Spheniscus demersus

Taxonomy

Assessment Information

Geographic Range

Population

Habitat and Ecology

Threats

Conservation Actions

Bibliography

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Taxonomy [top]

Kingdom	Phylum	Class	Order	Family
ANIMALIA	CHORDATA	AVES	SPHENISCIFORMES	SPHENISCIDAE

Scientific Name:

Spheniscus demersus

Species Authority:

(Linnaeus, 1758)

Common Name/s:

English	–	African Penguin, Black-footed Penguin, Jackass Penguin
French	–	Manchot du Cap
Spanish	–	Pingüino del Cabo

Assessment Information [top]

Red List Category & Criteria:

Endangered A2ace+3ce+4ace ver 3.1

Year Published:

2010

Assessor/s:

BirdLife International

Reviewer/s:

Butchart, S., Calvert, R. & Symes, A.

Contributor/s:

Crawford, R., Kemper, J., Pichegru, L., Simmons, R., Underhill, L. & Wanless, R.

Justification:
This species has been uplisted to Endangered because recent data has revealed that it is undergoing a very rapid population decline, probably as a result of commercial fisheries and shifts in prey populations. This trend currently shows no sign of reversing, and immediate conservation action is required to prevent further declines.

History:

2008	–	Vulnerable
2005	–	Vulnerable
2004	–	Vulnerable
2000	–	Vulnerable

Geographic Range [top]

Range Description:	Spheniscus demersus breeds at 25 islands and four mainland sites in Namibia and South Africa¹⁶. It has been recorded as far north as Gabon and Mozambique¹⁷. Breeding on Neglectus Island, Namibia, was confirmed in 2001, following the absence of confirmed breeding since 1952 at least, and an increase in numbers since 1995⁶. In 2003, there were thought to be 11 breeding pairs on the island⁶. In the 1980s, the species colonised Stony Point and Boulders Beach on the South African mainland, and recolonised Robben Island⁸. Immigration to mainland sites in recent years has been attributed to an eastward shift in the species's prey populations^12,13. Just seven islands now support 80% of the global population. Its population at the beginning of the 21st century had fallen to about 10% of its numbers 100 years before. The total population was estimated at 141,000 pairs in 1956-1957, 69,000 pairs in 1979-1980, 57,000 pairs in 2004-2005 and 36,000 pairs in 2006-2007¹⁶. Declines have continued, with the global population in 2009 estimated at just 25,262 pairs^25,28, equating to a decline of 60.5% over 28 years (three generations).
Countries:	Native: Angola; Mozambique; Namibia; South Africa; United States; United States Vagrant: Congo; Gabon
Range Map:	Click here to open the map viewer and explore range.

Population [top]

Population:	The Namibian population was estimated at c.5,000 breeding pairs in 2008, and the South African population at c.21,000 breeding pairs in 2009 (R. Crawford in litt. 2010), thus, the figure used here is 52,000 mature individuals.
Population Trend:	Decreasing

Habitat and Ecology [top]

