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, 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