Biology:African penguin

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Short description: Species of bird

African penguin
African penguin side profile.jpg
At Boulders Beach in Simon’s Town, South Africa
CITES Appendix II (CITES)[2]
Scientific classification edit
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Sphenisciformes
Family: Spheniscidae
Genus: Spheniscus
Species:
S. demersus
Binomial name
Spheniscus demersus
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  Extant (resident)
  Extant at-sea range (resident)
African penguin distribution en.jpg
Distribution of the African penguin
Synonyms

Diomedea demersa Linnaeus, 1758

The African penguin (Spheniscus demersus), also known as Cape penguin or South African penguin, is a species of penguin confined to southern African waters. Like all penguins, it is flightless, with a streamlined body and wings stiffened and flattened into flippers for a marine habitat. Adults weigh an average of 2.2–3.5 kg (4.9–7.7 lb) and are 60–70 cm (24–28 in) tall. The species has distinctive pink patches of skin above the eyes and a black facial mask. The body's upper parts are black and sharply delineated from the white underparts, which are spotted and marked with a black band.

The African penguin is a pursuit diver and feeds primarily on fish and squid. Once extremely numerous, the African penguin is declining rapidly due to a combination of several threats and is classified as endangered. It is a charismatic species and is popular with tourists. Other vernacular names of the species include black-footed penguin and jackass penguin, due to the species' loud, donkey-like noise[3] (although several related species of South American penguins produce the same sound). They can be found along the coast of South Africa and Namibia.

Taxonomy

In 1747, the English naturalist George Edwards included an illustration and a description of the African penguin in the second volume of his A Natural History of Uncommon Birds. He used the English name "The Black-Footed Penguins". Edwards based his hand-coloured etching on two preserved specimens that had been brought to London. He suspected that they had been collected near the Cape of Good Hope.[4] When in 1758 the Swedish naturalist Carl Linnaeus updated his Systema Naturae for the tenth edition, he placed the African penguin with the wandering albatross in the genus Diomedea. Linnaeus included a brief description, coined the binomial name Diomedea demersa and cited Edwards' work.[5] The African penguin is now placed with the banded penguins in the genus Spheniscus that was introduced in 1760 by the French zoologist Mathurin Jacques Brisson.[6][7] The genus name Spheniscus is from Ancient Greek word σφήν (sphēn) meaning "wedge" and is a reference to the animal's thin, wedge-shaped flippers. The specific epithet demersus is Latin meaning "plunging" (from demergere meaning "to sink").[8]

Banded penguins are found mainly in the Southern Hemisphere with the Humboldt penguin and Magellanic penguins found in southern South America and the Galápagos penguin found in the Pacific Ocean near the equator. All are similar in shape, colour and behaviour.

File:Penguin Colony Betty's Bay.ogv

Description

African penguin diving in Siam Center, Bangkok, Thailand
African penguin skeleton at the Museum of Osteology in Oklahoma City, Oklahoma, United States

African penguins grow to 60–70 cm (24–28 in) tall and weigh between 2.2–3.5 kg (4.9–7.7 lb).[9] The beak length of the African penguin varies, usually growing between 20–30 cm (7.9–11.8 in). They have a black stripe and black spots on the chest, the pattern of which is unique to each penguin, like human fingerprints. The sweat glands above the eyes cool the birds' blood and as the temperature rises, increased blood flow causes the glands to get pinker.[10] This species exhibits slight sexual dimorphism; the males are slightly larger than the females and have longer beaks.[11] Juveniles do not possess the bold, delineated markings of the adult, but instead have dark upperparts that vary from greyish-blue to brown; the pale underparts lack both spots and the band. The beak is more pointed than that of the Humboldt penguin. The African penguin's colouring is a form of protective colouration known as countershading. The white undersides of the birds are difficult to spot by predators under the water and the penguins' black backs blend in with the water when viewed from above.

African penguins resemble and are thought to be related to the Humboldt, Magellanic and Galápagos penguins.[12] African penguins have a very recognisable appearance, with a thick band of black that is in the shape of an upside-down horseshoe. They have black feet and black spots that vary in size and shape between individuals. Magellanic penguins share a similar bar marking that often confuses the two; the Magellanic has a double bar on the throat and chest, whereas the African has a single bar. These penguins have the nickname "jackass penguin", which comes from the loud penguin noises they make.

Distribution and habitat

The African penguin is only found on the southwestern coast of Africa, living in colonies on 24 islands between Namibia and Algoa Bay, near Port Elizabeth, South Africa.[1] It is the only penguin species that breeds in Africa and its presence gave name to the Penguin Islands.

