Justification of Red List Category
This species is classified as Endangered because 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. Recent count data for the number of breeding pairs suggests that the rate of decline may actually have increased in recent years. If the estimated rate of population decline is confirmed to have accelerated, the species may require uplisting.
In 2015, the overall number of pairs was about 20,850 pairs, or 41,700 mature individuals (Sherley et al. 2019a). This roughly equates to about 66,720 individuals in adult plumage based on the conversion factor of 3.2 for pairs to individuals (Crawford and Boonstra 1994).
The population in Namibia declined from 12,162 pairs in 1978 to an estimated 5,800 pairs in 2015. The South African population declined from c.70,000 pairs in 1978/1979 (Shelton et al. 1984) to 19,300 pairs in 2015. Decreases in both countries amount to > 50% in three generations (Kemper 2015, Hagen 2016).
Spheniscus demersus is endemic to southern Africa, where it breeds at 28 localities in Namibia and South Africa (Kemper et al. 2007b, Crawford et al. 2013, Kemper 2015). It has been recorded as far north as Gabon and Mozambique (Crawford et al. 2013).
In Namibia, Neglectus Islet and Penguin Island were recolonised in 2001 and 2006 respectively (Kemper et al. 2007a). In the 1980s, the species colonised Stony Point and Boulders Beach on the South African mainland and recolonised Robben Island, all in the southwest of the country (Underhill et al. 2006). A colony formed on the southern mainland at De Hoop in 2003, but disappeared after 2007. The northernmost colony at Lambert’s Bay became extinct in 2006 (Underhill et al. 2006, Crawford et al. 2011).
In 2015, the population for Namibia was estimated at 5,700 to 5,800 pairs (MFMR unpubl. data), the uncertainty in the estimate arising from a few islands that had not been counted for several years (J. Kemper pers. comm.). The most important colonies were Mercury Island: 2,646 pairs, Ichaboe Island: 488 pairs, Halifax Island: 1,092 pairs and Possession Island: 1,205 pairs (MFMR unpubl. data).
In 2019, c.13,300 pairs bred in South Africa: St Croix Island: 3,638 pairs, Bird Island (Algoa Bay): 2,378 pairs, Dassen Island: 1,912 pairs, Stony Point: 1,705 pairs , Robben Island: 1,190 pairs, Dyer Island: 1,071 pairs, Simonstown: 932 pairs (Department of Environmental Affairs, SANParks and CapeNature unpubl. data). Just seven colonies now support 97% of the South African population. Recent declines at South African colonies are coincident with changes in the abundance and availability of forage fish and an eastward movement of spawning forage fish (Crawford et al. 2011, Waller 2011, Sherley et al. 2014a).
Behaviour Adults are largely resident, but some movements occur in response to prey movements (Hockey et al. 2005). Adults generally remain within 400 km of their breeding locality, although they have been recorded up to 900 km away (Hockey et al. 2005, Roberts 2015). They breed and moult on land before taking to the sea, where they can remain for up to four months (Crawford et al. 2013, Roberts 2015). On gaining independence, juveniles disperse up to 2,000 km from their natal colonies, with those from the east heading west, and those from the west and south moving north (Sherley et al. 2013a, Sherley et al. 2017). Most birds later return to their natal colony to moult and breed (Randall et al. 1987, Sherley et al. 2014a), although the growth of some colonies has been attributed to the immigration of first-time breeders tracking food availability (Crawford 1998, Crawford et al. 2013). Adults nest colonially, but may also nest in isolation. At sea they forage singly, in pairs or sometimes co-operatively in small groups of up to 150 individuals (Wilson et al. 1986, Kemper et al. 2007b, Ryan et al. 2012, McInnes et al. 2019). African Penguins forage more successfully in groups when feeding on schooling fish (McInnes et al. 2017). The species breeds year round with peak months varying locally (Crawford et al. 2013). In the north-western 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 between April and June (Crawford et al. 2013). The average age at first breeding is thought to be 4-6 years (Whittington et al. 2005).
