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African Penguin (Spheniscus demersus) Fact Sheet: Bibliography & Resources

San Diego Zoo Wildlife Alliance Library fact sheet for the African penguin, Spheniscus demersus


Adams N. 1994. Patterns and impacts of oiling of African Penguins Spheniscus demersus: 1981-1991. Biological Conservation. 68: 35-41.

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].

Barham PJ, Underhill LG, Crawford RJM, Leshoro TM. 2007. Differences in breeding success between African Penguins (Spheniscus demersus) that were and were not oiled in the MV Treasure oil-spill in 2000. Emu.107: 7-13.

Berruti A. 1986. The predatory impact of feral cats Felis catus and their control on Dassen Island. South African Journal of Antarctic Research. 16(3): 123-7.

Bertelli S, Giannini NP. 2005. A phylogeny of extant penguins (Aves: Sphenisciformes) combining morphology and mitochondrial sequences. Cladistics. 21: 209-239.

BirdLife International [cited 07 Nov 2013]. Spheniscus demersus. In: IUCN 2013. IUCN Red List of threatened species. Version 2013.1. Available from:

BirdLife International. 2016. Spheniscus demersus. The IUCN Red List of Threatened Species 2016: e.T22697810A93641269. [accessed 2018 Jun 20]. <>.

BirdLife International. 2018. Spheniscus demersus. The IUCN Red List of Threatened Species 2016: e.T22697810A132604504. [accessed 2019 Apr 10]. <>.

BirdLife International. 2020. Spheniscus demersus. The IUCN red list of threatened species. Version 2023.1; e.T22697810A157423361 [species assessed 2019 Sep 09; page accessed 2024 Mar 22]. doi:10.2305/IUCN.UK.2020-3.RLTS.T22697810A157423361.en

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

Brown LH, Urban EK, Newman K. 1982. The Birds of Africa, Vol. I. Academic Press, New York.

Clark BD, Bemis W. 1979. Kinematics of swimming of penguins at the Detroit Zoo. Journal of Zoology. 188:411-428.

Clarke JA, Olivero EB, Puerta P. 2003. Description of the earliest fossil penguin from South America and first paleogene vertebrate locality of Tierra Del Fuego, Argentina. American Museum Novitates. 3423:1-18.

Cooper J. 1971. The Jackass Penguin. Marine Pollution Bulletin. 2(4):52-52

Cooper J. 1977. Jackass Penguins sunning at sea. Auk. 94: 586-587.

Cooper J. 1978. Moult of the black footed penguin. International Zoo Yearbook. 18:22-27.

Cott HB. 1953. The exploitation of wild birds for their eggs. The Ibis. 95(3): 409-449.

Crawford RJM. 2013. Long-term change in the population sizes and conservation status of South Africa’s seabirds. Ostrich. 84(2): v-ix.

Crawford RJM, Shelton PA.1978. Pelagic fishes and sea bird interactions off the coasts of south west and south Africa. Biological Conservation. 14:85-109.

Crawford RJM, Shelton PA. 1981. Population trends for some southern African seabirds related to fish availability. In: Proceedings of the Symposium on the birds of the sea and shore, 1979. Editors J Cooper, Cape Town, African Seabird Group. Pp 15-41.

Crawford RJM, Williams AJ, Randall RM, Randall BM, Berruti A, Ross GJB. 1990. Recent population trends of Jackass Penguins Spheniscus demersus off southern Africa. Biological Conservation. 52: 229-243.

Crawford RJM, Allwright DM, He l CW. 1992. High mortality of cape cormorants (Phalacrocorax capensis) off western South Africa in 1991 caused by Pasteurella multocida. Colonial Waterbirds. 15: 236–238.

Crawford RJM, Shannon LJ, Whittington PA. 1999. Population dynamics of the African Penguin Spheniscus demersus at Robben Island, South Africa. Marine Ornithology. 27: 139-147.

