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African Elephants (Loxodonta africana and L. cyclotis) Fact Sheet: Bibliography & Resources

Update in Progress

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This fact sheet, like an elephant, is aging gracefully. San Diego Zoo Wildlife Alliance is working to bring you an updated version of the African Elephants Fact Sheet with new science and conservation information. Thanks for your patience, as our tusks go to the ground and dig into this huge project. Please check back soon. SDZWA team members can email questions to

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Adams,J. 1981. Wild Elephants in Captivity. Carson: Center for the Study of Elephants.

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

Benedict FG. 1936. The Physiology of the Elephant. Washington, DC: Carnegie Institute.

Benirschke K., Roocroft A. 1992. Elephant Inflicted Injuries. Paper Presentation, Symposium on Erkrankungen der Zoo-und Wildtiere, Spain.

Blanc J. 2008. Loxodonta africana. The IUCN Red List of threatened species [Internet]. Version 2018-2; e.T12392A3339343. [species assessed 2008 Jun 30; page accessed 2018 Dec 18].  

Blanc JJ, Thouless CR, Hart JA, Dublin HT, Douglas-Hamilton I, Craig CG, Barnes RFW. 2003. African elephant status report 2002: an update from the African elephant database. Occasional Paper Series of teh IUCN Species Survival Commision, No. 29. IUCN/SSC African Elephant Specialist Group. IUCN, Gland (Switzerland).

Blanc JJ, Barnes RFW, Craig GC, Dublin HT, Thouless CR, Douglas-Hamilton I, Hart JA. 2007. African elephant status report 2007: an update from the African elephant database. Occasional Paper Series of the IUCN Species Survival Commision, No. 33. IUCN/SSC African Elephant Specialist Group. IUCN, Gland (Switzerland).

Brandt AL, Ishida Y, Georgiadis NJ, Roca AL. 2012. Forest elephant mitochondrial genomes reveal that elephantid diversification in Africa tracked climate transitions. Molecular Ecology. 21(5):1175-1189.

Buckley C, Collins D, Doyle C, Johnson G, Lehnhardt J, Redfox D. 1992. EMA Statement on Free/Protected Contact. Elephant Managers' Association Journal (EMA), Indianapolis, IN. pp. 9-12.

Caloi L, Palombo M. 1992. Dwarf elephants of the past, In:  J. Shoshani, ed., Elephants; Majestic Creatures of the Wild.  Emmaus, PN: Rodale Press.

Chadwick DH. 1992. The Fate of the Elephant. San Francisco: Sierra Club Books.

Chase M, Griffin C. 2009. Elephants caught in the middle: impacts of war, fences and people on elephant distribution and abundance in the Caprivi Strip, Namibia. African Journal of Ecology 47:223-233.

Chase MJ, Schlossberg S, Griffin CR, Bouché PJC, Djene SW, Elkan PW, Ferreira S, Grossman F, Kohi EM, Landen K, et al. 2016. Continent-wide survey reveals massive decline in African savannah elephants. PeerJ. 4:e2354. doi:10.7717/peerj.2354.

CITES Website 2010. Consideration of proposals for amendment of appendices I and II. From: Convention on international trade in endangered species of wild fauna and flora. Fifteenth meeting of the conference of the parties Dona (Qatar), 13-25 March 2010.​

Clubb R. Mason G. 2002. A Review of the Welfare of Zoo Elephants in Europe, A report Commissioned by the RSPCA.

Cutler A. 1985. The Craft of ivory. Sources, techniques and uses in the Mediterranean world: A.D. 200-1400. Washington D.C.: Dumbarton Oaks.

Daniel J. 1992. The Asian elephant population today, In: J. Shoshani, ed. Elephants; Majestic Creatures of the Wild. Emmaus, PN: Rodale Press.

Desmond T, Laule G. 1991. Protected Contact Elephant Training. American Association of Zoological Parks & Aquariums. National Conference, San Diego, CA.Conference Proceedings. (AAZPA) Wheeling, West Virginia. pp. 606-613.

Desmond T, Laule G. 1993. The Politics of Protected Contact. American Association of Zoological Parks and Aquariums. National Conference, Omaha, Nebraska. Conference Proceedings. (AAZPA) Wheeling, West Virginia. pp. 12-18.

Eltringham SK. 1982. Elephants. Dorsett: Blanford Press.

Eltringham SK. 1991. The Illustrated Encyclopedia of Elephants. New York: Crescent Books.

Eggert LS, Rasner CA, Woodruff DS 2002. The evolution and phylogeography of the African elephant inferred from mitochondrial DNA sequence and nuclear microsatellite markers. Proceedings of the Royal Society 269. DOI 10.1098/rspb.2002.2070.

Estes, R.D. 1993. The Safari Companion. Post Mill: Chelsea Green Publishing Co.

Estes RD. 1991. The Behavior Guide to African Mammals. Los Angeles: The University of California Press.

