taxonID	type	description	language	source
03928788FFEAFF8828B2FDA8F8E3C123.taxon	diagnosis	• Mediumto large-sized mammals with round or pointed ears relatively large in comparison to body size, exceptionally powerfuljaw muscles, hindquarters long and sloping; somewhat dog-like in overall appearance, very muscular. • 85 - 185 cm.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEAFF8828B2FDA8F8E3C123.taxon	distribution	• Africa and Middle East to India.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEAFF8828B2FDA8F8E3C123.taxon	diagnosis	• Mainly savannas, but other semi-arid and desert regions to edges of forests, both tropical and temperate zones. • 4 genera, 4 species, at least 5 extant taxa. • All Lower Risk; none Extinct since 1600.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEBFF8B2AA6F458FAD2C3D3.taxon	materials_examined	Eastern Cape Province, South Africa.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEBFF8B2AA6F458FAD2C3D3.taxon	discussion	Although previously placed in its own family (Protelidae), it is now considered a member of the family Hyaenidae. The Aardwolf belongs to the subfamily Protelinae, of which it is the only extant member. They occur in two distinct populations separated by about 1500 km. However, studies ofthe extent of genetic and morphological differences between these groups have not been conducted. Two subspecies are recognized.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEBFF8B2AA6F458FAD2C3D3.taxon	distribution	Subspecies and Distribution. P. c. cristata Sparrman, 1783 — E African coast (S Egypt, Sudan, Eritrea, Djibouti, Ethiopia, Somalia, Kenya, NE Uganda to C Tanzania). P. c. septentrionalis Rothschild, 1902 — most of S Africa (S Angola, S Zambia, SW Mozambique, Namibia, Botswana, Zimbabwe, Swaziland, Lesotho, and South Africa).	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEBFF8B2AA6F458FAD2C3D3.taxon	description	Descriptive notes. By far the smallest of the four hyaenid species. Head-body 55 - 80 cm, tail 20 - 30 cm, height at shoulder 45 - 50 cm; weight (adult) 8 - 12 kg with seasonal variation, and reported as high as 14 kg. No sexual size dimorphism. Superficially similar in appearance to the Striped Hyena, with dark vertical stripes on a buff, yellowish-white or rufous body, and irregular horizontal stripes on the legs. However, the Striped Hyena is more than twice as large with less regular striping. The Aardwolf’s coat is about 2 - 5 cm long, with longer hairs along the mane and in the bushy tail. The neck is long and the throat is a pale gray-white. The legs are long and slender and the striping terminates in black at the feet. As in the Striped and Brown Hyena, the Aardwolf has long, pointed ears and a long erectile mane extending the length ofits body. Like the other hyaenids, Proteles has a sloping back with the forelegs longer than the hindlegs, and a well-developed anal gland used for scent marking. Females have two pairs of teats. Uniquely among the hyaenids, Profeles has a number of adaptations for feeding exclusively on termites, including a long, spatulate tongue with large and varied papillae, and a large submaxillary gland which produces copious amounts of sticky saliva. Very small peg-like cheek teeth are widely spaced along thejaw margins, yet large canines have been retained for use in territorial disputes with other Aardwolves and defense againstjackals. Their skulls also feature a relatively broad, nearly parallel-sided palate, and extraordinarily large tympanic bulla.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEBFF8B2AA6F458FAD2C3D3.taxon	biology_ecology	Habitat. Aardwolves are primarily found on open, grassy plains or in bush country, but can live in a range of habitats with rainfall between 100 - 800 mm. They are most common where rainfall is 100 - 600 mm. They do not occur in forests or pure desert and are independent of drinking water. The northern subspecies occurs in grasslands and tree savannas of the Somali-Masai Arid Zone and the southern subspecies in the Southern Savanna and South-west Arid Zone.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEBFF8B2AA6F458FAD2C3D3.taxon	food_feeding	Food and Feeding. Aardwolves are solitary foragers and feed almost exclusively on Trinervitermes termites, usually on one species in each particular region: 1. bettionianus in East Africa, 1. rhodesiensis in Zimbabwe and Botswana and 71. trinervoides in South Africa. These termites are largely avoided by other termite-eating mammals due to the noxious terpenoid chemicals secreted by the soldier termites, to which the Aardwolfis uniquely tolerant. In addition to the lack of competition, Aardwolf preference for this termite genus is likely due to the fact that these termites regularly congregate at night in large aboveground foraging parties. The termites are licked directly from the soil surface, and are easily obtainable in large quantities. Also, unlike true harvester termites, Trinervitermes forage throughout the year, making them a dependable year-round food source. Due to the small size of Trinervitermes and the wide dispersion of colonies, female Aardwolves must forage for at least six hours a night, during which up to 300,000 termites / night (1 - 2 kg) are consumed. Foraging Aardwolves travel approximately 1 km / h, with their ears cocked forward and head bentslightly down, following an erratic zig-zag route. Because they often approach termite colonies from downwind, and approach with directed movement before termites could be seen, it appears that termite foraging parties are detected at least partially by smell. However, the hearing of the Aardwolf is particularly acute and is assumed to play a role in colony detection. The average time spent foraging at individual termite patches was 20 - 28 seconds in East Africa, but in drier Namibian grassland, Aardwolves spent an average of 1 - 8 and 9 - 2 minutes at each patch in consecutive years of observation. In South Africa, a newly weaned four-month old cub spent an average of only eleven seconds at each patch, and juveniles frequently are seen vomiting after feeding on Trinervitermes, indicating that tolerance to the chemical secretions of Trinervitermes soldiers increases with age. Even adult Aardwolves maintain some aversion to the terpene chemicals, because they will avoid feeding on mounds under repair, where typically only dense concentrations of soldiers are found at the surface. Other surface-foraging termites, particularly Hodotermes and Microhodotermes (South Africa), Odontotermes and Macrotermes species (East Africa), make up a larger proportion of the diet when Trinervitermes are seasonally uncommon or unavailable, as during winter (May-August) in South Africa, and during the rainy season in East Africa. However, these species forage aboveground in much smaller parties (10 - 20 individuals vs. 4000 in Trinervitermes) and the reduced winter availability of Trinervitermes in South Africa results in a significant seasonal reduction in Aardwolf body weight and field metabolic rate. Aardwolves here were found to consume only one-sixth the number of termites in winter that they did in summer. Winter is also the highest period of mortality in Aardwolf cubs, which are 7 - 10 months old at this time, further indicating that this is a period ofsignificant energetic stress. Other termites found in fecal samples have included Odontotermes, Macrotermes, and Lepidotermes that are not surface-foraging species and therefore not important components of the Aardwolf diet. Occasional additional food items include ants and Coleopterans, yet the Aardwolf appears to be surprisingly inefficient at foraging on non-termite insects. Due to the high degree of specialization of its tongue for licking small arthropods, and the almost complete degeneration ofits cheek teeth, it is thought that they are unable to handle larger food items, making the species highly dependent on Trinervitermes. This dependence is supported by the absence of Aardwolves from Zambia and central and western Africa, where surface foraging Trinervitermes are either uncommon or available only a small part of the year. Aardwolves defecate in middens (also called latrines). The first defecation occurs when they exit the burrow in the evening and is typically very large, weighing up to 1 kg. Defecations are typically covered with soil. This practice of concentrating and burying their faeces, which retains some of the terpene smell of the soldier termites, has been suggested as a way of reducing the probability that an Aardwolf will mistake its own faeces for a termite colony when it is foraging. Up to 20 middens may be located in a territory.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEBFF8B2AA6F458FAD2C3D3.taxon	activity	Activity patterns. Predominantly nocturnal, in South Africa Aardwolves are generally active for 8 - 9 hours a day in summer but only 3 - 4 hours a day in winter. In summer, they generally leave the den within an hour after sunset and return 1 - 4 hours before sunrise. However, during winter, some diurnal activity may be observed. Aardwolves typically become active up to an hour before sunset and return to their dens after 3 — 4 hours of foraging. A higher proportion of their time is spent feeding in winter than in summer, and a relatively large portion of the activity of both males and females in winter (12: 6 %) consists of breeding activities (e. g. courtship / copulation). Inactive hours during the day are spent in underground dens, which provide refuge from temperature extremes and predators, particularly Black-backed Jackals. Dens also function importantly in cub-rearing. Nighttime rest periods are also often spent near or inside dens. A territory may include up to ten different den holes, which are typically spring hare burrows that have been enlarged by Aardwolves.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEBFF8B2AA6F458FAD2C3D3.taxon	biology_ecology	Movements, Home range and Social organization. A social unit, which occupies a welldefined territory throughout the year, consists of a male-female pair and their most recent offspring. All natal animals disperse from the territory, usually 1 - 2 months before birth of the nextlitter. Pair bonds arefairly stable lasting 2 - 5 years. Males without mates (due to death or abandonment by females) establish pair bonds with adjacent females and may abandon their originalterritories. Territory size ranges from 1 - 5 km? in East Africa to 3 - 8 km? in South Africa, and appears to be negatively related to the density of available termite mounds. Territories in the Northern Cape Province of South Africa generally supported 3000 termite mounds with an average of 55,000 termites / mound. Aardwolf density reaches 1 adult / km? in optimal habitat. Territories are maintained primarily by scent marking, which is concentrated along territory boundaries, but direct interactions between neighboring residents also occasionally occur. Both males and females actively defend territorial boundaries. Chasing and fighting, with manes raised, occurs between same-sex individuals defending territories. Intruders encountered within the territory are usually chased to the boundary and mutual avoidance is generally practiced along boundary areas. If physical contact occurs both combatants drop to their carpals and bite at each others’ necks. Although territorial behavior is exhibited by males and females, it differs between mating (June andJuly in South Africa) and non-mating seasons. Direct fights between Aardwolves appear restricted to the mating season. Whereas females tend to stay within territory boundaries year round, male behavior undergoes a marked change at the start of the mating season. After an approximately one-month “ scouting ” period at the beginning of the mating season, when males make frequent extra-territorial movements, yet largely refrain from pasting outside the territory, they begin more aggressive extra-territorial pasting. Their movements outside the territory continue to increase, peaking in frequency about a week before females come into estrus. These excursions are suggested to be advertizements of quality to both males and females in surrounding territories. Males engage in consecutive over-markings by pasting on particular grass stalks; the less aggressive, and apparently less fit, individual will eventually cease pasting, thus “ losing ” the contest. During pre-estrus females also increase their rate of pasting, primarily along territory boundaries and just outside them, apparently to encourage visits by extra-pair males. Visiting males during this period frequently “ flirt ” with resident females and chase or fight males that they encounter. “ Flirting ” typically involves the male running toward the female, then veering off and prancing past with his tail raised. However, by the time the female is in estrus (lasting 1 - 3 