Penguin with chicks, Boulders Beach, South Africa

Two colonies were established by penguins in the 1980s on the mainland near Cape Town, namely Boulders Beach near Simon's Town and Stony Point in Betty's Bay. Mainland colonies likely became possible only in recent times due to the reduction of predator numbers, although the Betty's Bay colony has been attacked by leopards.[13][14] The only other mainland colony is in Namibia, but it is not known when it was established.

Boulders Beach is a tourist attraction due to the beach, swimming and the penguins.[15][16] The penguins will allow people to approach them as close as a meter.

Breeding populations of African penguins are being kept in numerous zoos worldwide. No colonies are known outside the southwestern coast of Africa, although vagrants (mostly juveniles) may occasionally be sighted beyond the normal range.

Population

Roughly 4 million African penguins existed at the beginning of the 19th century. Of the 1.5 million African penguins estimated in 1910, only some 10% remained at the end of the 20th century. African penguin populations, which breed in Namibia and South Africa, have declined by 95% since pre-industrial times.[17]

Today, their breeding is largely restricted to 24 islands from Namibia to Algoa Bay, South Africa,[18] with the Boulders Beach colony being an exception to this rule.

The total population fell to approximately 150,000–180,000 in 2000.[19][20] Of those, 56,000 belonged to the Dassen Island colony and 14,000 to the Robben Island colony.[21] The colony at Dyer Island in South Africa fell from 46,000 in the early 1970s to 3,000 in 2008.[22]

In 2008, 5,000 breeding pairs were estimated to live in Namibia.

In 2010, the total African penguin population was estimated at 55,000. At the rate of decline seen from 2000 to 2010, the African penguin is expected to be extinct in the wild by 2026.[23]

In 2012, about 18,700 breeding pairs were estimated to live in South Africa, with the majority on St. Croix Island in Algoa Bay.[1][24]

The total breeding population across both South Africa and Namibia fell to a historic low of about 20,850 pairs in 2019.[25]

Behaviour

Diet

African penguins forage in the open sea, where they feed on pelagic fish such as sardines (including the blue pilchard), Cape horse mackerels, red-eye round herrings[26] and anchovies (specifically the European anchovy and the Southern African anchovy)[27][26] and marine invertebrates such as squids and small crustaceans, primarily krills and shrimps.[28] Penguins normally swim within 20 km (12 mi) of the shore.[9] A penguin may consume up to 540 grams (1.19 lb) of prey every day,[29] but this may increase to over 1 kg (2.2 lb) when raising older chicks.[28]

Due to the marked decline of sardines in the waters near its habitat, African penguins' diet has shifted towards anchovies to some extent, although available sardine biomass is still a notable determinant of penguin population development and breeding success. While a diet of anchovies appears to be generally sufficient for the penguins, it is not ideal due to anchovies' lower concentrations of fat and protein. The Penguin's diet changes throughout the year; as in many seabirds, it is believed that the interaction of diet choice and breeding success helps the penguins maintain their population size. Although parent penguins are protective of their chicks, they will not incur nutritional deficits themselves if food is scarce and hunting requires a greater time or energy commitment. This may lead to higher rates of brood loss under poor food conditions.

When foraging, African penguins carry out dives that reach an average depth of 25 m (82 ft) and last for 69 seconds, although a maximum depth of 130 m (430 ft) and duration of 275 seconds has been recorded.[30]

Breeding

Egg from the collection of Museum Wiesbaden, Wiesbaden, Hesse, Germany
Nesting burrows of the African penguin, Boulders Beach (2017)

The African penguin is monogamous; it breeds in colonies and pairs return to the same site each year. It has an extended breeding season, with nesting usually peaking from March to May in South Africa and November to December in Namibia.[27] A clutch of two eggs is laid either in burrows burrowed in guano or nests in the sand under boulders or bushes. Incubation is undertaken equally by both parents for around 40 days. At least one parent guards the chicks for about one month, whereafter the chicks join a crèche with other chicks and both parents spend most of the day foraging in the sea.

