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Aptenodytes patagonicusking penguin

By Keena Edwards

Geographic Range

Aptenodytes patagonicus (king penguins) colonies are mainly located on islands surrounding Antarctica. Islands include Crozet, Falkland, Heard, Kerguelen, Macquarie, Prince Edward, South Georgia and South Sandwich. Although no colonies have been found south of latitude 60 degrees S, some non-breeding members have taken residence in southern Chile and southern Argentina. Some lone wanderers have been found as far north as Brazil and South Africa and as far south as the Antarctic Coast.

On South Georgia Island over 30 colonies of A. patagonicus patagonicus reside. Colony sizes range from approximately 39,000 breeding pairs at both Salisbury Plains and St. Andrews Bay to 9,000 pairs located at Royal Bay. This subspecies is only found on South Georgia and the Falkland Islands. Another genetically unique subspecies, A. patagonicus halli, is only found on Crozet, Heard, Kerguelen, Macquarie and Prince Edwards islands. (McGonigal and Woodworth, 2001; Shirihai, 2008)

Habitat

King penguins spend a lot of time in the ocean feeding, but their primary habitats are sparsely vegetated areas of islands in the southern oceans and sub-Antarctic. The islands south of the Polar Front are typically more glaciated and at higher altitudes. For example, Heard Island is 2745 meters above sea level. These islands are still out of reach of the Antarctic pack ice, but icy conditions are still prevalent. As in the case of South Georgia, the bays freeze over and the island is over half covered in ice during the winter months. In the locations not covered in ice, bryophytes are the primary vegetation. Islands like Macquarie (433 meters above sea level) have some flowering plants and ferns and the air temperature only varies a few degrees between the summer and winter seasons.

While king penguins prefer to live on islands south of the Polar Front, they prefer to fish in waters just north of it where surface air temperatures are around 4.5 degrees C. They have been known to dive to a maximum of 322 meters. (McGonigal and Woodworth, 2001)

  • Range elevation
    433 to 2745 m
    1420.60 to 9005.91 ft
  • Range depth
    0 to 322 m
    0.00 to 1056.43 ft

Physical Description

King penguins are the second largest of all penguin species. Females are noted to be slightly smaller than males. However, no specific female measurements have been recorded. Their documented height ranges from 85 to 95 cm and weight is between 9.3 and 17.3 kg. Average adult weight has been found to be 11.8 kg.

Although they are easily confused with emperor penguins (Aptenodytes forsteri), king penguins are more colorful and have a longer, more slender bill. This bill has a stripe on the lower mandible that ranges in color from pinkish-red to orange-yellow and exhibits ultraviolet (UV) reflectance. The function of this beak spot is unclear, but it is thought to signal sexual maturity, health and/or social standing. The spot, without UV reflectance, is found in juveniles. Neither the beak spot nor UV reflectance are seen in chicks. This UV reflective beak spot does not differ between sexes.

Adult king penguins have a dark, nearly black head with orange to orange-yellow, spoon-shaped spots on either side of the head and an orange area that is most intense at the throat and fades down the upper breast into pale yellow then finally a white ventral side. The dorsal side of the body and flippers consist of gray and black feathers with a silvery sheen. The sides are separated from the ventral side by a narrow, black line. The front edge of the flipper also has a black line that extends to a black tip. King penguins are sexually monomorphic in plumage, but males are slightly larger.

Juvenile king penguins are similar to adults, but their coloration isn't as vivid. They do not reach full adult coloration until three years of age. Prior to the first molt, the chicks are simply dark brown.

There is no evidence of physical differences between the two subspecies. (Jouventin, et al., 2005; McGonigal and Woodworth, 2001; Nolan, et al., 2010; Putz and Bost, 1994; Shirihai, 2008)

  • Sexual Dimorphism
  • sexes alike
  • male larger
  • Range mass
    9.3 to 17.3 kg
    20.48 to 38.11 lb
  • Average mass
    11.8 kg
    25.99 lb
  • Range length
    85 to 95 cm
    33.46 to 37.40 in
  • Average basal metabolic rate
    25.889 W
    AnAge

Reproduction

King penguins have a lower rate of monogamy than smaller penguin species. Currently, there is no definitive answer as to why this occurs; however, two explanations have merit: 1) mates not arriving to the colony at the same time, and 2) the amount of fat the penguin has stored. Fat storage plays a role in the low monogamy rate because if the penguins begin storing fat too early they become more vulnerable to predators. If they begin storing fat too late, they may not return to the colony at the same time as their mate. If both mates do not arrive at the colony at the same time, breeding can be delayed or a new mate may be chosen.