Crawford RJM, Shannon LJ, Whittington PA, Murison G. 2000a. Factors influencing growth of the African Penguin colony at Boulders, South Africa, 1985–1999. South African Journal of Marine Science. 22: 111–119.

Crawford RJM, Davis SA, Harding RT, Jackson LF, Leshoro TM, Meyer MA, Randall RM, Underhill LG, Upfold L, van Dalsen AP, van der Merwe E, Whittington PA, Williams AJ, Wolfaardt AC. 2000b. Initial impact of the Treasure oil spill on seabirds off western South Africa. South African Journal of Marine Science. 22: 157–176.

Crawford RJM, David JHM, Shannon LJ, Kemper J, Klages NTW, Roux J-P, Underhill LG, Ward VL, Williams AJ, Wolfaardt AC. 2001. African Penguins as predators and prey – coping (or not) with change. South African Journal of Marine Science. 23: 435–447.

Crawford RJM, Barham PJ, Underhill LG, Shannon LJ, Coetzee JC, Dyer BM, Leshoro TM, Upfold L. 2006. The influence of food availability on breeding success of African Penguins Spheniscus demersus at Robben Island, South Africa. Biological Conservation. 132(1): 119-125.

Crawford RJM, Altwegg R, Barham BJ, Barham PJ, Durant JM, Dyer BM, Geldenhuys D, Makhado .B, Pichegru L, Ryan PG, Underhill LG, Upfold L, Visagie J, Waller LJ, Whittington PA. 2011. Collapse of South Africa’s penguins in the 21st century: a consideration of food availability. African Journal of Marine Science. 33: 139–156.

Crissey S., Slifka K, McGill P. 2005. Chapter 5: diet and nutrition. In: Penguin Taxon Advisory Group: Penguin Husbandry Manual, 3rd Edition. American Association of Zoo and Aquarium Association.

David JHM. 1987. Diet of the South African fur seal (1974–1985) and an assessment of competition with fisheries in southern Africa. In: Payne AIL, Gulland JA, Brink KH, editors. The Benguela and Comparable Ecosystems. South African Journal of Marine Science. pp. 693–713.

David JHM, Cury P, Crawford RJM, Randall RM, Underhill LG, Meyer MA. 2003. Assessing conservation priorities in the Benguela ecosystem, South Africa: analyzing predation by seals on threatened seabirds. Biological Conservation. 114, 289–292.

Davis LS, Speirs EAH. 1990. Mate choice in penguins. In: Davis LS, Darby JT, editors. Penguin Biology. San Diego (California): Academic Press.

de V. Reitz JB. 1971. The Jackass Penguin of South Africa – a threatened species. Biological Conservation. 4(1): 73.

deKock AC, Randall RM. 1984. Organochlorine insecticide and polychlorinated biphenyl residues in eggs of coastal birds from the eastern cape, South Africa. 1984. Environmental Pollution Series-A, Ecological and Biological. 35: 193-201.

Dunning JB Jr. 2008. Body masses of birds of the world. In: Dunning JB Jr, editor. CRC Handbook of Avian Body Masses, 2nd Ed. New York (New York): CRC Press, Taylor and Francis Group. p. 13.

Dyer BM. 1996. Predation by snakes on seabirds at three South African islands. South African Journal of Marine Science. 17:309-313.

Eggleton P, Siegfried WR. 1979. Displays of the Jackass Penguin. Ostrich 50:139-167.

Ellis S. 2005. Chapter 3: behavior and social organization. In Penguin Taxon Advisory Group: Penguin Husbandry Manual, 3rd Edition. American Association of Zoo and Aquarium Association.

Falla RA, Mougin JL. 1979. Order Sphenisciformes. In: Peters JL, editor. Check-List of Birds of the World. 2nd Edition, vol. I. Cambridge (Massachusetts): Museum of Comparative Zoology. 121-133.

Fordyce RE, Jones CM. 1990. Penguin history and new fossil material from New Zealand. In: Davis LS, Darby JT, editors. Penguin Biology. San Diego (California): Academic Press. Pp 419-446.

