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114-727: Kirkdale Cave is a cave and fossil site located in Kirkdale near Kirkbymoorside in the Vale of Pickering , North Yorkshire , England . It was discovered by workmen in 1821, and found to contain fossilized bones of a variety of mammals from the Eemian interglacial (globally known as the Last Interglacial, ~130-115,000 years ago), when temperatures were comparable to contemporary times, including animals currently absent from Britain or globally extinct, including hippopotamuses (amongst

228-508: A pompom in appearance. Unusually among hyaenids, and mammals in general, the female spotted hyena is considerably larger than the male. Both sexes have a pair of anal glands which open into the rectum just inside the anal opening. These glands produce a white, creamy secretion which is pasted onto grass stalks by everting the rectum. The odour of this secretion is strong, smelling of boiling cheap soap or burning, and can be detected by humans several metres downwind. The spotted hyena has

342-500: A "tigerwolf", because the settlers in southern Africa did not know of hyenas, and thus labelled them as "wolves". Bosman and Kolben's descriptions went largely unnoticed until 1771, when the Welsh naturalist Thomas Pennant used the descriptions and his personal experience with a captive specimen as a basis for consistently differentiating the spotted hyena from the striped in his Synopsis of Quadrupeds . The description given by Pennant

456-779: A behaviour which does not occur when they target other prey species. One of the signs of social intelligence is the ability to have a keen olfactory sense or nasal recognition. Spotted hyenas have unique scent signatures that help them distinguish themselves from other clans and/or individuals (i.e., males or females conspecific), which enables them to mark their territories with secretions from their scent glands. Also, spotted hyenas have been recorded to utilise deceptive behaviour, including giving alarm calls during feeding when no enemies are present, thus frightening off other hyenas and allowing them to temporarily eat in peace. Similarly, mothers will emit alarm calls when attempting to interrupt attacks on their cubs by other hyenas. The spotted hyena

570-415: A body mass range of 56–182 kg (123–401 lb), with a mode of 102 kg (225 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 anti-predator behaviours. Small prey is killed by being shaken in

684-434: A distinct epoch has, as it were, been established in the history of the revolutions of our globe: a point fixed from which our researches may be pursued through the immensity of ages, and the records of animate nature, as it were, carried back to the time of the creation. The cave is a Site of Special Scientific Interest and a Geological Conservation Review site. The Saxon St Gregory's Minster with its unusual sundial

798-430: A dominant animal, subordinate spotted hyenas will walk on the knees of their forelegs in submission. Greeting ceremonies among clan-members consist of two individuals standing parallel to each other and facing opposite directions. Both individuals raise their hind legs and lick each other's anogenital area . During these greeting ceremonies, the penis or pseudo-penis often becomes erect, in both males and females. Erection

912-455: A firm grip on it. The head is wide and flat with a blunt muzzle and broad rhinarium . In contrast to the striped hyena, the ears of the spotted hyena are rounded rather than pointed. Each foot has four digits, which are webbed and have short, stout and blunt claws. The paw-pads are broad and flat, with the whole undersurface of the foot around them being naked. The tail is relatively short, being 300–350 mm (12–14 in) long, and resembles

1026-713: 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. The intelligence of the spotted hyena was attested to by Dutch colonists in 19th-century South Africa, who noted that hyenas were exceedingly cunning and suspicious, particularly after successfully escaping from traps. Spotted hyenas seem to plan on hunting specific species in advance; hyenas have been observed to engage in activities such as scent marking before setting off to hunt zebras,

1140-472: A hunt, spotted hyenas often run through ungulate herds to select an individual to attack. Once selected, their prey is chased over a long distance, often several kilometres, at speeds of up to 60 kilometres per hour (37 mph). The spotted hyena has a long history of interaction with humanity ; depictions of the species exist from the Upper Paleolithic period, with carvings and paintings from

1254-612: A kill by lions, spotted hyenas will either leave or wait patiently at a distance of 30–100 metres until the lions have finished eating. In some cases, spotted hyenas are bold enough to feed alongside lions, and may occasionally force lions off a kill. This mostly occurs during the nighttime, when hyenas are bolder. Spotted hyenas usually prevail against groups of lionesses unaccompanied by males if they outnumber them 4:1. In some instances they were seen to have taken on and routed two pride males while outnumbering them 5:1. The two species may act aggressively toward one another even when there

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1368-418: A large degree of morphological geographic variation, which led to an equally extensive set of specific and subspecific epithets. It was gradually realised that all of this variation could be applied to individual differences in a single subspecies. In 1939, biologist L Harrison Matthews demonstrated through comparisons between a large selection of spotted hyena skulls from Tanzania that all the variation seen in

1482-471: A male. There have also been cases in which a clan has been led by a male rather than a female. Cubs take the rank directly below their mothers at birth. So when the matriarch passes away (or, in rare instances, disperses into another clan ), their youngest female cub will take over as matriarch. It is typical for females to remain with their natal clan, thus large clans usually contain several matrilines , whereas males typically disperse from their natal clan at

1596-485: A massive loss of lowland habitats favoured by spotted hyenas, and a corresponding increase in mixed woodlands . Spotted hyenas, under these circumstances, would have been outcompeted by wolves and humans which were as much at home in forests as in open lands, and in highlands as in lowlands. Spotted hyena populations began to shrink roughly 20,000 years ago, completely disappearing from Western Europe between 14 and 11,000 years ago, and earlier in some areas. Historically,

