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Crocuta crocutaspotted hyena

By Jason Law

Geographic Range

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)

Habitat

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)

  • Range elevation
    4,000 (high) m
    ft

Physical Description

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)

  • Range mass
    45 to 80 kg
    99.12 to 176.21 lb
  • Range length
    95 to 150 cm
    37.40 to 59.06 in
  • Average length
    130 cm
    51.18 in

Reproduction

Mating in spotted hyaenas is polygynous. Males perform a bowing display to females before mating. The male lowers his muzzle to the ground, advances quickly toward the female, bows again, and then paws the ground close behind the female. The dominance of females assures that males are timid and will retreat immediately if the female shows any aggression. The female's reproductive tract makes mating somewhat difficult. Male hyaenas approach and slide their haunches under the female to achieve intromission. Once intromission is acheived they move to a more typical mating posture, with the male's underside resting on the female's back. The female phallus is completely slack during mating. (Estes, 1993; Frank, et al., 1995; Hofer, 2002; Kingdon, 1977; Nowak, 1999)

Crocuta crocuta clans are matrilinear and females are dominant over males. Juvenile males emigrate after puberty and join new clans where their position in the dominance hierarchy may increase over time. Females, however, have stable linear dominance hierarchies. In addition, rank is inherited from the mother so, these hierarchies remain stable for many generations. (Frank, et al., 1995)

Crocuta crocuta is highly polygynous and mating is aseasonal. Although all females produce litters, alpha females have a younger age at first breeding, shorter interbirth intervals, and increased survival of offspring. These benefits are passed directly to female offspring. The mechanism responsible for the increased fitness of high ranking females is probably the increased access to food that alpha females receive. In addition, high ranking female hyaenas seem to preferentially give birth to sons. Infanticide has been witnessed several times in the wild both by hyaenas from neighboring clans and also by females from the same clan. (Frank, et al., 1995; Hofer and East, 2003)

The gestation period is 4 months in C. crocuta. Females usually bear twins although 1 to 4 young are possible. The females give birth through their penis-like clitoris. During birth, the clitoris ruptures to allow the young to pass through. The resulting wound takes several weeks to heal. Cubs are not weaned until they are between 14 and 18 months of age. Females are capable of producing a litter every 11 to 21 months.

The newborns weigh from 1 to 1.6 kg and are quite precocious, being born with their eyes open. Newborns are almost entirely black. If siblings are the same sex, they begin fighting violently soon after birth, which usually results in the death of one of the two. Since single young receive more food and mature faster, this behavior is probably adaptive. Two to six weeks after birth, the mother transports young from the burrow in which they were born, often an abandoned aardvark, warthog or bat-eared fox burrow, to a communal den. The major source of food for the young during this time is milk from the mother.

Communal denning seems to be an important part of spotted hyaena social behavior, but no communal care of young takes place. One exception to this has been observed in the Kalahari during a particularly difficult period. (Frank, et al., 1995; Hofer, 2002; Kingdon, 1977; Kruuk, 1972; Nowak, 1999)

  • Breeding interval
    Spotted hyaenas breed about every 16 months with a range from 11 to 21 months.
  • Breeding season
    Breeding can occur throughout the year.
  • Range number of offspring
    1 to 4
  • Average number of offspring
    2
  • Average number of offspring
    2
    AnAge
  • Average gestation period
    110 days
  • Average gestation period
    110 days
    AnAge
  • Range weaning age
    12 to 16 months
  • Average time to independence
    18 months
  • Range age at sexual or reproductive maturity (female)
    21 to 48 months
  • Average age at sexual or reproductive maturity (female)
    40 months
  • Range age at sexual or reproductive maturity (male)
    2 (low) years
  • Average age at sexual or reproductive maturity (male)
    3 years

Crocuta crocuta has the highest parental investment of any carnivore for several reasons. First, spotted hyaena milk has extremely high energy content. The mean protein content is 14.9 %, and the mean fat content is 14.1 %. This is only exceeded by some bears and sea otters. Weaning occurs from about the age of 12 to 16 months, which is extremely late. By the weaning age, juvenile hyaenas already have completely erupted adult teeth, which is also very rare. The age at sexual maturity is about three years, although some males may be sexually active at two. C. crocuta females are very protective of their young and do not tolerate other hyaenas around them at first. Finally, females intervene on behalf of their daughters in antagonistic encounters and form coalitions with them to secure the place of the daughters in the dominance hierarchy immediately below that of the mother. Males have not been reported to have a role in parental care. (Frank, et al., 1995; Hofer, 2002; Kruuk, 1972)

  • Parental Investment
  • no parental involvement
  • precocial
  • female parental care
  • pre-fertilization
    • protecting
      • female
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female
  • pre-independence
    • provisioning
      • female
    • protecting
      • female
  • post-independence association with parents
  • extended period of juvenile learning
  • inherits maternal/paternal territory
  • maternal position in the dominance hierarchy affects status of young

Lifespan/Longevity

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)

Behavior

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)

  • Range territory size
    40 to 1000 km^2

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)

Communication and Perception

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.

