Animal Diversity Web

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)

• Biogeographic Regions
• ethiopian
  • native

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)

• Habitat Regions
• tropical
• terrestrial

• Terrestrial Biomes
• desert or dune
• savanna or grassland
• forest
• scrub forest
• mountains

• Other Habitat Features
• riparian

• 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)

• Other Physical Features
• endothermic
• homoiothermic
• bilateral symmetry

• Sexual Dimorphism
• female larger
• ornamentation

• 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)

• Mating System
• polygynous

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)

• Key Reproductive Features
• iteroparous
• year-round breeding
• gonochoric/gonochoristic/dioecious (sexes separate)
• sexual
• fertilization
• viviparous

• 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)

• Range lifespan
  Status: wild

  20 (high) years

• Range lifespan
  Status: captivity

  41 (high) years

• Average lifespan
  Status: captivity

  12 years

• Average lifespan
  Status: wild

  25.0 years

  Max Planck Institute for Demographic Research

• Average lifespan
  Status: captivity

  40.0 years

  Max Planck Institute for Demographic Research

• Average lifespan
  Status: captivity

  41.1 years

  Max Planck Institute for Demographic Research

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)

• Key Behaviors
• terricolous
• diurnal
• motile
• sedentary
• territorial
• social
• dominance hierarchies

• 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.

• Communication Channels
• visual
• tactile
• acoustic
• chemical

• Other Communication Modes
• scent marks

• Perception Channels
• visual
• tactile
• acoustic
• chemical

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)

• Primary Diet
• carnivore
  • eats terrestrial vertebrates
  • scavenger

• 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)

• Known Predators

  • lions (Panthera leo)
  • humans (Homo sapiens)

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)

• Ecosystem Impact
• keystone species

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)

• Positive Impacts
• food
• ecotourism
• source of medicine or drug

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)

• IUCN Red List

  Least Concern
  More information

• IUCN Red List

  Least Concern
  More information

• US Federal List

  No special status

• CITES

  No special status

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.

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.

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