year and reach sexual maturity at three years. The average lifespan in zoos is 12 years, with a maximum of 25.
The clan's social life revolves around a communal den that can have more than a dozen entrances. Tunnels narrow from an entrance of half a metre to one metre down to as little as 25 cm, too small for adults to use fully. Hyenas rarely dig, preferring the abandoned burrows of warthogs, springhares and jackals. The narrow channels likely shield cubs from predators while their mothers are away. Several females often share a den, and it is not uncommon to see up to 20 cubs at a single site.
A spotted hyena's heart makes up close to 1% of its body weight, while a lion's heart is only 0.45-0.57 percent. That large heart gives the animal endurance for long chases, which often cover several kilometres at speeds of up to 60 km/h. During a hunt, hyenas often run through herds of hoofed animals to pick out one to pursue. They hunt alone, in parties of 2-5, or in large groups, with more flexibility than other African carnivores.
The skull is one of the most powerfully built among the Carnivora. The bone-crushing premolars sit in front of the sharp carnassials, so the hyena can crack bone without blunting its cutting teeth. Large jaw muscles and a vaulted skull absorb the forces. The result is a bite of 80 kgf/cm2, which is 40% more force than a leopard can generate. Free-ranging hyenas have cracked open giraffe long bones 7 cm across, and one individual exerted 4,500 newtons on a measuring instrument.
Serengeti males weigh 40.5-55.0 kg, and females, which are larger than males, weigh 44.5-63.9 kg. In Zambia the averages climb to 67.6 kg for males and 69.2 kg for females. Weights of 81.7 kg and 90 kg are known. The rump is rounded rather than angular, which denies attackers from behind a firm grip.
For decades the hyena was seen as a scavenger. Wildlife ecologist Hans Kruuk showed otherwise in a 7-year study in the Serengeti and the Ngorongoro Crater, finding that spotted hyenas hunt as much as lions. Depending on habitat and the other predators present, hunting supplies anywhere from at least half to 98% of the diet, and the rest is scavenged. A hyena can eat and digest skin and bone, and it makes the most efficient use of animal matter of any African carnivore.
In Kenya's Masai Mara, 80% of the prey is topi and Thomson's gazelle, except during the four months when zebra and wildebeest herds pass through. In Namibia's Etosha National Park, springbok and kudu are the main prey. Hyenas favour animals weighing 56-182 kg and often target the young and the old. They shake small prey in the mouth and eat large prey alive. Unlike other big African carnivores, they have no preferred species, and only African buffalo and giraffe are significantly avoided.
In the Ngorongoro Crater, lions often live largely on kills stolen from hyenas. This reverses a common belief that hyenas are the thieves. Where the two species share ground, they fill the same niche, and their diets can overlap by as much as 68.8%. Field experiments played tape-recorded calls of hyenas feeding, and lions repeatedly approached.
Confronted by lions on a kill, hyenas either leave or wait 30-100 metres away until the lions finish. At night they are bolder, and they sometimes feed beside lions or force them off. They usually prevail against lionesses without males if they outnumber them 4:1. Hyenas have even routed two pride males while outnumbering them 5:1. The danger runs both ways. Lion predation accounts for up to 71% of hyena deaths in Etosha, and one male lion was filmed killing two hyenas without eating them. Captive hyenas with no experience of lions ignore the sight of them but react fearfully to the scent.
Cheetahs are easily intimidated and give up their kills. Leopards, especially males, may stand their ground, and some male leopards prey on hyenas. Still, hyenas are dangerous opponents, and one young adult male leopard died of sepsis from wounds a hyena inflicted.
African wild dogs lose out to hyenas in the long run. A solitary hyena approaching a dog kill creeps in cautiously and tries to leave with a piece of meat, and a pack can mob it. Groups of hyenas pirate more successfully, and wild dog numbers fall where hyena numbers are high. Jackals and African wolves may feed alongside hyenas but are chased if they come too close. Four hyenas reportedly took half an hour to eat a golden wolf.