Chicks fledge at 60 to 130 days, the timing depending on environmental factors such as the quality and availability of food. The fledged chicks then go to sea on their own, where they spend the next one to nearly two years. They then return to their natal colony to moult into adult plumage.[31]

When penguins moult, they are unable to forage in the sea as their new feathers are not yet waterproof; therefore, they fast over the entire moulting period.[31] African penguins typically take around three weeks to moult and lose about half of their body weight by burning up their fat reserves in the process.[32]

African penguins spend most of their lives at sea until it comes time for them to lay their eggs. Females remain fertile for about 10 years. Due to high predation on the mainland, African penguins will seek protection on offshore islands, where they are safer from larger mammals and natural challenges. These penguins usually breed during the winter when temperatures are cooler. African penguins often will abandon their eggs if they become overheated in the hot sun and abandoned eggs never survive the heat. The eggs are three to four times bigger than chicken eggs. Ideally, the eggs are incubated in a burrow dug into the guano layer (which provides suitable temperature regulation), but the widespread human removal of guano deposits has rendered this type of nest unfeasible in many colonies. To compensate, penguins burrow holes in the sand, nest under rocks or bushes or make use of nest boxes if they are provided. The penguins spend three weeks on land caring for their offspring, after which chicks may be left alone during the day while the parents forage. The chicks are frequently killed by predators or succumb to the hot sun. Parents usually feed hatchlings during dusk or dawn.

In 2015, when foraging conditions were favourable, more male than female African penguin chicks were produced in the colony on Bird Island. Male chicks also had higher growth rates and fledging mass and therefore may have higher post-fledging survival than females. This, coupled with higher adult female mortality in this species, may result in a male-biased adult sex ratio and may indicate that conservation strategies focused on benefiting female African penguins may be necessary.[33]

Predation

The average lifespan of an African penguin is 10 to around 25 years in the wild and up to 30 in captivity.[34]

The primary predators of African penguins at sea include sharks and fur seals. While nesting: kelp gulls, Cape genets, mongooses, caracals and domestic cats and dogs may prey on the penguins and their chicks.[35][36] Mortality from terrestrial predators is higher if penguins are forced to breed in the open in the absence of suitable burrows or nest boxes.

Threats and conservation

African penguin at the New England Aquarium, Boston, Massachusetts , United States

File:Spheniscus demersus 2010.ogv

Historical exploitation

African penguin eggs were considered a delicacy and were still being eaten and collected for sale as recently as the 1970s. In the 1950s, they were being collected from Dassen Island and sold in nearby towns.[37] In 1953, 12,000 eggs were collected.[38] In the late 1950s, some French chefs expressed interest in recipes including African penguin eggs collected from the islands off the west coast of South Africa and placed annual orders for small quantities.[39][40][41] In the mid-1960s, eggs were collected in the thousands and sold by the dozen,[42] with each customer limited to two dozen eggs in total.[43]

The practice of collecting African penguin eggs involved smashing those found a few days before a collecting effort to ensure that only freshly laid eggs were sold. This added to the drastic decline of the African penguin population around the Cape coast, a decline which was hastened by the removal of guano from islands for use as fertiliser, eliminating the burrowing material used by penguins.[44]

Oil spills

Penguins remain susceptible to pollution of their habitat by petrochemicals from spills, shipwrecks and cleaning of tankers while at sea. Accounts of African penguins impacted by oil date back to the 1930s.[45] African penguins' exposure to oil spills is both chronic (higher frequency small discharges of oil at sea) and acute (rare maritime disasters where large volumes of oil are released in a single event). Penguins of many species have been impacted by oil spills across the southern hemisphere.

In 1948, the tanker Esso Wheeling sank, subsequently oiling and killing thousands of penguins of the Dyer Island colony.[46] In 1953, dead penguins were among a range of dead birds, fish and other marine life that washed ashore after the tanker Sliedrecht was holed and spilled oil near Table Bay.[47] In 1971, the SS Wafra oil spill impacted the African penguin colony of Dyer Island. In 1972, oil spilt following the Oswego-Guardian and Texanita collision oiled roughly 500 penguins.[48] In 1975, newspapers reported that oil pollution from shipwrecks and the pumping of bilges at sea had killed tens of thousands of African penguins. At the time, the Dassen Island colony was being passed by 650 oil tankers each month[49] because the Suez Canal had become blocked with wrecked vessels, thus increasing maritime traffic past the Cape of Good Hope.[48]

In 1979, an oil spill prompted the collection and treatment of 150 African penguins from St. Croix Island near Port Elizabeth. The animals were later released at Robben Island and four of them promptly swam back to St. Croix Island, surprising scientists.[50][51]

In 1983, the exposure of penguins of Dassen Island to the oil slick from the Castillo de Bellver was also a topic of concern given the penguins' conservation status at the time, but owing to the prevailing wind and current, only gannets were oiled.[52]