1710-505: A mate. Hyenas travelling in another clan's home range typically exhibit bodily postures associated with fear, particularly when meeting other hyenas. An intruder can be accepted into another clan after a long period of time if it persists in wandering into the clan's territory, dens or kills. The spotted hyena is a non-seasonal breeder, though a birth peak does occur during the wet season . Females are polyestrous, with an estrus period lasting two weeks. Like many feliform species,

1824-505: A necessity, as is the case for brown and striped hyenas, and thus they became pack hunters as well as scavengers. They began forming increasingly larger territories, necessitated by the fact that their prey was often migratory and long chases in a small territory would have caused them to encroach into another clan's land. It has been theorised that female dominance in spotted hyena clans could be an adaptation in order to successfully compete with males on kills, and thus ensure that enough milk

1938-412: 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

2052-422: A proportionately large heart , constituting close to 1% of its body weight, thus giving it great endurance in long chases. In contrast, a lion's heart makes up only 0.45–0.57 percent of its body weight. The now extinct Eurasian populations were distinguished from the modern African populations by their shorter distal extremities and longer humerus and femur . The skull of the spotted hyena differs from that of

2166-425: A rabies vector, thus indicating that the species catches the disease from other animals rather than from intraspecifics. The microfilaria of Dipetalonema dracuneuloides have been recorded in spotted hyenas in northern Kenya. The species is known to carry at least three cestode species of the genus Taenia , none of which are harmful to humans. It also carries protozoan parasites of the genus Hepatozoon in

2280-558: A subspecies of Crocuta crocuta , known as cave hyenas , roamed across Eurasia for at least one million years until the end of the Late Pleistocene . The spotted hyena is the largest extant member of the Hyaenidae, and is further physically distinguished from other species by its vaguely bear -like build, rounded ears, less prominent mane, spotted pelt, more dual-purposed dentition, fewer nipples , and pseudo-penis . It

2394-559: A symbiotic relationship between spotted hyenas and their scent gland bacteria, where the differences in fatty acids can be attributed to fermentation by different microbes. Clan boundaries are usually respected; hyenas chasing prey have been observed to stop dead in their tracks once their prey crosses into another clan's range. Hyenas will however ignore clan boundaries in times of food shortage. Males are more likely to enter another clan's territory than females are, as they are less attached to their natal group and will leave it when in search of

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2508-576: Is nepotistic ; the offspring of dominant females automatically outrank adult females subordinate to their mother. However, rank in spotted hyena cubs is greatly dependent on the presence of the mother; low-ranking adults may act aggressively toward higher-ranking cubs when the mother is absent. Although individual spotted hyenas only care for their own young, and males take no part in raising their young, cubs are able to identify relatives as distantly related as great-aunts. Also, males associate more closely with their own daughters rather than unrelated cubs, and

2622-528: Is 12 years, with a maximum of 25 years. The clan's social life revolves around a communal den. While some clans may use particular den sites for years, others may use several different dens within a year or several den sites simultaneously. Spotted hyena dens can have more than a dozen entrances, and are mostly located on flat ground. The tunnels are usually oval in section, being wider than they are high, and narrow down from an entrance width of ½–1 metre (1.6–7.7 ft) to as small as 25 cm (9.8 in). In

2736-640: Is a stub . You can help Misplaced Pages by expanding it . Spotted hyena The spotted hyena ( Crocuta crocuta ), also known as the laughing hyena , is a hyena species, currently classed as the sole extant member of the genus Crocuta , native to sub-Saharan Africa . It is listed as being of least concern by the IUCN due to its widespread range and large numbers estimated between 27,000 and 47,000 individuals. The species is, however, experiencing declines outside of protected areas due to habitat loss and poaching. Populations of Crocuta , usually considered

2850-631: Is a highly successful animal, being the most common large carnivore in Africa. Its success is due in part to its adaptability and opportunism ; it is primarily a hunter but may also scavenge , with the capacity to eat and digest skin, bone and other animal waste. In functional terms, the spotted hyena makes the most efficient use of animal matter of all African carnivores. The spotted hyena displays greater plasticity in its hunting and foraging behaviour than other African carnivores; it hunts alone, in small parties of 2–5 individuals or in large groups. During

2964-424: Is at least one record of a young adult male leopard dying from a sepsis infection caused by wounds inflicted by a spotted hyena. Spotted hyenas will follow packs of African wild dogs to appropriate their kills. They will typically inspect areas where wild dogs have rested and eat any food remains they find. When approaching wild dogs at a kill, solitary hyenas will approach cautiously and attempt to take off with

3078-496: Is contracted by consuming already infected herbivores, rather than through direct infection from tsetse flies . It is known that adult spotted hyenas in the Serengeti have antibodies against rabies , canine herpes , canine brucellosis , canine parvovirus , feline calicivirus , leptospirosis , bovine brucellosis , rinderpest and anaplasmosis. During the canine distemper outbreak of 1993–94, molecular studies indicated that

3192-545: Is derived from the Sanskrit koṭṭhâraka , which in turn originates from kroshṭuka (both of which were originally meant to signify the golden jackal ). The earliest recorded mention of Κροκόττας is from Strabo 's Geographica , where the animal is described as a mix of wolf and dog native to Ethiopia. From Classical antiquity until the Renaissance , the spotted and striped hyena were either assumed to be

