Skip to Main Content
It looks like you're using Internet Explorer 11 or older. This website works best with modern browsers such as the latest versions of Chrome, Firefox, Safari, and Edge. If you continue with this browser, you may see unexpected results.
San Diego Zoo Wildlife Alliance logo
San Diego Zoo Wildlife Alliance Library logo

Koala (Phascolarctos cinereus) Fact Sheet: Bibliography & Resources


Adams-Hosking C, McBride MF, Baxter G, Burgman M, de Villiers D, Kavanagh R, Lawler I, Lunney D, Melzer A, Menkhorst P, et al. 2016. Use of expert knowledge to elicit population trends for the koala (Phascolarctos cinereus). Diversity and Distributions. 22(3):249–262. doi:10.1111/ddi.12400.
Agar J. 2008. Diprotodontia introduction. Tree of Life Web Project. Retrieved May 25 2010.

Archer M, Arena R, Bassarova M, Black K, Brammall J, et al. 1999. The evolutionary history and diversity of Australian mammals. Australian Mammalogy 21:1-45.

Archer MSJ, Hand H, Godthelp H. 1991. Riversleigh: The story of animals in ancient rainforests of inland Australia. Sydney: Reed Books. 264 pp.

Australian Government. 2012. Koala (Phasolarctos cinereus) listing. Retrieved 4 May 2012 from

Australian Government. 2022. Increased protection for koalas [media release]. Accessed 10 May 2022.

Australian Koala Foundation. 2010. Koala population estimates: explanation of methodology & recommendations to the threatened species scientific committee (TSSC) [PDF]. Australia Koala Foundation, Brisbane QLD, Australia.

Beck RMD. 2008. A dated phylogeny of marsupials using a molecular supermatrix and multiple fossil constraints. Journal of Mammalogy 89(1):175-189.

Beeby R. 2008. Koala retrovirus: a genome invasion in real time. Genome Biology 7(11):241. Retrieved 22 November 2011 from

Bercovitch FB, Tobey JR, Andrus CH, Doyle L. 2006. Mating patterns and reproductive success in captive koalas (Phascolarctos cinereus). Journal of Zoology 270:512-516.

Bingham J. 2005. Aboriginal art and culture. Chicago, Illinois: Raintree Press, Reed Elsevier. Inc. (young adult book). 56 pp.

Black K. 1999. Diversity and relationships of living and extinct koalas (Phascolarctidae, Marsupialia). Australian Mammalogy. 21(1):16.

Booth RJ, Blanshard WH. 1999. Diseases in koalas. In: Fowler ME, Miller RE (Eds.). Zoo and Wild Animal Medicine. Philadelphia: W.B. Saunders. pp. 321-33.

Brooker MIH et al. 2006. EUCLID: eucalypts of Australia [Database]. Collingswood, Australia: CSIRO Publishing. Sample page [cited 2012 Oct 12]. Available from:

Charlton BD, Ellis WAH, McKinnon AJ, Brumm J, Nilsson K, Fitch WT. 2011. Perception of male caller identity in koalas (Phascolarctos cinereus): Acoustic analysis and playback experiments. PLoS ONE 6(5):e20329.

Charlton BD, Frey R, McKinnon AJ, Fritsch G, Tecumseh-Fitch W, Reby D. 2013. Koalas use a novel vocal organ to produce unusually low-pitched mating calls. Current Biology. 23(23): R1035-R1036.

Choi CQ 2006. Retrovirus invading koalas: Insertion-in-process provides firsthand look at how retroviruses get incorporated into genomes. The Scientist. Retrieved 22 November 2011 from

Cork SJ, Sanson GD. 1990. Digestion and nutrition in the koala: a review. In: Lee AK, Handasyde KA, Sanson GD. (Eds.). Biology of the Koala. Sydney: Surrey, Beatty & Sons. pp. 129-144.

Crandall LS. 1964. The management of wild mammals in captivity. Chicago: The University of Chicago Press. pp. 29-31.