Frost PGH, Siegfried WR, Burger AE. 1976a. Behavioural adaptations of the Jackass Penguin Spheniscus demersus to a hot, arid environment. Journal of Zoology. 179: 165-187.

Frost PGH, Siegfried WR, Cooper J. 1976b. Conservation of the Jackass Penguin. Biological Conservation. 9: 79-99.

Hacket SJ, Kimball RT, Reddy S, Bowie RCK, Braun EL, Braun MJ, Chojnowski JL, Cox WA, Han KL, Harshman J, Huddleston CJ, Marks BD, Miglia KJ, Moore WS, Sheldon FH, Steadman DW, Witt CC, Yuri T. 2008. A phylogenomic study of birds reveals their evolutionary history. Science. 320:1763-1768.

Hagen C, McInnes A, Waller LJ, Stephenson PJ. 2021. Spheniscus demersus (Green Status assessment). The IUCN Red List of Threatened Species. Version 2023.1; e.T22697810A2269781020221 [species assessed 2021 Oct 31; page accessed 2023 Mar 22].

Handbook of the Birds of the World Alive. (HBW). [cited Oct 2013]. Available from:

Harrison JA, Allan DG, Underhill LG, Herremans M, Tree AJ, Parker V, Brown CJ. 1997. The atlas of southern African birds. Johannesburg: BirdLife South Africa.

Heath RGM, Randall RM. 1985. Growth of Jackass Penguin chicks (Spheniscus demersus) hand reared on different diets. Journal of Zoology. 205:91-105.

Heath RGM, Randall RM. 1989. Foraging ranges and movements of Jackass Penguins (Spheniscus demersus) established through radio telemetry. Journal of Zoology. 217: 367–379.

Henry L, Sirpenski G. 2005. Chapter 4: reproduction. In: Penguin Taxon Advisory Group: Penguin Husbandry Manual, 3rd Edition. American Zoo and Aquarium Association.

Hui CA. 1987. The porpoising of penguins: an energy-conservation behavior for respiratory ventilation? Canadian Journal of Zoology. 65: 209-211.

Integrated Taxonomic Information System database (ITIS). [cited Nov 2013]. Spheniscus demersus (Linnaeus, 1758). Available from:

International Tanker Owners Pollution Federation Limited (ITOPF). [cited Oct 2013]. Available from:

Kemper J, Underhill LG, Crawford RJM, Kirkman SP. 2007. Revision of the conservation status of seabirds and seals breeding in the Benguela ecosystem. In: Kirkman SP, editor. Final Report of the BCLME (Benguela Current Large Marine Ecosystem). Pp. 325-342.

Koenig R. 2007. African Penguin populations reported in a puzzling decline. Science. 315(5816): 120.

Koning C. 1997. Coastal oil spill: Apollo sea shipping disaster - June 1994. Journal of Contingencies and Crisis Management. 5(2): 188-122.

Ksepka DT, Bertelli S, Giannini NP. 2006. The phylogeny of the living and fossil Sphenisciformes (penguins). Cladistics. 22(5):412-441.

la Cock GD, Duffy DC, Cooper J. 1987. Population dynamics of the African Penguin Spheniscus demersus at Marcus Island in the Benguela upwelling ecosystem: 1979-85. Biological Conservation. 40: 117-126.

Livezey BC. 1989. Morphometric patterns in recent and fossil penguins (Aves, Sphenisciformes). Journal of Zoology. 219:269-307.

Louw GJ. 1992. Functional anatomy of the penguin flipper. Journal of the South African Veterinary Association. 63(3):113-120.

Makhado AB, Crawford RJM, Waller LJ, Underhill LG. 2013. An assessment of the impact of predation by Cape fur sealsArctocephalus pusillus pusilluson seabirds at Dyer Island, South Africa. Ostrich. 84(3):191-198.

