African Penguin is assessed as Critically Endangered because it is undergoing an extremely rapid population decline, probably principally because of the impacts of competition with commercial fisheries and climate-mediated shifts in prey populations. Recent, near-complete count data for the number of breeding pairs show an alarming acceleration in the rate of decline and the current and future projected population reduction exceeds 80% over three generations. This trend currently shows no sign of reversing, and immediate conservation action is required.

Spheniscus demersus is endemic to southern Africa, where it breeds at 26 localities in Namibia and South Africa (Crawford et al. 2013, Kemper 2015, Makhado et al. 2024). 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) and Sylvia Hill had 12 confirmed breeding pairs in 2022/2023. 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. It is currently being re-established by BirdLife South Africa through translocation of chicks bolstered from other colonies and there is active breeding since 2022 (four breeding pairs were counted in 2023: BirdLife South Africa, unpublished data).

The northernmost colony at Lambert’s Bay became extinct in 2006 (Underhill et al. 2006, Crawford et al. 2011) and the colony at Marcus Island c. 100 km to the south, which once held over 1,000 pairs, may also have gone extinct around 2018 (Makhado et al. 2024). Consequently there is an estimated continuing decline in the extent of occurrence and the area of occupancy.

The population in Namibia had been relatively stable at around 5,000 breeding pairs between 1997 and 2017 but had significantly reduced to an estimated 1,200 pairs in 2023 (Sherley et al. 2024). The most important colony, Mercury Island, which held over half of the Namibian population for much of the last four decades (Kemper et al. 2007a, Kemper 2015), had more than 2,500 pairs in 2015 but no breeding pairs in 2023 (Sherley et al. 2024). None of the other colonies, including historically important sites like Ichaboe Island (13 pairs), Halifax Island (603 pairs) and Possession Island (366 pairs), held more than 650 breeding pairs in 2023 (Sherley et al. 2024).

In South Africa, 8,536 breeding pairs were counted in 2023: Dassen Island, 2037 pairs; Bird Island (Algoa Bay), 1,722 pairs; Stony Point, 1,200 pairs; Robben Island, 931 pairs; Dyer Island, 903 pairs; Simon’s Town, 845 pairs; St Croix Island, 712 pairs; and 186 pairs at six other colonies that each held less than 65 pairs (Sherley et al. 2024). The seven named colonies support almost 98% of the South African population and nine colonies (with Halifax Island and Possession Island in Namibia) support >95% of the global population. Recent declines at South African colonies are coincident with changes in the abundance and availability of forage fish, competition with commercial fisheries and the establishment of ship-to-ship bunkering activities in Algoa Bay (Crawford et al. 2011, 2019; Waller 2011; Sherley et al. 2014a, 2018, Pichegru et al. 2022).

In 2023, the overall number of pairs was about 9,900 pairs, or 19,800 mature individuals (Sherley et al. 2024). This roughly equates to about 31,680 individuals in adult plumage based on the conversion factor of 3.2 for pairs to individuals (Crawford and Boonstra 1994). This compares to historic estimates of 141,000 pairs in 1956/57, 69,000 pairs in 1979/80, 57,000 pairs in 2004/05, 36,000 pairs in 2006/07 (Kemper et al. 2007a) and 17,700 pairs in 2019 (Sherley et al. 2020).

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, Carpenter-Kling et al. 2022). They breed and moult on land before taking to the sea, where they can remain for up to four months (Crawford et al. 2013, Carpenter-Kling et al. 2022). 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, Robinson 2015). 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, Ryan et al. 2012, McInnes et al. 2019). When feeding on schooling fish, African Penguins forage more successfully in groups of conspecifics (McInnes et al. 2017) or with other seabirds (Sutton et al. 2020). 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, 2013; Campbell et al. 2019; McInnes et al. 2019), 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) and, rarely, inside caves (Simmons and Kemper 2003). African Penguins are sometimes found close to the summit of islands and may move over a kilometre inland in search of breeding sites. 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 ~1,400 km from their natal colonies (Sherley et al. 2013a), 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 600 km from their colonies (Carpenter-Kling et al. 2022). 