3306-648: Is nearby. Kirkdale, North Yorkshire Kirkdale is a valley in North Yorkshire , England , which along with Sleightholmedale makes up the larger Bransdale and carries the Hodge Beck from its moorland source near Cockayne to the River Dove and onto the River Rye in the Vale of Pickering . Corallian Limestone which outcrops on the hills surrounding the Vale of Pickering runs across

3420-526: Is no food at stake. Lions may charge at hyenas and maul them for no apparent reason; one male lion was filmed killing two hyenas on separate occasions without eating them, 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. Experiments on captive spotted hyenas revealed that specimens with no prior experience with lions act indifferently to

3534-513: Is not uncommon to see up to 20 cubs at a single site. The general form of a spotted hyena den is tunnel-shaped, with a spacious end chamber used for sleeping or breeding. This chamber measures up to 2 metres (6.6 feet) in width, the height being rather less. Females generally give birth at the communal den or a private birth den. The latter is primarily used by low status females to maintain continual access to their cubs, as well as ensure that they become acquainted with their cubs before transferral to

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3648-453: Is particularly apparent in same sexed litters, and can result in the death of the weaker cub. This neonatal siblicide kills an estimated 25% of all hyenas in their first month. Male cubs which survive grow faster and are likelier to achieve reproductive dominance, while female survivors eliminate rivals for dominance in their natal clan. Lactating females can carry 3–4 kg (6.6–8.8 lb) of milk in their udders . Spotted hyena milk has

3762-477: Is produced for their cubs. Another theory is that it is an adaptation to the length of time it takes for cubs to develop their massive skulls and jaws, thus necessitating greater attention and dominating behaviours from females. Both Björn Kurtén and Camille Arambourg promoted an Asiatic origin for the species; Kurtén focussed his arguments on the Plio-Pleistocene taxon Crocuta sivalensis from

3876-556: Is scarce or absent in tropical rainforests and coastal areas. Its preferred habitats in west Africa include the Guinea and Sudan savannahs, and is absent in the belt of dense coastal forest. In the Namib Desert , it occurs in riverine growth along seasonal rivers, the sub-desertic pro-Namib and the adjoining inland plateau. In ideal habitats, the spotted hyena outnumbers other large carnivores, including other hyena species. However,

3990-439: Is superimposed on the back and hind quarters. The spots, which are of variable distinction, may be reddish, deep brown or almost blackish. The spots vary in size, even on single individuals, but are commonly 20 mm (0.79 in) in diameter. A less distinct spot pattern is present on the legs and belly but not on the throat and chest. Some research groups (such as Ngorongoro Crater Hyena Project and MSU Hyena Project ) often use

4104-729: Is the largest extant member of the Hyaenidae. Adults measure 95–165.8 cm (37.4–65.3 in) in body length, and have a shoulder height of 70–91.5 cm (27.6–36.0 in). Adult male spotted hyenas in the Serengeti weigh 40.5–55.0 kg (89.3–121.3 lb), while females weigh 44.5–63.9 kg (98–141 lb). Spotted hyenas in Zambia tend to be heavier, with males weighing on average 67.6 kg (149 lb), and females 69.2 kg (153 lb). Exceptionally large weights of 81.7 kg (180 lb) and 90 kg (200 lb) are known. It has been estimated that adult members of

4218-606: Is the most carnivorous member of the Hyaenidae. Unlike its brown and striped cousins, the spotted hyena is primarily a predator; this has been shown since the 1960s. One of the earliest studies to demonstrate their hunting abilities was done by wildlife ecologist Hans Kruuk, and he showed through a 7-year study of hyena populations in the Serengeti and Ngorongoro Crater that spotted hyenas hunt as much as lions, with later studies showing this tendency in many other areas of Africa. There are however large differences between populations, depending on their habitat, available prey species and

4332-485: Is the northernmost site in the world where hippopotamus remains have been found. It also included a considerable amount of fossilized hyena faeces. The fossilized remains were embedded in a silty layer sandwiched between layers of stalagmite . The discovery at Kirkdale occurred in the wake of new forms of stratigraphic dating developed during the Enlightenment. As was the case for many nineteenth century fossils,

4446-602: Is the only placental mammalian species where females have a pseudo-penis and lack an external vaginal opening. The spotted hyena is the most social of the Carnivora in that it has the largest group sizes and most complex social behaviours. Its social organisation is unlike that of any other carnivore, bearing closer resemblance to that of cercopithecine primates ( baboons and macaques ) with respect to group size, hierarchical structure, and frequency of social interaction among both kin and unrelated group-mates. However,

4560-473: Is the other way around, and lions will readily steal the kills of spotted hyenas. In the Ngorongoro Crater , it is common for lions to subsist largely on kills stolen from hyenas. Lions are quick to follow the calls of hyenas feeding, a fact demonstrated by field experiments, during which lions repeatedly approached whenever the tape-recorded calls of hyenas feeding were played. When confronted on

4674-427: Is traversed to its tip by a central urogenital canal, through which the female urinates, copulates and gives birth. The pseudo-penis can be distinguished from the males' genitalia by its somewhat shorter length, greater thickness, and more rounded glans . In both males and females, the base of the glans is covered with penile spines . The formation of the pseudo-penis appears largely androgen independent, as

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4788-580: Is usually a sign of submission, rather than dominance, and is more common in males than in females. It is said that feasting Hyaenas engage in violent fights, and there is such a croaking, shrieking and laughing at such times that a superstitious person might really think all the inhabitants of the infernal regions had been let loose. The spotted hyena has an extensive vocal range, with sounds ranging from whoops, fast whoops, grunts, groans, lows, giggles, yells, growls, soft grunt-laughs, loud grunt-laughs, whines and soft squeals. The loud who-oop call, along with