Degabriele R. 1980. The physiology of the koala. Scientific American 243(1):110-117.

Doyle L, Andrus C, Harris J, Moll J, Thompson V. 2002. The San Diego Zoo's koalas. ZooNooz 75(7): 8-13.

Dixon RMW, Moore B, Ramson WS, Thomas M. 2006. Australian Aboriginal words in English. Their origin and meaning. 2nd Edition. Oxford University Press. Pp. 22, 64-65, and 211-213.

Eberhard IH. 1978. Ecology of the koala, Phascolarctos cinereus (Goldfuss) Marsupialia: Phascolarctidae, in Australia. In: Montgomery GG (Ed.). The Ecology of arboreal folivores. Washington D.C.: Smithsonian Institution Press. pp. 315-327.

Ellis WA, Bercovitch F. 2007. Xena: Wanderer Princess. Zoonooz 79:8-9.

Ellis WA, Bercovitch F. 2011. Body size and sexual selection in the koala. Behavioral Ecology and Sociobiology 65:1229-1235.

Ellis W, Bercovitch F, FitzGibbon S, Melzer A, De Villiers D, Dique D. 2010a. Koala birth seasonality and sex ratios across multiple sites in Queensland, Australia. Journal of Mammalogy 91:177-182.

Ellis WA, Bercovitch F, FitzGibbon S, Roe P, Wimmer J, Melzer A, Wilson R. 2011. Koala bellows and their association with the spatial dynamics of free-ranging koalas. Behavioral Ecology 22:372-377.

Ellis WA, Hale PT, Carrick F. 2002a. Breeding dynamics of koalas in open woodlands. Wildlife Research 29(1):19-25.

Ellis WA, Hale PT, Carrick C, Hasegawa M, Nielsen M, Esser D. 2001. Aspects of the ecology of koalas at Blair Athol Coal Mine. In: Melzer A, Lamb D (Eds.). Proceedings of the Emerald Conference on the research and management of non-urban koala populations, 2-4 July 1997. pp. 127-138.

Ellis WA, Melzer A, Bercovitch FB. 2009. Spatiotemporal dynamics of habitat use by koalas: the checkerboard model. Behavioral Ecology and Sociobiology 63(8):1181-1188.

Ellis WA, Melzer A, Carrick FN, Hasegawa M. 2002b. Tree use, diet and home range of the koala (Phascolarctos cinereus) at Blair Athol, central Queensland. Wildlife Research 29:303-311.

Ellis WA, Melzer A, Clifton D, Carrick F. 2010b. Climate change and the koala Phascolarctos cinereus: water and energy. Australian Zoologist 35:(2): 369-377.

Fiebig UG,. Hartmann G, Bannert N, Kurth R, Denner J. 2006. Transspecies transmission of the endogenous koala retrovirus. Journal of Virology 80(11):5561-5654. doi:10.1128/JVI.02597-05.

Giezendanner T. 2011. Personal Communication regarding koala population at San Diego Zoo Global. 22 November 2011.

Godthelp H, Archer M, Cifelli R, Hand S, Gilkeson C. 1992. Earliest known Australian Tertiary mammal fauna. Nature 356:514-516.

Gordon GP, Hrdina F, Patterson R. 2006. Decline in the distribution of the koala Phascolarctos cinereus in Queensland. Australian Zoologist 33:345-358.

Gordon GP, McGreevy DG, Lawrie BC. 1990. Koala populations in Queensland: major limiting factors. In: Lee AK, Handasyde KA, Sanson GD (Eds.). Biology of the Koala. Sydney: Surrey, Beatty, & Sons. pp. 85-95.

Gordon GP, Menkhorst P, Robinson T, Lunney D, Martin R, Ellis M. 2008. Phascolarctos cinereus. In: IUCN 2010. IUCN Redlist of Threatened Species. Version 2010.1 Downloaded 24 June 2010.

Grand TI, Barboza PS. 2001. Anatomy and development of the koala, Phascolarctos cinereus: an evolutionary perspective on the superfamily Vombatoidea. Anatomy and Embryology 203:211-223.

