Penguin

For other uses, see Penguin (disambiguation).

Penguins (order Sphenisciformes, family Sphenisci-dae) are a group of aquatic, ightless birds living al-most exclusively in the Southern Hemisphere, especiallyin Antarctica. Highly adapted for life in the water, pen-guins have countershaded dark and white plumage, andtheir wings have evolved into ippers. Most penguinsfeed on krill, sh, squid and other forms of sealife caughtwhile swimming underwater. They spend about half oftheir lives on land and half in the oceans.Although all penguin species are native to the SouthernHemisphere, they are not found only in cold climates,such as Antarctica. In fact, only a few species of pen-guin live so far south. Several species are found in thetemperate zone, and one species, the Galpagos penguin,lives near the equator.The largest living species is the emperor penguin (Apten-odytes forsteri): on average adults are about 1.1 m (3 ft7 in) tall and weigh 35 kg (77 lb) or more. The small-est penguin species is the little blue penguin (Eudyptulaminor), also known as the fairy penguin, which standsaround 40 cm (16 in) tall and weighs 1 kg (2.2 lb). Amongextant penguins, larger penguins inhabit colder regions,while smaller penguins are generally found in temper-ate or even tropical climates (see also Bergmanns rule).Some prehistoric species attained enormous sizes, be-coming as tall or as heavy as an adult human. Thesewere not restricted to Antarctic regions; on the contrary,subantarctic regions harboured high diversity, and at leastone giant penguin occurred in a region not quite 2,000km south of the equator 35 mya, in a climate decidedlywarmer than today.

1 EtymologyThe word penguin rst appears in the 16th century asa synonym for great auk.[1] When European explorersdiscovered what are today known as penguins in theSouthern Hemisphere, they noticed their similar appear-ance to the great auk of the Northern Hemisphere, andnamed them after this bird, although they are not closelyrelated.[2]

The etymology of the word penguin is still debated. TheEnglish word is not apparently of French,[1] Breton[3]or Spanish[4] origin (the latter two are attributed to theFrench word pingouin "auk"), but rst appears in English

or Dutch.[1]

Some dictionaries suggest a derivation from Welsh pen,head and gwyn, white, including the Oxford En-glish Dictionary, the American Heritage Dictionary,[5]the Century Dictionary[5] and Merriam-Webster,[6] onthe basis that the name was originally applied to thegreat auk, either because it was found on White Head Is-land (Welsh Pen Gwyn) in Newfoundland, or because ithad white circles around its eyes (though the head wasblack). However, the Welsh word pen is also used tomean front, foremost part or extremity and there-fore white front is far more likely to be the sense inwhich Welsh sailors used the term when referring to thebird. Indeed the Welsh expression for bow or prowof a ship is pen blaen.[7]

An alternative etymology links the word to Latin pinguis,which means fat. In Dutch, the alternative word forpenguin is fat-goose (vetgans see: Dutch wiki or dic-tionaries under Pingun), and would indicate this bird re-ceived its name from its appearance.

2 Systematics and evolution

2.1 Living species and recent extinctions

Adlie penguin (Pygoscelis adeliae) feeding young. Like its rel-atives, a neatly bi-coloured species with a head marking.

The number of extant penguin species is debated.Depending on which authority is followed, penguinbiodiversity varies between 18 and 20 living species,all in the subfamily Spheniscinae. Some sources con-sider the white-ippered penguin a separate Eudyptulaspecies, while others treat it as a subspecies of thelittle penguin;[8][9] the actual situation seems to be more

1

2 2 SYSTEMATICS AND EVOLUTION

Magellanic penguins (Spheniscus magellanicus). The closedneck collar denotes this species.

Closeup of southern rockhopper penguin (Eudyptes chrysocome)

Two king penguins and one gentoo penguin walking on a beachon South Georgia, British overseas territory

complicated.[10] Similarly, it is still unclear whether theroyal penguin is merely a color morph of the macaronipenguin. The status of the rockhopper penguins is alsounclear.Updated after Marples (1962),[11] Acosta Hospitaleche(2004),[12] and Ksepka et al. (2006).[13]

Subfamily Spheniscinae Modern penguins

Aptenodytes Great penguins King penguin, Aptenodytes patagonicus Emperor penguin, Aptenodytes forsteri

Pygoscelis Brush-tailed penguins Adlie penguin, Pygoscelis adeliae Chinstrap penguin, Pygoscelis antarctica Gentoo penguin, Pygoscelis papua

Eudyptula Little penguins Little blue penguin, Eudyptula minor White-ippered penguin, Eudyptula albosig-

nata (provisional) Spheniscus Banded penguins

Magellanic penguin, Spheniscus magellanicus Humboldt penguin, Spheniscus humboldti Galapagos penguin, Spheniscus mendiculus African penguin, Spheniscus demersus

Megadyptes Yellow-eyed penguin, Megadyptes antipodes Waitaha penguin, Megadyptes waitaha (ex-

tinct) Eudyptes Crested penguins

Fiordland penguin, Eudyptes pachyrynchus Snares penguin, Eudyptes robustus Erect-crested penguin, Eudyptes sclateri Western rockhopper penguin, Eudyptes

chrysocome Eastern rockhopper penguin, Eudyptes lholi Northern rockhopper penguin, Eudyptes mose-

leyi Royal penguin, Eudyptes schlegeli (disputed) Macaroni penguin, Eudyptes chrysolophus Chatham penguin, Eudyptes chathamensis (ex-

tinct)

2.2 Fossil generaOrder Sphenisciformes

Basal and unresolved taxa (all fossil) Waimanu basal (Middle-Late Paleocene) Perudyptes (Middle Eocene of Atacama

Desert, Peru) basal?