In the Kalahari, brown hyenas outnumber spotted ones, yet the larger spotted hyenas usually take over carcasses. A brown hyena may raise its mane and growl, which often leaves its rival seemingly bewildered, though not always unharmed. Spotted hyenas avoid crocodile-infested water and keep their distance from Nile crocodiles. Recent observations show that African rock pythons can hunt adult spotted hyenas.
Territory size among spotted hyenas runs from under 40 km2 in the Ngorongoro Crater to more than 1000 km2 in the Kalahari. Clans defend their ground with vocal displays, scent marking and boundary patrols. They pile up scent in special latrines along the borders, by pasting or pawing. Each hyena also has anal glands that produce a creamy white secretion, which it pastes onto grass stalks. Humans can smell it several metres downwind.
One study of hyenas from three clans found a high degree of similarity in fatty acids among members of the same clan. Hyenas from different clans differed more. Researchers have proposed that bacteria in the scent glands ferment these fatty acids differently, which would make the relationship between hyena and microbe a symbiotic one. Within a clan, the secretions also signal sex and reproductive state.
Borders are mostly respected. Hyenas chasing prey have been seen to halt as soon as the prey crosses into another clan's range. In times of food shortage they ignore the line, and males wander across it more than females do. A hyena in foreign territory shows the postures of fear. An intruder that keeps returning to a clan's dens and kills may eventually be accepted.
The vocal range includes whoops, grunts, groans, giggles, yells, growls, whines and squeals. High-pitched calls usually signal fear or submission, and loud, lower calls signal aggression. The pitch of a laugh reveals the animal's age, and variations in the notes carry information about its social rank. In 1972 Hans Kruuk catalogued the calls. A giggle comes from a hyena that is being chased, usually over a kill. A yell comes when it is actually being bitten. A loud grunt-laugh occurs in encounters with lions or other clans.
A 2022 study found that whoops carry signatures unique to each individual. A random forest algorithm, a statistical method, matched a bout of whoops to the right hyena roughly six times more often than chance would predict. Hyenas may be able to identify one another by whoops alone.
Greetings are physical. Two hyenas stand side by side facing opposite directions, raise a hind leg and lick each other's anogenital area. An erection during the greeting is usually a sign of submission rather than dominance, more commonly in males. Subordinates approaching a dominant animal may walk on their forelegs' knees.
Deception is part of the repertoire. Hyenas have been recorded giving alarm calls during feeding when no enemy is present, which frightens off others and lets the caller eat in peace. Mothers use alarm calls to interrupt attacks on their own cubs by other hyenas.
The earliest recorded mention of the Greek name Κροκόττας (krokottas) appears in Strabo's Geographica, which describes a mix of wolf and dog native to Ethiopia. Scholars once thought the genus name Crocuta came from a Latin word meaning 'saffron-coloured one', for the animal's fur. That turned out to be wrong, because the Latin spelling would differ and Graeco-Roman writers never used the word that way. The Greek word traces back through Sanskrit to a term for the golden jackal.
Hiob Ludolf, in his Historia aethiopica, was the first to clearly separate Crocuta from Hyaena, though he took his accounts from an Ethiopian intermediary. European travellers in Ethiopia still called hyenas wolves. The Amharic word for hyena, ጅብ (jɨbb), is linked to the Arabic word for wolf. Willem Bosman, a Dutch tradesman on the Gold Coast from 1688 to 1701, gave the first detailed European descriptions. Peter Kolbe, a German mathematician and astronomer at the Cape of Good Hope from 1705 to 1713, called the animal a tigerwolf.
Their accounts went unnoticed until 1771, when the Welsh naturalist Thomas Pennant used them, plus a captive specimen, to separate the spotted from the striped hyena in his Synopsis of Quadrupeds. Johann Erxleben simply translated Pennant's text into Latin for his Systema regni animalis. Crocuta was recognised as a separate genus in 1828. In 1939 the biologist L Harrison Matthews compared many skulls from Tanzania and showed that one population held all the variation behind the many proposed subspecies.