1994 MV Apollo Sea disaster

African penguin casualties were significant following the sinking of the MV Apollo Sea and a subsequent oil slick in 1994. 10,000 penguins were collected and cleaned, of which less than half survived.[53]

2000 MV Treasure crisis

Disaster struck on 23 June 2000, when the iron ore tanker MV Treasure sank between Robben Island and Dassen Island, South Africa. It released 400–1,000 tonnes (390–980 long tons; 440–1,100 short tons) of fuel oil, causing an unprecedented coastal bird crisis and oiling 19,000 adult penguins at the height of the best breeding season on record for this vulnerable species.[54] The oiled birds were brought to an abandoned train repair warehouse in Cape Town to be cared for. An additional 19,500 un-oiled penguins were removed from Dassen Island and other areas before they became oiled and were released about 800 kilometres east of Cape Town. This gave workers enough time to clean up the oiled waters and shores before the birds could complete their long swim home (which took the penguins between one and three weeks). Some of the penguins were named and radio-tracked as they swam back to their breeding grounds. Tens of thousands of volunteers helped with the rescue and rehabilitation process, which was overseen by the International Fund for Animal Welfare (IFAW) and the South African Foundation for the Conservation of Coastal Birds (SANCCOB) and took more than three months to complete. This was the largest animal rescue event in history; more than 91% of the penguins were successfully rehabilitated and released – an amazing feat that could not have been accomplished without such a tremendous international response.[55]

Due to the positive outcome of African penguins being raised in captivity after tragedies such as the Treasure oil spill, the species is considered a good "candidate for a captive-breeding programme which aims to release offspring into the wild"; however, worry about the spread of new strains of avian malaria is a major concern in the situation.[56]

Bringing the birds inland led to the exposure of penguins to parasites and disease vectors such as mosquitoes carrying avian malaria, which has caused 27% of the rehabilitated penguin deaths annually.[57]

2016 & 2019 Port of Ngqura

Small-scale oil spills (of less than 400 litres (110 US gal)) have occurred at the Port of Ngqura since bunkering activities started there in 2016. Bunkering is a ship refuelling process that can result in oil spills and oil slicks entering the water. Hundreds of African penguins have been harmed following these spills[58] due to the port's close proximity to penguin rookeries on St. Croix Island and seabird habitat on neighbouring Jahleel and Brenton Islands.

Competition with fisheries

Commercial fisheries of sardines and anchovy, the two main prey species of the penguins, have forced these penguins to search for prey farther offshore, as well as having to switch to eating less nutritious prey.[17] Restricting commercial fishing near colony sites such as Robben Island for short periods (3 years) was shown to markedly improve penguin breeding success. Longer closure periods and closures near other colonies are being evaluated.[59][60][61]

Conservation status

The African penguin is one of the species to which the African-Eurasian Waterbird Agreement (AEWA) applies. In September 2010, it was listed as endangered under the US Endangered Species Act.[17] As of 2018, the African penguin is listed as endangered on the IUCN Red List. Its population is approximately 50,000 birds and declining.[1]

Mediation efforts

Many organisations such as SANCCOB, Dyer Island Conservation Trust, SAMREC, and Raggy Charters with the Penguin Research Fund in Port Elizabeth are working to halt the decline of the African penguin. Measures include: monitoring population trends, hand-rearing and releasing abandoned chicks, establishing artificial nests and proclaiming marine reserves in which fishing is prohibited.[1] Some colonies (such as on Dyer Island) are suspected to be under heavy pressure from predation by Cape fur seals and may benefit from the culling of individual problem animals,[60][62] which has been found effective (although requiring a large amount of management effort) in trials.[63]

Established in 1968, SANCCOB is currently the only organisation mandated by the South African government to respond to crises involving seabirds along South Africa's coastline and is internationally recognised for the role it played during the MV Treasure oil spill. A modelling exercise conducted in 2003 by the University of Cape Town's FitzPatrick Institute of African Ornithology found that rehabilitating oiled African penguins has resulted in the current population being 19% larger than it would have been in the absence of SANCCOB's rehabilitation efforts.[64]

In February 2015, the Dyer Island Conservation Trust opened the African Penguin and Seabird Sanctuary (APSS) in Gansbaai, South Africa.[65] The centre was opened by then-Department of Tourism minister Derek Hanekom[66] and will serve as a hub for seabird research carried out by the Dyer Island Conservation Trust. The centre will also run local education projects, host international marine volunteers and seek to improve seabird handling techniques and rehabilitation protocols.