Females appear to be more selective than males when choosing a mate, but both sexes seem to choose a mate based on their plumage. Early breeding pairs have higher ultraviolet reflectance of beak spots than those breeding later in the season. The plumage color on the breasts and auricular areas are thought to directly reflect the health of a king penguin’s immune system. The healthier it is, the brighter the plumage.

Males advertise for mates with a combination of vocalizations and visual displays. Male king penguins produce a trumpeting call and then stretch to full height with bills raised. Once a female accepts, the two face each other and continue to engage in physical displays including strutting, bowing, shaking, calling, and stretching to maximum height with bills in the air. (Bried, et al., 1999; Nolan, et al., 2010)

King penguins breed yearly on the flat shorelines of the sub-Antarctic islands. Their cycle beings with a 1-month molting stage for both parents, which is complete by the end of October. Once the molt is complete, the courtship stage can last for just over a month. The female lays a single, greenish-white egg in November or December. This is transferred to the male penguin's feet and is incubated for approximately 54 days under a pouch of belly skin that keeps it at the penguin’s internal body temperature. After laying the egg, the female leaves to feed and replenish the weight that was lost. When the female returns, the partners take turns in incubating the egg, with shifts ranging from 5 to 22 days. The average birth mass for king penguin chicks is 430 g. Post-hatching, the parents continue taking turns to incubate and feed the chick until May, when it is big enough to survive on its own.

By May, the chicks are fairly independent. They stay with the colony and survive off their stored fat until the following October. In these months, both parents leave to forage and return periodically to feed their chick. During that time the chicks live in crèches (groups of juveniles) until they have gained enough weight to become completely independent. Birth to independence takes 14 to 16 months. Juvenile king penguins do not reach reproductive maturity until 3 to 5 years of age.

Successful parents do not begin their next breeding cycle until their chick has successfully fledged. This causes a successful breeding pair to begin the next season late. The outcome is usually a failed cycle because an egg laid after December typically is not successful. However, this failure allows them to breed earlier the following season. The earlier that the breeding cycle begins the more likely it is to be successful. This biennial pattern to their breeding cycle makes king penguin reproduction unique.

Not all breeding pairs in a colony are on the same biennial cycle, and not all are guaranteed to follow the success-fail-success-fail pattern. It is most likely for them to follow this pattern or a success-fail-fail-success pattern. Some are on alternating cycles so that there are chicks born during every breeding season.

It is believed that food availability plays a role in the timing and success rate of the breeding cycle because it directly affects the health of the parent. (Aubin and Jouventin, 1998; Bried, et al., 1999; Cote, 2000; Dobson and Jouventin, 2003; Lockley, 1984; Nicolaus, et al., 2007; Olsson, 1996; Shirihai, 2008)

  • Breeding interval
    King penguins breed once yearly, but normally are successful twice over 3 calendar years.
  • Breeding season
    King penguins breed between October and December.
  • Average eggs per season
    1
  • Average time to hatching
    54 days
  • Range fledging age
    14 to 16 months
  • Range time to independence
    14 to 16 months
  • Range age at sexual or reproductive maturity (female)
    3 (low) years
  • Average age at sexual or reproductive maturity (female)
    5 years
  • Range age at sexual or reproductive maturity (male)
    3 (low) years
  • Average age at sexual or reproductive maturity (male)
    5 years

King penguin parents highly invest in their reproductive cycles. The males must begin the cycle with enough fat stored to sustain them through courtship, egg-laying and the first part of incubation. By the end of their first incubation shift the males have typically lost 30 percent of their body weight. A minimum body mass of 10 kg is considered to be a critical mass for male king penguins. When they are approaching 10 kg, the males must choose whether to abandon the egg or to continue waiting for the female to return and relieve them. Hatchlings are semi-altricial, and therefore have considerable development to achieve post-hatching. This requires a large parental investment to brood and nourish the young. (Olsson, 1996; Olsson, 1997)

