King penguin

King penguin
King penguin Fortuna Bay, South Georgia
Scientific classification
Kingdom:Animalia
Phylum:Chordata
Class:Aves
Order:Sphenisciformes
Family:Spheniscidae
Genus:Aptenodytes
Species: A. patagonicus
Binomial name
Aptenodytes patagonicus
Miller,JF, 1778
Red: Aptenodytes patagonicus patagonicus

Yellow: Aptenodytes patagonicus halli
Green: breeding areas

The king penguin (Aptenodytes patagonicus) is a large species of penguin, second only to the emperor penguin in size. There are two subspecies: A. p. patagonicus and A. p. halli; patagonicus is found in the South Atlantic and halli found at the Kerguelen Islands and Crozet Island, Prince Edward Islands, Heard Island and McDonald Islands, and Macquarie Island.[2]

King penguins eat small fish, mainly lanternfish, squid and krill. On foraging trips king penguins repeatedly dive to over 100 metres (300 ft), and have been recorded at depths greater than 300 metres (1,000 ft).[3]

King penguins breed on the subantarctic islands at the northern reaches of Antarctica, South Georgia, and other temperate islands of the region.

Appearance

The close-up of A. p. patagonicus subspecies in St. Andrews Bay, South Georgia, British Overseas Territories, UK

The king penguin stands at 70 to 100 cm (28 to 39 in) tall and weighs from 9.3 to 18 kg (21 to 40 lb).[4][5][6] Males are slightly larger than females. The mean body mass of adults from Marion Island was 12.4 kg (27 lb) for 70 males and 11.1 kg (24 lb) for 71 females. Another study from Marion Island found that the mean mass of 33 adults feeding chicks was 13.1 kg (29 lb). Thus the average weight of the king penguin is similar or just slightly higher than that of the largest living flying birds.[7][8]

At first glance, the king penguin appears very similar to the closely related emperor penguin, with a broad cheek patch contrasting with surrounding dark feathers and yellow-orange color at the top of the chest. However, the cheek patch of the adult king penguin is bright orange whereas that of the emperor penguins is white, while the chest orange tends to be more vivid and less yellowish in the king species. Both species have colorful markings along the side of their lower mandible, but these are pinkish in emperor penguins and orange in king penguins. Emperor and king penguins typically do not inhabit the same areas in the wild, with the possible exception of vagrants at sea, but the king can readily be distinguished by being noticeably sleeker, with a longer, straighter bill than the emperor. The chicks of both species are completely different from one another in appearance. Once fully molted of its heavy dark brown down, the juvenile king penguin resembles the adult but is somewhat less colorful.

Mounted skeleton of king penguin (Aptenodytes patagonicus)

King penguins often breed on the same large, circumpolar islands as at least half of all living penguins, but it is easily distinguished from co-occurring penguins by its much larger size and taller frame, distinctive markings and grizzled sooty-grayish rather than blackish back.[4][9][10]

Distribution and habitat

Great colony of king penguins on Salisbury Plain in South Georgia

King penguins breed on subantarctic islands between 45 and 55°S, at the northern reaches of Antarctica, as well as Tierra del Fuego, the Falkland Islands, and other temperate islands of the region. The total population is estimated to be 2.23 million pairs and is increasing.[4] The largest breeding populations are on the Crozet Islands, with around 455,000 pairs, 228,000 pairs on the Prince Edward Islands, 240,000–280,000 on the Kerguelen Islands and over 100,000 in the South Georgia archipelago. Macquarie Island has around 70,000 pairs. The non-breeding range is unknown due to many vagrant birds having been seen on the Antarctic peninsula as well as in South Africa, Australia and New Zealand.

King penguins appear to have suffered a massive population decline of nearly 90% on Île aux Cochons, or Pig Island in the Crozet Archipelago. Analysing new helicopter and satellite images from 2015 and 2017, the colony’s numbers have dropped to 60,000 breeding pairs.[11] The cause of this decline may be due to changes in the ecosystem related to climate change as their primary source of food is moving farther away from places where the penguins can breed. This may result in population declines and shifts in the locations of the King penguin breeding grounds.[12]

The Nature Protection Society released several king penguins in Gjesvær in Finnmark, and Røst in Lofoten in northern Norway in August 1936. Birds were seen in the area several times during the 1940s though none have been recorded since 1949.There were a few unconfirmed sightings of birds in the area during the early 1950s.[13]