The ancestors of Crocuta split from Hyaena, the genus of the striped and brown hyenas, about 10 million years ago. The earliest fossils of the genus come from early Pliocene Africa, roughly 3.63 to 3.85 million years ago. Pressure from other predators on carcasses likely pushed the ancestors of the spotted hyena to work in teams. Closely related species began forming clans about 350,000 years ago. Sharp carnassials behind the crushing premolars freed them from waiting for prey to die, so they became pack hunters as well as scavengers. One theory says female dominance helps females compete for kills and produce enough milk. Another ties it to the long time cubs need to grow their massive skulls and jaws.
Cave hyenas, the Eurasian populations of Crocuta, roamed Eurasia for at least one million years. Their range ran from the Iberian Peninsula to the Urals, and remains turn up in the Russian Far East. It has been theorised that hyenas there delayed the colonisation of North America. Carvings and paintings in the Lascaux and Chauvet Caves depict the species, and Paleolithic rock art shows they kept their spots. Eurasian animals were estimated at 102 kg. Their arrival in Europe and China coincided with the decline of the giant short-faced hyena, Pachycrocuta brevirostris, though a more recent analysis found no overlap in Europe. In Western Europe, hyena numbers shrank from about 20,000 years ago as grasslands declined, and hyenas disappeared between 14,000 and 11,000 years ago. Wolves and humans, at home in forests as well as open land, probably outcompeted them.
In Harar, Ethiopia, residents regularly feed spotted hyenas, believing the animals keep devils away and can foretell fortunes. Elsewhere in Africa the hyena is a more troubling figure. It is portrayed as sly, brutish, dangerous and ugly, yet it also stands for physical power and sacredness. Among the Tabwa, it is a solar animal that first brought the sun to warm the cold earth. In the Gelede cult of the Yoruba, a hyena mask signals the end of the èfè ceremony at dawn, because the hyena usually finishes the meals of other carnivores.
The Bambara of Mali, in their Korè cult, hold that the hyena is a hermaphrodite. Their rituals use its mask to combine male principles with femininity and make the initiate a complete moral being. The Beng of Côte d'Ivoire hold that anyone finding a freshly killed hyena with its anus everted must plug it back in, or be struck down with perpetual laughter. They also evacuate a village if a hyena relieves itself within its bounds.
Western beliefs go back to Aristotle's Historia Animalium, which called the hyena a corpse-eating, cowardly and potentially dangerous animal that uses retching noises to attract dogs. In On the Generation of Animals, Aristotle rejected the claim that it is a hermaphrodite, a belief likely born from the masculinised genitalia of females. Pliny the Elder backed Aristotle and added that hyenas imitate human voices and are held in high regard among the Magi. The Physiologus then called the animal a hermaphrodite and a grave-robber, with explicit judgment.
In the 20th century the Western and African stereotypes merged. Ernest Hemingway's Green Hills of Africa and Disney's The Lion King both add gluttony and comic stupidity to the Western image of a cowardly, ugly beast. After the film's release, hyena biologists protested. One researcher sued Disney for defamation of character, and another, who had arranged the animators' visit to observe captive hyenas at the University of California's field station, suggested a boycott.
Livestock losses are often overstated. A hyena seen at a carcass may be blamed for a kill it did not make. One study in northern Kenya found that 90% of hyena predation on livestock happened outside the protection of thorn fences. Attacks on people are rare but likely underreported. A pair of man-eating hyenas killed 27 people in Mlanje, Malawi, in 1962. They weighed 72 and 77 kg when shot. In Addis Ababa, which may hold up to a thousand resident hyenas, a baby boy was killed in 2013. That December marksmen killed ten hyenas on wasteland near the city centre.
On early Dutch farms in southern Africa, colonists trapped hyenas, which they called wolves, in a wolwehok, a rough stone or wooden pen baited with meat and fitted with a trap-door. Elsewhere, body parts are used in traditional medicine, including the tail in Burkina Faso and the paws in Mozambique. Oromo hunters go through ritual purification after killing one.