Fowler ME. 1993. Zoo & Wild Animal Medicine, Current Therapy 3. Philadelphia: W.B. Saunders.

Gobush KS, Edwards, CTT, Balfour D, Wittemyer G, Maisels F, Taylor RD. 2021. African savanna elephant, Loxodonta africana. The IUCN red list of threatened species. Version 2021-1; e.T181008073A181022663 (species assessed 2020 Nov 13; page accessed 2021 Mar 30). doi:10.2305/IUCN.UK.2021-1.RLTS.T181008073A181022663.en.

Gobush KS, Edwards, CTT, Maisels F, Wittemyer G, Balfour D, Taylor RD. 2021. African forest elephant, Loxodonta cyclotis. The IUCN red list of threatened species. Version 2021-1; e.T181007989A181019888 (species assessed 2020 Nov 13; page accessed 2021 Mar 30). doi:10.2305/IUCN.UK.2021-1.RLTS.T181007989A181019888.en.

Gove PB. 1971. Webster's third new international dictionary of the English language, unabridged. Springfield (MA): Merriam-Webster.

Green R, Olsen T, Ruhter D. 1992. Elephant Management at the Houston Zoo. American Association of Zoological Parks and Aquariums. Western Regional Conference, Tucson, Arizona. Conference Proceedings. (AAZPA) Wheeling, West Virginia. pp. 730-733.

Groves CP, Grubb P 2000. Do Loxodonta cyclotis and L. africana interbreed? Elephant 2:4-7.

Grubb P, Groves CP, Dudley JP, Shoshani J. 2000. Living African elephants belong to two species: Loxodonta africana (Blumenbach, 1797) and Loxodonta cyclotis (Matschie, 1900). Elephant 2(4):1-4.

Guerrero D. 1995. Elephant Management: The Evolution of Change. Elephant Managers' Association Journal (EMA), VI (2) Indianapolis, IN. pp. 43-52.

Guerrero D. 1995. Elephant & Human Social Styles: How to Apply Them to Training. Elephant Managers' Association Journal (EMA), VI (3) Indianapolis, IN. pp. 48-53.

Guerrero D. 1996. Environmentalism & Emotionalism: Operating Through Instead of Reacting To. Elephant Managers' Association Journal (EMA), VII (1) Indianapolis, IN. pp. 46-50.

Guerrero D. 1996. ABC's: Animal behavior concerns and solutions. Behavior evaluation: elephant (Elephas maximus indicus). Animal Keeper's Forum. 23 (9):482-485. American Association of Zookeepers, (AAZK) Topeka, KS. 

Heffner R,  Heffner H. 1980. Hearing in the elephant (Elephas maximus). Science, 208(4443):518-520.

Ishida Y, Oleksyk TK, Georgiadis NJ, David VA, Zhao K, Stephens RM, Kolokotronis S-O, Roca AL. 2011. Reconciling apparent conflicts between mitochondrial and nuclear phylogenies in African elephants. PloS one. 6(6):e20642.

IUCN. 2015. Updated African Elephant Database reveals declining elephant populations.

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Jackson P. 1991. Endangered Species: Elephants. London: Quintet Publishing Ltd.

Lee PC, S Sayialel, Lindsay WK, Moss CJ. 2012. African elephant age determination from teeth: validation from known individuals. African Journal of Ecology 50:9-20.

Lee PC, Fishlock V, Webber CE, Moss CJ. 2016. The reproductive advantages of a long life: longevity and senescence in wild female African elephants. Behavioral Ecology and Sociobiology 70:337-345.

Lehnhardt J. 1991. Elephant Handling - A Problem of Risk Management. American Association of Zoological Parks & Aquariums. National Conference, San Diego, CA. Conference Proceedings. (AAZPA) Wheeling, West Virginia. pp. 569-575.

Maddox S. 1992. Bull Elephant Management: A Safe Alternative. American Association of Zoological Parks & Aquariums. Central Regional Conference, Dallas, Texas. Conference Proceedings. (AAZPA) Wheeling, West Virginia. pp. 376-382.

Maisels F, Strindberg S, Blake S, Wittemyer G, Hart J, Williamson EA, Aba’a R, Abitsi G, Ambahe RD, Amsini F et al. 2013. Devastating decline of forest elephants in Central Africa. PloS One. 8(3):e59469.

Martin E. 1992. The rise and fall of the ivory market. In: J. Shoshani, ed. Elephants: Majestic Creatures of the Wild.  Emmaus, PN: Rodale Press.

McCarthy M. 2010. Trade in mammoth ivory 'is fueling slaughter of African elephants': conservationists fear that legal treade is eing used as a front for launderig of poached tusks. Independent/UK Sept. 29, 2010. Retrieved 6 July 2011 from:

Mennell KG, Scholes RJ, editors. 2008. Assessment of South African Elephant Management. Johannesburg: Witwatersrand University Press. 448 p. [PDF]

Mikota SK, Sargent EL  Ranglack GS.  1994. Medical Management of the Elephant. West Bloomfield, MI: Indira Publishing House.