  • Parental Investment
  • altricial
  • male parental care
  • female parental care
  • pre-fertilization
    • provisioning
    • protecting
      • female
  • pre-hatching/birth
    • provisioning
      • male
      • female
    • protecting
      • male
      • female
  • pre-weaning/fledging
    • provisioning
      • male
      • female
    • protecting
      • male
      • female
  • pre-independence
    • provisioning
      • male
      • female
    • protecting
      • male
      • female

Lifespan/Longevity

Aptenodytes patagonicus has been generically documented as a long-lived bird. However, no numerical lifespan data has been published. Captive king penguins can live up to 26 years. (Flower, 1938)

  • Range lifespan
    Status: captivity
    26 (high) years
  • Average lifespan
    Status: captivity
    26 years
    AnAge

Behavior

King penguins are one of only two penguin species that do not build nests. Instead of building a nest, king penguins carry their eggs and young around on their feet during development. The lack of a nest also adds to the noise level of the colonies because they use their unique voice system to call to both their young and their mates.

King penguins are very social birds, and live in colonies with populations of up to 39,000 breeding pairs. The colonies are generally found on large, sparsely vegetated areas of the islands they inhabit because king penguins avoid climbing. They have been seen huddling together during blizzards to conserve heat. Large colonies have communal crèches, or groups, of chicks that stay together for protection while both parents are simultaneously foraging.

They have been known to travel up to 500 km from their breeding grounds to forage, and will bathe in the ocean before fishing.

The social hierarchies of king penguin colonies are not entirely understood. The penguins compete for positions in the center of the colonies, and these birds are more aggressive and more successful at reproduction than ones further out. It is thought to possibly relate to the ultraviolet beak spots or plumage. However, no definitive conclusions have been made. (Aubin and Jouventin, 1998; Dobson and Jouventin, 2003; Lockley, 1984; McGonigal and Woodworth, 2001; Shirihai, 2008; Viera, et al., 2007; de Villiers and de Bruyn, 2004)

  • Average territory size
    0.5 m^2

Home Range

Breeding pairs defend an area that is approximately 0.5 square meters against neighboring king penguins and predators.

Communication and Perception

The primary form of communication in king penguins is a two-voice system that is produced by the syrinx, a two-part organ located where the bronchi join. Each part produces sound independently. In penguins, the syrinx is only found in the genus Aptenodytes. The UV reflective beak spots may be used for signaling, but the true use is unknown.

Due to the noisy environment of king penguin colonies, adults repeatedly call out 3 to 7 syllables of varying volumes with two frequencies to locate chicks. The chicks are thought to identify the calls by the lower of the two frequencies because they transmit farther in the seeming chaos of many adults calling at once. The higher frequency has no documented use. This ability has been termed "cocktail-party effect." The parental call is thought to be learned during the first five weeks of life and is important because of the lack of nests and landmarks in the king penguins' habitats.

At this time communication research is mainly focused on the parent/offspring connection, but it is believed that the two-voice systems are also used to locate mates.

King penguins also incorporate movements and behaviors into courtship rituals. Males will produce trumpeting calls and stretch to their maximum height to attract mates. Once a female accepts, the two will stand facing each other and will engage in a series of stretching, bobbing, shaking, bowing, and calling. Like all birds, king penguins perceive their environments through visual, auditory, tactile and chemical stimuli. (Aubin and Jouventin, 1998; Aubin, et al., 2000; Jouventin, et al., 2005; Jouventin, et al., 1999)

Food Habits

Aptenodytes patagonicus travels up to 500 km from its colony to the ocean to feed on cephalopods, small fish, and squid. They can remain underwater approximately 10 minutes and reach speeds of 12 km per hour while diving 25 to 322 m. Adults take turns returning at irregular intervals to supplement the young that stay in the breeding grounds. During their first winter the chicks predominantly survive from their stored fat. (Aubin and Jouventin, 1998; Lockley, 1984; McGonigal and Woodworth, 2001; Shirihai, 2008)

