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Giraffes (Giraffa spp.) Fact Sheet: Bibliography & Resources


Agaba M, Ishengoma E, Miller WC, McGrath BC, Hudson CN, Bedoya Reina OC, Ratan A, Burhans R, Chikhi R, Medvedev P et al. 2016. Giraffe genome sequence reveals clues to its unique morphology and physiology. Nature Communications. 7 (online). http:/

Baotic A, Sicks F, Stoeger AS. 2015. Nocturnal "humming" vocalizations: Adding a piece to the puzzle of giraffe vocal communication. BMC Research Notes. 8(1):1-11.

Bercovitch F, Beliveau S, and Bashaw M. 2003. A tall order: hormones and behavior in giraffes. ZOONOOZ. 76(4):24-25.

Bercovitch FB, Berry PSM. 2009a. Ecological determinants of herd size in the Thornicroft's giraffe of Zambia. African Journal of Ecology. 48(4):962-971.

Bercovitch FB, Berry PSM. 2009b. Reproductive life history of Thornicroft's giraffe in Zambia. African Journal of Ecology. 48(2):535-538.

Bercovitch FB, Berry PSM. 2012. Herd composition, kinship and fission-fusion social dynamics among wild giraffe. African Journal of Ecology. 51(2):206-216.

Bercovitch FB, Berry PSM. 2014. The composition and function of all-male herds of Thornicroft's giraffe, Giraffa camelopardalis thornicrofti, in Zambia. African Journal of Ecology. 53(2):167-174.

Bercovitch F, Carter K, Fennessy J, Tutchings A. 2018. Giraffa camelopardalis spp. thornicrofti. The IUCN red list of threatened species [Internet]. Version 2018.3; e.T88421020A88421024 [species assessed 2018 Feb 26; page accessed 2018 Nov 20].

Bercovitch FB, Deacon F. 2015. Gazing at a giraffe gyroscope: Where are we going? African Journal of Ecology. 53(2):135-146.

Berry PSM, Bercovitch FB. 2012. Darkening coat colour reveals life history and life expectancy of male Thornicroft's giraffes. Journal of Zoology. 287(3):157-160.

Bradshaw Foundation. "The Giraffe Carvings of the Tenere Desert." Accessed on 07 June 2016.

Brenneman RA, Bagine RK, Brown DM, Ndetei R, Louis EE. 2009. Implications of closed ecosystem conservation management: The decline of Rothschild’s giraffe (Giraffa camelopardalis rothschildi) in Lake Nakuru National Park, Kenya. African Journal of Ecology. 47(4):711-719.

Brown DM, Brenneman RA, Koepfli K-P, Pollinger JP, Milá B, Georgiadis NJ, Louis Jr EE, Grether GF, Jacobs DK, Wayne RK. 2007. Extensive population genetic structure in the giraffe. BMC Biology. 5:57. 10.1186/1741-7007-5-57

Cameron EZ, du Toit JT. 2005. Social influences on vigilance behaviour in giraffes, Giraffa camelopardalis. Animal Behaviour. 69:1337-1344.

Carter KD, Seddon JM, Frère CH, Carter JK, Goldizen AW. 2013. Fission–fusion dynamics in wild giraffes may be driven by kinship, spatial overlap and individual social preferences. Animal Behaviour. 85(2):385-394.

Ciofolo I, Le Pendu Y. Giraffa camelopardalis Giraffe. In: Kingdon J, Hoffman M, editors. Mammals of Africa: pigs, hippopotamuses, cherotain, giraffes, deer and bovids. London (UK): Bloomsbury Publishing; c2013. p. 98-110.

Coe MJ. 1967. Necking behaviour in the giraffe. Journal of Zoology. 151:313-321.

Colbert EH, Morales M. 1990. Evolution of the vertebrates: A history of the backboned animals through time. New York (NY): John Wiley & Sons, Inc.

Dagg AI. 1971. Giraffa camelopardalis. Mammalian Species. (5):1-8.

Dagg AI, Foster JB. 1976. The giraffe: its biology, behavior, and ecology. New York (NY): Van Nostrand Reinhold.

Dagg AI. 2014. Giraffe: biology, behaviour and conservation. Cambridge, UK: Cambridge University Press.

Davidson Z, Valeix M, Van Kesteren F, Loveridge AJ, Hunt JE, Murindagomo F, Macdonald DW. 2013. Seasonal diet and prey preference of the African lion in a waterhole-driven semi-arid savanna. PloS One. 8(2):e55182.