days) she is typically left with only her resident male, and potentially an aggressive neighbor. As in Striped Hyenas, there have been cases in which two male Aardwolves shared a territory with a female, both males mating with her and both guarding her cubs, but this appears to be exceptional and rare. Aardwolves are remarkably antisocial outside the breeding season. Members of resident mixed-sex pairs feed alone and typically ignore each other when they meet. Unlike the other three hyaenids, Aardwolves usually do not engage in greeting ceremonies between familiar individuals, with the exception of an occasional muzzle to muzzle sniff between mother and cub. In South Africa, nightly distance traveled by foraging females ranged from 1 - 5 to 9 - 1 km (average 4 - 2 km). Summer travel distances ranged from 8 - 12 km per night, whereas winter distances were highly variable (from less than 3 km to more than 24 km) depending on whether males were conducting extra-territorial mating forays. Travel speed is 2 - 3 km / h when not feeding, and about I km / h when feeding intensely. Aardwolves return to underground burrows during the day. There are typically 5 - 6 dens per territory. Dens are used for only 1 - 2 months at a time, and mates rarely use the same dens concurrently. Because Aardwolves rarely interact, the primary form of communication is olfactory. Like the other three hyaenids, the Aardwolf engages in scent marking behavior called pasting, during which a strong-smelling, yellowish-orange secretion (which quickly turns black) is deposited onto grass stalks from an extruded anal gland, located just above the anus. In addition to marking frequently at dens and latrines, which generally are not associated with territory boundaries, Aardwolves appear to use pasting as a means of territory defense. Boundary marking occurs most frequently, and is most concentrated along borders where neighboring Aardwolves maintain territories. Pasting is generally frequent, occurring about twice every 100 m of travel, and about 200 times per night, with males pasting more than females. Based on experiments with translocated scent marks, information conveyed in scent marks appears to include the sex, female reproductive state, and individual identity, at least in the case of resident neighbors, partners, and self-recognition. Outside the mating season, pasting outside territory boundaries is rare if not nonexistent, but this behavior, particularly by males, changes notably during mating periods. Even though direct interactions are rare, Aardwolves possess an impressive visual display, during which the hairs along the mane are erected, resulting in a near doubling of the apparent size of the animal. This is used in intraspecific aggressive interactions involving territory defense and in interspecific defensive interactions. Although generally a silent species, the most comprehensive analysis of the vocal repertoire of Proteles identified nine distinct sound types: “ purr ”, whine, jaw click, lip smack, growl, snarl, bark, squeal, and a whizzing sound which was only documented in one individual. Agonistic vocalizations are relatively diverse and increase in intensity in the following order: lip smack / jaw click, growl, snarl, bark. Squeals are heard only in cubs and appear to represent begging to mothers. The whineelicits a variety of reactions depending on the addressee and addressor, but likely functions as an appeasing or reassuring sound. As in striped and brown hyenas, Aardwolves lack a loud, longdistance vocalization like the whoop of spotted hyenas.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEBFF8B2AA6F458FAD2C3D3.taxon	breeding	Breeding. Monogamous, yet during the mating season extra-pair copulations can be common (40 % in South Africa). Strictly seasonal breeding in the Northern Cape where most mating occurs during the first two weeks ofJuly. Females give birth every year in early October to 1 - 4 cubs (average of 2 - 5), after a 90 day gestation period. In more northern parts of the range, breeding seasons appear to be less restricted. Estrus lasts 1 - 3 days but females may cycle again within two weeks if fertilization does not occur. Copulation lasts from 1 - 4 - 5 hours during which multiple ejaculations occur. Copulations may be interrupted by extra-pair males and in some cases females copulate with these new males. Cubs are born in a den and rarely emerge above ground during the first month. By three months, cubs have begun making short excursions from the den, usually accompanied by an adult. Weaning occurs around four months, and by this time cubs begin foraging alone within the territory. After weaning, cubs spend little time with their parents, are independent by about seven months, and are excluded from the territory soon after, usually by one year of age. Cubs grow quickly, reaching adult body mass by 3: 6 months. Thisis likely an adaptation to maximize survival of cubs through their first winter, when cub mortality is highest. Sexual maturity is reached by 1 - 8 years. Each resident breeding male guards the female’s cubs during the period of den dependence. This is energetically costly asit typically leaves the males only 2 - 3 hours of foraging time before sunrise, compared to at least six hours for females. Due to the frequency of extrapair copulations, cuckoldry appears to be an established aspect of the mating system (“ overt cuckoldry ”), and males are likely to frequently help raise litters of mixed paternity or sired entirely by other males. Currently this appears to be unique among mammals.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEBFF8B2AA6F458FAD2C3D3.taxon	conservation	Status and Conservation. Listed as a species of Least Concern on The IUCN Red List. Due to their shy and nocturnal nature, Aardwolves are probably more common than usually believed. That notwithstanding, Aardwolves in southern Africa generally occur outside of protected areas, and the primary threat in these locationsis indirect poisoning aimed at locust outbreaks. Poisoning events can result in the death of up to half the local adult population and all the cubs. Within protected areas, the most important mortality sources are severe drought and predation on cubs by Black-backed Jackals. Human-caused mortality also occurs as a result of direct persecution from farmers suspecting Aardwolf involvement in lamb predation, harvesting of Aardwolves as a food source, and indirect persecution during organized hunting for jackals. Aardwolves may also be killed by vehicles during the night. However, all these other mortality sources appear insignificant relative to poisoning, jackal predation and drought. Across its range, habitat fragmentation and isolation may be the most serious threat to long-term population viability; however, its dependence on habitat preferred for use in livestock grazing makes extensive habitat loss improbable.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFE9FF8D2ADAF6DDF8B1C136.taxon	materials_examined	Guinea, Aethiopia; restricted to “ Senegambia ”.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFE9FF8D2ADAF6DDF8B1C136.taxon	discussion	The earliest members of the genus Crocuta first appear in the fossil record of Africa in the early Pliocene, dated at roughly 3 - 7 million years ago. However, members of this genus soon dispersed out of Africa, and based on fossils from the period ofits greatest range expansion in the Pleistocene, the genus Crocuta occupied virtually all of Europe and Asia, as well as most of sub-Saharan Africa. When exactly modern C. crocuta arose is not entirely certain, but this species is clearly very recent. C. crocuta does not appear in the fossil record until sometime after 990,000 years ago, and probably substantially closer to the present, perhaps within the last 250,000 years. Modern Spotted Hyenas can be distinguished from members of the genus Crocuta found in the fossil record based on body size, limb length and stoutness, the length and shape of particular skull bones, and unique characteristics of the cheek teeth. In contrast to earlier members of the genus, including the Cave Hyenas of Europe and Asia (C. spelaea), modern Spotted Hyenas have a post-cranial skeleton that is modified for cursorial hunting. Currently only one subspeciesis recognized despite substantial variation in coloration and body mass throughout sub-Saharan Africa. For example, individuals from southern Africa are larger than those from eastern Africa. Monotypic.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFE9FF8D2ADAF6DDF8B1C136.taxon	distribution	Distribution. Most of Africa S of the Sahara Desert, except in lowland tropical rainforests. Spotted Hyenas have been extirpated from many areas of southern Africa.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFE9FF8D2ADAF6DDF8B1C136.taxon	description	Descriptive notes. Largest of the four hyaenid species. Head-body 125 - 160 cm, tail 22 - 27 cm, height at shoulder 77.3 - 80. 7 cm; weight 45 - 55 kg and up to 86 kg. Females approximately 10 % larger than males, although size distributions for males and females overlap. Degree of sexual dimorphism in body size varies geographically, being most pronounced in southern Africa. Its general color is sandy, ginger, or dull gray to reddish-brown, with black or dark brown spots on the back, flanks, rump, and legs. Spots may turn brown and fade with age. The fur is shorter in this species than in the other extant hyaenids. The head is large, rounded and powerful, with a short and blunt muzzle. In contrast to the other extant hyaenids, all of which have pointed ears, Spotted Hyenas have ears with rounded tops. The tail ends in a black, bushy tip, with approximately 12 cm of hair extending beyond the end of the tail bone. Like the other hyaenids, the Spotted Hyena has a sloping back because the forelegs are longer than the hindlegs, and a well-developed anal gland used for scent marking. The mane in this speciesis more poorly developed than in other hyaenids. The feet have four toes. Females usually have only two teats. The Spotted Hyena has long been considered a hermaphrodite in many parts of Africa because the external genitalia of the female are very similar to those of the male. The female has a peniform clitoris that is only a few mm shorter than the male’s penis, and is fully erectile. The sexes can be distinguished by the shape of the penile glans: the male glans is pointed whereas that of the female is blunt. A single urogenital canal traverses the enlarged clitoris; through this canal the female urinates, copulates and gives birth. There is no external vaginal opening as the outer labiae are fused to form a structure that resembles the scrotal sac of the male. The female’s pseudoscrotum has a bi-lobed appearance; the testes of the adult male make the scrotal sac larger and give it more distinctly rounded bulges. Thus scientists who study these animals can distinguish males from females even when the animals are lying down.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFE9FF8D2ADAF6DDF8B1C136.taxon	biology_ecology	Habitat. Spotted Hyenas occupy an extraordinarily diverse array of habitats, including savanna, semi-desert, swamps, woodland, and montane forest up to 4000 m of elevation, but are absent in lowland tropical rainforests, in alpine areas at high elevation, and in extreme desert conditions. Although they require water for drinking, they are able to make do with very little water, and seldom require access to it. Even lactating females can survive without water for over one week. The highest population densities reported for this species occur on the prey-rich plains of Kenya and Tanzania, and surprisingly, in the forests of the Aberdare Mountains in Kenya. In these areas, densities of Spotted Hyenas exceed one animal per square kilometer.