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Spotted Hyena (Crocuta crocuta) Fact Sheet: Bibliography & Resources


African Wildlife Foundation (AWF website). [cited 27 March 2014]. Available from:

AZA. 2023. Species Survival Statistics [data table report]. Chicago (IL): Lincoln Park Zoo/AZA Population Management Center; [accessed 2023 Apr 18].

AZA (Association of Zoos & Aquariums). 2024. Species Survival Statistics [data table report]. Chicago (IL): Lincoln Park Zoo/AZA Population Management Center; [accessed 2024 Feb 28; data table last updated Jan 2024].

Bearder SK. 1977. Feeding habits of spotted hyaenas in woodland habitat. East African Wildlife Journal. 15: 263-280.

Bearder SK, Randal RM. 1978. The use of fecal marking sites by spotted hyenas and civets. Carnivore. 1: 32-48.

Binder WJ, Van Valkenburgh B. 2000. Development of bite strength and feeding behaviour in juvenile spotted hyenas (Crocuta crocuta). Journal of Zoology, London. 252: 273-283.

Bohm T and Höner OR. 2015. Spotted hyaena Crocuta crocuta. The IUCN red list of threatened species [Internet]. Version 2019.1; e.T5674A45194782 [species assessed 2014 Apr 01; page accessed 2019 Apr 03]. doi:10.2305/IUCN.UK.2015-2.RLTS.T5674A45194782.en.

Botswana Predator Conservation Trust (BPCT website). [cited 27 March 2014]. Available from:

Boydston EE, Morelli TL, Holekamp KE. 2001. Sex differences in territorial behaviour exhibited by the spotted hyena (Hyaenidae, Crocuta crocuta). Ethology. 107: 369-385.

Boydston EE, Kapheim KM, Szykman M, Holekamp KE. 2003. Individual variation in space use by female spotted hyenas. Journal of Mammalogy. 84(3): 1006-1018.

Boydston EE, Kapheim KM, Van Horn RC, Smale L, Holekamp KE. 2005. Sexually dimorphic patterns of space use throughout ontogeny in the spotted hyena (Crocuta crocuta). Journal of Zoology. 267(3): 271-281.

Breuer T. 2005. Diet choice of large carnivores in northern Cameroon. African Journal of Ecology. 43: 97-106.

Brottman M. 2012. Hyena. London: Reaktion Books LTD.

Brown RW. 1956. Composition of Scientific Words. Published by the author.

Calderone JB, Reese BE, Jacobs, GH. 2003. Topography of photoreceptors and retinal ganglion cells in the spotted hyena (Crocuta crocuta). Brain, Behavior and Evolution. 62(4): 182-192.

Cooper SM. 1990. The hunting behaviour of spotted hyaenas (Crocuta crocuta) in a region containing both sedentary and migratory populations of herbivores. African Journal of Ecology. 28: 131-141.

Cooper SM, Holekamp KE, Smale L. 1999. A seasonal feast: long-term analysis of feeding behaviour in the spotted hyaena (Crocuta crocuta). African Journal of Ecology. 37: 149-160.

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

East ML, Hofer H. 1991a. Loud calling in a female-dominated mammalian society: I. Structure and composition of whooping bouts of spotted hyaenas, Crocuta crocuta. Animal Behaviour. 42(4): 637-649.

East ML, Hofer H. 1991b. Loud calling in a female-dominated mammalian society: II. Behavioral contexts and functions of whooping of spotted hyaenas, Crocuta crocuta. Animal Behaviour. 42(4): 651-669.

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

East ML, Hofer H. 2013. Crocuta crocuta spotted hyaena. In: Kingdon J, Hoffmann M (eds). Mammals of Africa. Volume V: carnivores, pangolins, equids and rhinoceroses. Bloomsbury.

East ML, Burke T, Wilhelm K, Greig C, Hofer H. 2003. Sexual conflicts in spotted hyenas: male and female mating tactics and their reproductive outcome with respect to age, social status and tenure. Proceedings of the Royal Society B. 270(1521): 1247-1254.