Hanger JJ, Bromham LD, McKee JJ, O'Brien TM, Robinson WF. 2000. The nucleotide sequence of koala (Phascolarctos cinereus) retrovirus: a novel type C endogenous virus related to Gibbon ape leukemia virus. Journal of Virology 74:4264-4272. Retrieved 22 November 2011 from

Henneberg MJ, Lambert KM, Leigh CM. 1997. Fingerprint homoplasy: koalas and humans. naturalSCIENCE 1(4).

Higgins AL, Bercovitch FB, Tobey JR, Andrus CH. 2011. Dietary specialization and Eucalyptus species preferences in Queensland koalas (Phascolarctors cinereus). Zoo Biology 30:52-58.

Hindell MA, Lee AK. 1990. Tree preferences of the koala. In: Lee AK, Handasyde KA, Sanson GD (Eds.). Biology of the koala. Sydney: Surrey Beatty & Sons. pp. 117-121.

Houlden BA, Costello BH, Melzer A, Ellis W, Carrick F, Baverstock PR, Elphinstone MS. 1999. Phylogeographic differentiation in the mitochondrial control region in the koala, Phascolarctos cinereus (Goldfuss 1817). Molecular Ecology 8:999-1011.

Hume ID. 1999. Marsupial nutrition. Cambridge, UK: Cambridge University Press. 434 pp..

Irvine G. 2011.Snugglepot and Cuddlepie: Headliners of the roaring 20s. San Diego Writing Women. Retrieved 22 November 2011 from:

Jackson S. 2015. Family Phascolarctidae (koala). In: Handbook of the mammals of the world: marsupials and monotremes. Barcelona (Spain): Lynx Edicions. (Wilson DE, Mittermeier RA, editors). p. 400-417.

Jackson S, Groves C. 2015. Phascolarctos cinereus. In: Taxonomy of Australian mammals. Collingwood, VIC (Australia): CSIRO Publishing; c2015. p. 96-97.

Jackson S, Perry L, O'Callaghan PO, Spittal D, Romer L, Reid K. Date Unknown. Koala Phascolarctos cinereus [PDF]. Captive Husbandry Guidelines. pp. 1-92.

Jackson S. 2003 Australian mammals: biology and captive management. Collingwood, Australia: CSIRO Publishing. 524 pp.

Jackson S. 2007. Koala: origins of an icon. New South Wales, Australia: Jacana Books. 337 pp.

Johnson RN, O’Meally D, Chen Z, Etherington GJ, Ho SYW, Nash WJ, Grueber CE, Cheng Y, Whittington CM, Dennison S, et al. 2018. Adaptation and conservation insights from the koala genome. Nature Genetics. 50(8):1102–1111. doi:10.1038/s41588-018-0153-5.
Johnston SD, M.R. McGowan, P. O'Callaghan, R. Cox, V. Nicolson 2000a. Studies of the oestrous cycle, oestrus and pregnancy in the koala (Phascolarctos cinerus). Journal of Reproduction and Fertility, 120(1):49-57.

Kirsch JAW, Lapointe J-F, Springer MS. 1997. DNA-hybridization studies of marsupials and their implications for metatherian classification. Australian Journal of Zoology 45:211-80.

Krockenberger AK. 1996. Composition of the milk of the Koala, Phascolarctos cinereus, an arboreal folivore. Physiological Zoology 69(3):701-718.

Lanyon JM, Sanson GD. 1986. Koala (Phascolarctos cinereus) dentition and nutrition. II: Implications of tooth wear in nutrition. Journal of Zoology (London). A. 209: 168-181.

Lee A, Martin R. 1988. The koala: a natural history. Kensington: New South Wales University Press. 102 pp.

Long JA, Archer M, Flannery TF, Hand SJ. 2002. Prehistoric mammals of Australia and New Guinea: one hundred million years of evolution. Sydney, New South Wales, Australia: UNSW Press. 240 pp.