Captivity

African penguin at the Dallas World Aquarium, Dallas, Texas , United States

African penguins are a commonly seen species in zoos across the world. Because they do not require particularly low temperatures, they are often kept in outside enclosures. They adapt fairly well to this captive environment and are rather easy to breed compared to other species of the family. In Europe, the breeding programme EAZA is regulated by Artis Royal Zoo in the Netherlands, whilst in the United States the SSP programme is cooperatively managed by the AZA. The idea is to create a backup captive population, as well as to aid in the conservation of the population in its natural habitat. Between 2010 and 2013, American zoos spent $300,000 on in situ (wild population) conservation.[67][68]

See also

Gallery

  • Nesting African penguin

  • Colony of African penguins nesting on Boulders Beach in Cape Town

  • African penguins on Boulders Beach

  • African penguins on a rock at Boulders Beach

  • African penguin part of the Betty's Bay colony

  • Adult (front) and juvenile (back)

References

  1. ↑ 1.0 1.1 1.2 1.3 1.4 BirdLife International (2020). "Spheniscus demersus". IUCN Red List of Threatened Species 2020: e.T22697810A157423361. doi:10.2305/IUCN.UK.2020-3.RLTS.T22697810A157423361.en. https://www.iucnredlist.org/species/22697810/157423361. Retrieved 11 September 2023. 
  2. ↑ "Appendices | CITES". https://cites.org/eng/app/appendices.php. 
  3. ↑ Favaro, Livio; Ozella, Laura; Pessani, Daniela; Pavan, Gianni (30 July 2014). "The Vocal Repertoire of the African Penguin (Spheniscus demersus): Structure and Function of Calls". PLOS ONE 9 (7): e103460. doi:10.1371/journal.pone.0103460. PMID 25076136. Bibcode: 2014PLoSO...9j3460F. 
  4. ↑ Edwards, George (1747). A Natural History of Uncommon Birds. Part II. London: Printed for the author at the College of Physicians. p. 94, Plate 94. https://www.biodiversitylibrary.org/page/50240832. 
  5. ↑ Linnaeus, Carl (1758) (in Latin). Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. 1 (10th ed.). Holmiae (Stockholm): Laurentii Salvii. p. 132. https://www.biodiversitylibrary.org/page/727037. 
  6. ↑ Brisson, Mathurin Jacques (1760) (in fr, la). Ornithologie, ou, Méthode contenant la division des oiseaux en ordres, sections, genres, especes & leurs variétés. 1. Paris: Jean-Baptiste Bauche. Vol. 1, p. 42, Vol. 6, p. 96. 
  7. ↑ Gill, Frank; Donsker, David; Rasmussen, Pamela, eds (July 2021). "Kagu, Sunbittern, tropicbirds, loons, penguins". IOC World Bird List Version 11.2. International Ornithologists' Union. https://www.worldbirdnames.org/bow/loons/. 
  8. ↑ Jobling, James A. (2010). The Helm Dictionary of Scientific Bird Names. London: Christopher Helm. pp. 132, 361. ISBN 978-1-4081-2501-4. 
  9. ↑ 9.0 9.1 Sinclair, Ian; Hockey, Phil; Tarboton, Warwick; Ryan, Peter (2011). Sasol Birds Of South Africa. Struik. p. 22. ISBN 9781770079250. 
  10. ↑ Mahard, Tyler (2012). "The Black-footed Penguin Spheniscus demersus". Wildlife Monthly. http://wildlifemonthly.blogspot.com/2012_02_01_archive.html. 
  11. ↑ "African Penguin (Spheniscus demersus)". Dyer Island Conservation Trust. http://www.dict.org.za/penguin.php. 
  12. ↑ a-z animals. "The African Penguin". a-z animals. http://a-z-animals.com/animals/african-penguin/. 
  13. ↑ "The African Penguin". Bettysbay. 2010-04-08. http://www.bettysbay.info/index.php?option=com_content&view=article&id=24&Itemid=26. 
  14. ↑ "CapeNature increases protection from predators at Stony Point". CapeNature. http://www.capenature.co.za/%E2%80%8Bcapenature-increases-%E2%80%8B%E2%80%8Bprotection-predators-stony-point/. 