4902-671: The Dutch West India Company at the Gold Coast (modern-day Ghana ) from 1688 to 1701, wrote of Jakhals, of Boshond (jackals or woodland dogs) whose physical descriptions match the spotted hyena. Kolben, a German mathematician and astronomer who worked for the Dutch East India Company in the Cape of Good Hope from 1705 to 1713, described the spotted hyena in great detail, but referred to it as

5016-580: The Eemian interglacial (also known as the Ipswichian in Britain or the Last Interglacial globally) around 130-115,000 years ago, when temperatures were comparable to those in Britain today, based uranium-series dating of a calcite layer above the bone deposits dating to 121,000 ± 4,000 years Before Present . Aside from the reported discovery of hippopotamus remains in Stockton-on-Tees, this

5130-477: The Lascaux and Chauvet Caves . The species has a largely negative reputation in both Western culture and African folklore . In the former, the species is mostly regarded as ugly and cowardly, while in the latter, it is viewed as greedy, gluttonous, stupid, and foolish, yet powerful and potentially dangerous. The majority of Western perceptions on the species can be found in the writings of Aristotle and Pliny

5244-630: The Russian Far East , and it has been theorised that the presence of hyenas there may have delayed the colonisation of North America . During the Last Glacial Maximum , the spotted hyena also roamed Southeast Asia . The causes of the species' extinction in Eurasia are still largely unknown. In Western Europe at least, the spotted hyena's extinction coincided with a decline in grasslands 12,500 years ago. Europe experienced

5358-522: The Siwaliks , a view defended by Arambourg, who nonetheless allowed the possibility of an Indo-Ethiopian origin. This stance was contested by Ficarelli and Torre, who referred to evidence of the spotted hyena's presence from African deposits dating from the early Pleistocene, a similar age to the Asian C. sivalensis . Modern scholarship supports an African origin for the genus, with the earliest fossils of

5472-671: The Transvaal Lowveld areas of South Africa . During the 1770s and 1780s the species was still widespread in southern and western South Africa, being recorded on the Cape Peninsula and Cape Flats , and near present-day Somerset West , Riviersonderend , Mossel Bay , George , Joubertina , Gamtoos River , Jansenville , Cannon Rocks , Alice , Onseepkans and Augrabies Falls . The species dwells in semi- deserts , savannah , open woodland , dense dry woodland, and mountainous forests up to 4,000 m in altitude. It

5586-680: The Yorkshire Philosophical Society ". While criticized by some, William Buckland 's analysis of Kirkland Cave and other bone caves was widely seen as a model for how careful analysis could be used to reconstruct the Earth's past, and the Royal Society awarded William Buckland the Copley Medal in 1822 for his Kirkdale paper. At the presentation the society's president, Humphry Davy , said: by these inquiries,

5700-446: The brown bear in bone-crushing ability, and free ranging hyenas have been observed to crack open the long bones of giraffes measuring 7 cm in diameter. A 63.1 kg (139 lb) spotted hyena is estimated to have a bite force of 565.7 newtons at the canine tip and 985.5 newtons at the carnassial eocene. One individual in a study was found to exert a bite force of 4,500 newtons on the measuring instruments. The spotted hyena

5814-508: The Elder , though in relatively unjudgmental form. Explicit, negative judgments occur in the Physiologus , where the animal is depicted as a hermaphrodite and grave-robber . The IUCN's hyena specialist group identifies the spotted hyena's negative reputation as detrimental to the species' continued survival, both in captivity and the wild. The spotted hyena's scientific name Crocuta

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5928-559: The Kalahari, where the brown outnumber the spotted. The two species typically encounter each other on carcasses, which the larger spotted species usually appropriate. Sometimes, brown hyenas will stand their ground and raise their manes while emitting growls. This usually has the effect of seemingly confusing spotted hyenas, which will act bewildered, though they will occasionally attack and maul their smaller cousins. Similar interactions have been recorded between spotted and striped hyenas in

6042-507: The Old Norse kirkja dalr (meaning "church valley"). The parish was composed of the townships of Beadlam , Bransdale , Westside, Muscoates , Nawton , North Holme, Skiplam, Welburn (which included the site of the parish church) and Wombleton . North Holme was transferred to the parish of Great Edstone in the 19th century. All these townships became separate civil parishes in 1866. This North Yorkshire location article

6156-643: The Serengeti, Kenya and South Africa. Spotted hyenas may act as hosts in the life-cycles of various parasites which start life in herbivores; Taenia hyaenae and T. olnogojinae occur in hyenas in their adult phase. Trichinella spiralis are found as cysts in hyena muscles. The spotted hyena's distribution once ranged in Europe from the Iberian Peninsula to the Urals , where it remained for at least one million years. Remains have also been found in

6270-490: The Serengeti, where prey is migratory, clans are smaller than those in the Ngorongoro Crater, where prey is sedentary. Spotted hyena clans are more compact and unified than wolf packs, but are not as closely knit as those of African wild dogs . Females usually dominate males, including in cases where low-ranking females generally dominate over high-ranking males, but they will also occasionally co-dominate with