East R. 1998. African Antelope Database. IUCN/SSC Antelope Specialist Group Report.

Estes RD.1991. The behavior guide of African mammals. Berkeley (CA): University of California Press.

Fennessy J. 2009. Home range and seasonal movements of Giraffa camelopardalis angolensis in the northern Namib Desert. African Journal of Ecology. 47(3):318.

Fennessy J, Marais A. 2018. Giraffa camelopardalis spp. antiquorum. The IUCN red list of threatened species [Internet]. Version 2018.3; e.T88420742A88420817 [species assessed 2018 Jan 24; page accessed 2018 Nov 20].

Fennessy J, Bidon T, Reuss F, Kumar V, Elkan P, Nilsson Maria A, Vamberger M, Fritz U, Janke A. 2016. Multi-locus analyses reveal four giraffe species instead of one. Current Biology.

Fennessy J, Leggett K, Schneider S. 2001. A preliminary study of giraffe (Giraffa camelopardalis). Desert Research Foundation of Namibia.

Fennessy J, Bock F, Tutchings A, Brenneman R, Janke A. 2013. Mitochondrial DNA analyses show that Zambia's South Luangwa Valley giraffe (Giraffa camelopardalis thornicrofti) are genetically isolated. African Journal of Ecology. 51(4):635-640.

Fennessy J, Brown D. 2010. Giraffa camelopardalis. In: IUCN Red List of Threatened Species 2010. Accessed on 28 March 2016.

Fennessy J, Marais A, Tutchings A. 2018. Giraffa camelopardalis spp. peralta. The IUCN red list of threatened species [Internet]. Version 2018.3; e.T136913A51140803 [species assessed 2017 Jan 15; page accessed 2018 Nov 20].

Fobar R. 2019. Masai Giraffes Declared Endangered. Wildlife Watch. 

Foster JB. 1966. The giraffe of Nairobi National Park: Home range, sex ratio, the herd, and food. East African Wildlife Journal. 4:139-148.

Foster JB, Dagg AI. 1972. Notes on the biology of the giraffe. East African Wildlife Journal. 10:1-16.

Gadjev I. 2015. Nature and nurture: Lamarck's legacy. Biological Journal of the Linnean Society. 114(1):242-247.

Giraffe Conservation Foundation. 2016. Africa's Giraffe: A conservation guide. 2nd ed. Downloaded on 08 May 2018.

Gotch AF. 1995. Latin names explained: A guide to the scientific classification of reptiles, birds, and mammals. London: Cassell PLC.

Guggisberg CA. 1969. Giraffes. London, UK: Arthur Barker Limited.

Hall-Martin AJ, Skinner JD, Van Dyk JM. 1975. Reproduction in the giraffe in relation to some environmental factors. East African Wildlife Journal. 13:237-248.

Hassanin A, Ropiquet A, Gourmand A-L, Chardonnet B, Rigoulet J. 2007. Mitochondrial DNA variability in Giraffa camelopardalis: Consequences for taxonomy, phylogeography and conservation of giraffes in West and central Africa. Comptes Rendus Biologies. 330(3):265-274.

Hayward MW, Kerley GI. 2005. Prey preferences of the lion (Panthera leo). Journal of Zoology. 267(03):309-322.

Innis AC. 1958. The behaviour of the giraffe, Giraffa camelopardalis, in the eastern Transvaal. Proceedings of the Zoological Society of London. 131:245-278.

Ishengoma E, Agaba M, Cavener DR. 2017. Evolutionary analysis of vision genes identifies potential drivers of visual differences between giraffe and okapi. PeerJ 5:e3145

IUCN SSC Giraffe & Okapi Specialist Group. 2016. Accessed on 28 March 2016.

Jones ML. 1982. Longevity of captive mammals. Zoologische Garten. 52:113-28.

Jones ML. 1993. Longevity of ungulates in captivity. International Yearbook. 32:159-169.

Kingdon J. 1979. East African mammals: An atlas of evolution in Africa, Vol III(B). Large Mammals. London: Academic Press.

Kingdon J. 1997. The Kingdon field guide to African mammals. San Diego: Academic Press.

Krecek RC, Boomker J, Penzhorn JL, and Scheepers L. 1990. Internal parasites of giraffes (Giraffa camelopardalis angolensis) from Etosha National Park, Namibia. Journal of Wildlife Diseases. 26(3):395-397.

Lackey BL, LaRue FV. 1997. North American regional giraffe studbook. Dallas: Dallas Zoo.