Habitat and Ecology:	Behaviour The adults of this species are largely sedentary but some movements occur in response to prey movements²². They generally remain within 400 km of their breeding locality, although they have been recorded up to 900 km away²². They breed and moult on land before taking to the sea where they remain for four months before returning to land for the next breeding season¹⁷. Small crèches of up to five juveniles may form at the breeding site¹⁷. On gaining independence, juveniles disperse up to 1,900 km from their natal colonies²², with those from the east heading west, and those from the west and south moving north¹⁷. Most birds later return to their natal colony to moult and breed¹⁷, although the growth of some island colonies has been attributed to the immigration of first-time breeders tracking food availability^22,23. Adults nest colonially²² and at sea forages singly, in pairs, or sometimes co-operatively in small groups if up to 150 individuals^17,18,22. It breeds year round with peak months varying locally¹⁷. In the north-west part of the range, peak laying occurs during the months of November to January, in the south-west it occurs between May and July and in the East Colonial between April and June¹⁷. Habitat This species is marine, and is usually found in seas within 40 km of the shore, coming ashore on inshore islands or isolated areas of the mainland coast^17,22 to breed, moult and rest²². Breeding Breeding habitats range from flat, sandy islands with sparse or abundant vegetation, to steep rocky islands with practically no vegetation, although the former is preferred²². It is sometimes found close to the summit of islands and may move over a kilometre inland in search of breeding sites¹⁸. Non-breeding At sea its distribution is restricted to the area influenced by the Benguela Current²⁴. It usually feeds within 12 km of the coastline¹⁶. Diet Adults feed on pelagic schooling fish of 50-120mm in length¹⁷. Important prey includes sardines (Sardinops sagax), anchovies (Engraulis capensis), pelagic goby Sufflogobius bibarbatus, and herring (Etrumeus teres)^17,19. In some localities cephalopods also represent an important food source¹⁹. Juveniles tend to prey on fish larvae¹⁸. Breeding site The nest is often built in burrows that are dug in guano or sand^15,20. Nests may also occur in depressions under large boulders or bushes¹⁸. Nesting in open areas has become increasingly common owing to the past harvesting of guano¹⁵. At some sites artificial nest-burrows made from pipes and boxes sunken into the ground have been regularly used by the species²¹. The average age at first breeding is thought to be 4-6 years⁷.
Systems:	Terrestrial; Marine

Habitat and Ecology:

Behaviour The adults of this species are largely sedentary but some movements occur in response to prey movements²². They generally remain within 400 km of their breeding locality, although they have been recorded up to 900 km away²². They breed and moult on land before taking to the sea where they remain for four months before returning to land for the next breeding season¹⁷. Small crèches of up to five juveniles may form at the breeding site¹⁷. On gaining independence, juveniles disperse up to 1,900 km from their natal colonies²², with those from the east heading west, and those from the west and south moving north¹⁷. Most birds later return to their natal colony to moult and breed¹⁷, although the growth of some island colonies has been attributed to the immigration of first-time breeders tracking food availability^22,23. Adults nest colonially²² and at sea forages singly, in pairs, or sometimes co-operatively in small groups if up to 150 individuals^17,18,22. It breeds year round with peak months varying locally¹⁷. In the north-west part of the range, peak laying occurs during the months of November to January, in the south-west it occurs between May and July and in the East Colonial between April and June¹⁷. Habitat This species is marine, and is usually found in seas within 40 km of the shore, coming ashore on inshore islands or isolated areas of the mainland coast^17,22 to breed, moult and rest²². Breeding Breeding habitats range from flat, sandy islands with sparse or abundant vegetation, to steep rocky islands with practically no vegetation, although the former is preferred²². It is sometimes found close to the summit of islands and may move over a kilometre inland in search of breeding sites¹⁸. Non-breeding At sea its distribution is restricted to the area influenced by the Benguela Current²⁴. It usually feeds within 12 km of the coastline¹⁶. Diet Adults feed on pelagic schooling fish of 50-120mm in length¹⁷. Important prey includes sardines (Sardinops sagax), anchovies (Engraulis capensis), pelagic goby Sufflogobius bibarbatus, and herring (Etrumeus teres)^17,19. In some localities cephalopods also represent an important food source¹⁹. Juveniles tend to prey on fish larvae¹⁸. Breeding site The nest is often built in burrows that are dug in guano or sand^15,20. Nests may also occur in depressions under large boulders or bushes¹⁸. Nesting in open areas has become increasingly common owing to the past harvesting of guano¹⁵. At some sites artificial nest-burrows made from pipes and boxes sunken into the ground have been regularly used by the species²¹. The average age at first breeding is thought to be 4-6 years⁷.