  15. ↑ "Table Mountain National Park". SANParks. http://www.sanparks.org/parks/table_mountain/tourism/attractions.php#boulders. 
  16. ↑ "Boulders Beach, Swimming with Penguins – Swimming with Penguins in South Africa". Goafrica. 2010-06-14. http://goafrica.about.com/od/capetownatravelguide/a/bouldersbeach.htm. 
  17. ↑ 17.0 17.1 17.2 Kilduff, Catherine (2010-09-28). "Vanishing African Penguin, Threatened by Climate Change and Fishing, Wins Protections". seaturtles.org. http://www.seaturtles.org/article.php?id=1800. 
  18. ↑ "Know your Georgia Aquarium penguins (2006)". The Atlanta Constitution: pp. G6. 2006-11-17. https://www.newspapers.com/clip/50561897/know-your-georgia-aquarium-penguins/. 
  19. ↑ "Penguins begin swim home (Treasure, 2000)". York Daily Record: pp. 6. 2000-10-12. https://www.newspapers.com/clip/50561727/penguins-begin-swim-home-treasure/. 
  20. ↑ "Penguins elude oil spill but fishermen open fire, South Africa (2000)". Calgary Herald: pp. 12. 2000-07-21. https://www.newspapers.com/clip/54568076/penguins-elude-oil-spill-but-fishermen/. 
  21. ↑ "Flight of the African penguin (2000)". Tampa Bay Times: pp. 2. 2000-07-08. https://www.newspapers.com/clip/50561218/flight-of-the-african-penguin-2000/. 
  22. ↑ "Experiments seek to house declining South African penguins (2009)". The Boston Globe: pp. A6. 2009-03-30. https://www.newspapers.com/clip/50561597/experiments-seek-to-house-declining/. 
  23. ↑ "African Penguin; Endangered; Cape Town". Globalpost.com. 2011-06-19. http://www.globalpost.com/dispatch/news/regions/africa/south-africa/110617/african-penguins-endangered-species-cape-town. 
  24. ↑ Makhado, A. B., B. N. Dyer, R. Fox, D. Geldenhuys, L. Pichegru, R. M. Randall, R .B. Sherley, L. Upfold, J. Visagie., L. J. Waller, P. A. Whittington, and R. J .M. Crawford. 2013. Estimates of numbers of twelve seabird species breeding in South Africa, updated to include 2012. Branch Oceans and Coasts, Department of Environmental Affairs, Cape Town.
  25. ↑ Sherley, R. B.; Crawford, R. J.; de Blocq, A. D.; Dyer, B. M.; Geldenhuys, D.; Hagen, C.; Kemper, J.; Makhado, A. B. et al. (2020). "The conservation status and population decline of the African penguin deconstructed in space and time.". bioRxiv 10 (15): 8506–8516. doi:10.1101/2020.01.15.907485. PMID 32788996. 
  26. ↑ 26.0 26.1 "Spheniscus demersus (Jackass penguin)". https://animaldiversity.org/accounts/Spheniscus_demersus/. 
  27. ↑ 27.0 27.1 "African penguin videos, photos and facts – Spheniscus demersus". ARKive. http://www.arkive.org/african-penguin/spheniscus-demersus/#text=Biology. 
  28. ↑ 28.0 28.1 "The African Penguin Simons Town". Simonstown.com. http://www.simonstown.com/tourism/penguins/penguins.htm. 
  29. ↑ Crawford, R. J. M.; Ryan, P. G.; Williams, A. J. (1991). "Seabird consumption and production in the Benguela and western Agulhas ecosystems". South African Journal of Marine Science 11 (1): 357–375. doi:10.2989/025776191784287709. 
  30. ↑ Ropert-Coudert Y; Kato A; Robbins A; Humphries GRW (October 2018). African Penguin (Spheniscus demersus). Unpublished. doi:10.13140/RG.2.2.32289.66406. http://www.penguiness.net. 
  31. ↑ 31.0 31.1 Wolfaardt, A. C.; Underhill, L. G.; Visagie, J. (2009). "Breeding and moult phenology of African penguins Spheniscus demersus at Dassen Island". African Journal of Marine Science 31 (2): 119–132. doi:10.2989/AJMS.2009.31.2.1.873. 
  32. ↑ "African Penguin". http://www.torontozoo.com/animals/African%20Penguin. 
  33. ↑ Spelt, A.; Pichegru, L. (2017). "Sex allocation and sex-specific parental investment in an endangered seabird". Ibis 159 (2): 272–284. doi:10.1111/ibi.12457. 