Habitat This species is marine and usually found within 40 km of the coast (Wilson et al. 1988, Petersen et al. 2006, Pichegru et al. 2009, 2012), coming ashore on islands or at non-contiguous areas of the mainland coast to breed, moult and rest (Hockey et al. 2005). Breeding: Breeding habitats range from flat, sandy islands with varying degrees of vegetation cover, to steep rocky islands with little vegetation (Hockey et al. 2005). African Penguins are sometimes found close to the summit of islands and may move over a kilometre inland in search of breeding sites (Hockey 2001). They usually feed within 20 km of the colony when breeding, although at some colonies the distance is greater (Pichegru et al. 2009, Waller 2011, Ludynia et al. 2012, Pichegru et al. 2012). Non-breeding: At sea, their distribution is mainly restricted to the greater Benguela Current region (Williams 1995). Juveniles have been observed to travel ~600 km from their natal colonies (Sherley et al. 2017), while immatures up to 700 km with an average of ~370 km from the colony (Grigg and Sherley 2019). Pre- and post moulting adults have been observed up to 550 km from their colonies (de Blocq et al. 2019).
Diet Adults feed predominantly on pelagic schooling fish of 50-120 mm length, with important prey including sardine Sardinops sagax, anchovy Engraulis capensis, bearded goby Sufflogobius bibarbatus and round herring Etrumeus whiteheadi (Crawford et al. 1985, Ludynia et al. 2010, Crawford et al. 2011). In some localities, cephalopods represent an important food source (Crawford et al. 1985, Connan et al. 2016). Juveniles are thought to prey on fish larvae (Wilson 1985).
Breeding site In the past, nests were usually built in burrows dug in guano or sand (Frost et al. 1976a, Shelton et al. 1984). Today, with the lack of guano at most colonies, nesting in open areas has become increasingly common (Kemper et al. 2007b, Sherley et al. 2012, Pichegru 2013). At some sites, artificial nest-burrows made from pipes and boxes sunken into the ground, and shelters shaped from dry vegetation have been regularly used by the species (Kemper et al. 2007a, Sherley et al. 2012, Pichegru 2013).
Population declines have been attributed to food shortages resulting from shifts in the distributions of prey species, competition with commercial purse-seine fisheries and environmental fluctuations (e.g. Crawford et al. 2011). A decrease in foraging effort at St Croix Island (Pichegru et al. 2010, 2012, Sherley et al. 2019b) and an increase in chick survival and chick condition at Robben Island (Sherley et al. 2015, 2018, 2019b) following the establishment of 20 km no-take zones provides some support for this theory. In the early 2000s, there was an eastward shift in sardine and anchovy stocks, with the mature biomass of these species decreasing near the breeding islands north of Cape Town (Crawford et al. 2011). The abundance of these prey species is known to influence foraging success (Campbell et al. 2019, McInnes et al. 2019), breeding success (Crawford et al. 2006, Sherley et al. 2013b), adult survival (Sherley et al. 2014a, Robinson et al. 2015), and juvenile survival (Weller et al. 2016; Sherley et al. 2017), all of which may often be too low off South Africa’s west coast to maintain population equilibrium (Weller et al. 2014, 2016). Western Cape populations declined by 69% between 2001-2009, considered at least partly due to this climate-induced shift in fish stocks. African penguin fledglings travelled to areas of low sea surface temperatures and high chlorophyll-a which were historically reliable cues for fish availability. Climate change and industrial fishing have depleted forage fish in these areas, resulting in an ecological trap for the species and associated low juvenile survival (Sherley et al. 2017). In Namibia, where sardine and anchovy are virtually absent from the foraging ranges of breeding penguins, breeding birds feed principally on the energy-poor Bearded Goby Sufflogobius bibarbatus (Ludynia et al. 2010). Limited penguin mortality in fishing nets may increase if gill-nets are set near colonies (Ellis et al. 1998, Crawford et al. 2017).