Louys J, Aplin K, Beck RMD, Archer M. 2009. Cranial anatomy of Oligo-Miocene koalas (Diprotodontia:Phascolarctidae): stages in the evolution of an extreme leaf-eating specialization. Journal of Vertebrate Paleontology 29(4):981-992.

Martin R.  2001. Koala. In: Macdonald D (Ed.). The encyclopedia of mammals. New York: Barnes and Noble. pp. 852-855.

Martin R. 1989. Draft management plan for the conservation of the koala (Phascolarctos cinereus) in Victoria. A report to the Department of Conservation, Forests and Lands, Victoria. Technical Report Series No. 99, Arthur Rylah Institute for Environmental Research.

Martin RW, Handasyde KA, Krockenberger A.. 2008. Koala, Phascolarctos cinereus. In: Van Dyck S, Strahan R (Eds.). The mammals of Australia. Third edition. Sydney, Australia: Reed New Holland. pp.198-201.

Martin R, Handasyde K. 1990. Population dynamics of the koala (Phascolarctos cinereus) in southeastern Australia. In: Lee AK, Handasyde K, Sanson GD (Eds.). Biology of the Koala. Sydney: Surrey Beatty & Sons. pp. 75-83.

Martin R, Handasyde K. 1999. The Koala: natural history, conservation, and management.  2nd ed.  Malabar, Florida: Krieger Publishing Company. pp. 64-65. 119 pp.

Maxwell S, Burbidge AA, Morris K. 1996. The 1996 Action Plan for Australian Marsupials and Monotremes. Australasian Marsupial and Monotreme Specialist Group, IUCN Species Survival Commission, Gland Switzerland.

McAlpine C, Melzer A, Lunney D, Foley B, Adams-Hosking C, Lawler I, Whisson D, Phillips S, Kavanagh R, Baxter G, et al. 2012. Conserving koalas in the 21st century: synthesising the dynamics of Australia’s Koala populations [working group report]. Australian Centre for Ecological Analysis & Synthesis. [PDF from Semantic Scholar].

McLean N, Handasyde K. 2006. Sexual maturity, factors affecting the breeding season and breeding in consecutive seasons in populations of overabundant Victorian koalas (Phascolarctos cinereus). Australian Journal of Zoology 54:385–392.

Melzer A, Ellis WA, Bercovitch F. 2010. Observations of male-on-male aggression among Queensland koalas (Phascolarctos cinereus) from Central Queensland. Queensland Naturalist 48(1-3):36-44.

Melzer A, Houston W. 2001. An overview of the understanding of koala ecology: how much more do we need to know? In: Lyons K, Melzer A, Carrick F, Lamb D. The research and management of non-urban koala populations. Koala Research Centre of Central Queensland. Rockhampton: Central Queensland University. pp. 98-114.

Meredith RW, Janecka JE, Gatesy, Ryder O, Fisher CA et al 2011. Impacts of the Cretaceous terrestrial revolution and KPg extinction on mammal diversification. Sciencexpress. doi:10.1126/science.1211028.

Mitchell P. 1988. Social organization of the koala. Phascolarctos cinereus. PhD Thesis, Monash University. Clayton, Victoria.

Mitchell P. 1990a. Social behavior and communication of koalas. In: Lee AK, Handasyde KA, Sanson GD (Ed.). Biology of the koala. Sydney: Surrey, Beatty & Sons. pp. 151-170.

Mitchell P. 1990b. The home ranges and social activity of koalas -a quantitative analysis. In: Lee AK, Handasyde KA, Sanson GD (Ed.). Biology of the koala. Sydney: Surrey, Beatty & Sons. pp. 171-187.

Mitchell P, Martin R. 1990. The structure and dynamics of koala populations - French Island in perspective. In: Lee AK, Handasyde KA, Sanson GD, (Eds.). Biology of the koala. Sydney: Surrey Beatty, & Sons. pp. 97-108.