Behavior

King penguin chick

American zoologist Gerry Kooyman revolutionized the study of penguin foraging behaviour in 1971 when he published his results from attaching automatic dive-recording devices to emperor penguins,[14] and recording a dive of 235 metres (771 ft) by a king penguin in 1982.[15] The current maximum dive recorded is 343 metres in the Falkland Islands region,[16] and a maximum time submerged of 552 seconds recorded at the Crozet Islands.[17] The king penguin dives to depths of 100–300 meters (350–1000 feet), spending around five minutes submerged, during daylight hours, and less than 30 metres (98 ft) at night.[18][19]

Sound from rookery at Lusitania Bay on Macquarie Island

The majority (around 88% in one study) of dives undertaken by king penguins are flat-bottomed; that is, the penguin dives to a certain depth and remains there for a period of time hunting (roughly 50% of total dive time) before returning to the surface. They have been described as U-shaped or W-shaped, relating to the course of the dive. The remaining 12% of dives have a V-shaped or "spike" pattern, in which the bird dives at an angle through the water column, reaches a certain depth, and then returns to the surface. In contrast, other penguins dive in this latter foraging pattern.[18][20] Observations at Crozet Islands revealed most king penguins were seen within 30 km (19 mi) of the colony.[21] Using the average swimming speed, Kooyman estimated the distance travelled to foraging areas at 28 km (17 mi).[18]

The king penguin's average swimming speed is 6.5–10 km/h (4–6 mph). On shallower dives under 60 m (200 ft), it averages 2 km/h (1.2 mph) descending and ascending, while on deeper dives over 150 m (490 ft) deep, it averages 5 km/h (3.1 mph) in both directions.[19][22] King penguins also "porpoise", a swimming technique used to breathe while maintaining speed. On land, the king penguin alternates between walking with a wobbling gait and tobogganing—sliding over the ice on its belly, propelled by its feet and wing-like flippers. Like all penguins, it is flightless.[23]

Diet

King penguin moulting, with pin feathers visible

King penguins eat various species of small fish, squid, and krill. Fish constitute roughly 80% of their diet, except in winter months of July and August, when they make up only around 30%.[19] Lanternfish are the main fish taken, principally the species Electrona carlsbergi and Krefftichthys anderssoni, as well as Protomyctophum tenisoni. Slender escolar (Paradiplospinus gracilis) of the Gempylidae, and Champsocephalus gunneri, is also consumed. Cephalopods consumed include those of the genus Moroteuthis, the hooked squid or Kondakovia longimana, the sevenstar flying squid (Martialia hyadesii), young Gonatus antarcticus and Onychoteuthis species.[19]

Predators

The king penguin's predators include other seabirds and aquatic mammals:

  • Giant petrels take many chicks of all sizes and some eggs. They have also been reported to occasionally kill adult king penguins, but very likely mostly sick or injured ones. Giant petrels will scavenge additional king penguin chicks which have already died from other causes.[24][25]
  • Skua species (Stercorarius spp.) take smaller chicks and eggs. Some studies may overemphasize the effect skua predation have on king penguin colonies, but large numbers of chicks and eggs are taken in areas where skua nest close to penguin colonies.[26][27][28]
  • The snowy sheathbill (Chionis alba) and kelp gull (Larus dominicanus) scavenge for dead chicks and unattended eggs.[29]
  • The leopard seal (Hydrurga leptonyx) takes adult birds at sea.[30]
  • Orcas also hunt king penguins.[31]
  • Antarctic fur seals on Marion Island have also been observed chasing king penguins on the beach, killing and eating them. It seems that especially males, and particularly pre-adult males, are involved.[32][33]

Courtship and breeding

A pair of king penguins performs a courtship ritual on Salisbury Plain, South Georgia.
Mating king penguins at Macquarie Island

The king penguin is able to breed at three years of age, although only a very small minority (5% recorded at Crozet Islands) actually do then; the average age of first breeding is around 6 years.[34] King penguins are serially monogamous. They have only one mate each year, and stay faithful to that mate. However, fidelity between years is only about 29%.[35] The long breeding cycle may contribute to this low rate.[36]

