Crocuta crocuta
spotted hyena

Through the end of the Pleistocene, spotted hyenas, Crocuta crocuta, ranged throughout Eurasia and the reasons for its extinction there are not certain. Until very recently, spotted hyaenas were a common species in most of sub-Saharan Africa. Since 1970, confirmed records of C. crocuta have been recorded in Tanzania, Kenya, Zambia, Zimbabwe, Mozambique, South Africa, Botswana, Namibia, Congo, Sudan, Central African Republic, Cameroon, Nigeria, Burkina Faso, Ivory Coast, Mali, Senegal, and Sierra Leone. Hyenas occur throughout sub-Saharan Africa, but their density varies widely across this area. Only occasional animals are seen in some areas (forests of Mt. Kenya) and they have been extirpated from most areas of South Africa, High densities occur in the Serengeti and especially the Ngorongoro crater in Tanzania. Crocuta crocuta is the most numerous large predator in the Serengeti. (Hofer, 2002; Kingdon, 1977; Nowak, 1999)

Crocuta crocuta is common in many types of open, dry habitat including semi-desert, savannah, acacia bush, and mountainous forest. The species becomes increasingly less common in dense forested habitat and is less common than Hyaena hyaena and Hyaena brunnea in desert habitats. Spotten hyaenas do not inhabit the coastal tropical rainforest of west or central Africa. In west Africa, the species prefers the Guinea and Sudan savannahs. Crocuta crocuta has been recorded from as high as 4,000 meters in east Africa and Ethiopia. (Hofer, 2002; Kingdon, 1977; Mills and Hes, 1997; Nowak, 1999)

Crocuta crocuta has a sandy, yellowish or gray coat with black or dark brown spots on the over most of the body. The spots are darkest in younger animals and can be almost completely absent in very old animals. The coat is very coarse and wooly. The body length from head to tail is about 95 to 150 cm and the height at the shoulder is reported from about 75 cm to 85cm. The tail is about 30 to 36 cm long and ends in a bushy black tip. About two-thirds of the tail is composed of bone with the other one-third being solely hair. Crocuta crocuta is sexually dimorphic with females weighing around 6.6 kg more than males. Male weight ranges from about 45 to 60 kg whereas females weigh 55 to over 70 kg. Crocuta crocuta is strongly built, with a massive neck and large head topped by rounded ears, unlike the other hyaenas. The jaws are probably the strongest in relation to size of any mammal. The front legs are longer than the hind legs, which gives the back of C. crocuta a slightly odd, downward slope. The feet have four digits with short, non-retractable claws and broad toe pads. (Hofer, 2002; Kingdon, 1977; Mills and Hes, 1997; Nowak, 1999)

C. crocuta females are extremely masculinated and the genitalia of females are almost indistinguishable from those of males. The clitoris is enlarged, looks like a penis, and is capable of erection. Females also have a pair of sacs in the genital region which are filled with fibrous tissue. These look much like a scrotum, but are covered with more hair than the male's scrotum. Thus, males and females look extremely similar. The female has no external vagina and must urinate, mate, and deliver young through the urogenital canal that exits through the pseudo-penis. High androgen levels were once thought to be a major cause of this masculinazation. One current hypothesis is that sexual mimicry is the driving force behind hyaena masculinization. Females that look like males may be protected from aggression from other females. (Hofer, 2002; Kingdon, 1977; Muller and Wrangham, 2002; Nowak, 1999)

Crocuta crocuta lives about 20 years in the wild. The longest known lifespan for this species in captivity is 41 years and 1 month. (Kruuk, 1972; Nowak, 1999)

Crocuta crocuta forms social groups called clans.l Clans may be composed of 3 to 80 members. Larger clans generally occur in prime territory with large prey concentrations, such as the Ngorongoro crater, whereas smaller clans occur in desert areas in southern Africa. All females are dominant to all males, and females remain in their natal clan for their entire lives. Males disperse upon reaching sexual maturity. Once a male joins another clan, he enters a dominance queue that the other males respect. As more males enter the queue and older males die, the male will move up through the social rank. Males spend a long time developing relationships with females in the clan. They follow females for periods of days or weeks and eventually gain favor with the females through this behavior. (East and Hofer, 2001; Hofer, 2002; Kruuk, 1972)

Although spotted hyaenas live in clans, the members of a clan are only observed all together in three circumstances: At kills, when defending the territory, and at a communal den. More often, the clan members forage alone or in small groups. Higher ranking females have been shown to associate more with kin than low ranking females. This behavior is beneficial to related females because they forage together and engage in coalitionary attacks against unrelated females when competing for food at a kill. Thus, females who associate with thier female kin are able to gather larger amounts of food more efficiently. In addition to allowing matrilines to defend their rank, close associations among female kin allow some of these kin groups to displace higher ranking matrilines under certain conditions. Finally, low ranking females preferentially associate with higher ranking females. It is hypothesized that these low ranking females receive benefits from high ranking females through reciprocal cooperation. (Holekamp et al., 1997; Kruuk, 1972)