2.2 Fossil genera 3

Spheniscidae gen. et sp. indet. CADIC P21 (Leticia Middle Eocene of Punta Torcida,Argentina)[14]

Delphinornis (Middle/Late Eocene? EarlyOligocene of Seymour Island, Antarctica) Palaeeudyptinae, basal, new subfamily 1?

Archaeospheniscus (Middle/Late Eocene Late Oligocene) Palaeeudyptinae? New sub-family 2?

Marambiornis (Late Eocene ? EarlyOligocene of Seymour Island, Antarctica) Palaeeudyptinae, basal, new subfamily 1?

Mesetaornis (Late Eocene ? EarlyOligocene of Seymour Island, Antarc-tica) Palaeeudyptinae, basal, new subfamily1?

Tonniornis (Late Eocene ? Early Oligoceneof Seymour Island, Antarctica)

Wimanornis (Late Eocene ? Early Oligoceneof Seymour Island, Antarctica)

Duntroonornis (Late Oligocene of Otago, NewZealand) possibly Spheniscinae

Korora (Late Oligocene of S Canterbury, NewZealand)

Kairuku (Late Oligocene of E South Island,New Zealand)

Platydyptes (Late Oligocene of New Zealand) possibly not monophyletic; Palaeeudyptinae,Paraptenodytinae or new subfamily?[15]

Spheniscidae gen. et sp. indet. (LateOligocene/Early Miocene of Hakataramea,New Zealand)[16]

Madrynornis (Puerto Madryn Late Mioceneof Argentina) possibly Spheniscinae

Pseudaptenodytes (Late Miocene/EarlyPliocene)

Dege (Early Pliocene of South Africa) pos-sibly Spheniscinae

Marplesornis (Early Pliocene) possiblySpheniscinae

Nucleornis (Early Pliocene of Duinfontain,South Africa) possibly Spheniscinae

Tereingaornis (Middle Pliocene of NewZealand)

Inguza (Late Pliocene) probably Sphenisci-nae; formerly Spheniscus predemersus

Palaeoapterodytes (Late Oligocene/EarlyMiocene of Argentina)

Family Spheniscidae

A reconstruction of the ancient penguin Icadyptes

Subfamily Palaeeudyptinae Giant pen-guins (fossil) Crossvallia (Cross Valley Late Paleocene

of Seymour Island, Antarctica) tenta-tively assigned to this subfamily

Anthropornis (Middle Eocene? EarlyOligocene of Seymour Island, Antarctica) tentatively assigned to this subfamily Nordenskjoelds giant penguin,

Anthropornis nordenskjoeldi Icadyptes (Late Eocene of Atacama

Desert, Peru) Palaeeudyptes (Middle/Late Eocene

Late Oligocene) polyphyletic; some be-long in other subfamilies

Pachydyptes (Late Eocene) Anthropodyptes (Middle Miocene) ten-

tatively assigned to this subfamily Subfamily Paraptenodytinae Stout-footed

penguins (fossil) Arthrodytes (San Julian Late

Eocene/Early Oligocene PatagoniaEarly Miocene of Patagonia, Argentina)

Paraptenodytes (Early LateMiocene/Early Pliocene)

Subfamily Palaeospheniscinae Slender-footed penguins (fossil) Eretiscus (Patagonia Early Miocene of

Patagonia, Argentina)

4 2 SYSTEMATICS AND EVOLUTION

Palaeospheniscus (Early? LateMiocene/Early Pliocene) includesChubutodyptes

The Early Oligocene genus Cruschedula was formerlythought to belong to Spheniscidae, however reexamina-tion of the holotype in 1943 resulted in the genus beingplaced in Accipitridae.[17] Further examination in 1980resulted in placement as Aves incertae sedis.[18]

2.3 TaxonomySome recent sources[13][14] apply the phylogenetic taxonSpheniscidae to what here is referred to as Sphenisci-nae. Furthermore, they restrict the phylogenetic taxonSphenisciformes to ightless taxa, and establish the phy-logenetic taxon Pansphenisciformes as equivalent to theLinnean taxon Sphenisciformes,[14] i.e., including any y-ing basal proto-penguins to be discovered eventually.Given that neither the relationships of the penguin sub-families to each other nor the placement of the pen-guins in the avian phylogeny is presently resolved, this isconfusing, so the established Linnean system is followedhere.

2.4 Evolution

Penguin tracks in the sand on Bruny Island, Tasmania

The evolutionary history of penguins is well-researchedand represents a showcase of evolutionary biogeography;though as penguin bones of any one species vary much insize and few good specimens are known, the alpha tax-onomy of many prehistoric forms still leaves much to bedesired. Some seminal articles about penguin prehistoryhave been published since 2005,[13][19][16][20] the evolu-tion of the living genera can be considered resolved bynow.The basal penguins lived around the time of theCretaceousPaleogene extinction event somewhere in thegeneral area of (southern) New Zealand and Byrd Land,Antarctica.[13] Due to plate tectonics, these areas were at

that time less than 1,500 kilometres (930 mi) apart ratherthan the 4,000 kilometres (2,500 mi) of today. The mostrecent common ancestor of penguins and their sister cladecan be roughly dated to the CampanianMaastrichtianboundary, around 7068 mya.[16][20][21] What can be saidas certainly as possible in the absence of direct (i.e., fos-sil) evidence is that, by the end of the Cretaceous, the pen-guin lineage must have been evolutionarily well distinct,though much less so morphologically; it is fairly likely thatthey were not yet entirely ightless at that time, as ight-less birds have generally low resilience to the breakdownof trophic webs that follows the initial phase of mass ex-tinctions because of their below-average dispersal capa-bilities (see also Flightless cormorant).