Systems:

Terrestrial; Marine

Threats [top]

Major Threat(s):	Population declines are largely attributed to food shortages, resulting from large catches of fish by commercial purse-seine fisheries, and environmental fluctuations. A decrease in foraging effort at the St Croix Island colony following the establishment of a 20 km no-take zone provides some support for this theory²⁶. An eastward shift in sardine and anchovy populations is also blamed, with the biomass of these species near the largest breeding islands west of Cape Town falling sharply since 2002¹². The abundance of these prey species is known to influence breeding success, which may often be too low to maintain population equilibrium¹⁴. Human disturbance and egg-collecting appear to have been additional factors in the species's declines². Tourists may cause nest-burrows to collapse, and their presence in large numbers may deter young birds from breeding. Mortality from oil spills is serious and may increase if proposed development of harbours close to colonies proceeds. In addition, most of the population is confined to just two areas, both near to major shipping ports⁵. There has been a dramatic increase in the number of birds oiled since 1990: two individual oil spills (in 1994 and 2000) have killed 30,000 individuals, despite successful rehabilitation programmes⁵. In addition, breeding success on Robben island fell to 0.23 chicks per pair in 2000, compared with an average of 0.62 ±0.19 over the other 15 years from 1989 to 2004¹⁴. Without continuing rehabilitation, the population is set to decrease 17-51% over the next 20 years⁵. However, rehabilitation does not necessarily prevent problems in the years after a spill. During 2001-2005, pairs involving at least one bird rehabilitated from the oil spill in 2000 achieved lower fledging success (43%), mostly owing to higher mortality in older chicks, compared to unaffected pairs (61%) and those involving at least one bird affected by a previous oil spill (71%)⁹. This may indicate physiological or behavioural problems that reduce the parents' ability to meet the food requirements of older chicks, perhaps owing to the toxicity of the heavy oil in the 2000 spill, or the effects of prolonged captivity and time between oiling and washing⁹. Guano collection has historically been a major cause of disturbance at many colonies and its removal has deprived penguins of nest-burrowing sites, causing birds to nest on open ground where they are more vulnerable to heat stress resulting in the abandonment of nests, flooding of nests by rain and increased predation²². The cape fur seal Arctocephalus pusillus competes with penguins for food, displaces them from breeding sites and is a periodic predator. Limited mortality in fishing nets may increase if gill-nets are set near colonies². Recently the potentially major effects of individual storms on breeding colonies at certain sites has been highlighted⁴. Sharks take birds at sea and Kelp Gulls Larus dominicanus and feral cats prey on eggs and chicks at colonies¹.

Major Threat(s):

Population declines are largely attributed to food shortages, resulting from large catches of fish by commercial purse-seine fisheries, and environmental fluctuations. A decrease in foraging effort at the St Croix Island colony following the establishment of a 20 km no-take zone provides some support for this theory²⁶. An eastward shift in sardine and anchovy populations is also blamed, with the biomass of these species near the largest breeding islands west of Cape Town falling sharply since 2002¹². The abundance of these prey species is known to influence breeding success, which may often be too low to maintain population equilibrium¹⁴. Human disturbance and egg-collecting appear to have been additional factors in the species's declines². Tourists may cause nest-burrows to collapse, and their presence in large numbers may deter young birds from breeding. Mortality from oil spills is serious and may increase if proposed development of harbours close to colonies proceeds. In addition, most of the population is confined to just two areas, both near to major shipping ports⁵. There has been a dramatic increase in the number of birds oiled since 1990: two individual oil spills (in 1994 and 2000) have killed 30,000 individuals, despite successful rehabilitation programmes⁵. In addition, breeding success on Robben island fell to 0.23 chicks per pair in 2000, compared with an average of 0.62 ±0.19 over the other 15 years from 1989 to 2004¹⁴. Without continuing rehabilitation, the population is set to decrease 17-51% over the next 20 years⁵. However, rehabilitation does not necessarily prevent problems in the years after a spill. During 2001-2005, pairs involving at least one bird rehabilitated from the oil spill in 2000 achieved lower fledging success (43%), mostly owing to higher mortality in older chicks, compared to unaffected pairs (61%) and those involving at least one bird affected by a previous oil spill (71%)⁹. This may indicate physiological or behavioural problems that reduce the parents' ability to meet the food requirements of older chicks, perhaps owing to the toxicity of the heavy oil in the 2000 spill, or the effects of prolonged captivity and time between oiling and washing⁹. Guano collection has historically been a major cause of disturbance at many colonies and its removal has deprived penguins of nest-burrowing sites, causing birds to nest on open ground where they are more vulnerable to heat stress resulting in the abandonment of nests, flooding of nests by rain and increased predation²². The cape fur seal Arctocephalus pusillus competes with penguins for food, displaces them from breeding sites and is a periodic predator. Limited mortality in fishing nets may increase if gill-nets are set near colonies². Recently the potentially major effects of individual storms on breeding colonies at certain sites has been highlighted⁴. Sharks take birds at sea and Kelp Gulls Larus dominicanus and feral cats prey on eggs and chicks at colonies¹.