Moore BD, Foley WJ. 2000. A review of feeding and diet selection in koalas (Phascolarctos cinereus). Australian Journal of Zoology, 48: 317-333

Moore BD, Foley WJ. 2005. Tree use by koalas in a chemically complex landscape. Nature 435:488-90

Natural Resource Management Ministerial Council. 2009. National Koala Conservation and Management Strategy 2009-2014. Retrieved 22 November 2011.

NCC. 2003? Noosa Council Conservation Action Plan Statement No. 1. Koalas. Draft.

Nowak RM. 1999. Walker's mammals of the world. Sixth Edition, Volume 1. Baltimore: The Johns Hopkins University Press.

Obendorf D. 1983. Causes of mortality and morbidity of wild koalas, Phascolarctos cinereus (Goldfuss), in Victoria, Australia. Journal of Wildlife Diseases 19(2):123-131.

Phillips S. 2000. Population trends and the koala conservation debate. Conservation Biology 14(3):650-659.

Pye GW. 2009. Shoulder dysplasia in koalas (Phascolarctos cinereus) at San Diego Zoo. Journal of Zoo and Wildlife Medicine 40(3):453-457. Retrieved 22 November 2011 from:

Pye GW, Hamlin-Andrus C, Moll J. 2008. Hip dysplasia in koalas (Phascolarctos cinereus) at the San Diego Zoo. Journal of Zoo and Wildlife Medicine. 39:61-68.

Reed AW. 2000. Aboriginal myths, legends and fables. United Kingdom: New Holland Books. 412 pp.

Russell EM. 1982. Patterns of parental care and parental investment in marsupials. Biological Review 57:423-486.

Savolainen P, Leitner T, Wilton AN, Matisoo-Smith E, Lundeberg J. 2004. A detailed picture of the origin of the Australian dingo, obtained from the study of mitochondrial DNA. PNAS 101(33):12387-12390.

Sharp A. 1995. The koala book. Gretna, LA: Pelican Publishing. 160 pp.

Sherwin WB, Timms P, Wilcken J, Houlden B. 2000. Analysis and conservation implications of koala genetics. Conservation Biology. 14(3):639.

Smith M. 1979a. Behaviour of the koala, Phascolarctos cinerus (Goldfuss), in captivity. I. Non-social behaviour. Australian Wildlife Research 6:117-128.

Smith M. 1980a. Behaviour of the koala, Phascolarctos cinerus (Goldfuss), in captivity. III. Vocalizations. Australian Wildlife Research 7:13-34.

Smith M. 1980b. Behaviour of the koala, Phascolarctos cinerus (Goldfuss), in captivity. IV. Scent-marking. Australian Wildlife Research 7:35-40.

Smith M. 1980c. Behaviour of the koala, Phascolarctos cinerus (Goldfuss), in captivity. V. Sexual behaviour. Australian Wildlife Research 7:41-51.

Smith M. 1987. Behaviour and ecology. In: Cronin L. (Ed.). Koala: Australia's endearing marsupial. Sydney: Reed Books. 136 pp.Takahashi M, Tobey JR, Andrus CH. 2007. Behavior of female koalas (Phascolarctos cinereus adustus) in a new exhibit at San Diego Zoo. Animal Keepers' Forum 35(12):509-514.

Sonntag CF 1921. The comparative anatomy of the koala (Phascolarctos cinereus) and vulpine phalanger (Trichosurus vulpecula). Proceeding of the Zoological Society of London 91(3):547-577.

Stoye JP. 2006. Koala retrovirus: a genome invasion in real time. Genome Biology 7(11):241. Retrieved 22 November 2011 from

Szalay FS. 1994. Evolutionary history of the marsupials and an analysis of osteological characters. New York: Cambridge University Press.

Takahashi M, Tobey JR, Pisacane CB, Andrus CH. 2009. Evaluating the utility of an accelerometer and urinary hormone analysis as indicators of estrus in a zoo-housed koala (Phascolarctors cinereus). Zoo Biology 28:59-68.

Takami K, Yoshida M, Yamamoto Y, Harada M, Furuyama J. 1998. Genetic variation of mitochondrial Cytochrome b genes among the subspecies of koala, Phascolarctos cinereus. Journal of Veterinary Medicine 60(10):1161-1163.

