Animal Diversity Web

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)

• Biogeographic Regions
• antarctica
  • native
• oceanic islands
  • native

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)

• Habitat Regions
• temperate
• terrestrial
• saltwater or marine

• Terrestrial Biomes
• tundra

• Aquatic Biomes
• pelagic

• 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)

• Other Physical Features
• endothermic
• homoiothermic
• bilateral symmetry

• 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)

• Mating System
• monogamous

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)

• Key Reproductive Features
• iteroparous
• seasonal breeding
• gonochoric/gonochoristic/dioecious (sexes separate)
• sexual
• oviparous

• 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)

• Key Behaviors
• terricolous
• natatorial
• diurnal
• motile
• sedentary
• colonial
• dominance hierarchies

• 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)

• Communication Channels
• visual
• tactile
• acoustic

• Perception Channels
• visual
• tactile
• acoustic
• chemical

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)

• Primary Diet
• carnivore
  • piscivore

• 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)

• Known Predators

  • Brown skuas (Catharacta lonnbergi)
  • Giant petrels (Macronectes giganteus)
  • Antarctic fur seals (Arctocephalus gazelle)
  • Sub-Antarctic fur seals (Arctocephalus tropicalis)
  • Leopard seals (Hydrurga leptonyx)
  • Killer whales (Orcinus orca)

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)

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)

• Positive Impacts
• ecotourism

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.

• IUCN Red List

  Least Concern
  More information

• IUCN Red List

  Least Concern
  More information

• CITES

  No special status

Con­trib­u­tors

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

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.