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFE9FF8D2ADAF6DDF8B1C136.taxon	food_feeding	Food and Feeding. The foraging behavior of Spotted Hyenas is remarkably flexible. Long believed to feed mainly on carrion, these animals are in fact efficient predators that kill 60 % to 95 % of their prey themselves. On average across populations in which the relative proportions of hunted and scavenged foods have been documented, two thirds of their diet is derived from kills they make themselves, and only one third from scavenged food items. In addition to being able to obtain food either by hunting or scavenging, Spotted Hyenas exhibit extraordinary plasticity with respect to their prey preferences. Spotted Hyenas have catholic tastes, they are extreme opportunists, and they are able to exploit a vast array of potential prey types, ranging from caterpillars to elephants; they may also occasionally consume some plant material. However, in most parts of Africa, Spotted Hyenas derive the large majority of their food intake from only a small subset of the prey species available to them locally. In most environments, they focus on the local mediumand large-sized ungulates, capture of which yields the greatest caloric return while demanding the least effort and the fewest risks. Thus, in eastern Africa, Spotted Hyenas prey most frequently on Blue Wildebeest, zebra, gazelles and Topi. In the arid parts of southern Africa, they prey most frequently on Gemsbok. In Kruger National Park, their most common prey is Impala, and in western Africa, common prey includes Red-fronted Gazelles and Hartebeest. Foraging behavior varies with the prey currently sought. Spotted Hyenas search for gazelle fawns by wandering upwind through open grassland in a zig-zag pattern. They may dig for crocodile eggs along large rivers, and snap flying termites out of the air with their jaws. When hunting, Spotted Hyenas modify their behavior to take advantage of the most abundant prey species, or the species thatis easiest to catch; these change seasonally in some localities with the migratory movements of particular ungulate species. Instead of using felid-like stealth as a primary hunting tactic, Spotted Hyenas rely on their extraordinary endurance for success in hunting. They can run at speeds of up to 55 km / h, but at slightly lower speeds, they can maintain a chase for several kilometres. If the antelope being chased becomes winded, and turns to defend itself with its horns, the Hyenas rush in and start tearing off pieces of the prey animal’s flesh. Like canids, Spotted Hyenas kill their prey by disembowelling and dismembering them rather than by using a particularkilling bite. Spotted Hyenas may hunt either solitarily or in groups; in the latter case group size varies with the type of prey sought. Mean hunting group sizes among Hyenas in Kenya are 1 - 2 for Topi, 1 - 7 for Impala, 2 - 08 for Thompson's Gazelle, 2 - 92 for Blue Wildebeest, and 9 - 1 for zebra. Thus only zebra hunts involve large groups of hunters, and most hunting parties contain only one or two Hyenas. Ungulates such as Topi and Blue Wildebeest weigh roughly three times as much as an adult Hyena, but solitary Hyenas routinely kill these antelope. Although hunting group size is often surprisingly small among Spotted Hyenas, the feeding groups formed by these animals are often very large once a prey animal has been killed. The noise produced by feeding Hyenas often draws members of the clan that were not involved in the hunt to the kill site. Feeding competition among the Hyenas present at a kill is usually very intense. In East Africa, often more than 30 Hyenas can be observed trying to feed from a single carcass. Because of this intense competition, each individual Hyena consumes as much food as possible in a very short period of time. A group of 20 - 30 hungry Hyenas can reduce an adult Blue Wildebeest to nothing more than a pile of rumen contents in only 13 minutes. It is estimated that an adult Spotted Hyena can consume a mass of food equal to 25 % - 30 % ofits body weight, and individual Hyenas have been observed to ingest up to 18 kg of meat and bone in one hour. However, as a result of limited access to carcasses, average food intake ranges only from 1 - 5 to 3 - 8 kg per day. Spotted Hyenas sometimes engage in kleptoparasitism, which is the aggressive acquisition of a fresh carcass from other predators. They have been observed displacingjackals, Striped Hyenas, Leopards, Cheetahs, and African Wild Dogs from kills. However Spotted Hyenas most frequently compete for kills with Lions. Spotted Hyenas and Lions occur sympatrically in many areas of Africa, and in most of these habitats, bi-directional food stealing has been observed between these two species. Dominance relations between Spotted Hyenas and competing species are not absolute but depend on the numerical presence of both parties. For instance, Lions usually displace Spotted Hyenas at kills. However, if Hyena group size is large and the ratio of Spotted Hyenas to female and subadult Lions exceeds four to one, Hyenas are often able to displace Lions from kills unless a male Lion is present. A single Spotted Hyena can usually dominate a Cheetah, Leopard, Striped Hyena, Brown Hyena, any species ofjackal, or an African Wild Dog. Spotted Hyenas have been observed caching surplus food in thickets and under water in ponds. These animals are very comfortable in water; they often play in seasonal pools, and lie in shallow water or wet mud to keep cool on hot days. Compared to the other bone-cracking hyenas, Spotted Hyenas rarely carry food to their young at dens. This appears to be because the risk of having one’s food stolen, even by much smaller hyenas, is very high at dens, particularly for low-ranking individuals.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFE9FF8D2ADAF6DDF8B1C136.taxon	activity	Activity patterns. Spotted Hyenas are predominantly nocturnal and crepuscular, although they may be active at midday when temperatures permit. Dens are typically modified holes dug by Aardvarks, although caves are used as den sites in some areas. Only cubslive in dens; adults sleep above ground, often in thickets, particularly when midday temperatures are high. Although Spotted Hyenas are active for roughly one third of each 24 hour cycle, their activity is not continuous. Instead, activity occurs in bouts interspersed with periods of rest. Hyenas in Kenya that were followed for complete 24 hour cycles spent 32 % of their time active, but 53 % of their active time occurred during hours of darkness.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFE9FF8D2ADAF6DDF8B1C136.taxon	biology_ecology	Movements, Home range and Social organization. On average, Spotted Hyenas in Kenya move 928 m per hour when active, and typically travel over 12 km during each 24 hour period, with males moving more than females. In Serengeti, daily movements may be much greater than this, as resident hyenas often commute 30 - 40 km in orderto feed on migratory herbivores. Spotted Hyenas live in social groups, called clans, which contain from ten to eighty members. Large clans contain multiple matrilines of related females and their offspring, as well as a number of adult immigrant males that are generally unrelated to one another. Small clans may contain only a single matriline and a single breeding male. Clan size appears to be determined by abundance of local prey animals: where these are plentiful, as on the prey-rich plains of eastern Africa, clans are typically very large, but in desert areas of southern Africa, clans may be tiny. Clans are fission-fusion societies. That is, all clan members know each other individually, occupy a common territory, and rear their cubs together at a communal den, yet they also spend much of their time alone or in small sub-groups. Spotted Hyena clans bear little resemblance to canid packs or Lion prides, but they are remarkably similar in their size, structure, and complexity to the societies of cercopithecine primates. Like troops of baboons and macaques, Hyena clans typically contain individuals from multiple overlapping generations, and clans are structured by clear linear dominance hierarchies in which an individual’s rank determinesits priority of access to food and other resources. In contrast to the situation characteristic of other hyaenids and most other mammals, female Spotted Hyenas are socially dominant to all adult immigrant males. Rank relationships among female clan-mates are usually stable for periods of many years. Average relatedness among females from different matrilines within a clan is extremely low. Like most primates, Spotted Hyenas produce tiny litters at long intervals, and their offspring require an unusually long period of nutritional dependence on the mother. Young Hyenas typically nurse for well over a year, and because it takes them years to become proficient at hunting and feeding, their mothers continue to help them gain access to food at ungulate kills long after weaning. Similar to female baboons, the social status of a female Hyena is determined not by hersize or fighting ability, but by her mother’s social rank. Indeed, the acquisition of social rank during early development occurs in a pattern identical to that seen in many monkey species, a pattern called “ maternal rank inheritance ” by primatologists even though no literal inheritance occurs involving genetic transfer of status from mother to offspring. Instead, in both Hyenas and baboons, maternal rank “ inheritance ” involves a great deal of important social learning that occurs during a protractedjuvenile period. Young Hyenas initially direct their aggressive behaviors equally at higherand lower-ranking individuals. But this changes rapidly during the first year of life as cubs learn to direct aggression only at animals lower in rank than their own mother. When youngsters become involved in disputes with group-mates, the mother intervenes on their behalf against all individuals lower-ranking than herself. Interventions by high-ranking mothers are more frequent and more effective than those by lowranking females. In addition, like young baboons, Hyena cubs are often joined in fights by coalition partners who may be either kin or unrelated animals. Along with maternal interventions, coalition formation functions importantly in rank acquisition. Thus the mechanisms by which youngsters acquire their social ranks are virtually identical in Hyenas and old-world monkeys. Patterns of competition and cooperation among Spotted Hyenas are also remarkably like those found in baboons. Although Hyenas compete intensively for food, they also rely heavily on cooperative interactions with group-mates, particularly their close kin, to acquire and defend both their social rank and such key resources as food and territory. Young Spotted Hyenas of both sexes “ inherit ” the social rank of their motherearly in life, and retain their maternal rank as long as they remain in the natal clan. However, whereas females remain in their natal group throughouttheir lives, virtually all males disperse after puberty tojoin a new clan. When a male immigrates into a new group, he entersas the lowest-ranking Hyena in the dominance hierarchy; he behaves submissively to all Hyenas he encounters in the new territory, regardless of their size, fighting ability, or social rank. This results in a society in which adult females and their cubs are dominant to all adult male immigrants. A male Hyena loses his maternal social rank and its associated feeding privileges when he disperses. In their new clans, immigrant males sometimes invest a great deal of time and energy in developing amicable relationships with resident adult females, as males engaging in these amicable relationships may enjoy a high probability ofsiring cubs. Due to the female’s male-like genitalia, coercive sex is impossible, so female choice of mates is an important sexually selected force in this species. Mate choice by female Spotted Hyenas apparently drives males to disperse: females strongly prefer to mate with immigrants, and they appear to discriminate against adult natal males. Therefore, almost all offspring are sired by immigrant males. Immigrants queue for status within the male hierarchy of the new clan; the highest-ranking males are those that immigrated first into the clan. Malesrise in rank only when higherranking immigrants die or engage in secondary dispersal; roughly 40 % of immigrants disperse again, although the potential benefits of secondary dispersal are unknown. Clan members defend group territories from neighboring Hyena groups. Territory size ranges from roughly 20 km? ® in East Africa to approximately 1500 km? in the desert regions of southern Africa, and is negatively related to the density of available prey. Territorial behavior is exhibited by both sexes, although females engage in these activities more frequently than males. Intruders encountered within the territory are usually chased to the territory boundary. Border clashes with neighboring clans, called “ clan wars ”, are most commonly observed in habitats containing high densities of Hyenas, where intrusion pressure is most intense. Territorial behavioris rarely observed among Spotted Hyenas inhabiting the vast desert regions of southern Africa, where prey are sparse, clan size is small, intrusion pressure is low, and the home ranges of resident hyenas are enormous. In some parts of Africa, where densities of resident prey may be low but where migratory herbivores are available as prey, Spotted Hyenas are known to adopt patterns of space-use that differ strikingly from those seen in areas with year-round resident prey. Specifically, Spotted Hyenas may frequently commute long distances from their defended territory to herds of migratory prey. In the unusual “ commuting system ” exhibited by Spotted Hyenas in the Serengeti, individuals travel long distances north or