6384-417: The Serengeti. Though they readily take to water to catch and store prey, spotted hyenas will avoid crocodile-inhabited waters, and usually keep a safe distance from Nile crocodiles . Recent observations shows that African rock pythons can hunt adult spotted hyenas. Spotted hyenas have a complex set of postures in communication. When afraid, the ears are folded flat, and are often combined with baring of

6498-466: The action of an universal deluge , challenging the belief that the bones were brought to the cave by Noah's flood and providing detailed evidence that instead hyenas had used the cave as a den into which they brought the bones of their prey. The specimens were an original part of the archaeology collection of the Yorkshire Museum and it is said that "the scientific interest aroused founded

6612-403: 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. Copulation in spotted hyenas is a relatively short affair, lasting 4–12 minutes, and typically only occurs at night with no other hyenas present. The mating process is complicated, as the male's penis enters and exits

6726-487: The age of 2½ years. Höner et al. say that when a male co-dominates with a female or is otherwise able to lead, this is because the male was born to the matriarch of the clan and has taken the rank directly below his mother. The clan is a fission–fusion society , in which clan-members do not often remain together, but may forage in small groups. High-ranking hyenas maintain their position through aggression directed against lower-ranking clan-members. Spotted hyena hierarchy

6840-441: The age of one month. Within ten days of birth, they are able to move at considerable speed. Cubs begin to lose the black coat and develop the spotted, lighter coloured pelage of the adults at 2–3 months. They begin to exhibit hunting behaviours at the age of eight months, and will begin fully participating in group hunts after their first year. Spotted hyenas reach sexual maturity at the age of three years. The average lifespan in zoos

6954-506: The age of the earliest Crocuta specimens in Eurasia, which are around 2  million years ago from China. The nuclear genome results also suggest that the European and Asian populations were distinct from each other, but were more closely related to each other overall than to African Crocuta populations. Analysis of the nuclear genome suggests that there had been interbreeding between African and Eurasian populations for some time after

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7068-399: The animals had lived in the local area, and that the hyenas had used the cave as a den and brought in remains of the various animals they fed on. This hypothesis was supported by the fact that many of the bones showed signs of having been gnawed prior to fossilization, and by the presence of objects which Buckland suspected to be fossilized hyena dung. Further analysis, including comparison with

7182-517: The bones in Kirkdale were originally found by local inhabitants. The entrance to the cave was found by limestone quarry workers in the summer of 1821. The quarry workers assumed that the abundant bones buried in the cave floor were the remains of cattle that had been dumped in the cave after dying from some past epidemic. They used some of the bones to fill potholes in a nearby road, where an amateur naturalist noticed them and realized that they were not

7296-479: The cave today. Although there is no village in the dale, Kirkdale was the centre of a large ancient parish . The Saxon parish church of St Gregory's Minster stands by the river. It was built in 1055 on the site of an earlier church, and has an 11th-century sundial bearing an Old English inscription. The church gave its name to the dale, first recorded in the Domesday Book of 1086, and derived from

7410-489: The communal den. Compared to other hyenas, the spotted hyena shows a greater relative amount of frontal cortex which is involved in the mediation of social behavior. Studies strongly suggest convergent evolution in spotted hyena and primate intelligence . A study done by evolutionary anthropologists demonstrated that spotted hyenas outperform chimpanzees on cooperative problem-solving tests; captive pairs of spotted hyenas were challenged to tug two ropes in unison to earn

7524-483: The den, consisting of small underground channels, is likely an effective anti-predator device which protects cubs from predation during the absence of the mother. Spotted hyenas rarely dig their own dens, having been observed for the most part to use the abandoned burrows of warthogs, springhares and jackals. Faeces are usually deposited 20 metres (66 feet) away from the den, though they urinate wherever they happen to be. Dens are used mostly by several females at once, and it

7638-633: The dung of modern spotted hyenas living in menageries , confirmed the identification of the fossilized dung. He published his analysis in an 1822 paper he read to the Royal Society . A few days before reading the formal paper, he gave the following colourful account at a dinner held by the Geological Society : The hyaenas, gentlemen, preferred the flesh of elephants, rhinoceros, deer, cows, horses, etc., but sometimes, unable to procure these, & half starved, they used to come out of

7752-595: The exact taxonomic nature of the hyena family in general, with most European travelers in Ethiopia referring to hyenas as "wolves". This partly stems from the Amharic word for hyena, ጅብ ( jɨbb ), which is linked to the Arabic word ذئب ( dhiʾb ) 'wolf'. The first detailed first-hand descriptions of the spotted hyena by Europeans come from Willem Bosman and Peter Kolbe . Bosman, a Dutch tradesman who worked for

7866-423: The eyes, the area around the rhinarium, the lips and the back portion of the chin are all blackish. The limbs are spotted, though the feet vary in colour, from light brown to blackish. The fur is relatively sparse and consists of two hair types; moderately fine underfur (measuring 15–20 mm (0.59–0.79 in)) and long, stout bristle hairs (30–40 mm (1.2–1.6 in)). European Paleolithic rock art depicting

7980-609: The farthest north any such remains have been found), straight-tusked elephants , the narrow-nosed rhinoceros , and cave hyenas . William Buckland analyzed the cave and its contents in December 1821 and determined that the bones were the remains of animals brought in by hyenas who used it for a den, and not a result of the Biblical flood floating corpses in from distant lands, as he had first thought. His reconstruction of an ancient ecosystem from detailed analysis of fossil evidence