The IUCN's hyena specialist group names the species' negative reputation as harmful to its survival, both in captivity and in the wild.
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Common questions
What is the spotted hyena?
The spotted hyena (Crocuta crocuta), also called the laughing hyena, is the sole living member of the genus Crocuta and is native to sub-Saharan Africa. It is the largest living member of the Hyaenidae and the most common large carnivore in Africa.
How many spotted hyenas are left in the wild?
Estimates put the spotted hyena at between 27,000 and 47,000 individuals, and the IUCN lists it as of least concern. Numbers are declining outside protected areas because of habitat loss and poaching.
Why do spotted hyena clans have female leaders?
Spotted hyena society is matriarchal because females are larger than males and usually dominate them. Cubs inherit their mother's rank, so a matriarch's daughters are ready to replace her. One theory says female dominance helps females compete on kills and produce enough milk for their cubs.
Does the spotted hyena hunt or only scavenge?
The spotted hyena is primarily a hunter. Hans Kruuk's 7-year study in the Serengeti and Ngorongoro Crater showed that spotted hyenas hunt as much as lions, and hunting supplies from at least half up to 98% of the diet depending on the population.
How strong is the spotted hyena's bite?
The spotted hyena's bite exerts a pressure of 80 kgf/cm2, which is 40% more force than a leopard can generate. Free-ranging hyenas have cracked giraffe long bones 7 cm across.
Do spotted hyenas steal kills from lions?
Lions steal kills from spotted hyenas more often than the reverse. In the Ngorongoro Crater, lions commonly live largely on meat taken from hyenas, and hyenas outnumbering lionesses 4:1 usually prevail.
When did spotted hyenas live in Europe?
Eurasian populations of Crocuta, known as cave hyenas, roamed Eurasia for at least one million years. They disappeared from Western Europe between 14,000 and 11,000 years ago.
What does the spotted hyena's laugh mean?
The spotted hyena's laugh carries information about the caller. Its pitch reveals the animal's age, and variations in the notes convey social rank. High-pitched calls usually signal fear or submission, and loud, lower calls signal aggression.
All sources
127 references cited across the entry
- 1Crocuta crocutaBohm, T. — 2015
- 2BookThe truth about animals: stoned sloths, lovelorn hippos, and other tales from the wild side of wildlifeLucy Cooke — Basic Books — 2018
- 3JournalThe presence of cave hyaena (Crocuta crocuta spelaea) in the Upper Palaeolithic rock art of EuropeN. Spassov et al. — 2004
- 5JournalHyenas and Humans in the Horn of AfricaD. W. Wade — 2006
- 6JournalThe population history of extant and extinct hyenasN Rohland et al. — 2005
- 7JournalWhen did spotted hyenas become social? Evidence from fossil endocastsTherese Flink et al. — 2025-11-15
- 8JournalThe Spotted Hyena (Crocuta crocuta) as a Model System for Study of the Evolution of IntelligenceKay E. Holekamp et al. — June 2007
- 9JournalPost-weaning maternal effects and the evolution of female dominance in the spotted hyenaHeather E. Watts et al. — 2009
- 11Ficarelli, G.; D. Torre. (1970), "Remarks on the taxonomy of hyaenids". Palaeontographia Italica 66:13–33.