Diet Adults feed predominantly on pelagic schooling fish of 50-120 mm length, with important prey including sardine Sardinops sagax, anchovy Engraulis capensis and round herring Etrumeus whiteheadi (Crawford et al. 1985, Crawford et al. 2011). In Namibia, where sardine and anchovy are now nearly absent, bearded goby Sufflogobius bibarbatus is the main prey (Ludynia et al. 2010) and 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 and the species regularly uses shelters shaped from dry vegetation (Kemper et al. 2007b,c, Sherley et al. 2012, Pichegru 2013). At most breeding sites, artificial burrows made from cement pipes, wooden boxes or fibreglass burrows are provided by managing institutions to increase breeding habitat availability and ultimately breeding success (Pichegru et al. in review).

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, 2019, 2022). 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; Sydeman et al. 2021) 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, 2019). The abundance or availability 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), chick growth (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; Leith et al. 2022; Sherley et al. 2024). West coast populations declined by 69% between 2001-2009, driven at least in part by demographic responses to this climate-induced shift in fish stocks (Robinson et al. 2015, Sherley et al. 2017). 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). Egg collection is now illegal. 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 (Weller et al. 2014). 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, Pichegru et al. 2022). 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). Overall, 71 oil spills along the coast of Namibia and South Africa affected 83,224 seabirds until 2021, 91% of which were African Penguins (Vanstreels et al. 2023). Although the magnitude of oil spills has decreased overtime, the numbers of birds affected have not (Vanstreels et al. 2023). Ship to ship bunkering activities off the south east coast in Algoa Bay between 2016 and 2023 resulted in 200 African penguins and 125 other seabirds oiled respectively (SANCCOB unpubl. data).  The number of oiled birds is relative to the proximity of the spill but not the volume spilled, underpinning the need for activities with risks of oil spills to be located away from penguin colonies (Vanstreels et al. 2023). Following a major oil spill, 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 oil toxicity; the effects of prolonged captivity; or the time between oiling and washing (Barham et al. 2007).

In addition to oil spill risks, increased maritime traffic linked to the initiation of ship-to-ship bunkering in Algoa Bay doubled underwater noise pollution, making Algoa Bay one of the noisiest bays globally (Pichegru et al. 2022). The neighbouring African Penguin colony on St Croix Island concomitantly collapsed, with a 90% decrease since 2015 (Sherley et al. 2024).

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 (Ludynia et al. 2014).

The potential effects of individual storms on breeding colonies at certain sites has been highlighted (de Villiers 2002, Russo et al. in press) 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 (Vanstreels et al. 2019, Rhoda 2022), whilst there are anecdotal reports of bees stinging penguins to death (SANParks unpublished data). 

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). Since 2021, an additional c. 230 African penguins died due to a new strain of avian influenza in South Africa (Western Cape State Veterinary Services, unpublished data). 

Previously egg-collecting and guano harvesting were carried out at a significant scale, but both have now ceased and are prohibited or illegal. The predominant use of the species now is as a tourist attraction, with visits to colonies a major draw for national and international wildlife tourism (Van Zyl and Kinghorn 2018). Colonies are also desirable subjects for the film industry, generating significant revenue.

Conservation Actions Underway
CITES Appendix II. CMS Appendix II. US Endangered Species Act. In South Africa, all breeding localities are under 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). 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. In 2023, additional purse-seine fishing exclusions were declared around some of the largest remaining African Penguin colonies (Bega 2023, Makhado et al. 2024), but again these are expected to be ineffective in protecting the birds’ foraging habitat due to their limited size and inadequate design (McInnes et al. 2023).

Continuous monitoring of population trends is carried out at all colonies annually in South Africa but less regularly in Namibia.

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: prohibition of fisheries from six major colonies, 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 all of its aims, especially regarding halting the decline of the species. It improved cohesion and collaborations across institutions to manage the conservation actions, as well as scientific research to assess these actions (Pichegru 2024). However, stronger measures especially around securing prey availability were not implemented due to a division between governmental departments and mandates (Pichegru 2024).