8094-410: The female's reproductive tract through her pseudo-penis rather than directly through the vagina, which is blocked by the false scrotum and testes. These unusual traits make mating more laborious for the male than in other mammals, and also make forced copulation physically impossible. The female retracts her clitoris before the male's penis enters it by sliding beneath it, an operation facilitated by

8208-463: The fossils in the cave were diluvial, that is that they had been deposited there by a deluge that had washed them from far away, possibly the Biblical flood. Upon further investigation, he realized the cave had never been open to the surface through its roof, and that the only entrance was too small for the carcasses of animals as large as elephants or hippos to have floated in. He began to suspect that

8322-439: The gazelle fawning season, as jackals and wolves are effective at tracking and catching young animals. Hyenas do not take to eating wolf flesh readily; four hyenas were reported to take half an hour in eating a golden wolf. Overall, the two animals typically ignore each other when there is no food or young at stake. Spotted hyenas dominate other hyena species wherever their ranges overlap. Brown hyenas encounter spotted hyenas in

8436-542: The genus Crocuta dating to the early Pliocene of Africa, around 3.63 to 3.85 million years ago. The earliest known fossil record of Crocuta crocuta in Africa is from the Cradle of Humankind , specifically from the Sterkfontein Member 4 which is dated approximately 2.1  million years ago in minimum, and it is also common in deposits dated between 2 to 1.5 million years ago . Analysis of

8550-513: The genus Crocuta still lacking. The ancestors of the genus Crocuta diverged from Hyaena (the genus of striped and brown hyenas ) 10 million years ago. The ancestors of the spotted hyena probably developed social behaviours in response to increased pressure from other predators on carcasses, which forced them to operate in teams. At one point in their evolution , spotted hyenas developed sharp carnassials behind their crushing premolars; this rendered waiting for their prey to die no longer

8664-419: The giant short-faced hyena. As there is no evidence of environmental change being responsible, it is likely that the giant short-faced hyena became extinct due to competition with the spotted hyena. The spotted hyena has a strong and well-developed neck and forequarters, but relatively underdeveloped hindquarters. The rump is rounded rather than angular, which prevents attackers coming from behind from getting

8778-488: The highest protein and fat content of any terrestrial carnivore. Cubs will nurse from their mother for 12–16 months, though they can process solid food as early as three months. Mothers do not regurgitate food for their young. Females are protective of their cubs, and will not tolerate other adults, particularly males, approaching them. Spotted hyenas exhibit adult behaviours early in life; cubs have been observed to ritually sniff each other and mark their living space before

8892-766: The latter favor their fathers by acting less aggressively toward them. Spotted hyena societies are more complex than those of other carnivorous mammals, and are remarkably similar to those of cercopithecine primates in respect to group size, structure, competition and cooperation. Like cercopithecine primates, spotted hyenas use multiple sensory modalities, recognise individual conspecifics, are conscious that some clan-mates may be more reliable than others, recognise third-party kin and rank relationships among clan-mates, and adaptively use this knowledge during social decision making. Also, like cercopithecine primates, dominance ranks in hyena societies are not correlated with size or aggression, but with ally networks. In this latter trait,

9006-573: The maniacal laughter, are among the most recognisable sounds of Africa. Typically, high-pitched calls indicate fear or submission, while loud, lower-pitched calls express aggression. 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. Dr. Hans Kruuk compiled the following table on spotted hyena calls in 1972: Spotted hyenas may contract brucellosis , rinderpest and anaplasmosis . They are vulnerable to Trypanosoma congolense , which

9120-551: The mitochondrial genomes of Eurasian Crocuta ( cave hyena ) specimens shows no clear separation from African lineages. It also suggests that modern spotted hyena populations descend from a widely ranging Afro-Eurasian population that was only recently reduced in range to Africa exclusively. However, analysis of the full nuclear genome suggests that Late Pleistocene African and Eurasian Crocuta populations were largely separate, having estimated to have diverged from each other around 2.5  million years ago , closely corresponding to

9234-470: The mouth, while large prey is eaten alive. Where spotted hyenas and lions occupy the same geographic area, the two species occupy the same ecological niche, and are thus in direct competition with one another. In some cases, the extent of dietary overlap can be as high as 68.8%. Lions typically ignore spotted hyenas, unless they are on a kill or are being harassed by them. There exists a common misconception that hyenas steal kills from lions, but most often it

9348-426: The narrow entrance of their cave in the evening down to the water's edge of a lake which must once have been there, & so helped themselves to some of the innumerable water-rats in wh[ich] the lake abounded. He developed these ideas further in his 1823 book Reliquiae Diluvianae; or, Observations on the organic remains contained in caves, fissures, and diluvial gravel, and on other geological phenomena, attesting

9462-436: The now extinct Eurasian populations weighed 102 kg (225 lb). Fur colour varies greatly and changes with age. Unlike the fur of the striped and brown hyena, that of the spotted hyena consists of spots rather than stripes and is much shorter, lacking the well defined spinal mane of the former two species. The base colour generally is a pale greyish-brown or yellowish-grey on which an irregular pattern of roundish spots

9576-517: The penis's upward angle. The hyenas then adopt a typical mammalian mating posture and usually lick their genitals for several minutes after mating. Copulation may be repeated multiple times during a period of several hours. The length of the gestation period tends to vary greatly, but the average length is 110 days. In the final stages of pregnancy, dominant females provide their developing offspring with higher androgen levels than lower-ranking mothers do. The higher androgen levels –