Human disturbance and egg-collecting were important factors in the decline of the species in the early 20th century (Frost et al. 1976b, Ellis et al. 1998, Shannon and Crawford 1999). While egg collection is now illegal. an incident of egg poaching was recorded in South Africa in 2016 (Brophy 2016). Guano collection was historically a major cause of disturbance at many colonies and the removal of guano 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 predation (Frost et al. 1976b, Shannon and Crawford 1999, Pichegru 2013, Kemper 2015). Guano harvesting is no longer practiced in South Africa, and, according to the Namibian Island’s Marine Protected Area Regulations, guano scraping is not permitted following the expiry of existing guano rights for Ichaboe Island in 2016 (MRA 27 of 2000).
Both chronic oil pollution and individual large oil spills appear to have long-term significant impacts on colonies. Past mortality from oil spills has been serious (Wolfaardt et al. 2009) and may increase if proposed development of harbours close to colonies proceeds. Most of the population is confined to areas that are near existing or planned major shipping ports (Nel and Whittington 2003, J. Kemper pers. comm.). There has been a dramatic increase in the number of birds oiled since 1990: two individual oil spills (in 1994 and 2000) killed 30,000 individuals, despite successful rehabilitation programmes (Nel and Whittington 2003, Wolfaardt et al. 2008, 2009). Ship to ship bunkering activities off the south east coast in Algoa Bay in 2016 and 2019, resulted in 200 African penguins and 125 seabirds oiled respectively (SANCCOB unpubl. data). 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 2004 (Crawford et al. 2006). 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%) compared to unaffected pairs (61%) and those involving at least one bird affected by a previous oil spill (71%), mostly owing to higher mortality in older chicks (Barham et al. 2007). 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; the effects of prolonged captivity; or the time between oiling and washing (Barham et al. 2007).
The Cape Fur Seal Arctocephalus pusillus competes with penguins for food, displaces them from breeding sites and imposes significant mortality at some colonies (Crawford et al. 1989, Makhado et al. 2013, Weller et al. 2016, MFMR unpubl. data). Modelling of the interaction of multiple pressures on the colonies at Robben and Dyer Islands indicate that predation by Cape Fur Seals is a key driver in current population declines at Dyer Island (Weller et al. 2016). However, this was found to be in addition to immature emigration, suggesting there may be additional bottom-up pressures impacting the viability of colonies.
The potential effects of individual storms on breeding colonies at certain sites has been highlighted (de Villiers 2002) and, as such, the increased frequency and severity of storms may cause localised losses. Sharks take some birds at sea and Kelp Gulls Larus dominicanus, dogs Canis familiaris and feral cats Felis catus prey on eggs and chicks at colonies (Underhill et al. 2006, Pichegru 2013, Weller et al. 2014, 2016). In some mainland colonies, predation by mongooses Herpestes spp., leopards Panthera pardus and caracals Caracal caracal, or illegal egg collection may have notable impacts (e.g. Underhill et al. 2006), with the Simonstown colony experiencing considerable mortality due to caracal (SANParks and City of Cape Town, unpubl. data, Vanstreels et al. 2019).
While a number of diseases have been documented in African penguins, few records of mass mortality through disease have been observed in the wild, up until 2018 and 2019, when a high pathogenic avian influenza (HPAI) strain killed approximately 100 penguins in South Africa and up to 600 in Namibia respectively (Khomenko et al. 2018, Molini et al. 2020)
Conservation Actions Underway
CITES Appendix II. CMS Appendix II. US Endangered Species Act. Continuous monitoring of population trends is carried out at all colonies annually in South Africa but less regularly in Namibia. In South Africa, most breeding localities are national parks or nature reserves. The colonies at Simonstown and Stony Point are in the process of receiving formal protection status. Collection of guano and eggs is prohibited within penguin colonies (Harrison et al. 1997, Currie et al. 2009).
The Namibian Islands’ Marine Protected Area (NIMPA), proclaimed in 2009, protects almost 10,000 km2 of ocean in southern Namibia, including all penguin breeding localities and key foraging habitats (Currie et al. 2009, Ludynia et al. 2012).
Oiled birds are rehabilitated with success (Barham et al. 2007, Wolfaardt et al. 2008). More than 80% of birds admitted for rehabilitation are returned successfully to the wild (Nel and Whittington 2003).