Food Habits

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, et al., 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)

  • Animal Foods
  • birds
  • mammals
  • fish
  • eggs
  • carrion
  • terrestrial non-insect arthropods
  • Other Foods
  • dung

Predation

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)

Ecosystem Roles

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)

Economic Importance for Humans: Positive

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)

Economic Importance for Humans: Negative

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)

  • Negative Impacts
  • injures humans
  • crop pest

Conservation Status

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)

Contributors

Nancy Shefferly (editor), Animal Diversity Web.

Jason Law (author), University of Michigan-Ann Arbor, Phil Myers (editor, instructor), Museum of Zoology, University of Michigan-Ann Arbor.

Glossary

Ethiopian

living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.

World Map

acoustic

uses sound to communicate

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

carnivore

an animal that mainly eats meat

carrion

flesh of dead animals.

chemical

uses smells or other chemicals to communicate

desert or dunes

in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.

diurnal
  1. active during the day, 2. lasting for one day.
dominance hierarchies

ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates

drug

a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease

ecotourism

humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals. Ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals.

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

fertilization

union of egg and spermatozoan

food

A substance that provides both nutrients and energy to a living thing.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

keystone species

a species whose presence or absence strongly affects populations of other species in that area such that the extirpation of the keystone species in an area will result in the ultimate extirpation of many more species in that area (Example: sea otter).

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

native range

the area in which the animal is naturally found, the region in which it is endemic.

polygynous

having more than one female as a mate at one time

riparian

Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).

scavenger

an animal that mainly eats dead animals

scent marks

communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them

scrub forest

scrub forests develop in areas that experience dry seasons.

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

sexual ornamentation

one of the sexes (usually males) has special physical structures used in courting the other sex or fighting the same sex. For example: antlers, elongated tails, special spurs.

social

associates with others of its species; forms social groups.

tactile

uses touch to communicate

terrestrial

Living on the ground.

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

year-round breeding

breeding takes place throughout the year

young precocial

young are relatively well-developed when born

References

Di Silvestre, I., O. Novelli, G. Bogliani. 2000. Feeding habits of the spotted hyaena in the Niokolo Koba National Park, Senegal. African Journal of Ecology, 38: 102-107.

East, M., H. Hofer. 2001. Male spotted hyenas (Crocuta crocuta) queue for status in social groups dominated by females. Behavioral Ecology, 12/5: 558-568.

Eltringham, S. 1979. Ecology and Conservation of Large African Mammals. London: MacMillann Press Ltd.

Engh, A., K. Esch, L. Smale, K. Holekamp. 2000. Mechanisms of maternal rank ‘inheritance’ in the spotted hyaena, Crocuta crocuta . Animal Behaviour, 60: 323-332.

Estes, R. 1993. The Safari Companion: A Guide to Watching African Mammals. White River Junction: Chelsea Green Publishing Company.

Frank, L., K. Holekamp, L. Smale. 1995. Dominance, demography, and reproductive success of female spotted hyenas. Pp. 364-384 in A Sinclair, P Arcese, eds. Serengeti II: Dynamics, Management, and Conservation of an Ecosystem. Chicago: University of Chicago Press.

Hofer, H. 2002. "Spotted Hyaena" (On-line). IUCN Species Survival Commission Hyaenidae Specialist Group. Accessed March 31, 2004 at http://www.hyaena.ge/spotted.htm.

Hofer, H., M. East. 2003. Behavioral processes and costs of co-existence in female spotted hyaenas: a life history perspective. Evolutionary Ecology, 17: 315-331.

Hofer, H., M. East. 1995. Population dynamics, population size, and the commuting system of Serengeti spotted hyenas. Pp. 332-363 in A Sinclair, P Arcese, eds. Serengeti II: Dynamics, Management, and Conservation of an Ecosystem. Chicago: University of Chicago Press.

Holekamp, K., S. Cooper, C. Katona, N. Berry, L. Frank, L. Smale. 1997. Patterns of association among female spotted hyenas (Crocuta crocuta). Journal of Mammalogy, 78/1: 55-64.

Kingdon, J. 1977. East African Mammals: An Atlas of Evolution in Africa. New York: Academic Press.

Kruuk, H. 1972. The Spotted Hyena: A Study of Predation and Social Behavior. Chicago: University of Chicago Press.

Mills, G., L. Hes. 1997. The Complete Book of Southern African Mammals. Cape Town: Struik Winchester.

Muller, M., R. Wrangham. 2002. Sexual mimicry in hyenas. The Quarterly Review of Biology, 77/1: 3-16.

Nowak, R. 1999. Walker's Mammals of the World, Sixth Edition. Baltimore and London: The Johns Hopkins University Press.

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