south from their centrally-located clan territories in order to feed on migratory herbivores. Intruders are tolerated by territory residents when the intruders are merely passing through, although residents behave aggressively toward intruders found hunting or feeding. In Namibia, Spotted Hyenas defend territories that expand and contract in size seasonally, as migratory prey change locations. Territory boundaries are visited sporadically by multiple clan members performing border patrols, during which boundaries are marked by pasting. A strong-smelling, yellowish buttery secretion is deposited from the anal glands onto grass stalks during border patrols. Spotted Hyenas also commonly paste deep inside their territories, although the frequency with which this occurs is generally far less than in the other hyaenid species. The paste transmits information about an individual’s identity, sex, reproductive state, and clan membership. Young Hyenas engage in pasting behavior long before there is any paste in their anal sacs, suggesting that this behavior enables cubs to acquire group odors from sites where clan-mates had pasted earlier. Spotted Hyenas engage in ritualized greeting ceremonies in which two individuals stand parallel and face in opposite directions. Both individuals usually lift the hindleg nearest to the other and sniff or lick the anogenital region of the other. The unique aspect of their greetingsis the prominent role of the erect “ penis ” in animals of both sexes. This is used to signal submission. Greetings occur between hyenas of all ages and both sexes, although greetings between adult females and adult males are uncommon and restricted to high-ranking males. Cubs can erect their penis or clitoris and engage in greeting ceremonies as early as four weeks after birth. Spotted Hyenas recognize their group mates based on visual cues, odors, and individually distinctive vocalizations. These animals are well known fortheirrich vocal repertoire. They emit deep groansto call their cubs out of dens, high-pitched whines to beg for food or milk, and cattle-like lowing sounds to bring group-mates to a common state of high arousal. The sound most frequently heard during the night throughout much of sub-Saharan Africa is the long-distance vocalization of the Spotted Hyena, called a whoop. This loud call can be heard over several kilometers. Whoopsclearly serve a variety of functions. They can be rallying calls to gather scattered clan members together to defend territory boundaries, food resources, the communal den, or clan-mates in danger. Mothers whoop to locate their wandering cubs, and hungry cubs whoop to call their mothers so they can nurse. Spotted Hyenas sometimes whoop to recruit hunting partners. Whoops are also used as a form of individual display, particularly by males of high rank. Adult males whoop more frequently than females, and high-ranking males whoop more often than lower ranking males. Finally, Spotted Hyenas are well known for their laugh or giggle, which sounds much like maniacal human laughter. This vocalization is a signal of submission. A submissive individual giggles to signal to another Hyena thatit accepts a lower status.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFE9FF8D2ADAF6DDF8B1C136.taxon	breeding	Breeding. Females bear young throughout the year in most parts of Africa, although there are distinct birth peaks and troughs in some populations. Both sexes mate promiscuously with multiple partners. Courtship by male Spotted Hyenas is unusual among mammals because it appears to reflect such extreme conflicting desires to approach the female and also to flee from her. Males often engage in approach-avoid and bowing displays, both of which appear to reflect strong motivational conflict and hesitancy on the part of the male. Their behavior suggests that interactions with females may be unusually risky for males in this species, and that males fear females. In general, the female seems to take little notice of the male hyena’s sexual advances. Estrus lasts 1 - 3 days, but the length of the female's cycle, and whether ovulation is spontaneous or induced, are not known. Copulation involves multiple mounts, intromissions, and ejaculations. Female receptivity is indicated by inhibited aggression toward the male and by assumption of a distinctive receptive stance in which the female lowers her head and keeps her mouth near the ground. The only behavior indicative of a female's interest in mating is that she may follow a male. Some males who sire cubs form consortships with females, but others do not, suggesting that individual male Hyenas may adopt multiple alternative reproductive tactics to attract and acquire mates. That is, male Hyenas may sometimes “ shadow ” or “ guard ” their mates, but intensive mate-guarding is not required to ensure that a male will sire the cubs of a particular female. Females have been observed mating with one to four males within a single estrous period, and multiple paternity has been documented to occur in 25 - 30 % of twin litters. Many copulations among Crocuta appear to be infertile. Female Spotted Hyenas are exposed to high concentrations of androgens in utero, and this early androgen exposure may have negative effects on female fertility by altering ovarian histology or other mechanisms. It has recently been determined that early androgen exposure is not necessary for formation of the female's peniform clitoris. Females give birth through their penis-like clitoris. During parturition, the clitoris tears to permit the passage of the young, creating a large bleeding wound on the posterior surface that may take weeks to heal. Females usually produce litters of two, although singletons are also common, and triplets are observed occasionally. Cubs weigh roughly 1 kg at birth. They are born with their eyes open, their deciduous canine and incisor teeth fully erupted, and they are capable of remarkably coordinated movement immediately after birth. They are thus relatively precocial compared to cubs in other hyaenid species or in most other carnivores. Their coats are pure black at birth; cubs start to molt at 5 - 6 weeks of age, and the natal coat is completely replaced by an adult-colored, spotted pelage by 4 - 5 months of age. The spots never change except to fade a bit with age. Cubs are usually born in an isolated natal den and are transferred to the clan’s communal den when they are 2 - 5 weeks old. They remain at the communal den until they are 8 - 12 months old, and then begin traveling around the clan’s territory, initially with their mothers and later alone. As in the other bone-cracking hyenas, weaning occurs surprisingly late, usually around 13 - 14 months of age, but twin litters borne by low-ranking females may be nursed as long as two years. Fifty percent of cubs die before puberty, and mortality rates are generally highest immediately after weaning. Males reach reproductive maturity at around two years of age, and most females start bearing young in their third or fourth year. However, age at first parturition varies between two and six years. All females in a clan reproduce, and females rear their young together in the communal den. Therefore occupied dens may contain up to 30 young of different ages from up to 20 litters. Females nurse only their own cubs and usually reject approaches by other cubs. The milk of Spotted Hyenas has the highest protein content (mean 14 - 9 %) recorded for any terrestrial carnivore, a fat content (mean 141 %) exceeded only by that of palaearctic bears and the sea otter, and a higher gross energy density than the milk of most other terrestrial carnivores. Due to the high energy content of their milk, and the long nursing period, Spotted Hyenas have the highest energetic investment per litter of any carnivore. Reproductive success in both sexes is related to dominancestatus, although this relationship is stronger among females than males. High-ranking females enjoy greater reproductive success than low-ranking females because they have longer reproductive life spans and shorter inter-birth intervals, and because their cubs experience lower mortality than do cubs of low-ranking females. Sex ratios among adults are usually slightly female-biased. Reproductive success among males varies with intra-sexual rank, although alpha males fare more poorly than would be expected based on social status alone. As most males disperse from their natal clan when they are at least two years old, most breeding males are immigrants. Spotted Hyenas are sometimes referred to as the “ Cain and Abel ” of the animal world because of the common belief that they routinely kill their siblings shortly after birth. Although littermates do engage in aggressive interactions within minutes after birth, and although this can result in obvious scarring of the subordinate littermate, these aggressive interactions seldom result in the death of a sibling. These early fights quickly lead to the establishment of a dominance relationship that allows the dominant cub to control access to the mother’s milk. Siblicide in the Spotted Hyena is facultative in that it occurs only in some twin litters rather than routinely. The purpose of the early fighting is to establish an unambiguous dominance relationship within the litter. It appears that the relative costs and benefits of killing one’s sibling vary with current socio-ecological conditions: a cub that killsits sibling may obtain significant benefits if its mother is unable to support multiple cubs. However, mothers can usually support two cubs in many parts of Africa without undue difficulty.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFE9FF8D2ADAF6DDF8B1C136.taxon	conservation	Status and Conservation. Listed as Least Concern on The IUCN Red List. The total world population of the Spotted Hyena is well above 10,000 individuals, several subpopulations exceed 1000 individuals and its range well exceeds 20,000 km ®. The rapid decline of populations outside conservation areas due to persecution and habitat loss makes the species increasingly dependent on the continued existence of protected areas. Spotted Hyenas have been extirpated in Algeria and Lesotho, and they are listed as threatened in Benin, Burundi, Cameroon, Mauritania, Niger, Nigeria, Rwanda, and Sierra Leone. The largest remaining populations are found in Kenya, Tanzania, South Africa, and Namibia. Most adult mortality is caused directly by lions and humans, although disease is an important mortality source in some areas. Human-caused mortality is common even inside protected areas. Spotted Hyenas may attack livestock, particularly where natural prey are usually or seasonally sparse. Often in response to confirmed or assumed livestock depredation, Spotted Hyenas are shot, snared, speared, or poisoned in many parts of their range by ranchers and pastoralists. Spotted Hyenas are also commonly killed on motor-ways. Habitat fragmentation and reduction are also having significant negative effects on the size of many Spotted Hyena populations. Finally, one of the most important threats to the conservation of Spotted Hyenas is their negative public image. Many people apparently believe these animals are not worth conserving. Educating the public about these complex and fascinating animals is expected to have a substantial positive effect on conservation efforts.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEFFF8F2FA5F9F5F9C3C731.taxon	materials_examined	India.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEFFF8F2FA5F9F5F9C3C731.taxon	discussion	Of the extant hyaenids, the Striped Hyena is most closely related to the Brown Hyena, and its lifestyle, reproduction, and social behavior more closely resemble those of Brown Hyenas than those of Spotted Hyenas or Aardwolves. Some authorities provisionally recognize five subspecies, distinguished mainly by differences in size and pelage (hyaena from India, barbara from NW Africa, dubbah from NE Africa, sultana from the Arabian Peninsula, and syriaca from Syria, Asia Minor and the Caucasus). However, other authorities argue that current morphological data and other evidence do not support multiple subspecies.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEFFF8F2FA5F9F5F9C3C731.taxon	distribution	Distribution. The Striped Hyena has a very large range, covering much of Africa and western Asia. Although they do not occur in the central Sahara, these animals occur at low density in patches throughout eastern, western and northern Africa, including Algeria, Benin, Burkina Faso, Cameroon, Chad, Djibouti, Egypt, Ethiopia, Ghana, Kenya, Libya, Mali, Nigeria, Mauritania, Morocco, Niger, Senegal, Tanzania, and Tunisia. Striped Hyenas also occur in the Middle East and Central Asia. Middle Eastern and Asian countries included in the modern distribution of the Striped Hyena are Afghanistan, Armenia, Azerbaijan, Georgia, India, Iran, Iraq, Israel, Jordan, Lebanon, Nepal, Oman, Pakistan, Saudi Arabia, Tajikistan, Turkey, Turkmenistan, Uzbekistan, and Yemen. The current distribution of this species is patchy, and usually appears to occur in small, isolated populations.