9690-427: The position of which allows it to crush bone with its premolars without blunting the carnassials. Combined with large jaw muscles and a special vaulting to protect the skull against large forces, these characteristics give the spotted hyena a powerful bite which can exert a pressure of 80  kgf/cm2 (1140 lbf/in² ), which is 40% more force than a leopard can generate. The jaws of the spotted hyena outmatch those of

9804-588: The presence of other predators. Thus the estimated contribution from hunting to their diet ranges from at least half up to 98%, the remainder being from scavenging. Wildebeest are the most commonly taken medium-sized ungulate prey item in both Ngorongoro and the Serengeti, with zebra and Thomson's gazelles , coming close behind. Cape buffalo are rarely attacked due to differences in habitat preference, though adult bulls have been recorded to be taken on occasion. In Kruger National Park, blue wildebeest , cape buffalo, plains zebra , greater kudu and impala are

9918-562: The pseudo-penis appears in the female fetus before differentiation of the fetal ovary and adrenal gland. When flaccid, the pseudo-penis is retracted into the abdomen, and only the prepuce is visible. After giving birth , the pseudo-penis is stretched, and loses many of its original aspects; it becomes a slack-walled and reduced prepuce with an enlarged orifice with split lips. Spotted hyenas are social animals that live in large communities (referred to as " clans ") which can consist of at most 80 individuals. Group-size varies geographically; in

10032-618: The raising of young. Giving birth is difficult for female hyenas, as the females give birth through their narrow clitoris, and spotted hyena cubs are the largest carnivoran young relative to their mothers' weight. During parturition, the clitoris ruptures to facilitate the passage of the young, and may take weeks to heal. Cubs are born with soft, dark brown hair, and weigh 1.5 kg on average. Unique among carnivorous mammals, spotted hyenas are also born with their eyes open and with 6–7 mm long canine teeth and 4 mm long incisors. Also, cubs will attack each other shortly after birth. This

10146-420: The region, and this appears as an aquifer in Kirkdale swallowing most of the water from Hodge Beck , which reappears further downstream. During summer months the river bed often runs dry as most of the water takes a subterranean passage. Kirkdale is noted for a bone cave , an ancient animal den, into which scavengers such as hyenas dragged the remains of many other animals. Numerous bones can still be found in

10260-521: The remains of livestock. This attracted the attention of numerous fossil collectors. Some of the fossils were sent to William Clift the curator of the museum of the Royal College of Surgeons ; he identified some of the bones as the remains of hyenas larger than any of the modern species. At the same time, William Buckland was told about the cave and shown some of the fossils by a colleague at Oxford. Buckland began his investigation believing that

10374-506: 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. The average litter consists of two cubs, with three occasionally being reported. Males take no part in

10488-402: The rocky areas of East Africa and Congo, spotted hyenas use caves as dens, while those in the Serengeti use kopjes as resting areas in daylight hours. Dens have large bare patches around their entrances, where hyenas move or lie down on. Because of their size, adult hyenas are incapable of using the full extent of their burrows, as most tunnels are dug by cubs or smaller animals. The structure of

10602-447: The same clan, differences in scent gland compositions can help individuals differentiate the reproductive states and sex of their members. One study analyzed the scent gland secretions of several hyenas belonging to three different clans and discovered a high degree of similarity in fatty acid composition among members of the same clans, while hyenas of different clans had more dissimilar scent gland secretions. Further studies have proposed

10716-460: The same species, or distinguished purely on geographical, rather than physical, grounds. Hiob Ludolf , in his Historia aethiopica , was the first to clearly distinguish the Crocuta from Hyaena on account of physical and geographical grounds, though he never had any first hand experience of the species, having gotten his accounts from an Ethiopian intermediary. Confusion still persisted over

10830-504: The sight of them, but will react fearfully to the scent. Although cheetahs and leopards preferentially prey on smaller animals than those hunted by spotted hyenas, hyenas will steal their kills when the opportunity presents itself. Cheetahs are usually easily intimidated by hyenas, and put up little resistance, while leopards, particularly males, may stand up to hyenas. There are records of some male leopards preying on hyenas. Hyenas are nonetheless dangerous opponents for leopards; there

10944-443: The social system of the spotted hyena is openly competitive rather than cooperative, with access to kills, mating opportunities and the time of dispersal for males depending on the ability to dominate other clan -members. Females provide only for their own cubs rather than assist each other, and males display no paternal care. Spotted hyena society is matriarchal ; females are larger than males, and dominate them. The spotted hyena

11058-473: The species indicates that the Eurasian populations retained the spots of their modern-day African counterparts. The genitalia of the female closely resembles that of the male; the clitoris is shaped and positioned like a penis, a pseudo-penis , and is capable of erection . The female also possesses no external vagina (vaginal opening), as the labia are fused to form a pseudo- scrotum . The pseudo-penis

11172-507: The split, which likely explains the discordance between the nuclear and mitochondrial genome results, with the mitochondrial genomes of African and European Crocuta more closely related to each other than to Asian Crocuta, suggesting gene flow between the two groups after the split between the Asian and European populations. Its appearance in Europe and China coincided with the decline and eventual extinction of Pachycrocuta brevirostris ,

11286-410: The spot patterns to help identify individual hyenas. A set of five, pale and barely distinct bands replace the spots on the back and sides of the neck. A broad, medial band is present on the back of the neck, and is lengthened into a forward facing crest. The crest is mostly reddish-brown in colour. The crown and upper part of the face is brownish, save for a white band above both eyes, though the front of