The king penguin has an unusually prolonged breeding cycle, taking some 14–16 months from laying to offspring fledging.[37] Although pairs will attempt to breed annually, they are generally only successful one year in two, or two years in three in a triennial pattern on South Georgia.[30] The reproductive cycle begins in September to November, as birds return to colonies for a prenuptial moult. Those that were unsuccessful in breeding the previous season will often arrive earlier. They then return to the sea for three weeks before coming ashore in November or December.[38]

Aptenodytes patagonicus egg

The female penguin lays one pyriform (pear-shaped) white egg weighing 300 g (⅔ lb).[39] It is initially soft, but hardens and darkens to a pale greenish colour. It measures around 10 cm × 7 cm (3.9 in × 2.8 in).[39] The egg is incubated for around 55 days with both birds sharing incubation in shifts of 6–18 days each.

Hatching may take up to 2–3 days to complete, and chicks are born semi-altricial and nidicolous. In other words, they have only a thin covering of down, and are entirely dependent on their parents for food and warmth.[40] The guard phase starts with the birth of the chick. Like the closely related emperor penguin, The young king penguin chick spends its time balanced on its parents' feet, sheltered by a pouch formed from the abdominal skin of the latter.[40] During this time, the parents alternate every 3–7 days, one guarding the chick while the other forages. The guard phase lasts for 30–40 days. By then the chick has grown much bigger, can keep itself warm and protect itself against most predators. It becomes more curious and starts to explore its surroundings. It ends up forming a group with other chicks, a so-called crèche. Crèches are guarded by only a few adult birds; most parents can leave their chick to forage for themselves and their chick. Other species of penguins also practice this method of communal care for offspring.

Close-up of king penguin chick.
Unlike emperor penguin chick, covered with brown down.
King penguin chicks

By April the chicks are almost fully grown, but lose weight by fasting over the winter months, gaining it again during spring in September. Fledging then takes place in late spring/early summer.

King penguins form huge breeding colonies; for example, the colony on South Georgia Island at Salisbury Plain holds over 100,000 breeding pairs and the one at St. Andrew's Bay over 100,000 birds. Because of the long breeding cycle, colonies are continuously occupied year-round with both adult birds and chicks.

The king penguin feeds its chicks by eating fish, digesting it slightly and regurgitating the food into the chick's mouth.

Because of their large size, king penguin chicks take 14–16 months before they are ready to go to sea. This is markedly different from smaller penguins, who rear their chicks through a single summer when food is plentiful. King penguins time their mating so the chicks will develop over the harshest season for fishing. In this way, by the time the young penguins are finally mature enough to leave their parents, it is summer when food is plentiful and conditions are more favorable for the young to survive alone.

Relationship with humans

A pair of king penguin at Edinburgh Zoo

Considered a flagship species, 176 individuals were counted in captivity in North American zoos and aquaria in 1999.[41] The species has been bred in captivity at SeaWorld San Diego. The species is exhibited at SeaWorld Orlando, Indianapolis Zoo,[42] Detroit Zoo, Saint Louis Zoo,[43] Kansas City Zoo, Newport Aquarium in Newport, Kentucky, Edinburgh Zoo in Scotland, Berlin Zoological Garden in Germany, Zurich Zoo in Switzerland, Diergaarde Blijdorp in the Netherlands, Antwerp Zoo in Belgium, 63 Seaworld in Seoul, South Korea, Melbourne Aquarium in Australia, Mar del Plata Museum of the Sea in Argentina, Loro Parque in Spain and Ski Dubai in United Arab Emirates, Calgary Zoo in Canada, Odense Zoo in Denmark and many other collections.

Notable king penguins

  • Brigadier Sir Nils Olav, the Edinburgh-based mascot and colonel-in-chief of the Royal Norwegian Guard
  • Misha, a central character and metaphor in two novels by Ukrainian writer Andrey Kurkov
  • The king penguin is also the species of penguin represented by the popular character Pondus, an image found on various paraphernalia in many retail stores throughout Canada. Pondus originates in Danish children's books written and photographed by Ivar Myrhøj and published in 1997 by Lademann publisher in the late 1960s. These penguins appeared in the production of Batman Returns.
  • Lala the Penguin became a viral video star after an Animal Planet special featured him venturing to a nearby market in Japan to fetch a fish with a specially made backpack.[44]