Marks MA, Brooke RK, Gildenhuys AM. 1997. Cape fur seal Arctocephalus pusillus predation on cape cormorants Phalacrocorax capensis and other birds at Dyer Island, South Africa. Marine Ornithology. 25, 9–12.

Martinez I. 1992. Order Sphenisciformes. In: Handbook of the Birds of the World, vol. 1: Ostrich to Ducks. Barcelona (Spain): Lynx Edicions.

Martínez I, Christie DA, Jutglar F, Garcia EFJ. 2017. African Penguin (Spheniscus demersus). In: del Hoyo J, Elliott A, Sargatal J, Christie DA, de Juana E (eds.). Handbook of the birds of the world alive. Lynx Edicions, Barcelona. Accessed 2017 Feb 01;

Mori Y. 1999. A note on swimming group size in captive African Penguins (Spheniscus demersus) in relation to weather conditions. Applied Animal Behavior Science. 62: 359-364.

Morris W, Morris M. 1988. Morris dictionary of word and phrase origins, second edition. New York (New York): Harper and Row Publishers.

Nagy KA, Siegfried WR, Wilson RP. 1984. Energy utilization by free-ranging Jackass Penguins, Spheniscus demersus. Ecology. 65(5): 1648-1655.

Nel DC, Whittington PA. 2003. Rehabilitation of oiled African Penguins: a conservation success story. Cape Town: BirdLife South Africa and Avian Demography Unit.

Paredes R, Zavalaga CB. 2001. Nesting sites and nest types as important factors for the conservation of Humboldt penguins (Spheniscus humboldti). Biological Conservation. 100:  199-205.

Petersen SL, Ryan PG, Gremillet D. 2006. Is food availability limiting African Penguins Spheniscus demersus at Boulders? A comparison of foraging effort at mainland and island colonies. Ibis. 148: 14-26.

Pichegru L, Gremillet D, Crawford RJM, Ryan PG. 2010. Marine no-take zone rapidly benefits endangered penguin. Biology Letters. 6: 498-501.

Rand RW. 1959. The cape fur seal, Arctocephalus pusillus, distribution, abundance and feeding habits off the south western coast of the Cape Province. Division of Sea Fisheries, South Africa, Investigational Report. 34: 1–75.

Randall RM, Randall BM. 1986. The diet of Jackass Penguins Spheniscus demersus in Algoa Bay, South Africa, and its bearing on population declines elsewhere. Biological Conservation. 37: 119-134.

Randall BM, Randall RM, Compagno LJV. 1988. Injuries to Jackass Penguins (Spheniscus demersus) - evidence for shark involvement. Journal of Zoology. 214(4):589-599.

Roux J-P, Kemper J, Bartlett PA, Dyer BM, Dundee BL. 2003. African Penguins Spheniscus demersus recolonize a formerly abandoned nesting locality in Namibia. Marine Ornithology. 31: 203-205.

Rutgers A, Norris KA. 1970. Encyclopaedia of Aviculture, vol. I. London: Blandford Press. Pp 65-69.

Seddon PJ, van Heezik YM. 1991a. Hatching asynchrony and brood reduction in the Jackass Penguin: an experimental study. Animal Behaviour. 42(3): 347-356.

Seddon PJ, van Heezik Y. 1991b. Effects of hatching order, sibling asymmetries, and nest site on survival analysis of Jackass Penguin chicks. The Auk. 108(3): 548.

Seddon PJ, van Heezik Y. 1991c. Patterns of nest relief during incubation by Jackass Penguins Spheniscus demersus. Ostrich. 62(1-2):82-83.

Seddon PJ, van Heezik Y. 1993a. Chick creching and intraspecific aggression in the Jackass Penguin. Journal of Field Ornithology. 64(1):90-95.

Seddon PJ, van Heezik Y. 1993b. Parent-offspring recognition in the Jackass Penguin. Journal of Field Ornithology. 61(4): 27-31.

Seddon PJ, van Heezik Y. 1993c. Behaviour of the Jackass Penguin chick. Ostrich.64: 8-12.