2.4.1 The basal fossils

The oldest known fossil penguin species is Waimanumanneringi, which lived in the early Paleocene epochof New Zealand, or about 62 mya.[20] While they werenot as well-adapted to aquatic life as modern penguins,Waimanu were generally loon-like birds but alreadyightless, with short wings adapted for deep diving.[20]They swam on the surface using mainly their feet, butthe wings were as opposed to most other diving birds(both living and extinct) already adapting to underwaterlocomotion.[22]

Perudyptes from northern Peru was dated to 42 mya.An unnamed fossil from Argentina proves that, by theBartonian (Middle Eocene), some 3938 mya,[23] prim-itive penguins had spread to South America and were inthe process of expanding into Atlantic waters.[14]

2.4.2 Palaeeudyptines

During the Late Eocene and the Early Oligocene (4030 mya), some lineages of gigantic penguins existed.Nordenskjoelds giant penguin was the tallest, growingnearly 1.80 meters (5.9 feet) tall. The New Zealand gi-ant penguin was probably the heaviest, weighing 80 kg ormore. Both were found on New Zealand, the former alsoin the Antarctic farther eastwards.Traditionally, most extinct species of penguins, giant orsmall, had been placed in the paraphyletic subfamilycalled Palaeeudyptinae. More recently, with new taxabeing discovered and placed in the phylogeny if possi-ble, it is becoming accepted that there were at least twomajor extinct lineages. One or two closely related onesoccurred in Patagonia, and at least one otherwhich isor includes the paleeudyptines as recognized today oc-curred on most Antarctic and Subantarctic coasts.But size plasticity seems to have been great at this initialstage of penguin radiation: on Seymour Island, Antarc-tica, for example, around 10 known species of penguinsranging in size from medium to huge apparently coexistedsome 35 mya during the Priabonian (Late Eocene).[24] It

2.4 Evolution 5

is not even known whether the gigantic palaeeudyptinesconstitute a monophyletic lineage, or whether gigan-tism was evolved independently in a much restrictedPalaeeudyptinae and the Anthropornithinae whetherthey were considered valid, or whether there was a widesize range present in the Palaeeudyptinae as delimited asusually done these days (i.e., including Anthropornis nor-denskjoeldi).[13] The oldest well-described giant penguin,the 5-foot (1.5 m)-tall Icadyptes salasi, actually occurredas far north as northern Peru about 36 mya.In any case, the gigantic penguins had disappeared bythe end of the Paleogene, around 25 mya. Their de-cline and disappearance coincided with the spread of theSqualodontoidea and other primitive, sh-eating toothedwhales, which certainly competed with them for food,and were ultimately more successful.[16] A new lineage,the Paraptenodytes, which includes smaller but decidedlystout-legged forms, had already arisen in southernmostSouth America by that time. The early Neogene saw theemergence of yet another morphotype in the same area,the similarly sized but more gracile Palaeospheniscinae,as well as the radiation that gave rise to the penguinbiodiversity of our time.

2.4.3 Origin and systematics of modern penguins

Modern penguins constitute two undisputed clades andanother two more basal genera with more ambiguousrelationships.[19] The origin of the Spheniscinae lies prob-ably in the latest Paleogene, and geographically it musthave been much the same as the general area in whichthe order evolved: the oceans between the Australia-NewZealand region and the Antarctic.[16] Presumably diverg-ing from other penguins around 40 mya,[16] it seems thatthe Spheniscinae were for quite some time limited totheir ancestral area, as the well-researched deposits ofthe Antarctic Peninsula and Patagonia have not yieldedPaleogene fossils of the subfamily. Also, the earliestspheniscine lineages are those with the most southern dis-tribution.The genus Aptenodytes appears to be the basalmost di-vergence among living penguins[13][25] they have brightyellow-orange neck, breast, and bill patches; incubate byplacing their eggs on their feet, and when they hatch thechicks are almost naked. This genus has a distributioncentered on the Antarctic coasts and barely extends tosome Subantarctic islands today.Pygoscelis contains species with a fairly simple black-and-white head pattern; their distribution is intermedi-ate, centered on Antarctic coasts but extending some-what northwards from there. In external morphology,these apparently still resemble the common ancestor ofthe Spheniscinae, as Aptenodytes ' autapomorphies arein most cases fairly pronounced adaptations related tothat genus extreme habitat conditions. As the formergenus, Pygoscelis seems to have diverged during the

Bartonian,[26] but the range expansion and radiation thatled to the present-day diversity probably did not occur un-til much later; around the Burdigalian stage of the EarlyMiocene, roughly 2015 mya.[16]

The genera Spheniscus and Eudyptula contain specieswith a mostly Subantarctic distribution centered on SouthAmerica; some, however, range quite far northwards.They all lack carotenoid coloration, and the former genushas a conspicuous banded head pattern; they are uniqueamong living penguins by nesting in burrows. This groupprobably radiated eastwards with the Antarctic Circum-polar Current out of the ancestral range of modern pen-guins throughout the Chattian (Late Oligocene), startingapproximately 28 mya.[16] While the two genera sepa-rated during this time, the present-day diversity is theresult of a Pliocene radiation, taking place some 42mya.[16]

The MegadyptesEudyptes clade occurs at similarlatitudes (though not as far north as the Galapagospenguin), has its highest diversity in the New Zealandregion, and represents a westward dispersal. They arecharacterized by hairy yellow ornamental head feathers;their bills are at least partly red. These two generadiverged apparently in the Middle Miocene (Langhian,roughly 1514 mya), but again, the living species ofEudyptes are the product of a later radiation, stretchingfrom about the late Tortonian (Late Miocene, 8 mya) tothe end of the Pliocene.[16]