- 12JournalHyena paleogenomes reveal a complex evolutionary history of cross-continental gene flow between spotted and cave hyenaMichael V. Westbury et al. — 2020-03-13
- 13JournalWhere the Wild Things Were: Spatial and Temporal Distribution of Carnivores in the Cradle of Humankind (Gauteng, South Africa) in Relation to the Accumulation of Mammalian and Hominin AssemblagesReynolds, S.C. — 2010
- 14JournalPleistocene C hinese cave hyenas and the recent E urasian history of the spotted hyena, C rocuta crocutaGui-Lian Sheng et al. — 29 October 2014
- 15Kurtén, Björn (1988) On evolution and fossil mammals, Columbia University Press, pp. 238–242,
- 16JournalThe extinction of the giant hyena Pachycrocuta brevirostris and a reappraisal of the Epivillafranchian and Galerian Hyaenidae in Europe: Faunal turnover during the Early–Middle Pleistocene TransitionAlessio Iannucci et al. — 15 November 2021
- 18JournalSKULL SHAPE EVOLUTION IN DUROPHAGOUS CARNIVORANS: EVOLUTION IN DUROPHAGESBorja Figueirido et al. — 1 July 2013
- 19Hunter, Luke & Hinde, Gerald (2005) Cats of Africa, Struik,
- 20Savage, R. J. G. (1955) Giant deer from Lough Beg. The Irish Naturalists' Journal 11d: 1–6.
- 21JournalOf arcs and vaults: the biomechanics of bone-cracking in spotted hyenas (Crocuta crocuta): FE ANALYSIS OF THE SPOTTED HYENA SKULLJaime B. Tanner et al. — 2 October 2008
- 22JournalBite Forces and Evolutionary Adaptations to Feeding Ecology in CarnivoresPer Christiansen et al. — February 2007
- 23JournalDevelopment of Bite Strength and Feeding Behaviour in Juvenile Spotted Hyenas (Crocuta crocuta)Wendy J. Binder et al. — 2000
- 27JournalÜber die weiblichen Begattungsorgane der gefleckten HyäneB. Schmotzer et al. — Gustav Fischer — 15 April 1922
- 28JournalCourtship and mating in free-living spotted hyenasM. Szykman et al. — 2007
- 30JournalUrogenital system of the spotted hyena (Crocuta crocuta Erxleben): a functional histological studyGerald R. Cunha — 2003
- 33BookThe Behavior Guide to African Mammals: Including Hoofed Mammals, Carnivores, PrimatesEstes — University of California Press — 1991
- 34JournalSocial support drives female dominance in the spotted hyaenaColin Vullioud et al. — 19 November 2018
- 35JournalA king among queensEve Davidian et al. — 2021
- 36JournalRank-dependent social inheritance determines social network structure in spotted hyenasAmiyaal Ilany et al. — 16 July 2021
- 37JournalAge Estimation and Dispersal in the Spotted Hyena (Crocuta crocuta)Russell C. Van Horn et al. — 2003-08-29
- 38JournalMaternal effects on offspring social status in spotted hyenasMarion L. East et al. — 1 May 2009
- 39Drea CM, Frank LG (2003) The social complexity of spotted hyenas. In: Animal Social Complexity: Intelligence, Culture, and Individualized Societies (eds de Waal FBM, Tyack PL). pp. 121–148, Harvard Un
- 40JournalSocial intelligence in the spotted hyena (Crocuta crocuta)KE Holekamp et al. — 2007
- 41JournalJuvenile rank acquisition is associated with fitness independent of adult rankEli D. Strauss et al. — 2020-03-04
- 42JournalTopological effects of network structure on long-term social network dynamics in a wild mammalA Ilany et al. — 2015
- 43JournalSocioecological variables predict telomere length in wild spotted hyenasN. Lewin — 25 February 2015
- 44JournalSocioecological predictors of immune defenses in a wild spotted hyenasA.S. Flies — 30 September 2016
- 45Chemical Signals in Vertebrates 11Nicole Burgener et al. — Springer New York — 2008
- 46JournalSymbiotic bacteria appear to mediate hyena social odorsKevin R. Theis et al. — 2013-11-11
- 49JournalSexual conflicts in spotted hyenas: male and female mating tactics and their reproductive outcome with respect to age, social status and tenureMarion L. East — 2003
- 50Nelson RJ. 2005. Introduction to Behavioral Endocrinology. Sinauer Associates: Massachusetts. p. 115.