Tarlinton RE, Meers J, Hanger J, Young P. 2005. Real-time reverse transcriptase PCR for the endogenous koala retrovirus reveals an association between plasma viral load and neoplastic disease in koalas. Journal of General Virology 86:783-787. Retrieved 22 November 2011 from:

Tarlinton RE, Meers J, Young PR. 2006. Retroviral invasion of the koala genome. Nature. 44(216):79-8.

Thompson VD. 1987. Parturition and development in the Queensland koala at San Diego Zoo. International Zoo Yearbook 26:217-222.

Tobey JR, Andrus CH, Doyle L, Thompson VD, Bercovitch FB. 2006. Maternal effort and joey growth in koalas (Phascolarctos cinereus) Journal of Zoology 268:423-431.

Tobey JR, Nute TR, Becovitch FB 2009. Age and seasonal changes in the semiochemicals of the sternal gland secretions of male koalas (Phascolarctos cinereus). Australian Journal of Zoology 57: 111-118.

Tuttle RH. 1975. Primate functional morphology and evolution. The Netherlands: Mouton Publishers p.166.

Tyndale-Biscoe CH. 2005. Life of marsupials. Collingwood, Australia: CSIRO Publishing. pp. 219-265.

Ullrey DE, Robinson PT, Whetter PA. 1981a. Composition of preferred and rejected Eucalyptus browse offered to captive koalas, Phascolarctos cinereus (Marsupialia). Australian Journal of Zoology 29(6):839-846.

Ullrey DE, Robinson PT, Whetter PA. 1981b. Eucalyptus digestibility and digestible energy requirements of adult male koalas, Phascolarctos cinereus (Marsupialia). Australian Journal of Zoology 29(6):847-852.

UNEP. 2018. Species+ [online database]. Nairobi, Kenya: UNEP [compiled by UNEP-WCMC, Cambridge, UK]. [accessed 2018 Sep 11].

U.S. Fish and Wildlife Service Federal Register 2000. Endangered and threatened species: Australian koala.Volume 65(90):26762-26771 Retrieved 22 November from:

U.S. Fish & Wildlife Service. 2018. ECOS: Environmental Conservation Online System [Internet database/portal]. Washington D.C.: U.S. Fish & Wildlife Service [accessed 2018 Sep 11].;jsessionid=67343079F1C4D65CDC4FB43A19AD663C?spcode=A0HG.

Weisbecker V, Sánchez-Villagra MR. 2006. Carpal evolution in diprotodontian marsupials. Zoological Journal of the Linnean Society 146:369-384.

White NA, Kunst ND. 1990. Aspects of the ecology of the koala in southeastern Queensland. In: Lee AK, Handasyde KA, Sanson GD (Eds.). Biology of the koala. Sydney: Surrey, Beatty & Sons. pp. 109-116

Wilson DE, Mittermeier RA, editors. Handbook of the mammals of the world, volume 5. Barcelona (Spain): Lynx Edicions; 2015. Order Diprotodontia, Monotremes and marsupials; p. 399-417.

Woinarski J, Burbidge AA. 2016. Phascolarctos cinereus. The IUCN red list of threatened species [Internet]. Version 2018.1; e.T16892A21960344 [species assessed 2014 May 27; page accessed 2018 Sep 11]. doi:10.2305/IUCN.UK.2016-1.RLTS.T16892A21960344.en.

Woinarski J, Burbidge AA. 2020. Phascolarctos cinereus. The IUCN red list of threatened species [Internet]. Version 2020.2; e.T16892A166496779 [species assessed 2014 May 27; page accessed 2020 Oct 27]. doi:10.2305/IUCN.UK.2020-1.RLTS.T16892A166496779.en..

Wroe S. 2004. Killer kangaroos and other murderous marsupials. Scientific American, Special Volume 14:48-55. Retrieved 22 November 2011 from:

Additional Internet Resources

SDZWA Library Links