The geographical and temporal pattern or spheniscineevolution corresponds closely to two episodes of globalcooling documented in the paleoclimatic record.[16] Theemergence of the Subantarctic lineage at the end of theBartonian corresponds with the onset of the slow periodof cooling that eventually led to the ice ages some 35 mil-lion years later. With habitat on the Antarctic coasts de-clining, by the Priabonian more hospitable conditions formost penguins existed in the Subantarctic regions ratherthan in Antarctica itself.[27] Notably, the cold AntarcticCircumpolar Current also started as a continuous circum-polar ow only around 30 mya, on the one hand forcingthe Antarctic cooling, and on the other facilitating theeastward expansion of Spheniscus to South America andeventually beyond.[16] Despite this, there is no fossil ev-idence to support the idea of a crown radiation from theAntarctic continent in the Paleogene.[27]

Later, an interspersed period of slight warming was endedby the Middle Miocene Climate Transition, a sharp dropin global average temperature from 1412 mya, and sim-ilar abrupt cooling events followed at 8 mya and 4 mya;by the end of the Tortonian, the Antarctic ice sheet wasalready much like today in volume and extent. The emer-gence of most of todays Subantarctic penguin species al-most certainly was caused by this sequence of Neogeneclimate shifts.

6 3 ANATOMY AND PHYSIOLOGY

2.4.4 Relationship to other bird orders

Penguin ancestry beyond Waimanu remains unknownand not well-resolved by molecular or morphologicalanalyses. The latter tend to be confounded by thestrong adaptive autapomorphies of the Sphenisciformes;a sometimes perceived fairly close relationship betweenpenguins and grebes is almost certainly an error basedon both groups strong diving adaptations, which arehomoplasies. On the other hand, dierent DNA se-quence datasets do not agree in detail with each other ei-ther.

Humboldt penguins in an aquarium. The penguin is an accom-plished swimmer, having ippers instead of wings.

What seems clear is that penguins belong to a cladeof Neoaves (living birds except paleognaths and fowl)that comprises what is sometimes called higher wa-terbirds to distinguish them from the more ancientwaterfowl. This group contains such birds as storks,rails, and the seabirds, with the possible exception of theCharadriiformes.[28]

Inside this group, penguin relationships are far less clear.Depending on the analysis and dataset, a close relation-ship to Ciconiiformes[20] or to Procellariiformes[16] hasbeen suggested. Some think the penguin-like plotopterids(usually considered relatives of anhingas and cormorants)may actually be a sister group of the penguins, and thatpenguins may have ultimately shared a common ancestorwith the Pelecaniformes and consequently would have tobe included in that order, or that the plotopterids werenot as close to other pelecaniforms as generally assumed,which would necessitate splitting the traditional Pelecan-iformes in three.[29]

A 2014 analysis of whole genomes of 48 representativebird species has concluded that penguins are the sistergroup of Procellariiformes,[30] from which they divergedabout 60 million years ago (95% CI, 56.8-62.7).[31]

3 Anatomy and physiologyPenguins are superbly adapted to aquatic life. Theirvestigial wings have become ippers, useless for ight in

Orcas swim by an iceberg with Adelie penguins in the Ross Sea,Antarctica. The Drygalski ice tongue is visible in the background.

the air. In the water, however, penguins are astonishinglyagile. Penguins swimming looks very similar to birdsight in the air.[32] Within the smooth plumage a layer ofair is preserved, ensuring buoyancy. The air layer alsohelps insulate the birds in cold waters. On land, penguinsuse their tails and wings to maintain balance for their up-right stance.All penguins are countershaded for camouage that is,they have black backs and wings with white fronts.[33]A predator looking up from below (such as an orca or aleopard seal) has diculty distinguishing between a whitepenguin belly and the reective water surface. The darkplumage on their backs camouages them from above.Diving penguins reach 6 to 12 km/h (3.7 to 7.5 mph),though there are reports of velocities of 27 km/h (17mph) (which are more realistic in the case of startledight). The small penguins do not usually dive deep; theycatch their prey near the surface in dives that normally lastonly one or two minutes. Larger penguins can dive deepin case of need. Dives of the large emperor penguin havebeen recorded reaching a depth of 565 m (1,854 ft) forup to 22 minutes.Penguins either waddle on their feet or slide on their bel-lies across the snow, a movement called tobogganing,which conserves energy while moving quickly. They alsojump with both feet together if they want to move morequickly or cross steep or rocky terrain.Penguins have an average sense of hearing for birds;[34]this is used by parents and chicks to locate one another incrowded colonies.[35] Their eyes are adapted for underwa-ter vision, and are their primary means of locating preyand avoiding predators; in air it has been suggested thatthey are nearsighted, although research has not supportedthis hypothesis.[36]

Penguins have a thick layer of insulating feathers thatkeeps them warm in water (heat loss in water is muchgreater than in air). The emperor penguin has the largestbody mass of all penguins, which further reduces relativesurface area and heat loss. They also are able to controlblood ow to their extremities, reducing the amount of

7Gentoo penguin swimming underwater at Nagasaki PenguinAquarium.

blood that gets cold, but still keeping the extremities fromfreezing. In the extreme cold of the Antarctic winter, thefemales are at sea shing for food leaving the males tobrave the weather by themselves. They often huddle to-gether to keep warm and rotate positions to make sure thateach penguin gets a turn in the center of the heat pack.Calculations of the heat loss and retention ability of ma-rine endotherms

8 6 PENGUINS AND HUMANS

Chinstrap penguins in Antarctica.