Conservation Actions [top]

Conservation Actions:	Conservation Actions Underway CITES Appendix II. CMS Appendix II. In South Africa, all breeding localities are national parks, nature reserves or otherwise protected. Collection of guano and eggs is prohibited within penguin colonies³. Oiled birds are rehabilitated with great success⁵. More than 80% of birds admitted for rehabilitation are returned successfully to the wild⁵. In one study it was found that the age at first breeding of five oiled and rehabilitated birds did not differ significantly from populations on Robben and Dassen Islands⁷. Since 1995 there has been a captive breeding programme at an aquarium in Cape Town, where around six birds are bred annually, and a total of 14 have been released nearby¹⁰. This programme has contributed to public awareness and education¹⁰. Research into feeding behaviour involving the use of satellite-tracking and transponders is ongoing¹¹. On Dyer Island, the effects of guano collection in the past are being mitigated by the installation of small fiberglass igloos as nest-sites, with 200 put in place by 2006 and an eventual target of 2,000, roughly one for every pair in the colony¹⁵. These artificial nest-sites are used by pairs very soon after installation, and if the programme is successful it will be expanded to other colonies in South Africa¹⁵. A research project into the potential positive impacts of small marine no-take zones surrounding breeding colonies is underway, with results in the first year showing a decrease in adult foraging effort^26,27. Conservation Actions Proposed Monitor population trends at all colonies². Continue and maintain the SANCCOB oil spill rehabilitation centre. Initiate research into the impacts of fishing and predation². Protect Namibian breeding localities². Develop plans to conserve pelagic fish resources³, namely through management of the purse-seine fishery¹⁴. Prevent oilspills from illegal cleaning of ship tanks³. Eliminate feral cats from Bird, Dassen and Robben Islands and implement measures to preclude the introduction of rats to any colonies^2,14. Investigate reintroduction techniques². Assess whether climate change is a factor in the shifting of prey populations¹¹. Consider the idea of establishing no-fishing zones around breeding islands^11,13. Consider translocating birds in reaction to shifts in food availability¹³. Maintain suitable breeding habitat¹⁴. Control the spread of disease¹⁴. Establish and then monitor 'trial colonies' close to current concentrations of food resources²⁹.

Conservation Actions:

Conservation Actions Underway
CITES Appendix II. CMS Appendix II. In South Africa, all breeding localities are national parks, nature reserves or otherwise protected. Collection of guano and eggs is prohibited within penguin colonies³. Oiled birds are rehabilitated with great success⁵. More than 80% of birds admitted for rehabilitation are returned successfully to the wild⁵. In one study it was found that the age at first breeding of five oiled and rehabilitated birds did not differ significantly from populations on Robben and Dassen Islands⁷. Since 1995 there has been a captive breeding programme at an aquarium in Cape Town, where around six birds are bred annually, and a total of 14 have been released nearby¹⁰. This programme has contributed to public awareness and education¹⁰. Research into feeding behaviour involving the use of satellite-tracking and transponders is ongoing¹¹. On Dyer Island, the effects of guano collection in the past are being mitigated by the installation of small fiberglass igloos as nest-sites, with 200 put in place by 2006 and an eventual target of 2,000, roughly one for every pair in the colony¹⁵. These artificial nest-sites are used by pairs very soon after installation, and if the programme is successful it will be expanded to other colonies in South Africa¹⁵. A research project into the potential positive impacts of small marine no-take zones surrounding breeding colonies is underway, with results in the first year showing a decrease in adult foraging effort^26,27.