Population reinforcement through hand rearing of abandoned chicks and eggs, or chicks removed from nests in compromised areas where survival was unlikely, added over 8,000 fledglings to the population between 2001 and 2022. These hand-reared fledglings survive and recruit into breeding populations at similar rates to their wild counterparts (Sherley et al. 2014b, SANCCOB unpublished data). Attempts are made to decrease predation of eggs, chicks and grown birds by managing kelp gulls, seals and terrestrial mammal predation (e.g., Makhado et al. 2013; Pichegru 2013, 2024). Lost nesting habitat has been augmented using artificial nests at a number of colonies in South Africa and Namibia for over 30 years with varying success (Sherley et al. 2012, Pichegru 2013, Lei et al. 2014). Recent results, however, demonstrated an overall increase by 16% in African Penguins breeding success in artificial nests compared to natural nests, but optimum nest designs were colony-specific (Pichegru et al. in review). Maintenance of natural breeding habitat takes place where possible.
Oiled birds are rehabilitated with success (Barham et al. 2007, Wolfaardt et al. 2008, Vanstreels et al. 2023). More than 80% of birds admitted for rehabilitation are returned successfully to the wild (Nel and Whittington 2003).
A national (South African) capture-mark-recapture programme using Passive Integrated Transponders has been implemented to monitor survival, recruitment and movements amongst colonies (e.g., Leith et al. 2022) and is being extended to Namibia. A disease risk assessment was conducted in 2016 (Parsons and Vanstreels 2016) and disease surveillance is ongoing (Roberts et al. 2023b, SANCCOB unpubl. data). The use of vaccinations against avian influenza has been trialled in African penguins (Roberts et al. 2023a).
Research into foraging behaviour using biologging technology (GPS and satellite-transmitters) is ongoing (Pichegru et al. 2010, Waller 2011, Ludynia et al. 2012, Pichegru et al. 2012, Sherley et al. 2017, Campbell et al. 2019, Carpenter-Kling et al. 2022). In South Africa, a large-scale experiment into the potential positive impacts of small marine no-take zones surrounding breeding colonies was conducted between 2008 and 2020. 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; Sydeman et al. 2021, 2022).

Translocation Guidelines for African Penguins were published in 2019 (Waller et al. 2019). Norms and Standards for Rehabilitation Centres for Seabirds were gazetted for public comments in 2019 and expected to be published in 2024. A National Oil Spill Contingency Plan was published in 2022, while the National Oiled Wildlife Contingency Plan is currently being finalised.

Conservation Actions Proposed
A revised African penguin Biodiversity Management Plan (BMP) for South Africa has been prepared for the next 10 years and is expected to be approved for implementation in 2024. Threats such as predation management and disaster prevention and mitigation are addressed in this plan as well as conservation translocations, ongoing habitat improvement and essential population monitoring and disease surveillance. Importantly, this includes actions dealing with the food availability threat through protecting at-sea habitat and the management of resources that are critical for the penguin’s survival at all phases in its life-cycle.

Ongoing monitoring of population trend and health status of all colonies is crucial. Continued research is required into impacts of threats, including fishing, predation, climate change and effects of underwater noise on the species is required. Further investigations into the effects of fishing exclusion zones are also necessary, and into joining up research on life-cycle movements and feeding habitat usage. Ensure all colonies are protected from development and impacts from adjacent shipping activity, and protect foraging habitat around the seven largest colonies, including establishing no purse-seine fishing zones. Instigate spatial management of fisheries in the non-breeding season to ensure sufficient prey availability. 
Continued managed at colonies is needed including the continuation of the reintroduction programme at the De Hoop colony, further improvements to artificial burrows, addressing predation by individual problematic animals, controlling disease outbreaks and acting to rescue and rehabilitate oiled individuals through maintaining rehabilitation centres. Establish a Risk Assessment for translocation and reintroduction of captive bred birds hatched from wild birds that could not be released.
With the deterioration in conservation status there is a need to intensify and co-ordinate awareness raising and communication initiatives. There is a need to lobby for legislation in order to successfully implement an ecosystem approach to fisheries, as well as ensuring the formal establishment of MPAs around breeding colonies that include the foraging habitat of breeding birds, while colony-specific oiled wildlife contingency plans are also required. Enforcing existing legislation on the cleaning of oil tanks of ships is vital to prevent many oil spills, and national oil contingency plans may need improving.