though the rate the same pair recouples varies drasti-cally. Most penguins lay two eggs in a clutch, althoughthe two largest species, the emperor and the king pen-guins, lay only one.[54] With the exception of the emperorpenguin, where the male does it all, all penguins share theincubation duties.[55] These incubation shifts can last daysand even weeks as one member of the pair feeds at sea.Penguins generally only lay one brood; the exception isthe little penguin, which can raise two or three broods ina season.[56]

Penguin eggs are smaller than any other bird species whencompared proportionally to the weight of the parent birds;at 52 g (2 oz), the little penguin egg is 4.7% of its moth-ers weight, and the 450 g (1 lb) emperor penguin egg is2.3%.[54] The relatively thick shell forms between 10 and16% of the weight of a penguin egg, presumably to min-imize the risk of breakage in an adverse nesting environ-ment. The yolk, too, is large, and comprises 2231% ofthe egg. Some yolk often remains when a chick is born,and is thought to help sustain the chick if the parents aredelayed in returning with food.[57]

When mothers lose a chick, they sometimes attempt tosteal another mothers chick, usually unsuccessfully asother females in the vicinity assist the defending motherin keeping her chick. In some species, such as emperorpenguins, young penguins assemble in large groups calledcrches.

6 Penguins and humansPenguins seem to have no special fear of humans, andhave approached groups of explorers without hesitation.This is probably because penguins have no land predatorsin Antarctica or the nearby oshore islands. Dogs preyedupon penguins while they were allowed in Antarctica dur-ing the age of early human exploration as sled dogs, butdogs are now banned from Antarctica.[58] Instead, adultpenguins are at risk at sea from predators such as sharks,the orca, and the leopard seal. Typically, penguins do not

A cook on the Endurance preparing a penguin for consumption

A penguin encounters a human during Antarctic summer.

approach closer than about 3 meters (9.8 feet) at whichpoint they become nervous. This is also the distance thatAntarctic tourists are told to keep from penguins (touristsare not supposed to approach closer than 3 meters, but arenot expected to withdraw if the penguins come closer).In June 2011, a penguin came ashore on New ZealandsPeka Peka Beach, 3200 km o course on its journey toAntarctica.[59] Nicknamed Happy Feet, after the movie ofthe same name, it was suering from heat exhaustion andhad to undergo a number of operations to remove objectslike driftwood and sand from its stomach.[60] Happy Feetwas a media sensation, with extensive coverage on TVand the web, including a live stream that had thousandsof views[61] and a visit from English actor Stephen Fry.[62]

Once he had recovered, Happy Feet was released backinto the water south of New Zealand.[63]

6.1 In popular culture

Main article: Cultural depictions of penguinsPenguins are popular around the world, primarily for

9Tux the Linux kernel mascot

their unusually upright, waddling gait and (compared toother birds) lack of fear of humans. Their striking black-and-white plumage is often likened to a white tie suit.Mistakenly, some artists and writers have penguins basedat the North Pole. This is incorrect, as there are almostno wild penguins in the Northern Hemisphere, except thesmall group on the northernmost of the Galpagos. Thecartoon seriesChillyWilly helped perpetuate this myth, asthe title penguin would interact with northern-hemispherespecies, such as polar bears and walruses.Penguins have been the subject of many books and lms,such as Happy Feet, Surfs Up and The Penguins of Mada-gascar, all CGI lms; March of the Penguins, a docu-mentary based on the migration process of the emperorpenguin; and a parody titled Farce of the Penguins. Mr.Poppers Penguins is a childrens book written by Richardand Florence Atwater; it was named a Newbery HonorBook in 1939. Penguins have also found their way intoa number of cartoons and television dramas; perhaps themost notable of these is Pingu, created by Silvio Maz-zola in 1986 and covering more than 100 short episodes.At the end of 2009, Entertainment Weekly put it on itsend-of-the-decade, best-of list, saying, Whether theywere walking (March of the Penguins), dancing (HappyFeet), or hanging ten (Surfs Up), these oddly adorablebirds took ight at the box oce all decade long. [64]

A video game called Pengo was released by Sega in 1982.Set in Antarctica, the player controls a penguin characterwho must navigate mazes of ice-cubes. The player is re-warded with cut-scenes of animated penguins marching,dancing, saluting and playing peekaboo. Several remakesand enhanced editions have followed, most recently in2012.

Several pro, minor, college and high school sport teamshave named themselves after the species, with thePittsburgh Penguins team in the National Hockey Leagueand the Youngstown State Penguins being the most rec-ognizable.The tendency of penguins to form large groups feeds thestereotype that they all look exactly alike, a popular notionexploited by cartoonists such as Gary Larson.Penguins featured regularly in the cartoons of UK car-toonist Steve Bell in his strip in The Guardian Newspa-per, particularly during and following the Falklands War,and the well-known Opus the Penguin from the cartoonsof Berkeley Breathed, is also described as hailing fromthe Falklands. Opus was a comical, existentialist pen-guin character in the cartoons Bloom County, Outland andOpus. He was also the star in the Christmas show AWishfor Wings That Work.In the mid-2000s, penguins became one of the most pub-licized species of animals that form lasting homosexualcouples. A childrens book, And Tango Makes Three, waswritten about one such penguin family in the New YorkZoo.

7 References[1] PINGOUIN : Etymologie de PINGOUIN. Centre Na-

tional de Ressources Textuelles et Lexicales. RetrievedJanuary 25, 2010.

[2] Croord, Emily (1989). Gone Forever: The Great Auk.New York: Crestwood House. p. 10. ISBN 0-89686-459-6.

[3] Harper, Douglas. penguin. Online Etymology Dictio-nary.

[4] pingino. Diccionario de la lengua espaola. rae.es

[5] penguin. Wordnik.com. Retrieved October 7, 2014.

[6] Penguin Denition. Merriam-webster.com. August31, 2012. Retrieved September 8, 2013.