References

Citations

  1. ↑ BirdLife International. (2016). Aptenodytes patagonicus. The IUCN Red List of Threatened Species doi:10.2305/IUCN.UK.2016-3.RLTS.T22697748A93636632.en
  2. ↑ "Aptenodytes patagonicus (King Penguin)". www.iucnredlist.org.
  3. ↑ Culik, B. M; K. PÜTZ; R. P. Wilson; D. Allers; J. LAGE; C. A. BOST; Y. LE MAHO (January 1996). "Diving Energetics in King Penguins (Aptenodytes patagonicus)" (PDF). Journal of Experimental Biology. 199 (4): 973–983.
  4. 1 2 3 Shirihai, Hadoram (2002). A Complete Guide to Antarctic Wildlife. Alula Press. ISBN 951-98947-0-5.
  5. ↑ McGonigal, D., L. Woodworth. (2001). Antarctica and the Arctic: The Complete Encyclopedia. Ontario: Firefly Books.
  6. ↑ Cherel, Y; Leloup, J; Le Maho, Y (1988). "Fasting in king penguin. II. Hormonal and metabolic changes during molt". The American Journal of Physiology. 254 (2 Pt 2): R178–84. doi:10.1152/ajpregu.1988.254.2.R178. PMID 3278625.
  7. ↑ Dunning, John B. Jr. (ed.) (2008). CRC Handbook of Avian Body Masses, 2nd Ed. CRC Press. ISBN 978-1-4200-6444-5.
  8. ↑ Adams, N. J.; Klages, N. T. (1987). "Seasonal variation in the diet of the king penguin (Aptenodytes patagonicus) at sub-Antarctic Marion Island". Journal of Zoology. 212 (2): 303. doi:10.1111/j.1469-7998.1987.tb05992.x.
  9. ↑ Nolan, Paul M.; Stephen Dobson, F.; Nicolaus, Marion; Karels, Tim J.; McGraw, Kevin J.; Jouventin, Pierre (2010). "Mutual Mate Choice for Colorful Traits in King Penguins". Ethology. doi:10.1111/j.1439-0310.2010.01775.x.
  10. ↑ Jouventin, Pierre; Nolan, Paul M.; Örnborg, Jonas; Dobson, F. Stephen (2005). "Ultraviolet Beak Spots in King and Emperor Penguins". The Condor. 107: 144. doi:10.1650/7512. JSTOR 3247764.
  11. ↑ "Enormous penguin population crashes by almost 90%". Nature. 2018-07-30. doi:10.1038/d41586-018-05850-2. Retrieved 2018-07-31.
  12. ↑ Cristofari, Robin; Liu, Xiaoming; Bonadonna, Francesco; Cherel, Yves; Pistorius, Pierre; Le Maho, Yvon; Raybaud, Virginie; Stenseth, Nils Christian; Le Bohec, Céline (2018-02-26). "Climate-driven range shifts of the king penguin in a fragmented ecosystem". Nature Climate Change. 8 (3): 245–251. doi:10.1038/s41558-018-0084-2. ISSN 1758-678X.
  13. ↑ Long, John L. (1981). Introduced Birds of the World: The worldwide history, distribution and influence of birds introduced to new environments. Terrey Hills, Sydney: Reed. p. 30. ISBN 0-589-50260-3.
  14. ↑ Kooyman GL, Drabek CM, Elsner R, Campbell WB (1971). "Diving behaviour of the Emperor Penguin Aptenodytes forsteri". Auk. 88 (4): 775–95. doi:10.2307/4083837. JSTOR 4083837.
  15. ↑ Kooyman GL, Davis RW, Croxall JP, Costa DP (1982). "Diving depths and energy requirements of the King Penguins". Science. 217 (4561): 726–27. doi:10.1126/science.7100916. PMID 7100916.
  16. ↑ Pütz, K.; Cherel, Y. (2005). "The diving behaviour of brooding king penguins (Aptenodytes patagonicus) from the Falkland Islands: variation in dive profiles and synchronous underwater swimming provide new insights into their foraging strategies". Marine Biology. 147 (2): 281. doi:10.1007/s00227-005-1577-x.
  17. ↑ Putz, K.; Wilson, R. P.; Charrassin, J.-B.; Raclot, T.; Lage, J.; Maho, Y. Le; Kierspel, M. A. M.; Culik, B. M.; Adelung, D. (1998). "Foraging strategy of king penguins (Aptenodytes patagonicus) during summer at the Crozet Islands". Ecology. 79 (6): 1905. doi:10.2307/176698. JSTOR 176698.