Eloff FC. 1964. Predatory habits of lions and hyenas. Koedoe. 7: 105-112.

Frank LG. 1986a. Social organization of the spotted hyaena (Crocuta crocuta). I. Demography. Animal Behaviour. 34: 1500-1509.

Frank LG. 1986b. Social organization of the spotted hyaena (Crocuta crocuta). II. Dominance and reproduction. Animal Behaviour. 34: 1510-1527.

Frank LG, Glickman SW, Zabel CJ. 1989. Ontogeny of female dominance in the spotted hyena: perspectives from nature and captivity. Symposia of the Zoological Society of London. 61: 127-146.

Genter H. 2014. Spotted hyena, Crocuta crocuta, North American Regional Studbook. Denver (CO): Denver Zoolological Foundation.

Glickman SE. 1995. The spotted hyaena from Aristotle to the Lion King: reputation is everything. Social Research. 62:501-537.

Gotch AF. 1979. Mammals: their Latin names explained. Blandford Press.

Gould SJ. 1981. Hyena myths and realities. Natural History. 90(2): 16-24.

Hayward MW, Hayward GJ, Druce DJ, Kerley GIH. 2009. Do fences constrain predator movements on an evolutionary scale? Home range, food intake and movement patterns of large predators reintroduced to Addo Elephant National Park, South Africa. Biodiversity Conservation. 18: 887-904.

Henschel JR, Skinner JD. 1990b. Parturition and early maternal care of spotted hyaenas Crocuta crocuta: a case report. Journal of Zoology (London). 222: 702-704.

Hofer H, East ML. 1993a. The commuting system of Serengeti spotted hyaenas: how a predator copes with migratory prey. I Social organization. Animal Behaviour. 46(3): 547-557.

Hofer H, East ML. 1993b. The commuting system of Serengeti spotted hyaenas: how a predator copes with migratory prey. II. Intrusion pressure and commuters' space use. Animal Behaviour. 46(3): 559-574.

Hofer H, East ML. 1993c. The communing system of Serengeti spotted hyaenas: how a predator copes with migratory prey. III. Attendance and maternal care. Animal Behaviour. 46(3): 575-589.

Holekamp K, Smale L, Szykman M. 1996. Rank and reproduction in female spotted hyena. Journal of Reproduction and Fertility. 108(2): 229-237.

Honer OP, Wachter B, East ML, Runyoro V, Hofer H. 2005. The effect of prey abundance and foraging tactics on the population dynamics of a social territorial carnivore, the spotted hyena. Oikos. 108: 544-554.

Honer O, Holekamp KE, Mills G. 2008. Crocuta crocuta. In: IUCN 2013. IUCN Red List of Threatened Species. Version 2013.2. [Cited 06 February 2014]. Available from:

Integrated Taxonomic Information System database (ITIS). [cited 27 March 2014]. Crocuta crocuta (Erxleben, 1777). Available from:

​IUCN Hyaena Specialist Group. [cited 27 March 2014]. Available from:

Koepfli KP, Jenks S, Eizirik E, Zahirpour T, Van Valkenburgh B, Wayne RK. 2006. Molecular systematics of the Hyaenidae: Relationships of a relictual lineage resolved by a molecular supermatrix. Molecular Phylogenetics and Evolution. 38: 603-620.

Krell FT, Korb J, Walter P. 2003. The beetle fauna of hyaena latrines: coprocenoses consisting of necrophagous beetles (Coleoptera Trogidae Scarabaeidae). Tropical Zoology. 16(2): 145-152.

Kruuk H. 1972. The spotted hyena: a study of predation and social behavior. Chicago: University of Chicago Press.

Mills MGL.1984a. Prey selection and feeding habits of the large carnivores in the southern Kalahari. Koedoe (Suppl): 281-294.

Mills MGL. 1984b. The comparative behavioural ecology of the brown hyaena Hyaena brunnea and the spotted hyaena Crocuta crocuta in the Southern Kalahari. Koedoe (Suppl). 27: 237-274.