[7] Pen - Denition. Prifysgol Cymru. Retrieved February18, 2015.

[8] Williams

[9] Davis; Lloyd S. & Renner; M. (1995). Penguins. London:T & A D Poyser. ISBN 0-7136-6550-5

[10] Banks, Jonathan C.; Mitchell, Anthony D.; Waas, JosephR. & Paterson, Adrian M. (2002). An unexpected pat-tern of molecular divergence within the blue penguin (Eu-dyptula minor) complex (PDF). Notornis 49 (1): 2938.

[11] Marples, B. J. (1962): Observations on the history of pen-guins. In: Leeper, G. W. (ed.), The evolution of livingorganisms. Melbourne, Melbourne University Press. pp.408416.

10 7 REFERENCES

[12] (Spanish) Acosta Hospitaleche, Carolina (2004) Lospinginos (Aves, Sphenisciformes) fsiles de Patagonia.Sistemtica, biogeografa y evolucin. Doctoral thesis, De-partment of Natural Sciences and Museum, UniversidadNacional de La Plata. La Plata, Argentina.

[13] Ksepka, D. T. B., Sara; Giannini, Norberto P; (2006).The phylogeny of the living and fossil Sphenisci-formes (penguins)". Cladistics 22 (5): 412441.doi:10.1111/j.1096-0031.2006.00116.x.

[14] Clarke, Julia A.; Olivero, Eduardo B. & Puerta, Pablo(2003). Description of the earliest fossil penguinfrom South America and rst Paleogene vertebrate lo-cality of Tierra Del Fuego, Argentina (PDF). Ameri-can Museum novitates 3423: 118. doi:10.1206/0003-0082(2003)4232.0.co;2.

[15] Platydyptes novaezealandiae; holotype. Collections On-line. Museum of New Zealand Te Papa Tongarewa. Re-trieved July 16, 2010.

[16] Baker, A. J; Pereira, S. L.; Haddrath, O. P; Edge, K.-A. (2006). Multiple gene evidence for expansion of ex-tant penguins out of Antarctica due to global cooling.Proceedings of the Royal Society B: Biological Sciences273 (1582): 117. doi:10.1098/rspb.2005.3260. PMC1560011. PMID 16519228.

[17] Simpson, G.G. (1946). Fossil penguins (PDF). Bulletinof the American Museum of Natural History 81.

[18] Olson, S.L. (1985). Faunal Turnover in South Amer-ican Fossil Avifaunas: The Insuciencies of the Fos-sil Record (PDF). Evolution 39 (5): 11741177.doi:10.2307/2408747. JSTOR 2408747.

[19] Bertelli, S.; Giannini, N. P. (2005). A phylogeny of ex-tant penguins (Aves: Sphenisciformes) combining mor-phology and mitochondrial sequences. Cladistics 21 (3):209. doi:10.1111/j.1096-0031.2005.00065.x.

[20] Slack, Kerryn E.; Jones, Craig M.; Ando, Tatsuro;Harrison, G.L. Abby"; Fordyce, R. Ewan; Arnason,Ulfur; Penny, David (2006). Early Penguin Fossils,plus Mitochondrial Genomes, Calibrate Avian Evolu-tion. Molecular Biology and Evolution 23 (6): 11441155. doi:10.1093/molbev/msj124. PMID 16533822.Supplementary Material

[21] The exact divergence dates according to Baker et al.(2006) mentioned in this section are not as precisely re-solved as it appears to be due to uncertainties of themolecular clock used.

[22] Elliott, K. H.; Ricklefs, R. E.; Gaston, A. J.; Hatch, S.A.; Speakman, J. R.; Davoren, G. K. (2013). High ightcosts, but low dive costs, in auks support the biomechan-ical hypothesis for ightlessness in penguins. Proceed-ings of the National Academy of Sciences 110 (23): 9380.doi:10.1073/pnas.1304838110.

[23] Contra Baker et al. (2006).

[24] Jadwiszczak, Piotr (2006). Eocene penguins of SeymourIsland, Antarctica: taxonomy (PDF). Polish Polar Re-search 27 (1): 362.

[25] Christidis L, Boles WE (2008). Systematics and Taxon-omy of Australian Birds. Canberra: CSIRO Publishing.p. 97. ISBN 978-0-643-06511-6.

[26] It is likely that, during the Bartonian, there was a near-synchronous but allopatric split between the ancestors ofAptenodytes, Pygoscelis, and the common ancestor of allremaining genera (Baker).

[27] Baker, A., Pereira, SL, Haddrath, OP, Edge, KA (2006).Multiple gene evidence for expansion of extant penguinsout of Antarctica due to global cooling. Proceedings ofthe Royal Society B-Biological Sciences 273 (1582): 117. doi:10.1098/rspb.2005.3260. PMC 1560011. PMID16519228.

[28] Fain, M. G.; Houde, P. (2004). Parallel Radiations in thePrimary Clades of Birds (PDF). Evolution 58 (11): 2558.doi:10.1554/04-235. PMID 15612298.

[29] Mayr, G. (2005). Tertiary plotopterids (Aves, Plotopteri-dae) and a novel hypothesis on the phylogenetic relation-ships of penguins (Spheniscidae)" (PDF). Journal of Zo-ological Systematics and Evolutionary Research 43: 61.doi:10.1111/j.1439-0469.2004.00291.x.

