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).

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).

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.