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEFFF8F2FA5F9F5F9C3C731.taxon	description	Descriptive notes. The Striped Hyena is the smallest of the three bone-cracking hyaenids, but is substantially larger than the Aardwolf. Head-body 100 - 115 cm, tail 30 - 40 cm, shoulder height between 66 - 75 cm; weight 26 - 41 kg (males) and 26 - 34 kg (females). Among the provisional subspecies, body mass and body size are only well studied in syriaca in Israel and dubbah in Kenya. In these populations, there was no significant sexual dimorphism in body size. However, in one recent study in Israel, sexual dimorphism accounted for 39 % of the variation in adult body size. Like the other hyaenids, the Striped Hyena has a sloping back because the forelegs are longer than the hindlegs, and has well-developed anal glands used for scent marking. Large ducts from the anal glands open into an anal pouch dorsal to the anus. As in the other bonecracking Hyenas, the head, neck, and shoulders are relatively massive and powerful. The fur is pale gray or straw-colored, with black vertical stripes on the sides of the body. Like the Brown Hyena, the Striped Hyena has longer fur than the Spotted Hyena, giving it a rather shaggy appearance. The Striped Hyena has a black muzzle and a black patch on the throat. It has five to nine vertical stripes on the flanks, two cheek stripes, and clear black transverse and horizontal stripes on all four legs. The head is roundish with a pointed muzzle and long, pointed ears. It has a gray or blond mane that runs along its dorsal midline from the ears to the tail; the mane can be erected to increase the animal’s apparentsize by over 30 %. The mane in this species is more pronounced than that in any other hyaenid, with hairs up to 20 cm long. The black and white tail is long and bushy, with hair that is generally coarse and long. Females have two or three pairs of teats, but if they have three, only the caudal two pairs are functional. Juvenile females have well-defined labia-like folds anterior to the vagina. These ridges are hairless and darker and rougher than the surrounding tissue. Juvenile males have smaller, smooth, hairless skin folds along the middle septum close to, but anterior to, the scrotum. Unlike Spotted Hyenas, these genital characteristics are not severe enough to confuse sexing of juveniles, and adult genitalia appear normal. Subspecies descriptions are based on limited data except for syriaca in Israel and dubbah in Kenya. In general animals living in the northern parts of the range tend to be slightly larger than those living in southern regions. Variation in pelage color appearsslight, although the Lebanese population is reported to have a reddish coat color, and hyenas on the Arabian Peninsula are described as having a yellow mark below the eyes and a mixed gray and black dorsal crest.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEFFF8F2FA5F9F5F9C3C731.taxon	biology_ecology	Habitat. In most ofits range the Striped Hyena occurs in rugged, arid habitat or light thorn bush country. These animals drink regularly where wateris available, but they can also survive in many waterless areas. In North Africa they prefer open woodlands and bushy and mountainous regions. The central Arabian and Sahara Deserts are not suitable habitat. In central Asia, Striped Hyenas avoid high altitudes and dense thickets and forests. The maximum elevations recorded are 2250 m in Iran, 2500 m in India and 3300 m in Pakistan. In the Caucasus region, Turkmenistan, Tadzhikistan, and Uzbekistan, prime habitats include savannah and semi-desert regions up to an elevation of 2100 m, mountain areas with strong relief, valleys with abundant caves or other resting sites, and riverine areas. The Striped Hyena avoids areas with minimum temperatures ofless than — 15 ° C to — 20 ° C and more than 80 - 120 days offrost per year. In Israel, Striped Hyenas are present even close to dense human settlements. In West Africa, they occur in dry scrub savanna and Sahel woodland, particularly in the belt of Acacia raddiana woodland that extends from Senegal to Chad. In eastern Africa, Striped Hyenas are found in a variety of habitats ranging from open savanna to rugged, bush-covered mountain terrain.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEFFF8F2FA5F9F5F9C3C731.taxon	food_feeding	Food and Feeding. The diet of Striped Hyenas apparently varies considerably from one part of their range to another, but these animals are clearly scavengers with catholic tastes. They are primarily carrion-eaters; their diet consists mainly of dried flesh and bones from carcasses of large vertebrates. They scavenge carrion and the remains of kills made by other predators, including Spotted Hyenas, Cheetahs, Leopards, Lions, and Tigers. The Hyena’s massive cheek teeth and supporting musculature easily permit the gnawing and breaking of bones, as well as the carapaces of tortoises and turtles. Striped Hyenas have also been reported to consume a wide variety of invertebrates, vegetables, fruit, garbage, and small vertebrates that the Hyenas hunt themselves. In central Kenya, analysis of bone fragments and hairs from fecal samples indicated that hyenas regularly consume small mammals and birds that are unlikely to be scavenged. The limited available diet data may underestimate the importance of active hunting in the lives of these animals. In various parts of eastern Africa, Striped Hyenas are reported to supplement their diet with Balanites fruits. The proportion of scavenged and killed prey items in the diet is still a matter of debate as there has been no detailed research on these Hyenas’ food intake. Some authors suggest that only individuals from the three larger subspecies, barbara, syriaca and hyaena, kill large prey, including livestock, as there is no evidence that the smaller subspecies, dubbah and sultana, attack large herbivores. In Turkmenistan the Striped Hyena has been reported feeding on Wild Boar, Kulan, porcupine, and particularly tortoises. In Uzbekhistan and Tadzhikistan, seasonal abundance of oil willow fruits (Eleagnus angustifolia) is reflected in the diet; in the Caucasus region the diet includes abundant grasshoppers. In Israel the Striped Hyena feeds on garbage, carrion, and fruits, particularly dates and melons. In eastern Jordan, the main sources of food are carcasses of feral horses and water buffalo and refuse from localvillages. The Striped Hyena can drink water of very variable quality, from fresh water to soda and salt water, but it may also satisfy its water requirements with melons or otherfruits. Very little is currently known about the hunting behavior of Striped Hyenas, but those few hunts that have been observed involved simple chases and grabs at prey. Seasonal influxes of Striped Hyenas follow migrations of large herds of domestic and wild ungulates in Turkmenistan, suggesting that the Hyenas cover long distances on foraging trips. In Egypt they are known to move along ancient caravan roads where the chance of locating dead camels is high. In Serengeti and in southern Kenya, they spend most of the night actively searching for food or moving between established foraging sites. Striped Hyenas apparently can remember the locations offruiting trees, garbage dumps and other established feeding sites, although the routes taken to re-visit such food sources are seldom repeated on consecutive foraging trips. They are also able to locate tortoises in their hiding places during periods of aestivation and hibernation. Striped Hyenas frequently cache bones or pieces of skin, using their snouts to push these items deep into clumps of grass or stands of dense shrubs. They may also carry food items back to their dens. Bone collections are common at den sites used by Striped Hyenas, although it is often unclear whether these collections represent scavenged or killed prey, and whether the bones collected play a significant nutritional role in the lives of these animals. Several studies have inferred diet by combining data from bone collections and fecal samples. In central Kenya, however, bone collections indicated a much broader range of prey than did scat analysis, and significant portions of bone assemblages were very old bones unlikely to represent material scavenged from fresh kills. From fecal analysis alone, several researchers have found remains of prey items that are more likely to have been scavenged than hunted, and larger mammals are represented far less often in the analysis of hairs in fecal material than would be expected based only on bone collections at dens. Striped Hyenas appear to be strictly solitary foragers, although multiple individuals occasionally gather at rich food sources such as large carcasses or refuse pits. These animals are sometimes found in small groups while resting, but there is no indication that they ever forage cooperatively. Genetic relatedness among members of groups seen feeding together has not been investigated. Foraging activity in Kenya and Tanzania is restricted to hours of darkness except during rain or unusually cloudy weather. Under those circumstances, Striped Hyenas may return to previously visited kills or carcasses, but do not embark on full foraging forays. In many areas, and for many centuries, Striped Hyenas have been described as raiding human graves and carrying away bones. Fruit and vegetable crop raiding by Striped Hyenas is currently considered a serious problem in some parts of Israel. While foraging, Striped Hyenas zigzag across the landscape and do not appear to follow set routes, even when returning to the same food source on multiple nights. Minimum mean distance travelled per night is 19 km at speeds of 2 - 4 km / h, occasionally trotting at speeds of up to 8 km / h, or running at a maximum of 50 km / h. Overall, the evidence indicates that Striped Hyenas are solitary foragers for which carrion, insects, fruits and vegetable matter represent significant portions of the diet.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEFFF8F2FA5F9F5F9C3C731.taxon	activity	Activity patterns. The Striped Hyena is almost strictly nocturnal, although it does occasionally engage in some activity after dawn and before dusk. Some authorities suggest Striped Hyenas may be moststrictly nocturnal in areas characterized by relatively intensive anthropogenic activity, and where they are directly persecuted by humans.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEFFF8F2FA5F9F5F9C3C731.taxon	biology_ecology	Movements, Home range and Social organization. In Serengeti, Striped Hyenas travel an average of 19 km per night (range 7 - 27 km), either following established animal tracks or zig-zagging cross-country. A similar pattern was observed in southern Kenya, where Striped Hyenas followed by human observers covered large distances, but stopped frequently to paste or to investigate grass clumps, carcasses, and other things found on the ground along the way. Home range sizes of one female and one male in the Serengeti were 44 km * and 72 km? respectively. There was little evidence of territorial behavior. Home range size was calculated for a single female in the Negev Desert in Israel to be approximately 61 km?; this range partly overlapped those of two other individuals. In the Laikipia District of Kenya, the mean home rangesize for 12 males was 82 km?, and for eight females was 71 km? with no significant difference in home range sizes between sexes. No evidence ofterritorial defense has been recorded in any studied population, but in some populations these hyenas are known to scent-mark frequently within their home ranges while traveling, and also to defecate in “ latrines ” near feeding sites and along travel routes. Striped Hyenas are the least well-studied of the extant hyaenids, and their social behavior is very poorly understood. They are most often reported to be solitary. Nevertheless, there appears to be considerable variability with respect to patterns of social grouping among Striped Hyena populations. In some areas, such as central Asia, these animals are reported to form short-term monogamous pair bonds for breeding, with a resulting family unit that may endure for several years. Such family units may sometimes contain offspring from multiple litters. Under these circumstances, both parents and the older offspring may be observed provisioning den-dwelling cubs. Typical group sizes are one or two in all subspecies, but groups of up to seven have been reported in Libya. In Israel, Striped Hyenas are generally solitary, but occasionally several are seen together at a carcass, including both males and females, or