[30] Jarvis, E. D.; Mirarab, S.; Aberer, A. J.; Li, B.; Houde,P.; Li, C.; Ho, S. Y. W.; Faircloth, B. C.; Nabholz, B.;Howard, J. T.; Suh, A.; Weber, C. C.; Da Fonseca, R. R.;Li, J.; Zhang, F.; Li, H.; Zhou, L.; Narula, N.; Liu, L.;Ganapathy, G.; Boussau, B.; Bayzid, M. S.; Zavidovych,V.; Subramanian, S.; Gabaldon, T.; Capella-Gutierrez,S.; Huerta-Cepas, J.; Rekepalli, B.; Munch, K. et al.(2014). Whole-genome analyses resolve early branchesin the tree of life of modern birds (PDF). Science346 (6215): 13201331. doi:10.1126/science.1253451.PMID 25504713.

[31] Li, C.; Zhang, Y.; Li, J.; Kong, L.; Hu, H.; Pan, H.;Xu, L.; Deng, Y.; Li, Q.; Jin, L.; Yu, H.; Chen, Y.;Liu, B.; Yang, L.; Liu, S.; Zhang, Y.; Lang, Y.; Xia,J.; He, W.; Shi, Q.; Subramanian, S.; Millar, C. D.;Meader, S.; Rands, C. M.; Fujita, M. K.; Greenwold, M.J.; Castoe, T. A.; Pollock, D. D.; Gu, W. et al. (2014-12-12). Two Antarctic penguin genomes reveal insightsinto their evolutionary history and molecular changes re-lated to the Antarctic environment. GigaScience 3: 27.doi:10.1186/2047-217X-3-27.

[32] Penguin swimming under water, Galapagos.Youtube.com. April 14, 2008. Retrieved Septem-ber 8, 2013.

[33] Buskey, Theresa. The Antarctic Polar Region. In AlanChristopherson, M.S. The Polar Regions. LIFEPAC. 804N. 2nd Ave. E., Rock Rapids, IA: Alpha Omegan Publi-cations, Inc. ISBN 978-1-58095-156-2.

[34] Wever, E. G.; Herman, P. N.; Simmons, J. A.; Hert-zler, D. R. (1969). Hearing in the blackfooted pen-guin, Spheniscus demersus, as represented by the cochlearpotentials. Proceedings of the National Academy ofSciences of the United States of America 63 (3): 67680. doi:10.1073/pnas.63.3.676. JSTOR 59401. PMC223504. PMID 5259756.

11

[35] Jouventin, P; Aubin, T; Lengagne, T (1999). Finding aparent in a king penguin colony: The acoustic system ofindividual recognition. Animal Behaviour 57 (6): 11751183. doi:10.1006/anbe.1999.1086. PMID 10373249.

[36] Sivak, J; Howland, H. C.; McGill-Harelstad, P (1987).Vision of the Humboldt penguin (Spheniscus humboldti)in air and water. Proceedings of the Royal Society of Lon-don. Series B, Containing papers of a Biological charac-ter. Royal Society (Great Britain) 229 (1257): 46772.JSTOR 36191. PMID 2881308.

[37] Downhower, J.F.; Blumer, L.S. (1988). Calculating justhow small a whale can be. Nature 335 (6192): 675675.doi:10.1038/335675b0.

[38] Williams, T.D. (1995). The penguins: Spheniscidae. NewYork: Oxford University Press.

[39] Thomas, D.B.; Fordyce, R.E. (2007). The heterothermicloophole exploited by penguins. Australian Journal ofZoology 55: 317321. doi:10.1071/zo07053.

[40] Thomas, D.B.; Fordyce, R.E. (2012). Biological plas-ticity in penguin heat-retention structures. AnatomicalRecord-Advances in Integrative Anatomy and EvolutionaryBiology 295: 249256. doi:10.1002/ar.21538.

[41] Animal Fact Sheets. Retrieved July 21, 2006.

[42] Humboldt Penguin: Saint Louis Zoo. Retrieved July 21,2006.

[43] van der Merwe, H.J. African Penguins and Penguins ofthe World. iafrica.com. Retrieved July 21, 2006.

[44] Convergence and divergence in the evolution of aquaticbirds by Marcel Van Tuinen, Dave Brian Butvill, John A.W. Kirsch and S. Blair Hedges.

[45] Piper, Ross (2007), Extraordinary Animals: An Encyclo-pedia of Curious and Unusual Animals, Greenwood Press.

[46] Ashton, K. (2002). Patterns of within-species bodysize variation of birds: strong evidence for Bergmannsrule. Global Ecology and Biogeography 11 (6): 505523.doi:10.1046/j.1466-822X.2002.00313.x.

[47] Meiri, S; Dayan, T. (2003). On the validity ofBergmanns rule (PDF). Journal of Biogeography 30 (3):331351. doi:10.1046/j.1365-2699.2003.00837.x.

[48] Clarke, J. A.; Ksepka; Stucchi; Urbina; Giannini;Bertelli; Narvez; Boyd (2007). Paleogene equa-torial penguins challenge the proposed relationshipbetween biogeography, diversity, and Cenozoic cli-mate change. Proceedings of the National Academyof Sciences of the United States of America 104(28): 1154511550. Bibcode:2007PNAS..10411545C.doi:10.1073/pnas.0611099104. PMC 1913862. PMID17601778.

[49] Gohlich, U. B. (2007). The oldest fossil record of theextant penguin genus Spheniscus a new species fromthe Miocene of Peru. Acta Palaeontologica Polonica 52:285298.

[50] Penguins of Australia and New Zealand. Archive.is.Retrieved September 8, 2013.

[51] Jadwiszczak, P. (2009). Penguin past: The current stateof knowledge. Polish Polar Research 30: 328.

[52] Williams, p. 17

[53] Williams, p. 57

[54] Williams, p. 23

[55] Numata, M.; Davis, L. S.; Renner, M. (2000). Prolongedforaging trips and egg desertion in little penguins (Eudyp-tula minor)". New Zealand Journal of Zoology 27 (4):277. doi:10.1080/03014223.2000.9518236.