- 52JournalPhylogenetic comparisons implicate sex hormone-binding globulin in "masculinization" of the female spotted hyena (Crocuta crocuta)Geoffrey L. Hammond et al. — March 2012
- 54JournalSocial intelligence in the spotted hyena ( Crocuta crocuta )Kay E Holekamp et al. — 2007-04-29
- 55JournalFecal glucocorticoids reflect socio-ecological and anthropogenic stressors in the lives of wild spotted hyenasPage E. Van Meter et al. — 2009-02-01
- 56JournalAltered behaviour in spotted hyenas associated with increased human activityErin E. Boydston et al. — 2003
- 57JournalMeasuring salivary cortisol in wild carnivoresTracy M. Montgomery et al. — 2022
- 58JournalDynamics of social and energetic stress in wild female chimpanzeesMelissa Emery Thompson et al. — 2010-08-01
- 59JournalSocial, state-dependent and environmental modulation of faecal corticosteroid levels in free-ranging female spotted hyenasWolfgang Goymann et al. — 2001
- 60JournalEarly-life social experience affects offspring DNA methylation and later life stress phenotypeZachary M. Laubach et al. — 2021-07-20
- 61JournalCauses, Consequences, and Conservation of Ungulate MigrationMatthew J. Kauffman et al. — 2021-11-02
- 62JournalThe effect of urbanization on innovation in spotted hyenasLily Johnson-Ulrich et al. — 2021
- 63JournalAnimal and human innovation: novel problems and novel solutionsSimon M. Reader et al. — 2016-03-19
- 64JournalFitness Consequences of Innovation in Spotted HyenasLily Johnson-Ulrich et al. — 2019-11-22
- 65JournalFacultative predation and scavenging by mammalian carnivores: seasonal, regional and intra-guild comparisonsL. M. Pereira et al. — 2014
- 66JournalFood acquisition by spotted hyaenas in Etosha National Park, Namibia: predation versus scavengingW. C. Gasaway et al. — 1991
- 67JournalPrey preferences of the spotted hyaena (Crocuta crocuta)M. W. Hayward — 2006
- 69JournalOptimal hunting group size: the need for lions to defend their kills against loss to spotted hyaenasCooper, S. — 1991
- 71VideoEternal Enemies: Lions and HyenasNational Geographic — 1992
- 72JournalCompetitive interactions between spotted hyenas and lions in the Etosha National Park, NamibiaTrinkel, Martina et al. — 2005
- 73JournalThe Lion King and the Hyaena Queen: large carnivore interactions and coexistenceS. Périquet et al. — 2015
- 74JournalIn the absence of a "landscape of fear": How lions, hyenas, and cheetahs coexistA. Swanson et al. — 2016
- 75BookBig Cat Diary: LeopardJonathan Scott — HarperCollins Publishers Limited — 2006
- 76JournalMortality in a protected Leopard population, Phinda Private Game Reserve, South Africa: A population in decline?Guy Balme et al. — 2004
- 77Creel, Scott; Creel, Nancy Marusha (2002), The African wild dog: behavior, ecology, and conservation, Princeton University Press, pp. 253–254,
- 78BookAfrican PredatorsMartin Harvey et al. — Smithsonian Books — October 2001
- 80BookWhy Men Won't Ask for Directions: The Seductions of SociobiologyRichard C. Francis — Princeton University Press — 2004
- 82JournalLong-distance vocalizations of spotted hyenas contain individual, but not group, signaturesKenna D. S. Lehmann et al. — 2022-07-27
- 84JournalTesting the savannah corridor hypothesis during MIS2: The Boh Dambang hyena site in southern CambodiaAnne-Marie Bacon et al. — 15 January 2018
- 87BookThe zoological exploration of southern Africa, 1650–1790L. C. Rookmaaker — A. Balkema — 1989
- 88JournalThe Magicality of the Hyena: Beliefs and Practices in West and South AsiaJürgen W. Frembgen — 1998
- 90Goodrich, S.G. and Winchell, A., Johnson's Natural History (New York: A.J. Johnson & Company, 1885), p. 248.