  • Animal Foods
  • fish
  • mollusks
  • other marine invertebrates

Predation

Antarctic fur seals, sub-Antarctic fur seals, leopard seals, and killer whales regularly prey on adult king penguins. Also, brown skuas and giant petrels prey on king penguin chicks. (Charbonnier, et al., 2009; Emslie, et al., 1995)

Ecosystem Roles

King penguins occasionally serve as prey for Antarctic fur seals (Arctocephalus gazelle), subantarctic fur seals (Arctocephalus tropicalis), brown skuas (Catharacta lonnbergi) and giant petrels (Macronectes giganteus). None of these predators appear to control the king penguin population because they are not a primary food source, and the king penguin population is steadily growing. King penguins act as predators for cephalopods, small fish and squid found in their geographic range, but there is no documentation stating if the populations are controlled by this predation. (Aubin and Jouventin, 1998; Charbonnier, et al., 2009; Emslie, et al., 1995; Lockley, 1984; McGonigal and Woodworth, 2001; Shirihai, 2008)

The hard tick, Ixodes uriae, is a parasite that infests king penguins. The mortality rate of adult king penguins due to hard ticks is unknown; however, there is documentation of death due to hyperinfestation of hard ticks. A bird louse, Austrogoniodes pauliani, is also a common parasite found on these penguins. (Gauthier-Clerc, et al., 1998)

Commensal/Parasitic Species
  • Hard ticks (Ixodes uriae)
  • Bird louse (Austrogoniodes pauliani)

Economic Importance for Humans: Positive

King penguins are part of the ecotourism business. During a study of nature-based tourism on the Falkland Islands, king penguins were the most popular of the penguins in the area. All of the tourists visited the king penguin colony, versus only half of the tourists visiting the second most popular penguins species. The tourists also stayed to view the king penguins for the longest time frame at a mean of 63 minutes versus only 19 minutes for the second most popular penguin species. (Otley, 2005)

Economic Importance for Humans: Negative

There are no known negative economic effects of king penguins on humans.

Conservation Status

The IUCN Red List states that king penguins have a status of Least Concern based on the following criteria: geographic range is greater than 20,000 square kilometers, population size is greater than 10,000 individuals, and the population size seems to be increasing.

Contributors

Keena Edwards (author), Radford University, Karen Powers (editor), Radford University, Rachelle Sterling (editor), Special Projects.

Glossary

Antarctica

lives on Antarctica, the southernmost continent which sits astride the southern pole.

acoustic

uses sound to communicate

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

carnivore

an animal that mainly eats meat

chemical

uses smells or other chemicals to communicate

colonial

used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.

diurnal
  1. active during the day, 2. lasting for one day.
dominance hierarchies

ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates

ecotourism

humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals. Ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals.

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

male parental care

parental care is carried out by males

monogamous

Having one mate at a time.

motile

having the capacity to move from one place to another.

natatorial

specialized for swimming

native range

the area in which the animal is naturally found, the region in which it is endemic.

oceanic islands

islands that are not part of continental shelf areas, they are not, and have never been, connected to a continental land mass, most typically these are volcanic islands.

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

pelagic

An aquatic biome consisting of the open ocean, far from land, does not include sea bottom (benthic zone).

piscivore

an animal that mainly eats fish

saltwater or marine

mainly lives in oceans, seas, or other bodies of salt water.

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

tundra

A terrestrial biome with low, shrubby or mat-like vegetation found at extremely high latitudes or elevations, near the limit of plant growth. Soils usually subject to permafrost. Plant diversity is typically low and the growing season is short.

visual

uses sight to communicate

References

Aubin, T., P. Jouventin. 1998. Cocktail-party effect in king penguin colonies. Proceedings of the Royal Society B: Biological Sciences, 265: 1665-1673.

Aubin, T., P. Jouventin, C. Hildebrand. 2000. Penguins use the two-voice system to recognize each other. Proceedings of the Royal Society B: Biological Sciences, 267: 1081-1087.