Conservation Actions Proposed
Monitor population trends at all colonies². Continue and maintain the SANCCOB oil spill rehabilitation centre. Initiate research into the impacts of fishing and predation². Protect Namibian breeding localities². Develop plans to conserve pelagic fish resources³, namely through management of the purse-seine fishery¹⁴. Prevent oilspills from illegal cleaning of ship tanks³. Eliminate feral cats from Bird, Dassen and Robben Islands and implement measures to preclude the introduction of rats to any colonies^2,14. Investigate reintroduction techniques². Assess whether climate change is a factor in the shifting of prey populations¹¹. Consider the idea of establishing no-fishing zones around breeding islands^11,13. Consider translocating birds in reaction to shifts in food availability¹³. Maintain suitable breeding habitat¹⁴. Control the spread of disease¹⁴. Establish and then monitor 'trial colonies' close to current concentrations of food resources²⁹.

Bibliography [top]

	Barham, P. J. ; Underhill, L. G.; Crawford, R. J. M.; Leshoro, T. M. 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. Barnes, K. N. 2000. The Eskom Red Data Book of birds of South Africa, Lesotho and Swaziland. BirdLife South Africa, Johannesburg. Crawford, R. 1998. Revised Red Data Book. Bird Numbers 7(2): 8-9. Crawford, R.J.M.; Barham, P.J.; Underhill, L. G.; Shannon, L.J.; Coetzee, J.C.; Dyer, B.M.; Leshoro, T.M.; 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, R.J.M., Dyer, B.M. and Brooke, R.K. 1994. Breeding nomadism in southern African seabirds: constraints, causes and conservation. Ostrich 65(2): 231-246. del Hoyo, J.; Elliot, A.; Sargatal, J. 1992. Handbook of the Birds of the World, vol. 1: Ostrich to Ducks. Lynx Edicions, Barcelona, Spain. de Villiers, M. 2002. Effect of a storm on breeding African Penguins Spheniscus demersus at Foxy Beach, Boulders Penguin Colony, Simon's Town. Bird Numbers 11: 7-9. Ellis, S.; Croxall, J. P.; Cooper, J. 1998. Penguin conservation assessment and management plan: report from the workshop held 8-9 September 1996, Cape Town, South Africa. IUCN/SSC, Apple Valley, USA. Hanes, S. 2006. Penguins get a helping home from fiberglass 'igloos'. Available at: http://www.usatoday.com/tech/science/2006-10-22-penguins-africa_x.htm#http://www.usatoday.com/tech/science/2006-10-22-penguins-africa_x.htm#. Harrison, J. A.; Allan, D. G.; Underhill, L. G.; Herremans, M.; Tree, A. J.; Parker, V.; Brown, C. J. 1997. The atlas of southern African birds. BirdLife South Africa, Johannesburg. Hockey, P. 2001. Struik Winchester, Cape Town, South Africa. Hockey, P. A. R.; Dean, W. R. J.; Ryan, P. G. 2005. Roberts birds of southern Africa. Trustees of the John Voelcker Bird Book Fund, Cape Town, South Africa. Kemper, J.; Underhill, L. G.; Crawford, R. J. M.; Kirkman, S. P. 2007. Revision of the conservation status of seabirds and seals breeding in the Benguela Ecosystem. In: Kirkman, S. P. (ed.), 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. Nel, D. C.; Whittington, P. A. 2003. Rehabilitation of oiled African Penguins: a conservation success story. Petersen, S. L.; Ryan, P. G.; 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.; Grémillet, D.; Crawford, R. M. J.; Ryan, P. G. 2010. Marine no-take zone rapidly benefits Endangered penguin. Biology Letters http://dx.doi.org/10.1098/rsbl.2009.0913. Roux, J.-P.; Kemper, J.; Bartlett, P. A.; Dyer, B. M.; Dundee, B. L. 2003. African penguins Spheniscus demersus recolonise a fomerly abandoned nesting locality in Namibia. Marine Ornithology 31: 203-205. Shelton, P.A., Crawford, R.J.M., Cooper, J., Brooke, R.K. 1984. Distribution, population size and conservation of the Jackass Penguin Spheniscus demersus. South African Journal of Marine Science 2: 217-257. 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, T. D. 1995. The penguins Spheniscidae. Oxford University Press, Oxford.