- 93JournalLivestock Depredation of the Spotted Hyena (Crocuta crocuta) in Southern Tigray, Northern EthiopiaYirga, Giday et al. — 2010
- 95Kruuk, Hans (2002) Hunter and hunted: relationships between carnivores and people Cambridge University Press, pp. 64–65,
- 96Roosevelt, Theodore (1910) African Game Trails: An Account of the African Wanderings of an American Hunter, Naturalist, New York, C. Scribner's sons
- 97BookNatural Enemies: People-Wildlife conflicts in Anthropological PerspectiveKnight, John — Psychology Press — 2000
- 98Marlowe, Frank (2010), The Hadza: hunter-gatherers of Tanzania, University of California Press,
- 99Kapuściński, Ryszard, The Emperor: Downfall of an Autocrat. 1978.
- 100Donham, Donald Lewis (1999) Marxist modern: an ethnographic history of the Ethiopian revolution, University of California Press, p. 135,
- 101Copson, Raymond W. (1994) Africa's wars and prospects for peace, M.E. Sharpe, p. 6,
- 102NewsThe urban hyenas that attack rough sleepersMartin Fletcher — BBC News — 23 February 2014
- 104Tjader, Richard (1910) The big game of Africa: with many illustrations from photographs by the author, D. Appleton and company in New York and, London
- 106Beinart, William (2008) The Rise of Conservation in South Africa: Settlers, Livestock, and the Environment 1770–1950, Oxford University Press US,
- 107Moodie, John Wedderburn Dunbar (1835) Ten Years in South Africa, Vol. 1, Richard Bentley
- 108Daniell, Samuele; Barrow, Sir John; Somerville, William (1820), Sketches representing the native tribes, animals, and scenery of southern Africa: from drawings made by the late Mr. Samuel Daniell, W.
- 109JournalMarkedly Elevated Antibody Responses in Wild versus Captive Spotted Hyenas Show that Environmental and Ecological Factors Are Important Modulators of ImmunityA.S. Flies — 7 October 2015
- 112Bingley, William (1829), Animal biography, or, Popular zoology
- 113Kruuk, Hans (1975) Hyaena. Oxford University Press, London
- 114BookBrehm's Life of AnimalsAlfred Edmund Brehm — Chicago: A. N. Marquis & Company — 1895
- 116BookThe safari companion: a guide to watching African mammals, including hoofed mammals, carnivores, and primatesRichard Estes — Chelsea Green Publishing — 1998
- 117BookHyaena: On the Naming and Localisation of an Enigmatic AnimalHolger Funk — GRIN Verlag — 2010
- 118BookEast African mammals: an atlas of evolution in Africa, Volume 3, Part 1Jonathan Kingdon — University of Chicago Press — 1988
- 119BookThe Spotted Hyena: A Study of Predation and Social BehaviourHans Kruuk — University of California Press — 1972
- 120BookPleistocene mammals of EuropeBjörn Kurtén — Weidenfeld and Nicolson — 1968
- 121BookThe Game Animals of AfricaRichard Lydekker — London, R. Ward, limited — 1908
- 122BookThe Velvet Claw: A Natural History of the CarnivoresDavid Macdonald — New York: Parkwest — 1992
- 123BookHyaenas: status survey and conservation action planGus Mills et al. — IUCN/SSC Hyena Specialist Group — 1998
- 124BookHyena Nights & Kalahari DaysGus Mills et al. — Jacana Media — 2011
- 125BookAfrican Game Trails: An Account of the African Wanderings of an American Hunter, NaturalistTheodore Roosevelt — New York, C. Scribner's sons — 1910
- 126BookThe carnivores of West AfricaDonovan Reginald Rosevear — London : Trustees of the British Museum (Natural History) — 1974
- 127BookThe Serengeti Lion: A Study of Predator-Prey RelationsGeorge B. Schaller — University of Chicago Press — 1976
- 128BookAnimal life in Africa, Vol. 1James Stevenson-Hamilton — London : William Heinemann — 1917