females with large cubs. Age-specific foraging data are extremely limited, but cubs have occasionally been observed accompanying their mothers on foraging trips away from the den by 6 - 12 months of age. Otherwise, foraging is strictly solitary. Almost invariably described as solitary in sub-Saharan Africa, it was recently discovered that Striped Hyenas in the Laikipia District of Kenya are behaviorally solitary but exhibit a polyandrous system of space use. These animals form groups of up to four adults; each group contains one adult female and one to three adult males. Females in both wild and captive populations appear to be highly intolerant of one another, starting around the time they reach puberty. The overall adult sex ratio in the Laikipia population was three males to two females. Members of these groups share a common home range and may be found resting together during daylight hours. The home ranges of group-mates exhibit 85 % overlap, whereas their ranges overlap only 22 % with those of animals in other groups. Individual group members spend more than 90 % of their time alone. Adult male group-mates included both closely-related and distantly related individuals. In contrast to spatial patterns of relatedness documented in other carnivores, pairs in non-adjacent groups tended to be more closely related genetically than pairs living in adjacent groups. This was true for females as well as males. As these animals are usually found alone, very little has been recorded regarding direct social interactions except for captive situations. In this species, males are slightly larger than females, and males also appear to be socially dominant to females in resource competition. The long dorsal hairs of the mane may be erected to enhance the apparentsize of the individual during confrontations with conspecifics. Both mane and tail hairs are erected when the animal assumes a defensive posture, but also when it adopts an aggressive stance. The mane is also commonly bristled whenever the animal pastes. When Striped Hyenas fight they bite at the throat and legs, rather than at the mane. During an agonistic interaction, the subordinate individual may hunch its body, lower its mane, and swing or turn its head from side to side whereas the dominant animal remains bristled and stands erect. The Striped Hyena exhibits a number of visual displays, the moststriking of which is the erection of the mane and bristling ofthe tail like a bottle brush. The mane and tail thus serve as signalling devices during social interactions. When members of the same social group meet after being separated, they engage in “ meeting ceremonies ”, which involve investigation and licking of the mid-back region and sniffing of the nose and extruded anal pouch. The tail is often held vertically during meeting ceremonies. Meeting ceremonies may also involve repeated pawing of the throat of the greeting partner. The well-developed anal pouch is inverted during scent marking, called pasting, and also during greetings. In scent marking, the anal pouch produces a pungent yellow to beige paste which is deposited at nose-height on grass stalks, stones, tree-trunks, or sticks. Foraging Striped Hyenas pause to paste at frequent intervals, and these scent marks appear to be deposited throughout the home range rather than exclusively at its borders. Pasting has also been observed at large carcasses in the wild, and, in captivity, on food bowls. Some Striped Hyena vocalizations resemble those of Spotted Hyenas, although calls emitted by Striped Hyenas tend to be much softer, and the sounds carry shorter distances. The vocal repertoire of the Spotted Hyenais also far more elaborate than that of the Striped Hyena. Most ofits vocalizations are uttered in the presence of conspecifics. Cubs whine while they are nursing. Giggling or yelling may occur when a Striped Hyena is frightened or being chased by another predator. A long, drawn-out lowing sound sometimes accompanies the defensive posture, and growling may occur during fighting or play-fighting with conspecifics.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEFFF8F2FA5F9F5F9C3C731.taxon	breeding	Breeding. Females are polyestrous and breed throughout the year. Estrus is reported to last one day. No detailed descriptions of sexual behavior in the wild have been reported, but during matings in captivity, females may mate several times at intervals of at least 15 minutes. In the wild, litter size varies from one to four (median of three), after a gestation period of 90 - 91 days. Average litter size in captivity is 2 - 4, with a range of one to five. Parturition is preceded by intensive digging behavior by the female. Cubs weigh approximately 700 g at birth; they have adult-like markings but lack manes, and instead have only black spinal stripes. They are born with eyes and ears closed, and they are barely able to crawl, so they are far more altricial than Spotted Hyena cubs at birth. Their eyesfirst open after five to nine days, and cubs may emerge from the den at around two weeks of age. Deciduous teeth start to erupt on day 21. Cubs begin to eat meat at the age of 30 days. In the wild cubs are known to nurse for over one year. They reach reproductive maturity during the second year oflife. The mating system is promiscuous or polyandrous. In the Laikipia population in Kenya, females appear to mate with both group males and males that reside elsewhere. It is not known whether sires contribute in any way to parental care in this population, but lactating females are usually found alone at dens with their cubs, males do not spend significant periods of time at dens, and females appear to be solely responsible for care of young. Multiple paternity in this population occurred in half of sampled litters, and extra-group males sired roughly one third of the cubs born to group females. Striped Hyenas usually use caves, ravines or other sheltered rocky places as dens, although earthen dens may also be used. Den entrances are fairly narrow and may be hidden by large boulders. Two dens were measured in the Karakum Desert. The entrances were 0 - 67 m and 0 - 72 m wide. The dens sloped downward 3 m and 2 bm and were 4 - 15 m and 5 m long, with no lateral extensions or special chambers. These simple constructions contrast with much more elaborate dens found in Israel, which can exceed 27 m in length.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEFFF8F2FA5F9F5F9C3C731.taxon	conservation	Status and Conservation. Listed as Near Threatened on The IUCN Red List. Despite their broad distribution, the basic biology of Striped Hyenas, including their abundance in most parts of their range, remains very poorly known. Throughout its range, the Striped Hyena occurs at low densities. There have been only two local estimates of Striped Hyena density in Africa, and it is considered either threatened or data deficient throughout its African range. In Serengeti and Laikipia, density was estimated to be 0 - 02 per km? and 0 - 03 adults per km?, respectively. Remarkably little information is available on the species. This is undoubtedly due to its shy, nocturnal, mostly solitary nature, its apparent affinity for rugged terrain, its generally negative reputation, and frequent confusion with, or lack of differentiation from, Spotted Hyenas where the two species overlap. Most adult mortality is directly caused by Lions and humans. Striped Hyenas, particularly those inhabiting areas where natural prey are usually or seasonally sparse, may attack livestock, and as a result they are shot, snared, speared, or poisoned in many parts of their range by ranchers and pastoralists. It appears that the Striped Hyena is already extinct in many localities, and that populations are generally declining. The major reasonsfor this decline appear to be decreasing natural and domestic sources of carrion due to declines in the populations of other large carnivores and their prey, and changes in livestock practices. Moreover, the low densities and associated large home ranges of these animals are likely to increase the chances that populations will become fragmented into small, non-viable units. This must be considered a key problem if these animals are to be protected. The Striped Hyena evokes many superstitious fears because of putative and documented cases of grave-robbing and attacks on humans. In addition, its body parts are widely exploited as aphrodisiacs, and are utilized in folk medicine. Striped Hyenas are often killed because of suspected or real damage inflicted on agricultural produce and livestock, and they are often shot by livestock ranchers. These animals have also been widely hunted through poisoning, baiting traps, pits, or with the help of dogs. The Striped Hyena does sometimes cause damage to crops, and may sometimes also attack domestic animals, predominantly goats, sheep, dogs, and poultry. In many cases of damage to livestock, it is unclear whether the targeted individual was adult or young, healthy or sick, so Striped Hyenas may be blamed for livestock mortality for which they are not truly responsible. In any case, the records suggest that attacks on livestock by Striped Hyenas usually occur at very low frequencies. Tentative estimates of the total worldwide population size range from 5000 to 14,000 individuals. Fragmentation into many subpopulations is suspected even though the actual degree of fragmentation is unknown. In addition, habitat loss and declining population size are occurring at unknown rates. The Striped Hyena is considered threatened in all parts of its African range where data are sufficient to evaluate the local status, except in Ethiopia and Kenya, whereit is considered at lowerrisk. As we currently know so little about the biology ofthis species, one of the most pressing conservation concerns is to study these animals in a number of different locales. In addition to knowing very little about their behavioral ecology, we also know virtually nothing about their abundance and population dynamics. Before we can make management decisions in the best interest of Striped Hyenas, we need more information. Fortunately, studies of these animals are currently underway in East Africa, India, and the Middle East.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEDFF812FA7FBEBF8DECE95.taxon	materials_examined	South Africa, Western Cape Province, Cape of Good Hope.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEDFF812FA7FBEBF8DECE95.taxon	distribution	Distribution. Namibia, Botswana, W & S Zimbabwe, S Mozambique, Swaziland, W Lesotho, and South Africa. Records from the SW of Angola are all before 1970.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEDFF812FA7FBEBF8DECE95.taxon	description	Descriptive notes. Head-body 110 - 136 cm; average 123 cm (males), 117 cm (females), tail 18.7 - 26. 5 cm, average height at shoulders 79 cm (males) and 74 cm (females); weight (adult) varies somewhat regionally, ranges from 28 to 47 - 5 kg and averages about 40 kg. Most studies show some sexual size dimorphism, but it is often minimal, with males slightly heavier and longer than females. Has a typical hyena appearance with front legs longer and more robust than the rear legs, a broad head and short muzzle, thick neck and short tail. Like the Striped Hyena it has large pointed ears and course shaggy fur that is longest along the back and on the tail. The general color is dark brown with lighter tawny hair on the neck and shoulders. The legs are banded with dark horizontalstripes and the front feet are large and well-developed for digging. Like the Spotted and Striped Hyenas, the Brown Hyena possesses the bonecrushing third premolar that is unique to this family. In contrast to Spotted Hyenas, there is no masculinization of the female genitalia in Brown Hyenas. Females have two to six pairs ofteats, but only the two most caudal pairs are functional.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEDFF812FA7FBEBF8DECE95.taxon	biology_ecology	Habitat. Brown Hyenas are found in a variety of relatively arid habitats from open desert or semi-desert in the Namib and Kalahari, to dry open scrub and woodland savannah, mopani scrub and tree savannah, as well as the bushveld of the northern Transvaal. They do not need drinking water and inhabit areas where annual rainfall may be even lower than 100 mm, up to about 650 mm.