Barham, P. J. ; Underhill, L. G.; Crawford, R. J. M.; Leshoro, T. M. 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.

Barnes, K. N. 2000. The Eskom Red Data Book of birds of South Africa, Lesotho and Swaziland. BirdLife South Africa, Johannesburg.

Crawford, R. 1998. Revised Red Data Book. Bird Numbers 7(2): 8-9.

Crawford, R.J.M.; Barham, P.J.; Underhill, L. G.; Shannon, L.J.; Coetzee, J.C.; Dyer, B.M.; Leshoro, T.M.; 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, R.J.M., Dyer, B.M. and Brooke, R.K. 1994. Breeding nomadism in southern African seabirds: constraints, causes and conservation. Ostrich 65(2): 231-246.

del Hoyo, J.; Elliot, A.; Sargatal, J. 1992. Handbook of the Birds of the World, vol. 1: Ostrich to Ducks. Lynx Edicions, Barcelona, Spain.

de Villiers, M. 2002. Effect of a storm on breeding African Penguins Spheniscus demersus at Foxy Beach, Boulders Penguin Colony, Simon's Town. Bird Numbers 11: 7-9.

Ellis, S.; Croxall, J. P.; Cooper, J. 1998. Penguin conservation assessment and management plan: report from the workshop held 8-9 September 1996, Cape Town, South Africa. IUCN/SSC, Apple Valley, USA.

Hanes, S. 2006. Penguins get a helping home from fiberglass 'igloos'. Available at: http://www.usatoday.com/tech/science/2006-10-22-penguins-africa_x.htm#http://www.usatoday.com/tech/science/2006-10-22-penguins-africa_x.htm#.

Harrison, J. A.; Allan, D. G.; Underhill, L. G.; Herremans, M.; Tree, A. J.; Parker, V.; Brown, C. J. 1997. The atlas of southern African birds. BirdLife South Africa, Johannesburg.

Hockey, P. 2001. Struik Winchester, Cape Town, South Africa.

Hockey, P. A. R.; Dean, W. R. J.; Ryan, P. G. 2005. Roberts birds of southern Africa. Trustees of the John Voelcker Bird Book Fund, Cape Town, South Africa.

Kemper, J.; Underhill, L. G.; Crawford, R. J. M.; Kirkman, S. P. 2007. Revision of the conservation status of seabirds and seals breeding in the Benguela Ecosystem. In: Kirkman, S. P. (ed.), 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.

Nel, D. C.; Whittington, P. A. 2003. Rehabilitation of oiled African Penguins: a conservation success story.

Petersen, S. L.; Ryan, P. G.; 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.; Grémillet, D.; Crawford, R. M. J.; Ryan, P. G. 2010. Marine no-take zone rapidly benefits Endangered penguin. Biology Letters http://dx.doi.org/10.1098/rsbl.2009.0913.

Roux, J.-P.; Kemper, J.; Bartlett, P. A.; Dyer, B. M.; Dundee, B. L. 2003. African penguins Spheniscus demersus recolonise a fomerly abandoned nesting locality in Namibia. Marine Ornithology 31: 203-205.

Shelton, P.A., Crawford, R.J.M., Cooper, J., Brooke, R.K. 1984. Distribution, population size and conservation of the Jackass Penguin Spheniscus demersus. South African Journal of Marine Science 2: 217-257.

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, T. D. 1995. The penguins Spheniscidae. Oxford University Press, Oxford.

Citation:	BirdLife International 2010. Spheniscus demersus. In: IUCN 2011. IUCN Red List of Threatened Species. Version 2011.2. <www.iucnredlist.org>. Downloaded on 22 May 2012.
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