Crocuta crocuta
Creators: name: Don E. Wilson
name: Russell A. Mittermeier
Publisher: name: Lynx Edicions
Year: 2009
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Mammalia
Carnivora
Hyaenidae
Crocuta
Crocuta crocuta

description: 2.
Spotted Hyena
Crocuta crocuta
French: Hyene tachetée / German: TlUpfelhyane / Spanish: Hiena manchada
Taxonomy. Canis crocuta Erxleben, 1777,
Guinea, Aethiopia; restricted to “Senegambia”.
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.
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.
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.
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.
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.
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 24hour cycle, their activity is not continuous. Instead, activity occurs in bouts interspersed with periods of rest. Hyenas in Kenya that were followed for complete 24hour cycles spent 32% of their time active, but 53% of their active time occurred during hours of darkness.
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 24hour 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.
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.
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.
Bibliography. Bearder (1977), Bearder & Randall (1978), Binder & Van Valkenburgh (2000), Binford et al. (1988), Boydston, Kapheim & Holekamp (2006), Boydston, Kapheim, Van Horn et al. (2005), Boydston, Morelli & Holekamp (2001), Cooper (1989, 1990, 1991, 1993), Cooper et al. (1999), Cunha et al. (2005), Di Silvestre et al. (2000), Drea & Frank (2003), Drea, Coscia & Glickman (1999), Drea, Place et al. (2002), Drea, Vignieri, Cunningham & Glickman (2002), Drea, Vignieri, Kim et al. (2002), Drea, Weldele et al. (1998), East & Hofer (1991a, 1991b, 2001, 2002), East, Burke et al. (2003), East, Hofer & Wickler (1993), Eloff (1964, 1975), Engh, Esch et al. (2000), Engh, Funk et al. (2002), Frank (1986a, 1986b, 1994, 1997), Frank & Glickman (1991, 1994), Frank, Davidson & Smith (1985), Frank, Glickman & Licht (1991), Frank, Glickman & Powch (1990), Frank, Glickman & Zabel (1989), Frank, Holekamp & Smale (1995), Frank, Weldele & Glickman (1995), Gasaway et al. (1989, 1991), Glickman, (1995), Glickman, Cunha et al. (2006), Glickman, Frank et al. (1993), Golla et al. (1999), Hamilton et al. (1986), Harvey (1992), Hayward (2006), Henschel & Skinner (1987, 1990a, 1990b, 1991), Henschel, & Tilson (1988), Hofer & East (1993a, 1993b, 1993c¢, 1995, 1996, 1997, 2003, 2008), Hofer, Campbell et al. (1996), Hofer, East & Campbell (1993), Holekamp & Smale (1990, 1993, 2000), Holekamp, Boydston & Smale (2000), Holekamp, Boydston, Szykman et al. (1999), Holekamp, Sakai & Lundrigan (2007), Holekamp, Smale, Berg & Cooper (1997), Holekamp, Smale & Szykman (1996), Holekamp, Szykman et al. (1999), Honer, Wachter, East & Hofer (2002), Honer, Wachter, East, Streich et al. (2007), Koepfli et al. (2006), Kolowski & Holekamp (2006), Kolowski et al. (2007), Kruuk (1972, 1977), Lewis & Werdelin (2000) Licht et al. (1992), Lindeque & Skinner (1982), Matthews (1939), Mills, (1985, 1989, 1990), Mills & Gorman (1987), Mills & Hofer (1998), Neaves et al. (1980), Pienaar (1969), Pournelle (1965), Rensberger (1999), Rohland et al. (2005), Rosevear (1974), Sillero-Zubiri & Gottelli (19924, 1992b), Skinner, Funston et al. (1992), Skinner, Henschel & Van Jaarsveld (1986), Smale, Frank & Holekamp (1993), Smale, Holekamp & White (1999), Smale, Nunes & Holekamp (1997), Sutcliffe, (1970), Szykman, Engh et al. (2001), Szykman, Van Horn et al. (2007), Theis, Greene et al. (2007), Theis, Heckla et al. (2008), Tilson & Hamilton (1984), Tilson & Henschel (1984, 1986), Tilson, von Blottnitz & Henschel (1980), Trinkel & Kastberger (2005), Trinkel et al. (2004), Van Horn, McElhinny & Holekamp (2003), Van Horn, Wahaj & Holekamp (2004), Van Horn, Engh et al. (2004), Van Jaarsveld et al. (1988), Wachter et al. (2002), Wahaj & Holekamp (2006), Wahaj, Place et al. (2007), Wahaj, Van Horn et al. (2004), Watts (2007), Werdelin & Solounias (1991), Whateley (1980, 1981), Whateley & Brooks (1978).
description: Published as part of Don E. Wilson & Russell A. Mittermeier, 2009, Hyaenidae, pp. 234-260 in Handbook of the Mammals of the World – Volume 1 Carnivores, Barcelona :Lynx Edicions on pages 254-256, DOI: 10.5281/zenodo.5676766