  34. ↑ "ADW: Spheniscus demersus: Information". Animaldiversity.ummz.umich.edu. 2010-02-01. http://animaldiversity.ummz.umich.edu/site/accounts/information/Spheniscus_demersus.html. 
  35. ↑ Whittington, P. A.; Hofmeyr, J. H.; Cooper, J. (1996). "Establishment, growth and conservation of a mainland colony of Jackass Penguins Spheniscus demersus at Stony Point, Betty's Bay, South Africa". Ostrich 67 (3–4): 144–150. doi:10.1080/00306525.1996.9639700. 
  36. ↑ "African penguin". Neaq.org. http://www.neaq.org/animals_and_exhibits/animals/african_penguin/index.php. 
  37. ↑ "Dassen Island African penguins (1955)". The Pantagraph: pp. 10. 1955-01-09. https://www.newspapers.com/clip/50560484/dassen-island-african-penguins-1955/. 
  38. ↑ "Penguin playmates & egg exports, Dassen Island (1954)". Press and Sun-Bulletin: pp. 4. 1954-10-23. https://www.newspapers.com/clip/50560421/penguin-playmates-egg-exports-dassen/. 
  39. ↑ "French Gourmet Finds Penguin Egg Recipes (1959)". The Racine Journal-Times Sunday Bulletin: pp. 17. 1959-08-09. https://www.newspapers.com/clip/50557919/french-gourmet-finds-penguin-egg/. 
  40. ↑ "Penguin-egg eating may spread (1959)". The Troy Record: pp. 14. 1956-07-23. https://www.newspapers.com/clip/50558213/penguin-egg-eating-may-spread-1959/. 
  41. ↑ "Penguin eggs sent to French eaters (1959)". The Daily Oklahoman: pp. 48. 1959-05-24. https://www.newspapers.com/clip/50559867/penguin-eggs-sent-to-french-eaters/. 
  42. ↑ "Penguins' Eggs Again To Grace Gourmets' Tables (1965)". Arizona Daily Star: pp. 32. 1965-05-30. https://www.newspapers.com/clip/50561011/penguins-eggs-again-to-grace-gourmets/. 
  43. ↑ "Must Ration Penguin Eggs at $2.10 a Dozen (1965)". Sioux City Journal: pp. 49. 1965-05-09. https://www.newspapers.com/clip/50561159/must-ration-penguin-eggs-at-210-a/. 
  44. ↑ Shannon, L. J.; Crawford, R. J. M. (1999). "Management of the African penguin Spheniscus demersus — insights from modelling". Marine Ornithology 27: 119–128. 
  45. ↑ "Oily penguin, South Africa (1936)". Hartford Courant: pp. 70. 1936-08-02. https://www.newspapers.com/clip/50561469/oily-penguin-south-africa-1936/. 
  46. ↑ "Oil wrecks penguins of Dyer Island (Esso Wheeling, 1949)". The Morning Call: pp. 22. 1949-04-29. https://www.newspapers.com/clip/54879646/oil-wrecks-penguins-of-dyer-island/. 
  47. ↑ "Ship Oil Kills Fish, Penguins". Newcastle Sun (NSW: 1918-1954): pp. 8. 1953-11-03. http://nla.gov.au/nla.news-article161508121. 
  48. ↑ 48.0 48.1 "Brief History of Penguin Oiling in South African Waters | Animal Demography Unit" (in en). http://www.adu.uct.ac.za/adu/projects/sea-shore-birds/oil-spill/history. 
  49. ↑ "South Africa's penguins die from oil pollution effects (1975)". Arizona Republic: pp. 1. 1975-05-01. https://www.newspapers.com/clip/50559428/south-africas-penguins-die-from-oil/. 
  50. ↑ ""Homing" Penguins Travel 500 Miles (1979)". The Charlotte News: pp. 4. 1979-09-17. https://www.newspapers.com/clip/50558328/homing-penguins-travel-500-miles/. 
  51. ↑ "Penguins baffle scientists (1979)". The Daily News: pp. 27. 1979-09-18. https://www.newspapers.com/clip/50559316/penguins-baffle-scientists-1979/. 
  52. ↑ "Split Supertanker Gets Slow Tow Out To Sea (Castillo de Bellver oil spill, 1983)". The Baytown Sun: pp. 6. 1983-08-08. https://www.newspapers.com/clip/50557292/split-supertanker-gets-slow-tow-out-to/. 
  53. ↑ Wolfaardt, A. C.; Underhill, L. G.; 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. doi:10.2989/ajms.2008.30.2.14.564. 
  54. ↑ "Penguins, rescue, rehabilitation, research, internships". Sanccob. http://www.sanccob.co.za/?m=2&s=5&idkey=625. 