Baxter, W. 1974. People or Penguins: The Case for Optimal Pollution. New York: Columbia University Press.

Bried, J., F. Jiguet, P. Jouventin. 1999. Why do Aptenodytes penguins have high divorce rates?. The Auk, 116: 504-512.

Charbonnier, Y., K. Delord, J. Thiebot. 2009. King-size fast food for Antartic fur seals. Polar Biology, 33: 721-724.

Cote, S. 2000. Aggressiveness in king penguins in relation to reproductive status and territory location. Animal Behaviour, 59: 813-821.

Dobson, F., P. Jouventin. 2003. Use of the nest site as a rendezvous in penguins. Waterbirds: The International Journal of Waterbird Biology, 26: 409-415.

Emslie, S., N. Karnovsky, W. Trivelpiece. 1995. Avian predation at penguin colonies on King George Island, Antarctica. The Wilson Bulletin, 107: 317-327.

Flower, 1938. Furter notes on the duration of life in animals. IV. Birds. Proceedings of the Zoological Society of London, Ser. A: 195-235.

Gauthier-Clerc, M., Y. Clerquin, Y. Handrich. 1998. Hyperinfestation by ticks Ixodes uriae: a possible cause of death in adult king penguins, a long-lived seabird. Colonial Waterbirds, 21: 229-233.

Jouventin, P., T. Aubin, T. Lengagne. 1999. Finding a parent in a king penguin colony: the acoustic system of individual recognition. Animal Behavior, 57: 1175-1183.

Jouventin, P., P. Nolan, J. Ornborg, F. Dobson. 2005. Ultraviolet beak spots in king and emperor penguins. The Condor, 107: 144-150.

Kooyman, G., Y. Cherel, Y. Le Maho, J. Croxall, P. Thorson, V. Ridoux, C. Kooyman. 1992. Diving behavior and energetics during foraging cycles in king penguins. Ecological Monographs, 62: 143-163.

Lockley, R. 1984. Seabirds of the World. New York, NY: Facts on File.

McGonigal, D., L. Woodworth. 2001. Antarctica and the Arctic: The Complete Encyclopedia. Ontario: Firefly Books.

Nelson, B. 1979. Seabirds, Their Biology and Ecology. New York, NY: A & W Publishers Inc..

Nicolaus, M., C. Le Bohec, P. Nolan, M. Gauthier-Clerc, Y. Le Maho, J. Komdeur, P. Jouventin. 2007. Ornamental colors reveal age in the king penguin. Polar Biology, 31: 53-61.

Nolan, P., F. Dobson, M. Nicolaus, T. Karels, K. McGraw, P. Jouventin. 2010. Mutual mate choice for colorful traits in king penguins. Ethology, 116: 635-644.

Olsson, O. 1997. Clutch abandonment: a state-dependent decision in king penguins. Journal of Avian Biology, 28: 264-267.

Olsson, O. 1996. Seasonal effects of timing and reproduction in the king penguin: a unique breeding cycle. Journal of Avian Biology, 27: 7-14.

Otley, H. 2005. Nature-based tourism: Experiences at the volunteer point penguin colony in the Falkland Islands. Marine Ornithology, 33: 181-187.

Parmelee, D. 1992. Antarctic Birds: Ecological and Behavioral Approaches. Minneapolis: University of Minnnesota Press.

Putz, K., C. Bost. 1994. Feeding behavior of free-ranging king penguins (Aptenodytes patagonicus). Ecology, 75: 489-497.

Richdale, L. 1951. Sexual Behavior in Penguins. Lawrence: University of Kansas Press.

Shirihai, H. 2008. The Complete Guide to Antarctic Wildlife: Birds and Marine Mammals of the Antarctic Continent and the Southern Ocean. Princeton, NJ: Princeton University Press.

Viera, V., P. Nolan, S. Cote, P. Jouventin, R. Groscolas. 2007. Is territory defence related to plumage ornaments in the king penguin Aptenodytes patagonicus?. Ethology, 114: 146-153.

de Villiers, M., P. de Bruyn. 2004. Stone-swallowing by three species of penguins at sub-antarctic Marion Island. Marine Ornithology, 32: 185-186.

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