  18. 1 2 3 Kooyman GL, Cherel Y, Le Maho Y, Croxall JP, Thorson PH, Ridoux V (1992). "Diving behaviour and energetics during foraging cycles in King Penguins". Ecological Monographs. 62 (1): 143–63. doi:10.2307/2937173. JSTOR 2937173.
  19. 1 2 3 4 Williams, p. 147
  20. ↑ Williams, pp. 87–88
  21. ↑ Ridoux V, Jouventin P, Stahl JC, Weimerskirch H (1988). "Ecologie alimentaire comparée des manchots nicheurs aux Iles Crozet". Revues Ecologie (in French). 43: 345–55.
  22. ↑ Adams, NJ (1987). "Foraging ranges of King Penguins Aptenodytes patagonicus during summer at Marion Island". Journal of Zoology. 212 (3): 475–82. doi:10.1111/j.1469-7998.1987.tb02918.x.
  23. ↑ Williams, p. 3
  24. ↑ Hunter, Stephen (2008). "The impact of avian predator-scavengers on King Penguin Aptenodytes patagonicus chicks at Marion Island". Ibis. 133 (4): 343. doi:10.1111/j.1474-919X.1991.tb04581.x.
  25. ↑ Le Bohec, Céline; Gauthier-Clerc, Michel; Gendner, Jean-Paul; Chatelain, Nicolas; Le Maho, Yvon (2003). "Nocturnal predation of king penguins by giant petrels on the Crozet Islands". Polar Biology. 26 (9): 587. doi:10.1007/s00300-003-0523-y.
  26. ↑ Descamps, Sébastien; Gauthier-Clerc, Michel; Le Bohec, Céline; Gendner, Jean-Paul; Le Maho, Yvon (2004). "Impact of predation on king penguin Aptenodytes patagonicus in Crozet Archipelago". Polar Biology. 28 (4): 303. doi:10.1007/s00300-004-0684-3.
  27. ↑ Emslie, S. D.; Karnovsky, N. & Trivelpiece, W. (1995). "Avian predation at penguin colonies on King George Island, Antarctica" (PDF). The Wilson Bulletin. 107: 317–327. JSTOR 4163547.
  28. ↑ Young, E. (2005). Skua and penguin: predator and prey. Cambridge University Press.
  29. ↑ Williams, p. 40
  30. 1 2 Stonehouse, B (1960). "The King Penguin Aptenodytes patagonicus of South Georgia I. Breeding behaviour and development". Falkland Islands Dependencies Survey Scientific Report. 23: 1–81.
  31. ↑ Condy, P. R.; Aarde, R. J. Van; Bester, M. N. (2009). "The seasonal occurrence and behaviour of killer whales Orcinus orca, at Marion Island". Journal of Zoology. 184 (4): 449. doi:10.1111/j.1469-7998.1978.tb03301.x.
  32. ↑ Walker, Matt. "King penguins become fast food for Antarctic fur seals". Retrieved 28 September 2012.
  33. ↑ Charbonnier, Yohan; Delord, Karine; Thiebot, Jean-Baptiste (2009). "King-size fast food for Antarctic fur seals". Polar Biology. 33 (5): 721. doi:10.1007/s00300-009-0753-8.
  34. ↑ Williams, p. 151
  35. ↑ Williams, p. 54
  36. ↑ Williams, p. 152
  37. ↑ Williams, p. 148
  38. ↑ Williams, p. 149
  39. 1 2 Williams, p. 150
  40. 1 2 Williams, p. 28
  41. ↑ Diebold EN, Branch S, Henry L (1999). "Management of penguin populations in North American zoos and aquariums" (PDF). Marine Ornithology. 27: 171–76. Retrieved 31 March 2008.
  42. ↑ "Penguin Feed/Chat". Indianapolis Zoo website. Indianapolis Zoo. Archived from the original on 2011-08-28. Retrieved 2011-12-01.
  43. ↑ "King Penguin". Saint Louis Zoo website. Saint Louis Zoo. 2009. Retrieved 3 September 2016.
  44. ↑ "Lala Penguin Goes Shopping". Animal Planet. 2011.

Bibliography

  • Williams, Tony D. (1995). The Penguins. Oxford, England: Oxford University Press. ISBN 019854667X.
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