Shelton PA, Crawford RJM, Cooper J, Brooke RK. 1984. Distribution, population size and conservation of the Jackass Penguin Spheniscus demersus. South African Journal of Marine Science. 2: 217-257.

Sherley RB, Barham BJ, Barham PJ, Leshoro TM, Underhill LG. 2012. Artificial nests enhance the breeding productivity of African Penguins (Spheniscus demersus) on Robben Island, South Africa. Emu. 112: 97-106.

Sherley RB, Underhill LG, Barham BJ, Barham PJ, Coetzee JC, Crawford RJM, Dyer BM, Leshoro TM, Upfold L. 2013. Influence of local and regional prey availability on breeding performance of African penguins Spheniscus demersus. Marine Ecology Progress Series 473:291-301.

Sibley CG, Monroe BL Jr. 1990. Distribution and taxonomy of birds of the world. New Haven (Connecticut): Yale University Press. Pp 318-320.

Siegfried WR, Frost PGH, Kinahan JB, Cooper J. 1975. Social behaviour of Jackass Penguins at sea. Zoology Africa. 10: 87-100.

Simmons RE, Kemper J. 2003. Cave breeding by African Penguins near the northern extreme of their range: Sylvia Hill, Namibia. Ostrich. 74(3-4):217-221.

Simpson GG. 1975. Fossil penguins. In: Bernard Stonehouse, editor. The Biology of Penguins. Baltimore (Maryland): University Park Press. Pp: 19-41.

South African Foundation for the Conservation of Coastal Birds (SANCCOB) [website]. [accessed Oct 2013].

South African National Biodiversity Institute (SANBI); Department of Environmental Affairs and Tourism, Republic of South Africa. [accessed Nov 2013]. National Environmental Management Act.

Sparks J, Soper T. 1987. Penguins. New York (New York): Facts on File Publications.

Spellerberg IF. 1975. The predators of penguins. In: Bernard Stonehouse, editor. The Biology of Penguins. Baltimore (Maryland): University Park Press. Pp: 413-434.

UNEP. 2018. Species+: Spheniscus demersus [online database]. Nairobi, Kenya: UNEP [compiled by UNEP-WCMC, Cambridge, UK]. [accessed 2018 Jun 26].

Whittington PA, Dyer BM, Klages NTW. 2000. Maximum longevities of African Penguins Spheniscus demersus based on banding records. Marine Ornithology 28(1): 81–82.

Whittington P, Klages N, Crawford R, Wolfaardt A, Kemper J. 2005. Age at first breeding of the African Penguin. Ostrich 76(1&2): 14-20.

Williams TD. 1995. The penguins Spheniscidae. Oxford: Oxford University Press.

Williams TD, Cooper J. 1984. Aspects of the breeding biology of the Jackass Penguin Spheniscus demersus. Proceedings of the Fifth Pan-African Ornithological Congress. Pp 841-853.

Wilson RP, Wilson MPT. 1990. Foraging ecology of breeding Spheniscus penguins. In: Davis LS, Darby JT, editors. Penguin Biology. San Diego (California): Academic Press. Pp 181-206.

Wilson RP, Wilson MPT, McQuaid L. 1986. Group-size in foraging African Penguins (Spheniscus demersus). Ethology. 72(4):338-341.

Wolfaardt AC, Underhill LG, Altwegg R, Visagie J. 2008. Restoration of oiled African penguins Spheniscus demersus a decade after the Apollo Sea spill. African Journal of Marine Science. 30(2):421-436.

Wolfaardt AC, Underhill LG, Visagie J. 2009. Breeding and moult phenology of African Penguins Spheniscus demersus at Dassen Island. African Journal of Marine Science. 31(2): 119-132.

Yom-Tov Y, Wilson R, Amos AR. 1986. Water loss from Jackass Penguin Spheniscus demersus eggs during natural incubation. Ibis. 128: 1-8.

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