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEDFF812FA7FBEBF8DECE95.taxon	food_feeding	Food and Feeding. These hyenas forage alone at night and do not cooperate in hunting or in feeding, although group members tolerate each other at large food items. Although not competent hunters, Brown Hyenas are extremely efficient scavengers with an omnivorous diet. They are opportunistic feeders on a range of vertebrates, primarily mammals, the vast majority of which are scavenged, often from the kills of other carnivores. Fruits, insects and reptiles can be important supplementary foods when carcasses are rare. In one population 58 different food items were identified from fecal analysis. Brown Hyenas in the southern Kalahari spent 30 % of their feeding time eating carrion, 28 % on vegetable matter, 4 - 5 % on small mammals, 1 - 5 % on birds’ eggs, and 29 % on unknown items. Only 5 - 8 % of food they were seen eating was killed by the hyenas themselves. In the central Kalahari 35 - 9 % ofall observed feeding bouts were on fresh scavenged kills, 33 - 9 % on old carcasses, 16 % on their own kills, and 12: 5 % on vegetative food sources. Brown Hyenas do not depend on standing water. In the central Kalahari, no free water orrain is typically found for eight months of the year. Although they will drink on a daily basis when water is present, much of the water during dry seasons is obtained from Cucurbitaceae fruits such as the tsama melon, gemsbok cucumber, and Hookeri melon, which can compose significant proportions of the diet in these seasons. Brown Hyenas inhabiting the Namibian coast feed almost exclusively on subadult Brown Fur Seals. The majority of these seals are thought to be scavenged, although Brown Hyenas have been seen hunting seal pups. In fact, hunting efficiency on seal pups during the peak pupping season can be as high as 47 %, and an average of almost five seals per day may be killed from a single colony. Small rodents and seabirds make up the rest of this unique diet. Elsewhere hunting attempts by Brown Hyenas are opportunistic and directed at small mammals such as Springbok lambs, spring hares, Bat-eared Foxes, and ground nesting birds. The hunting technique of the Brown Hyena is unspecialized, rarely successful (except in the case of Brown Fur Seal pups), and may include a brief lunge at a surprised prey, a prolonged chase of up 1 km, or an attempt to dig up a burrowed animal. The percent of observed hunting attempts that were successful was 4: 7 % in the southern Kalahari and 13 - 7 % in the central Kalahari. The hyenas generally feed where they find food, but food from larger carcasses is frequently cached nearby in a clump of grass or under a bush. Considerable time is often spent at carcasses removing limbs for this purpose and one animal may remove and cache up to three legs before any competitors arrive. Sites where food is cached are scent-marked and may be re-visited over multiple days. In a remarkable example of food caching in the southern Kalahari, a Brown Hyena arrived at an abandoned ostrich (Struthio camelus) nest with 26 eggs, which are prized food items. The hyena spent four hours carrying 14 eggs distances of 150 - 600 m from the nest, some of which it simply dropped in the open. It ate only three eggs during this period. Brown Hyenas also carry food back to cubs at the den. This provisioning of cubs can result in significant bone accumulations at densites. In the southern Kalahari the average distance moved between significant meals was 7 - 2 km, and the average nightly distance traveled was 31 - 1 km. During the dry season, nightly movements were longer and were recorded as high as 54 - 4 km. During the wet season movements were reduced and ranged from 10 - 20 km. In Namibia, daily distances traveled ranged from 15 - 47 km. When foraging, Brown Hyenas move at a pace of about 4 km / h, often walking in a zig-zag pattern, probably to maximize their chance of coming across food items. They use smell to locate much of their food, as evidenced by frequent sniffing and moving upwind toward food sources. Their hearing appears to be acute as well and is likely also used in foraging. The Brown Hyena is subordinate at kills to Spotted Hyenas, Lions and African Wild Dogs, although it appears to dominate Leopards in most situations. While they always dominate Cheetahs, they compete heavily with the much smaller Black-backed Jackal, which is often able to steal scraps from hyenas at carcasses. There is some evidence that, where they are sympatric, Brown Hyenas avoid areas frequented by Spotted Hyenas, potentially to avoid direct aggression and competition. For example, dens in the southern Kalahari were rarely found in the prey-rich riverine habitat where most Spotted Hyena dens occurred. Although their presence at livestock carcasses has resulted in much antagonism toward, and persecution of, Brown Hyenas by livestock owners, predation on livestock by these animals appears to be done by a small number of individuals. However, these hyenas, which typically target cow calves and sheep, can account for a large number of kills. Removal of these problem individuals appears effective at halting stock losses.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEDFF812FA7FBEBF8DECE95.taxon	activity	Activity patterns. Primarily a nocturnal animal, although activity is occasionally observed during the day, particularly on cool, cloudy days during the rainy season. There are typcally two peaks of activity, from 19: 30 h to 24: 00 h and 2: 30 h to 6: 00 h, with a rest period in between. Radio-collared adults in the southern Kalahari were active for 42.6 % of the 24 h period, and 80.2 % of the period between 18: 00 h and 06: 00 h. In Namibia, three males with satellite collars spent an average of 57 - 1 - 72 - 3 % of 24 h active. Brown Hyenas typically rest during the day in a hole, or undera large tree or bush.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEDFF812FA7FBEBF8DECE95.taxon	biology_ecology	Movements, Home range and Social organization. Approximately 65 % of Brown Hyenas in a population are members of small social groups called clans, with the remaining individuals living as nomads. Clan size ranges from 4 - 14 individuals, including cubs, and clans defend large stable territories. In the southern Kalahari far-ranging nomadic males (8 % of the adult population) were the only males observed to breed with clan-living females, yet in the central Kalahari, breeding also occurs with resident immigrant males. In the southern Kalahari territories averaged 308 km? (range: 215 - 461 km?) with never more than 20 % overlap between territories. Brown Hyena density there was calculated to be 1 - 8 hyenas / 100 km? ®. In the central Kalahari territories averaged 170 km?, but varied greatly with annualrainfall, reaching a maximum of 400 km?. In Namibia, where Brown Hyenas depend almost entirely on Brown Fur Seals along the coast, territories of two clans in one study were 31 - 9 km? and 220 km? In another study the home ranges of three males ranged from 420 - 1460 km? with the largest home range being an inland location. Density in this latter Namibian study ranged from 1 - 0 - 2 - 9 / 100 km ®. In general, group size appears to be correlated with food abundance and quality within the territory, whereas territory size is influenced by the distribution of food resources. Territories are maintained primarily through scent marking behavior (called “ pasting ”) and aggression toward intruders. Clan structure appears to vary across regions, but always includes 1 - 5 breeding females and their subadult offspring. In the central Kalahari, groups often also include at least one adult resident immigrant male. Mean clan size in the southern Kalahari was 3 - 7 adults and subadults, and total clan size ranged from 4 - 14. In the central Kalahari, a well-studied clan contained 13 members including cubs. Because adult females and their offspring are the core of a social group, the majority of clan members are related. However, dispersal from and immigration into the clan occurs. Although subadults of both sexes may disperse from their natal clan, males do so more often than females and most males disperse by 36 - 40 months of age. In two reported cases of female emigration in the central Kalahari, the number of resident adult females was atits zenith (five) and the dispersing females both were targets of severe aggression from other resident females prior to dispersal. In both cases, dispersal appeared to be prompted by conflicts with established adults of the same sex. The central Kalahari and southern Kalahari locales also apparently differ with respect to clan social hierarchies. In the central Kalahari a linear within-sex dominance hierarchy was apparent, and at carcasses with more than one hyena, rank determined priority of access to food. Although immigrant males were dominantto all natal males, the highest ranking male and female appeared to be of equal status. Adult females were typically dominant to natal males of less than 36 months of age. These natal males were tolerated until about 24 months, when aggression gradually increased until their dispersal. However, in one case, a natal male remained in the clan and eventually dominated the clan females. In the southern Kalahari no dominance hierarchy was apparent, with no sex, age-class, or individual consistently winning fights or monopolizing food resources in clans. Differences in the breeding systems and the existence or lack of a hierarchy are thought to be related to significant differences in Brown Hyena density in the two locales. In the central Kalahari, 37 - 81 % of observations involve the association of two or more hyenas. In the southern Kalahari contact between group members appears to be less frequent. Although there is typically aggression between hyenas of the same sex from different groups when they meet, the level of aggression within clans appears to vary between the southern and central Kalahari. In the central Kalahari, neck-biting appears to be used to maintain rank relationships within the clan and is observed with some frequency, while in the southern Kalahari, fighting within the clan is rare, with clan members seldom interacting at all. Here, the only aggression observed is between same-sex members of neighboring clans, and this is extremely infrequent. Interestingly, in the southern Kalahari, where resident males do not breed with clan females, these males show little aggression to nomadic males, who are responsible for mating with groupliving females, suggesting that Brown Hyenas can differentiate between neighboring males and nomadic males. When they meet after being separated, Brown Hyenas from the same group engage in a greeting ceremony in which each animal in turn crouches and presents its extruded anal pouch to the other. This is accompanied by a lowering of the ears and a “ grin ” (teeth exposed by pulling lips up and corners of the mouth back) by the subordinate animal whenit is greeting a dominant. Greetings can last as long as five minutes. Two additional behaviors that appear to be important in Brown Hyena society are neck-biting and muzzle-wrestling. Neck-biting is a purely agonistic interaction (though cubs may engage in it during play) and is primarily intrasexual. In the southern Kalahari, this behavioris largely restricted to interactions between members of neighboring clans, whereas in the central Kalahari, it can be seen more frequently between clan members and is thought to function in maintenance of a dominance hierarchy. Neck-biting behavioris a ritualized, somewhat elaborate interaction in which dominant and subordinate animals are clear from the start. The submissive animal approaches a standing dominant individual grinning and with its mane and tail raised. Either before or at its approach, the dominant seizes the neck of the subordinate, holding the skin and hair of the neck with its incisors and one or both canines, and vigorously shakes the victim from side to side. This type of interaction typically lasts less than five minutes, and only rarely does the subordinate flee at its conclusion. Muzzle wresting may be observed anywhere in the territory and is exhibited by all clan members. However, adults rarely engage in this behavior with other adults, though they will do so with cubs with some frequency. Most muzzle-wrestling occurs between cubs and subadults. The two participants stand face to face, and attemptto bite each other on the jowls or along the side of face. Their heads pitch rapidly from side to side with mouths open, and they often growl softly throughout. One or both hyenas may be crouched on their carpals, and in some cases one may lie beneath the other. This behavior is clearly less aggressive than neck biting and may often be play, although it can escalate into true aggression. There is no clear loser, and animals typically remain with each other after muzzle-wrestling, which may last from a few minutes to an hour. The most striking visual display of the Brown Hyena is pilo-erection of the long hairs along its neck and back, which is observed in situations calling for either an attack or flight response. Despite its rather elaborate social interactions, Brown Hyenas spend the vast majority of their time alone, and the primary form of communication between hyenas is olfactory. They convey information to conspecifics