11400-415: The spotted hyena further show parallels with primates by acquiring rank through coalition . However, rank reversals and overthrows in spotted hyena clans are rare. The social network dynamics of spotted hyenas are determined by multiple factors. Environmental factors include rainfall and prey abundance; individual factors include preference to bond with females and with kin; and topological effects include

11514-411: The spotted hyena is promiscuous , and no enduring pair bonds are formed. Members of both sexes may copulate with several mates over the course of several years. Males will show submissive behaviour when approaching females in heat, even if the male outweighs his partner. Females usually favour younger males born or joined into the clan after they were born. Older females show a similar preference, with

11628-516: The spotted hyena was widespread throughout Sub-Saharan Africa . It is present in all habitats save for the most extreme desert conditions, tropical rainforests and the top of alpine mountains. Its current distribution is patchy in many places, especially in West Africa. Populations are concentrated in protected areas and surrounding land. There is a continuous distribution over large areas of Ethiopia , Kenya , Tanzania , Botswana , Namibia and

11742-615: The spotted hyena's most important prey, while giraffe , impala, wildebeest and zebra are its major food sources in the nearby Timbavati area. Springbok and kudu are the main prey in Namibia 's Etosha National Park , and springbok in the Namib . In the southern Kalahari , gemsbok , common eland , lechwe , waterbuck , wildebeest and springbok are the principal prey. In Chobe , the spotted hyena's primary prey consists of migratory zebra and resident impala. In Kenya 's Masai Mara , 80% of

11856-618: The spotted hyena's prey consists of topi and Thomson's gazelle, save for during the four-month period when zebra and wildebeest herds migrate to the area. Bushbuck , suni and buffalo are the dominant prey items in the Aberdare Mountains , while Grant's gazelle , gerenuk , sheep , goats and cattle are likely preyed upon in northern Kenya. Unlike other large African carnivores, spotted hyenas do not preferentially prey on any species, and only African buffalo and giraffe are significantly avoided. Spotted hyenas prefer prey with

11970-560: The striped hyena by its much greater size and narrower sagittal crest . For its size, the spotted hyena has one of the most powerfully built skulls among the Carnivora. The dentition is more dual purposed than that of other modern hyena species, which are mostly scavengers; the upper and lower third premolars are conical bone-crushers, with a third bone-holding cone jutting from the lower fourth premolar. The spotted hyena also has its carnassials situated behind its bone-crushing premolars,

12084-424: 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

12198-411: 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,

12312-743: The tendency to close triads in the network. Female hyenas are more flexible than males in their social bonding preferences. Higher ranking adult spotted hyenas tend to have higher telomere length and higher levels of some immune defense proteins in their blood serum. Territory size is highly variable, ranging from less than 40 km in the Ngorongoro Crater to over 1,000 km in the Kalahari. Home ranges are defended through vocal displays, scent marking and boundary patrols. Clans mark their territories by either pasting or pawing in special latrines located on clan range boundaries. Spotted hyenas use scent gland secretions to distinguish between members of their own clan and members of neighboring clans. Within

12426-530: The then recognised subspecies could also be found in a single population, with the only set of characters standing out being pelage (which is subject to a high degree of individual variation) and size (which is subject to Bergmann's Rule ). When fossils are taken into consideration, the species displayed even greater variation than it does in modern times, and a number of these named fossil species have since been classed as synonymous with Crocuta crocuta , with firm evidence of there being more than one species within

12540-461: The viruses isolated from hyenas and lions were more closely related to each other than to the closest canine distemper virus in dogs. Evidence of canine distemper in spotted hyenas has also been recorded in the Masai Mara. Exposure to rabies does not cause clinical symptoms or affect individual survival or longevity. Analyses of several hyena saliva samples showed that the species is unlikely to be

12654-412: The whole, the relationship between the two species is a one sided benefit for the hyenas, with wild dog densities being negatively correlated with high hyena populations. Medium-sized canids like Black-backed and side-striped jackals , and African wolves will feed alongside hyenas, though they will be chased if they approach too closely. Spotted hyenas will sometimes follow jackals and wolves during

12768-588: Was admired at the time, and considered to be an example of how geo-historical research should be done. The cave was extended from its original length of 175 metres (574 ft) to 436 metres (1,430 ft) by Scarborough Caving Club in 1995. A survey was published in Descent magazine. The fossil bones found in the cave included those of straight-tusked elephants , hippopotamuses , the narrow-nosed rhinoceros , cave hyena , cave lion , Irish elk , brown bears , red deer and fallow deer . These have been dated to

12882-537: Was once widely thought to be derived from the Latin loanword crocutus , which translates as 'saffron-coloured one', in reference to the animal's fur colour. This was proven to be incorrect, as the correct spelling of the loanword would have been Crocāta , and the word was never used in that sense by Graeco-Roman sources. Crocuta actually comes from the Ancient Greek word Κροκόττας ( krokottas ), which

12996-489: Was precise enough to be included by Johann Erxleben in his Systema regni animalis by simply translating Pennant's text into Latin. Crocuta was finally recognised as a separate genus from Hyaena in 1828. Unlike the striped hyena, for which a number of subspecies were proposed in light of its extensive modern range, the spotted hyena is a genuinely variable species, both temporally and spatially. Its range once encompassed almost all of Africa and Eurasia , and displayed

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