Langman VA. 1977. Cow-calf relationships in giraffe (Giraffa camelopardalis giraffa). Ethology. 43(3):264-286.

Le Pendu Y, Ciofolo I, Gosser A. 2000. The social organization of giraffes in Niger. African Journal of Ecology. 38(1):78-85.

Lee DE, Bond ML, Bolger DT. 2017. Season of birth affects juvenile survival of giraffe. 59(1):45-54.

Leroy R, Visscher M-N, Halidou O, Boureima A. 2009. The last African white giraffes live in farmers’ fields. Biodiversity and Conservation. 18(10):2663-2677.

Leuthold BM. 1979. Social organization and behavior of giraffe in Tsavo East National Park. African Journal of Ecology. 17:19-34.

Leuthold BM, Leuthold W. 1978. Ecology of the giraffe in Tsavo East National Park, Kenya. East African Wildlife Journal. 16:1-20.

Maa Dictionary (online). 2008. Maa Maasai Language Project, University of Oregon.

MacClintock D. 1973. A natural history of giraffes. New York: Charles Scribner's Sons.

Marais A, Fennessy J, Fennessy S, Brand R, Carter K. 2018. Giraffa camelopardalis spp. angolensis. The IUCN red list of threatened species [Internet]. Version 2018.3; e.T88420726A88420729 [species assessed 2018 Feb 22; page accessed 2018 Nov 20].

Meester J, Setzer HW. 1977. The mammals of Africa: An identification manual. Washington, D.C.: Smithsonian Institution Press.

Mitchell G. 2009. The origins of the scientific study and classification of giraffes. Transactions of the Royal Society of South Africa. 64(1):1-13.

Mitchell G, Skinner JD. 2009. An allometric analysis of the giraffe cardiovascular system. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology.154(4):523-529.

Mitchell G, Roberts DG, van Sittert SJ, Skinner JD. 2013. Orbit orientation and eye morphometrics in giraffes (Giraffa camelopardalis). African Zoology. 48(2):333-339.

Mramba RP, Mayhenya O, Siyaya a, Mathisen KM, Andreassen, HP, Skarpe C. 2017. Sexual segregation in foraging giraffe. Acta Oecologica 79:26-35.

Muller Z, Bercovitch F, Brand R, Brown D, Brown M, Bolger D, Carter K, Deacon F, Doherty J et al. 2016. Giraffa camelopardalis (amended version of 2016 assessment) . The IUCN red list of threatened species 2018 [Internet]. Versions 2019.2.

Muller Z, Bercovitch F, Brand R, Brown D, Bogler D, Carter K, Deacon F, Doherty JB, Fennessy J, Fennessy S et al. 2018. Giraffa camelopardalis. The IUCN red list of threatened species [Internet]. Version 2018.3; e.T9194A136266699 [species assessed 2016 Jul 09, published 2018, as an amended version of 2016 assessment; page accessed 2018 Nov 20].

Muneza A, Doherty JB, Hussein A, Fennessy J, Marais A, O'Connor D, Wube T. 2018. Giraffa camelopardalis reticulata. The IUCN red list of threatened species [Internet]. Version 2018.3; e.T88420717A88420720 [subspecies assessed 2018 Mar 16; page accessed 2018 Nov 20].

Nowak RM.. 1999. Walker's mammals of the world, Volume II. 6th ed. Baltimore (MD): The Johns Hopkins University Press.

O'Connor DA, Butt B, Foufopoulos JB. 2015. Foraging ecologies of giraffe (Giraffa camelopardalis reticulata) and camels (Camelus dromedarius) in northern Kenya: effects of habitat structure and possibilities for competition? African Journal of Ecology. 53(2):183-193.

O’Connor D, Stacy‐Dawes J, Muneza A, Fennessy J, Gobush K, Chase MJ, Brown MB, Bracis C, Elkan P, Zaberirou ARM, et al. 2019. Updated geographic range maps for giraffe, Giraffa spp., throughout sub‐Saharan Africa, and implications of changing distributions for conservation. Mam Rev.:mam.12165. doi:10.1111/mam.12165.

Parker DM, Bernard RTF. 2005. The diet and ecological role of giraffe (Giraffa camelopardalis) introduced to the Eastern Cape, South Africa. Journal of Zoology. 267(2):203-210.

Pellew RA. 1984. The feeding ecology of a selective browser, the giraffe (Giraffa camelopardalis tippelskirchi). Journal of Zoology. 202(1):57-81.