Mills MGL. 1989. The comparative behavioral ecology of hyenas: the importance of diet and food dispersion. In: JL Gittleman (ed). Carnivore behaviour, ecology and evolution. London: Chapman and Hall. Pp 125-142.

Mills MGL. 2013. Hyaena brunnea, brown hyaena (standwolf). In: Kingdon J, Hoffmann M (eds). Mammals of Africa. Volume 5. Carnivores, pangolins, equids and rhinoceroses. London: Bloomsbury. Pp. 263-267.

Online Etymology Dictionary (OED). [cited 27 March 2014]. Hyena. Available from:

Palmqvist P, Martinez-Navarro B, Perez-Claros J, Torregrosa V, Figueirido B, Jimenez-Arenas JM, Espigares MP, Ros-Montoya S, De Renzi M. 2011. The giant hyena Pachycrocuta brevirostris. Modeling the bone-cracking behavior of an extinct carnivore. Quaternary International. 243(1): 61-79.

Parrinder G. 1986. African mythology. New York (New York): Peter Bedrick Books. Pp. 38-40, 132.

Pournelle GH. 1965. Observations of the birth and early development of the spotted hyaena. Journal of Mammalogy. 46(3): 503.

Racey PA, Skinner JD. 1979. Endocrine aspects of sexual mimicry in spotted hyenas, Crocuta crocuta. Journal of Zoology. 187: 315-326.

Rensberger JM, Stefen C. 1995. Enamel specialization in hyenas: a structure useful for assessing bone-eating behavior in carnivorous mammals. Journal of Vertebrate Paleontology. 15(3 suppl): 49A.

Sillero-Zubiri C, Gottelli D. 1992a. Population ecology of spotted hyaena in an equatorial mountain forest. African Journal of Ecology. 30: 292-300.

Stefen C, Rensberger JM. 1999. The specialized structure of hyaenid enamel: description and development within the lineage-including percrocutids. Scanning Microscopy. 13(2-3): 363-380.

Stratford KJ, Stratford SMC. 2011. Fine-scale movements and use of space by spotted hyaena (Crocuta crocuta) on Ongava Game Reserve, Namibia. African Journal of Ecology. 49(3): 343-352.

Szykman M, Engh AL, Scribner KT, Van Horn RC, Runk SM, Holekamp KE. 2001. Association patterns among male and female spotted hyenas (Crocuta crocuta) reflect male mate choice. Behavioral Ecology and Sociobiology. 50(3): 231-238.

Theis KR, Venkataraman A, Dycus JA, et al. 2013. Symbiotic bacteria appear to mediate hyena social odors. Proceedings of the National Academy of Sciences of the United States of America. 110(49): 19832-19837.

Tilson RL, Henschel RJ. 1986. Spatial arrangements of spotted hyena groups in a desert environment in Namibia. African Journal of Ecology. 24: 173-180.

Turner A, Anton M, Werdelin L. 2008. Taxonomy and evolutionary patterns in the fossil Hyaenidae of Europe. Geobios. 41(5): 677-687.

UNEP. 2019. Species+ [online database]. Nairobi, Kenya: UNEP [compiled by UNEP-WCMC, Cambridge, UK]. [accessed 2019 Apr 03].

Wagner AP. 2013. Hyaena hyaena, striped hyaena. In: Kingdon J, Hoffmann M (eds). Mammals of Africa. Volume 5. Carnivores, pangolins, equids and rhinoceroses. London: Bloomsbury. Pp. 267-272.

Watts HE. 2007. Social and ecological influences on survival and reproduction in the spotted hyena, Crocuta crocuta [Ph.D. dissertation]. East Lansing (MI): Michigan State University. 166 pp.

Werdelin L. 2013. Family Hyaenidae. In: Kingdon J, Hoffmann M (eds). Mammals of Africa. Volume 5: Carnivores, pangolins, equids and rhinoceroses. London: Bloomsbury. Pp. 260-261, 272.

Werdelin L, Solounias N. 1991. The Hyaenidae: taxonomy, systematics and evolution. Fossils and Strata. 30: 1-104.

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