Lost nesting habitat has been augmented using artificial nests at a number of colonies. Some designs have proved successful, increasing breeding success (Kemper et al. 2007a, Sherley et al. 2012). At other locations, the same designs have not been as successful (Pichegru 2013, Lei et al. 2014). The optimal design of artificial nests is currently being researched. Maintenance of natural breeding habitat takes place where possible.
Research into foraging behaviour using biologging technology (GPS and satellite-transmitters) is ongoing (Ludynia 2007, Pichegru et al. 2010, Waller 2011, Ludynia et al. 2012, Pichegru et al. 2012, Waller 2011, Sherley et al. 2013a, Campbell et al. 2019, BirdLife South Africa unpubl. data). In South Africa, a research project into the potential positive impacts of small marine no-take zones surrounding breeding colonies is underway. Results suggest a decrease in adult foraging effort and increases in chick survival and condition, but not uniformly across the colonies involved (Pichegru et al. 2010, 2012; Sherley et al. 2015, 2018, 2019b). South Africa declared new Marine Protected Areas in 2019, including around some of the seabird colonies, but they are largely ineffective in protecting penguin foraging habitat.
Population reinforcement through hand rearing of abandoned chicks, or chicks removed from nests in compromised areas where survival was unlikely, added over 7000 fledglings to the population between 2001 and 2019. These hand-reared fledglings survive and recruit in to breeding populations at similar rates to their wild counterparts (Sherley et al. 2014b). Attempts are made to decrease predation of eggs, chicks and grown birds (e.g. Makhado et al. 2013, Pichegru 2013). A national (South African) capture-mark-recapture programme using Passive Integrated Transponders has been implemented to monitor survival, recruitment and movements amongst colonies.
Conservation Actions Proposed
In South Africa, the African penguin Biodiversity Management Plan (BMP), gazetted in 2013, guided the conservation actions to be implemented with the aim to halt the decline of the species. This 5-year BMP included the above actions and identified additional ones, such as: ensuring adequate prey for penguins during the breeding and non-breeding seasons; spatial management of the pelagic fishery; investigating conservation translocations in this species; improving the disaster response to oiling, disease and fire; establish minimum standards for rehabilitation and rehabilitation facilities; improving penguin numbers through targeted interventions at existing but declining breeding localities where the reasons for the decline can be addressed.
This plan did not achieve its aims, and a revised plan has been prepared for the next 5 years and is expected to be approved for implementation in 2020. Threats such as predation and disaster prevention and mitigation are addressed in this plan as well as conservation translocations, habitat improvement and ongoing essential population monitoring and disease surveillance. Critically, actions dealing with the food availability threat with protecting at sea habitat and the management of resources that are critical for the penguin’s survival at all phases in its life-cycle are included.
60-70 cm. Medium-sized, black-and-white penguin. Adult black above, white below with variable amount of black spotting on breast and belly. Broad, black breast-band and black-and-white facial pattern diagnostic. Whitish bare skin over the eyes becomes bright pinkish-red in very hot conditions. Male has deeper, more robust bill. Juvenile initially dark slaty-blue above, turning browner and, in second and third year, shows varying amount of adult facial pattern. Similar spp. Very rarely, some individuals show a double black breast-band - indicative of Magellanic Penguin S. magellanicus, which has never been positively recorded in Africa.
Text account compilers
Martin, R., Shutes, S., Symes, A., Taylor, J., Moreno, R., Pearmain, L., Everest, J., Waller, L.
Barham, P., de Blocq, A., Crawford, R.J.M., García Borboroglu, P., Hagen , C., Kemper, J., Ludynia, K., Makhado, A., McInnes, A., Morris, T., Pichegru, L., Sherley, R.B., Simmons, R.E., Steinfurth, A., Underhill, L., Waller, L., Wanless, R. & van der Spuy, S.
BirdLife International (2021) Species factsheet: Spheniscus demersus. Downloaded from http://www.birdlife.org on 21/10/2021. Recommended citation for factsheets for more than one species: BirdLife International (2021) IUCN Red List for birds. Downloaded from http://www.birdlife.org on 21/10/2021.