Rentsch D, Packer C. 2015. The effect of bushmeat consumption on migratory wildlife in the Serengeti ecosystem, Tanzania. Oryx. 49(2):287-294.

Ritter C. 1998. Getting to know giraffes. ZOONOOZ. 71(7):14-19.

Rubanza CDK, Shem MN, Bakengesa SS, Ichinohe T, Fujihara T. 2007. The content of protein, fibre and minerals of leaves of selected Acacia species indigenous to north-western Tanzania. Archives Of Animal Nutrition. 61(2):151-156.

Sathar F, Badlangana NL, Manger PR. 2010. Variations in the thickness and composition of the skin of the giraffe. Anatomical Record. 293(9):1615-1627.

Seeber PA, Ciofolo I, Ganswindt A. Behavioural inventory of the giraffe (Giraffa camelopardalis). BMC Research Notes. 2012(5):650.

Seeber PA, Ciofolo I, Ganswindt A. Behavioural inventory of the giraffe (Giraffa camelopardalis). BMC Research Notes. 2012(5):650. Additional files 1-8 Tables S1-S8.

Sherr L. 1997. Tall blondes: A book about giraffes. Kansas City (MI): Andrews McMeel Publishing.

Shorrocks B, Croft DP. 2009. Necks and networks: a preliminary study of population structure in the reticulated giraffe (Giraffa camelopardalis reticulata de Winston). African Journal of Ecology. 47(3):374-381.

Skinner JD, Hall-Martin AJ. 1975. A note on foetal growth and development of the giraffe Giraffa camelopardalis giraffa. Journal of Zoology. 177(1):73-79.

Spinage CA. 1968. The book of giraffe. London: Collins.

Spinage CA. 1993. The median ossicone of Giraffa camelopardalis. Journal of Zoology. 230(1):1-5.

Solounais N, Tang N. 1990. The two types of cranial appendages in Giraffa camelopardalis (Mammalia, Artiodactyla). Journal of Zoology. 222(2):293-302.

Strauss MKL, Kilewo M, Rentsch D, Packer C. 2015. Food supply and poaching limit giraffe abundance in the Serengeti. Population Ecology. 57(3):505-516.

Strauss MKL, Packer C. 2013. Using claw marks to study lion predation on giraffes of the Serengeti. Journal of Zoology. 289(2):134-142.

Suraud JP, Fennessy J, Bonnaud E, Issa AM, Fritz H, Gaillard JM. 2012. Higher than expected growth rate of the endangered West African giraffe Giraffa camelopardalis peralta: A successful human–wildlife cohabitation. Oryx. 46(4):577-583.

Toon A and Toon SB. Okapis and Giraffes (Giraffidae). In: Hutchins M, Garrison RW, Geist V, Loiselle PV, Schlager N, McDade MC, ...Duellman WE, editors. Grzimek's Animal Life Encyclopedia, Volume 15. 2nd ed. Detroit (MI): Gale; c2004. p.399-409. Retrieved from on 04 May 2016.

Trust for African Rock Art. Accessed on 07 June 2016.

UNEP. 2019. Species + [online database]. Nairobi, Kenya: UNEP [compiled by UNEP-WCMC, Cambridge, UK]. [accessed 2019 May 21].

UNEP. 2020. Species+: Giraffa camelopardalis [online database]. Nairobi, Kenya: UNEP [compiled by UNEP-WCMC, Cambridge, UK]. [accessed 2020 Nov 25].

Wexo JB. 1982. Giraffes. Zoobooks. San Diego: Wildlife Education Limited.

Wilson DE, Mittermeier RA, editors. Handbook of the mammals of the world, volume 2. Barcelona (Spain): Lynx Edicions; c2011. Family Giraffidae, Hoofed Mammals; p. 788-802.

Wube T, Doherty JB, Fennessy J, Marais A. 2018. Giraffa camelopardalis spp. camelopardalis. The IUCN red list of threatened species [Internet]. Version 2018.3; e.T88420707A88420710 [species assessed 2018 Feb 12; page accessed 2018 Nov 20].

VanderWaal KL, Wang H, McCowan B, Fushing H, Isbell LA. 2014. Multilevel social organization and space use in reticulated giraffe (Giraffa camelopardalis). Behavioral Ecology. 25(1):17-26.

Additional Internet Resources

Muzzle Touch

A giraffe calf getting kissed by mom

Belart, a Masai giraffe calf just four days old, is attended by his mother, Harriet, at the San Diego Zoo. Photo date: 19 June 2014.

Image credit: © San Diego Zoo Wildlife Alliance. All rights reserved.


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