Home Range

Crocuta crocuta clans defend group territories. However, they make frequent long distance foraging trips to the nearest ungulate herds to hunt. This system has been labeled a “commuting” system. It allows C. crocuta to live at much higher population densities than their clan territories would support. Territory size is extremely variable ranging from as small as 40 km^2 in the Ngorongoro crater to 1000 km^2 in the Kalahari. Territories are defended using vocal displays and scent marking. Scent marks are deposited from a secretion of the anal gland and from a secretion of glands on the feet. In addition, spotted hyaenas use communal latrines which also serve to mark territory boundaries. (Hofer and East, 1995; Hofer, 2002)

The commuting system of C. crocuta depends on the ability of hyaena clans to differentiate between "commuting" groups and groups that are actively searching for food. Aggression is rare between resident clan members and commuting individual hyaenas. Non-resident hyaenas typically defer to resident clan members at a kill, however this situation may also result in aggression. (Hofer and East, 1995; Hofer, 2002)

Crocuta crocuta is well known for the wide variety of vocal communication used. Groans and soft squeals are emitted during hyaena greetings. A whoop is used as a contact call in addition to a fast whoop which is used by excited hyaenas at a kill. Males give the fast whoop more often than females but are generally ignored. Female calls generally elicit much more of a reaction. Finally, a lowing call is used by impatient hyaenas who are kept waiting at a kill. (Estes, 1993; Kingdon, 1977; Nowak, 1999)

In addition to the previously mentioned calls, hyaenas give several calls related to aggression. These include grunting, giggling, growling, yelling, and a rattling growl. These calls are given in various aggressive interactions with clan members, other clans, or other species. The giggling is the trademark "laughing" call of the hyaena. Is associated with fear or excitement and is often given when an individual is being chased. (Estes, 1993)

Spotted hyaenas also perform a phallic inspection as a greeting. Two individuals stand head to tail, lift the rear leg closest to the other and then sniff and touch each other's extended phallus for up to 30 seconds. Females usually do not greet males in this manner, and if they do it is usually only the highest ranking males. Cubs can perform this ritual within the first month of life. (Estes, 1993; Hofer, 2002)

Chemical communication occurs because of the use of common latrine areas, as well as in scent marking. Tactile communication is involved in the genital investigation greeting, as well as between mothers and their young, rival young, and mates.

Hyaenas have a reputation for being mostly scavengers, however, this is not accurate. A study in the Kalahari found that 70 % of the diet was composed of direct kills. Typically, clans split up into hunting groups of 2 to 5 individuals, although zebra are hunted in larger groups. In the Serengeti and Ngongoro crater, Tanzania, C. crocuta was observed eating a wide variety of items including wildebeest, zebra, Thompson's gazelle, Grant's gazelle, topi, kongoni, waterbuck, eland, Cape buffalo, impala, Warthog, hare, springhare, ostrich eggs, bat-eared fox, golden jackal, porcupine, puff adder, domestic animals, lion, other hyaenas, termites, and afterbirth. Fecal analysis in these same two areas revealed that about 80 % of the samples contained wildebeest, zebra, and various gazelle species. In a study in Senegal, Hyaenas were found to prey on large herbivores such as buffalo, hartebeest, kob, warthog, bushbuck. In addition, C. crocuta has been known to prey on the young of giraffe, hippopotamus and rhinoceros. (Di Silvestre, Novelli, and Bogliani, 2000; Kruuk, 1972)

Crocuta crocuta uses its keen senses of sight, hearing and smell to hunt live prey and to detect carrion from afar. It often chases its prey long distances at speeds up to 60 km/hr. A chase in the Kalahari lasted 24 km before the prey, an eland, was captured. In the Serengeti, many prey species of ungulates are migratory and are not found in the clan territory during some parts of the year. When this occurs, the clan goes on hunting trips to the nearest concentrations of prey. The average round trip for these trips is about 80 km and a lactating female can make 40 to 50 trips per year for a total of 2800 to 3600 km per year. (Hofer and East, 2003; Hofer, 2002; Kruuk, 1972)

Crocuta crocuta is one of the top predators in Africa. However, there are several species which may kill them. In one study 13 of 24 hyaena carcasses found were killed by lions. Hyaenas and lions compete directly for food and often scavenge each other's kills. This competition often leads to antagonistic encounters that may result in death. In addition, humans often kill hyaenas in numerous ways. Through the early 1960's, hyaenas were shot on sight in numerous parks and game reserves in East Africa. Otherwise, this species is free of predators. (Eltringham, 1979; Kruuk, 1972)

Hyaenas are the most numerous large predator in Africa in areas where ungulates are common. Thus, they are an extremely important component of this ecosystem. Hyaenas utilize almost every part of their prey except for horns and rumen, and scavenge often. (Kruuk, 1972; Nowak, 1999)

Crocuta crocuta is a common predator on domestic livestock in Africa. In addition, they have also been known to attack and kill humans, especially during human disease outbreaks. (Hofer, 2002; Kingdon, 1977)

Crocuta crocuta is a large, common carnivore in many parts of Africa and so it is quite a resource for safari companies and should be considered an important part of the tourist industry. The species is sport hunted in some places in Africa, although hyaenas are not much in demand from trophy hunters because they are not viewed as very attractive. They are also hunted sometimes for food or medicine. (Hofer, 2002; Kruuk, 1972)

Crocuta crocuta has been categorized as a "Lower Risk" species by the IUCN Hyaena Specialist Group. In addition, the group has identified this species as "Conservation dependent". This means that there is currently a conservation program aimed at this species, but without this program the species would most likely be eligible for threatened status within 5 years. (Hofer, 2002)