  55. ↑ ""Jackass Penguins Freed after Rehab", National Geographic's Video News, June 17, 2009". National Geographic. 2009-06-17. http://news.nationalgeographic.com/news/2009/06/090617-SAF-penguinsfree-video-ap.html. 
  56. ↑ Barham, P. J.; L. G. Underhill; R. J. M. Crawford; R. Altewegg; T. M. Leshoro; D. A. Bolton; B. M. Dyer; L. Upfold (2008). "The efficacy of hand-rearing penguin chicks: evidence from African penguins (Spheniscus demersus) orphaned in the Treasure oil spill". Bird Conservation International 18 (2). doi:10.1017/S0959270908000142. 
  57. ↑ Grim, C. (2003). "Plasmodium juxtanucleare associated with mortality in black-footed penguins (Spheniscus demersus) admitted to a rehabilitation center". Journal of Zoo and Wildlife Medicine 34 (3): 250–5. doi:10.1638/02-070. PMID 14582786. 
  58. ↑ "Refuelling under scrutiny as S. Africa penguins hit by oil spill" (in en). https://phys.org/news/2019-07-refuelling-scrutiny-safrica-penguins-oil.html. 
  59. ↑ Sherley, R. B.; Underhill, L. G.; Barham, B. J.; Barham, P. J.; Coetzee, J. C.; Crawford, R. J.; Dyer, B. M.; Leshoro, T. M. et al. (2013). "Influence of local and regional prey availability on breeding performance of African penguins Spheniscus demersus". Marine Ecology Progress Series 473: 291–301. doi:10.3354/meps10070. Bibcode: 2013MEPS..473..291S. 
  60. ↑ 60.0 60.1 Weller, Florian; Sherley, Richard B.; Waller, Lauren J.; Ludynia, Katrin; Geldenhuys, Deon; Shannon, Lynne J.; Jarre, Astrid (2016). "System dynamics modelling of the Endangered African penguin populations on Dyer and Robben islands, South Africa". Ecological Modelling 327: 44–56. doi:10.1016/j.ecolmodel.2016.01.011. 
  61. ↑ Sherley, Richard B. (2018). "Bayesian inference reveals positive but subtle effects of experimental fishery closures on marine predator demographics". Proceedings of the Royal Society B 285 (1871): 20172443. doi:10.1098/rspb.2017.2443. PMID 29343602. 
  62. ↑ Ludynia, K.; L. J. Waller; R. B. Sherley; F. Abadi; Y. Galada; D. Geldenhuys; R. J. M. Crawford; L. J. Shannon et al. (2014). "Processes influencing the population dynamics and conservation of African penguins on Dyer Island, South Africa". African Journal of Marine Science 36 (2): 253. doi:10.2989/1814232X.2014.929027. 
  63. ↑ Makhado, A. B.; M. A. Meÿer; R. J. M. Crawford; L. G. Underhill; C. Wilke (2009). "The efficacy of culling seals seen preying on seabirds as a means of reducing seabird mortality". African Journal of Ecology 47 (3): 335. doi:10.1111/j.1365-2028.2008.00966.x. 
  64. ↑ Nel, D. C.; Crawford, R. J. M.; Parsons, N. J. (2003). "The conservation status and impact of oiling on the African penguin". in Nel, D. C.. Rehabilitation of Oiled African Penguins: a Conservation Success Story. Cape Town: BirdLife South Africa and Avian Demography Unit. pp. 1–7. 
  65. ↑ "First Class African Penguin and Seabird Sanctuary Opens in Kleinbaai, Overberg". SceinicSouth. 2015-02-26. http://scenicsouth.co.za/first-class-african-penguin-and-seabird-sanctuary-opens-in-kleinbaai-overberg/. 
  66. ↑ ""Rehab center goes to the birds", iol SciTech, March 3, 2015". iol.co.za. 2015-03-03. http://www.iol.co.za/scitech/science/environment/rehab-centre-goes-to-the-birds-1.1826123#.VQGFMfzF-So. 
  67. ↑ "African Penguin". aza.org. https://www.aza.org/SAFE-african-penguin. 
  68. ↑ "Nature conservation project African penguin". artis.nl. https://www.artis.nl/en/footer/about-artis/nature-conservation/nature-conservation-project-african-penguin/. 

External links

Wikidata ☰ Q244813 entry



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