with latrines, which have accumulations of feces, and grass stalks on which they have deposited a strong-smelling white secretion and a smaller black secretion. Both secretions are deposited during pasting from an extruded anal scent pouch located between the rectum and base ofthe tail. Although all four hyaenid species paste, the deposition of two different secretionsis unique to Brown Hyenas. Whereas the lipid-rich white secretion is discernible to the human nose for well over 30 days, the more watery black secretion appears unscented after a few hours. This black secretion is thought to convey information relating to the time elapsed since it was deposited, and therefore signal that a hyena has recently foraged in the area. It is suggested that this allows other group members to avoid unproductive areas, and minimizes competition between group members for limited resources. The longer-lasting white secretion is thought to function in territory marking and defense. Pasting is done throughout a clan’s territory. Although most pasting occurs in the central part of the territory, where residents spend most of their time, frequency of pasting and over-pasting (deposition on an existing mark) is highest when individuals visit territory boundaries. Very little pasting is done by residents when they are outside of their territory. Pasting during traveling / foraging movements can be quite frequent, with ten individuals averaging a paste every six minutes. However, this is highly variable, with some individuals pasting only once or twice during a night-time observation period. Males and females do not differ in rate of pasting during their travels. At the den, adults and subadults frequently paste soon after arriving and before departing. At least in the southern Kalahari, the perimeter of hyena territories is thought to be too large to make strict border marking possible or effective. Instead, marks are scattered throughout the residents’ territory. This is known as hinterland marking. Given the frequency of pasting, and how widely pastes are deposited across a territory, simulations indicate that in the southern Kalahari, hinterland marking is effective. Intruders would likely encounter resident paste marks very soon after entering a territory. Indeed, individual hyenas are estimated to deposit some 29,000 paste marks in a year. Experiments with translocated pasted grass stalks indicate that hyenas can distinguish between pastes of group and non-group members and that over-pasting is more commonly done on pastings from non-group hyenas. Chemical analysis of white and black paste suggests that the scent of these substances probably varies between individuals, allowing for identification of the paster. In the southern Kalahari, latrines are not as regularly spaced as pastings, and show a clumped distribution, largely around primary foraging areas that occur along the territory border. Latrines are often associated with landmarks such as trees, bushes or roads, and those along the border are visited more frequently than those in the interior. The vocal repertoire of the Brown Hyena is relatively small, as in the Striped Hyena, and consists of eight vocalizations: a yell, a hoot, two whines and four growls, none of which functions as a longdistance communication. Some authors group these into five calls, the squeal / whine, squeak, scream, yell, and growl / grunt. The squeal is a shrill sharp cry emitted by a juvenile or other subordinate while approaching to greet or beg food from a dominant individual. The squeak is a hoarse rasping cry of abject submission associated with carpal crawling. A scream is a high-pitched, cackling shriek given by a hyena whose neck is being bitten. The yell is a loud, abrupt high-pitched call associated with defensive threat. A growl / grunt is low-pitched, breathless and throaty, and is given while muzzle-wrestling. All but the growl appear to indicate submission or appeasement in social contexts of varying intensity.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEDFF812FA7FBEBF8DECE95.taxon	breeding	Breeding. Brown Hyenas are polyestrous, non-seasonal breeders. Litters range from 1 - 4 cubs with a modallitter size of three. Estrus lasts approximately one week but mating in captivity occurs over a 15 day period. Based on six observed mating bouts in the southern Kalahari, mating associations consist of multiple copulation attempts over a 5 - 90 minute period, and may be preceded by extended courtship, during which both animals may show aggression and there are mutual approaches and retreats. Gestation in captivity was 96 days. Interbirth intervals appear to range widely. In the southern Kalahari they were as short as a year and as long as 41 months apart, although lost litters in the interim could not be ruled out. Cubs are born with their eyes closed and their ears bent forward. Their fur is similar in color to that of adults. Their eyes begin to open at eight days and are completely open at 14 days. Unlike Spotted Hyenas, Brown Hyena cubs are born without teeth. As in the other bone-cracking hyaenids, den dependenceis long and weaning occurs late. Cubs from 0 - 3 months of age rarely leave the den hole except when their mother or another adult is present. During this period, mothers attend the den frequently, often at sunrise and sunset and cubs rely completely on their mother’s milk. At four months, visits by the mother become less frequent, with mothers visiting about once a night, but suckling periods are longer. At this time, mothers and other group members begin bringing food to the den for the cubs. Weaning normally occurs at 12 - 16 months of age, yet weaning conflicts have been observed at ten months. Starting at ten months, cubs begin extensive, and very often solitary, foraging movements away from the den. Length of den residence is variable however, ranging from 8 - 15 months. Regardless of their dependence on the den, by 16 months weaning has occurred and full adult dentition is present. As they mature, subadults themselves begin to bring food back to the den for younger cubs. This has been observed in subadults as young as 22 months. Adult size is reached at 30 months. The earliest breeding recorded in the wild is 35 months; breeding continues until at least ten years of age. Cubs are raised in underground, sometimes extensive, tunnels, always small enough to prevent adults and potential predators from entering. They are easily distinguished by accumulations of bones, hair, feathers, horns, pieces of hide, and hyena feces. In the southern Kalahari Brown Hyena dens appear to be used only rarely by multiple females at once, and cubs are typically raised in the same den in which they are born. However, communal denning appears to be common in the central Kalahari, with cubs of multiple females, and of different ages, raised together at a single den location. In this system, cubs are born in a solitary den and transported to the communal den sometime before they are four months of age. Throughout their development, den moves are common and cubs may reside at as many as seven different dens, though distances between dens are typically not large. In the central Kalahari, where a social rank system is evident, dominant females enjoy greater reproductive success in terms of number of surviving offspring, yet the number oflitters does not vary based on rank. There appears to be variation, both regionally and temporally, in the mating system of Brown Hyenas. In one system, females breed only with nomadic males that range over wide areas without defended territories or family groups. In the other, females breed not only with nomadic males but with resident immigrant males as well. These residents are members of the clan and assist in territory defense, yet their tenure, at least in the central Kalahari, is relatively short (less than three years). In this area, where both nomadic males and resident immigrant males are present, dominant clan males were observed to copulate with resident females more frequently than nomadic males. In the southern Kalahari, the only mating observed involved nomadic males, and no resident males, either immigrant or natal, were observed to mate. Although males known to be natal showed little sexual interest at all in any resident females, researchers were unable to observe immigrant males long enough to ascertain sexual interest in resident females. The source of the variation in the mating system of Brown Hyenas is unclear, although it may be related to dispersion of food. In the central Kalahari, where both systems were observed over time, mating with nomadic males was restricted to the dry season, when resident clan males would likely have difficulty maintaining contact with clan females (some were separated by 22 km). Because territories are very large in the southern Kalahari and only breeding with nomadic malesis seen, the food dispersion theory is further supported. Individual reproductive patterns in males are also likely to be influenced by individual status and the behavior of other males in the population. In either case, natal males are never observed to mate with females in their natal clan. Communal care of cubs is better developed among Brown Hyenas than in any of the other three hyaenid species. Non-parental aid in cub rearing includes communal suckling (although preference in nursing one’s own cubs has been shown), food provisioning, den maintenance, defense against predators, play, and adoption of orphans. Subadult and adult females ofall social ranks and reproductive states bring food items to the den for cubs. However, the extent of involvement in provisioning by adult males seems to vary by region. In the central Kalahari, subadult males provision cubs, but to a lesser degree than females, and they only bring food to closely related cubs. Neither immigrant nor natal adult males were seen provisioning cubs. It has been suggested thatthis is because males do not benefit from an increased group size, as they are likely to emigrate from their natal clan. In the southern Kalahari, however, group-living males and females, both adult and subadult, were observed to provision cubs regularly. Average distance in this population from which food was carried back to the den was 6 - 4 km.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
03928788FFEDFF812FA7FBEBF8DECE95.taxon	conservation	Status and Conservation. Listed as Near Threatened on The IUCN Red List. In 1994 the species was down-listed from Appendix I status, which was afforded the species in 1975, to Appendix II by the IUCN and it has since been deleted from CITES listing altogether. [tis generally considered to be widespread yet rare. The total population is estimated to be 5000 to 8000, but this may be an underestimate due to the secretive nature and nocturnal habits of this animal. It is estimated that areas in excess of 1000 km? ” are required to maintain a viable population of Brown Hyenas. These populations currently exist in the Kalahari Gemsbok National Park, South Africa and the adjacent Gemsbok National Park, Botswana, the Central Kalahari Game Reserve, Botswana, and the coastal regions of the southern Namib Desert. These are also the sites of the primary research projects that provide much of what we know about this species in the wild. Much of the habitat where Brown Hyenas occur outside protected areas is used for livestock ranching, and the hyenas are heavily persecuted (shot, poisoned, trapped, and hunted with dogs) in these areas because they are assumed to be livestock predators. This persecution, and habitat loss and fragmentation, are the primary threats to persistence of Brown Hyena. Because they are scavengers, many livestock carcasses where they are seen feeding are likely not to have been killed by Brown Hyenas. Although the species can be involved in depredation, this is usually restricted to a few individuals. Regardless, management of Brown Hyenas on ranchlands must address livestock losses. Typically, removal of individual problem hyenas ends the depredation. Because there is evidence that Brown Hyenas may be limited by the presence of Spotted Hyenas and perhaps other large predators, which are often absent from ranches, these ranchlands have the potential to be developed as Brown Hyena conservation areas, given proper management and conservation education efforts. Brown Hyenas are uncommon in captivity and traditionally do not breed well in confinement. Due to difficulties in captive breeding, the international studbook was discontinued in 1993 and as of 1995 there were only 16 specimens in nine collections. There is no known illegal trade in the species.	en	Don E. Wilson, Russell A. Mittermeier (2009): Hyaenidae. In: Handbook of the Mammals of the World – Volume 1 Carnivores. Barcelona: Lynx Edicions: 234-260, ISBN: 978-84-96553-49-1, DOI: 10.5281/zenodo.5676766