Miller G, Wilcox C, Latores S, Upchurch B. 1993. "C" is for Cindy: A Case History. American Association of Zoological Parks & Aquariums. National Conference, Omaha, Nebraska. Conference Proceedings. (AAZPA) Wheeling, West Virginia. pp. 32-34.

Moss C. 1998. Elephant Memories: Thirteen Years in the Life of an Elephant Family. New York: William Morrow & Co. 336 pp.

Moss CJ. 2001. The demography of an African elephant (Loxodonta africana) population in Amboseli, Kenya. Journal of Zoology 255: 145-156.

Moss CJ, Croze H, Lee PC (Eds.). 2011 The Amboseli elephants: a long-term perspective on a long-lived mammal. Chicago, Illinois: The University of Chicago Press.

Nikaido, M., H. Nishihara, Y. Hukumoto, N. Okada 2003. Ancient SINEs from African Endemic Mammals. Molecular Biology and Evolution 20(4):522-527i

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

Nowak, R.M. 2018. Proboscidea: Elephantidae. In: Walker's mammals of the world: monotremes, marsupials, afrotherians, xenartrans, and sundatherians. Baltimore (MD): Johns Hopkins University Press; p. 463-536.

Olson DJ 1997. North American Regional Studbook for the African Elephant. Indianapolis, IN: Indianapolis Zoo.

Olson D. 2014. African elephant studbook: North American region. Azle (TX): International Elephant Foundation. 127 p.

Orenstein R. 1992. The 1989 CITES meeting in Lausanne, Switzerland. In: J. Shoshani, ed. Elephants. London: Simon & Schuster Ltd.

O'Sullivan T. 1993. Protected Contact at the St. Louis Zoo. Paper presentation, Elephant Managers' Association Conference. Vallejo, CA.

Palombo, M. R. and P. Villa 2001. Schreger lines as support in the Elephantidae identification. The World of Elephants - International Congress, Rome [PDF]

Parker ISC, Graham AD. 2017. The african elephants’ toe nails. Journal of East African Natural History. 106(1):47–51. doi:10.2982/028.106.0101.

Parrott,J. 1993. A New Modified Protected Contact Elephant Management Program to Maximize Keeper Safety. American Association of Zoological Parks & Aquariums. National Conference, Omaha, Nebraska. Conference Proceedings. (AAZPA) Wheeling, West Virginia. pp. 35-38.

Payne K, Langbauer W. 1992.  Elephant communication. In: J. Shoshani ed., Elephants. London: Simon & Schuster Ltd.

Priest G. 1994. Answers to questions about protected contact elephant management. Animal Keeper's Forum. 21(3): 80-91. American Association of Zookeepers, (AAZK) Topeka, KS.

Priest G. 1994. Introducing change is never easy. At the San Diego Wild Animal Park, the elephants caught on more quickly than their trainers.  Magazine. October 1994. pp.26-28

Roca AL, Geogiadis N, Pecon-Slattery J, O'Brien SJ 2001. Genetic evidence for two species of elephant in Africa. Science 293:1473-1477.

Roca AL, O'Brien SJ. 2005. Genomic inferences from Afrotheria and the evolution of elephants. Curr. Opin Genet. Dev. 15(6):652-659.

Roca AL, Georgiadis N, O'Brien SJ. 2005. Cytonuclear genomic dissociation in African elephant species. Nature Genetics. 37(1):96-100.

Rohland N, Malaspinas A, Pollack J, Slatkin M, Matheus P, Hofreiter M. 2007. Proboscidean mitogenomics: chronology and mode of elephant evolution using mastodon as outgroup. Public Library of Science (PLOS) Biology 5(8) [PDF]

Rholand N, Reich D, Mallick S, Meyer M, Green RE, Georgiadis NJ, Roca AL, Hofreiter M. 2010. Genomic DNA sequences from mastodon and wolly mammoth reveal deep speciation of forest and savanna elephants. Public Library of Science (PLOS) Biology. 8(12):e31000564.

Roocroft A., Zoll DA.. 1994. Managing Elephants: An Introduction to Their Training and Management. Ramona, CA: Fever Tree Press (ISBN 0-9640-0730-4).

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Shoshani, J. 1992. Comparing the living elephants. In: Elephants: Majestic Creatures of the Wild, p 36-51 . Emmaus, PN: Rodale Press.

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Shoshani J. 2006. Taxonomy, Classification, History, and Evolution of the Elephants. In: Biology, Medicine and Surgery of Elephants. Ames, IA: Blackwell Publishing. p. 3-14.

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Stoinski TS, Daniel E, Maple TL.. 2000. A preliminary study of the behavioral effects of feeding enrichment on African elephants. Zoo Biology 19(6):485-493.

Sukumar R. 2003. The Living Elephants. Oxford UK: Oxford University Press.

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