[56] Reilly PN, Balmford P (1975). A breeding study ofthe little penguin, Eudyptula minor, in Australia. InStonehouse, Bernard. The Biology of Penguins. London:Macmillan. pp. 16187. ISBN 0-333-16791-0.

[57] Williams, p. 24

[58] Removal of the sledge dogs from Antarctica. BritishAntarctic Survey

[59] Emperor penguin not out of the woods yet. 3 News NZ.June 25, 2011.

[60] Happy Feet pulls through third operation. 3 News NZ.June 27, 2011.

[61] The truth about Happy Feets home. 3 News NZ. August8, 2011.

[62] Stephen Fry visits Happy Feet. 3 News NZ. August 11,2011.

[63] VIDEO: Happy Feet starts the long journey home. 3News NZ. August 29, 2011.

[64] Geier, Thom; Jensen, Je; Jordan, Tina; Lyons, Margaret;Markovitz, Adam; Nashawaty, Chris; Pastorek, Whit-ney; Rice, Lynette; Rottenberg, Josh; Schwartz, Missy;Slezak, Michael; Snierson, Dan; Stack, Tim; Stroup,Kate; Tucker, Ken; Vary, Adam B.; Vozick-Levinson, Si-mon; Ward, Kate, The 100 Greatest Movies, TV Shows,Albums, Books, Characters, Scenes, Episodes, Songs,Dresses, Music Videos, and Trends That Entertained UsOver the Past 10 Years. Entertainment Weekly. Decem-ber 11, 2009 (1079/1080):74-84

8 Bibliography

Williams; Tony D. (1995). The Penguins Sphenis-cidae. Oxford: Oxford University Press. ISBN 0-19-854667-X.

12 9 EXTERNAL LINKS

9 External links Two new fossil penguin species found in Peru.

news.nationalgeographic.com Information about penguins at pinguins.info Integrated Taxonomic Information System Penguin information on 70South Penguin research projects on the web Penguin videos and photos on the Internet Bird Col-

lection

Penguin World Penguins in Te Ara the Encyclopedia of New

Zealand Seaworld Penguin Information Lessons in a Land of Wind and Ice from National

Wildlife Magazine 1/15/2010

Live 24/7 camera inside a penguin habitat

13

10 Text and image sources, contributors, and licenses10.1 Text

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14 10 TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

isdabomb, Crouchingtigershiningdragon, Mehmet Karatay, Mishappy, Hawkeye2010, SieBot, Kkgirl94, Pallab1234, Miettarox, Hansonton,Mbz1, X-Fi6, MeegsC, Tom116p, Whiteghost.ink, Bladesonic, Kdp1992, FunkMonk, Toddst1, , Neopet910, Mas-tercoz, Sanoah200, Superniol, Zariya, Jasjamlew, Baseball Bugs, Callumw 88, Jwakizzle, Lightmouse, Mesoso2, Callidior, Danimations,WacoJacko, TheAbsoluteTruth, Opium Dave, Freepic, Dabomb87, Adallow, Dolphin51, Monkeyshgirl, Kach'i kayo, Tripod86, ImageR-emovalBot, SallyForth123, ClueBot, Awlvrn, Rjd0060, Jure Grm, Findfunds, Fenwayguy, AnthonyB415, Magelanic, Neverquick, Pork-tan, AllTheWayGuster, DragonBot, Maxjlegs, Jusdafax, Bingodile, Nicols10~enwiki, NiciVampireHeart, Jotterbot, Psinu, SchreiberBike,Mike7772, Melon247, DerBorg, Ashzekool, Tezero, 10ngalinski, Bighoots15, Rosebudsled, Minkyman, Dfysgrvty, Beefrocks, XLinkBot,Knab5711, JuicyPink1200, Bjpeewee1, Dthomsen8, WikHead, Rabblewabble, Salokineragn, Dman8929, MystBot, WeareND0220, Is-landbaygardener, Addbot, Kemrin45, Wonderful2091, Pvanrompay, Hayl zb, Brandizzo, CarsracBot, Ccacsmss, Z. Patterson, Mfrank9635,Debresser, Kyle1278, OurBoyJake, Tide rolls, Lightbot, Krano, Thisismadness, MuZemike, Luckas-bot, Yobot, TaBOT-zerem, Vincnet,Tearanz, Bility, Craigmclark, Phoenix B 1of3, Galoubet, Xufanc, Materialscientist, Citation bot, Stronach, Quebec99, Obersachsebot,Xqbot, Gigemag76, Khajidha, Gilo1969, Djlund999, GrouchoBot, Ute in DC, Titi2009, Mandrake76, DasallmchtigeJ, Leslyou99, Stat-icVision, Nojan, MGA73bot, Rgbroitman, Citation bot 1, DigbyDalton, Lucky55399, Focus, Secret Saturdays, Jauhienij, Maksa, FoxBot,Vrenator, Zvn, GreenUpGreenOut, Pbrower2a, Sgt. R.K. Blue, Reaper Eternal, Innotata, Tbhotch, Nuyguy123, DARTH SIDIOUS 2,Bitblaster, RjwilmsiBot, Bento00, Alph Bot, Thiridaz, Mandolinface, EmausBot, Rainy1315, Abby 92, Tommy2010, TeleComNasSprVen,Blessing-light, Vilamm, Bryce Carmony, Hazard-SJ, Ocaasi, Ocean Shores, Citron, , L Kensington, Kittenono, Superbowldreamer19,Pun, Skatesmurf, Spamelgoog, ClueBot NG, Jnorton7558, Delusion23, North Atlanticist Usonian, Helpful Pixie Bot, BG19bot, Phnom-Pencil, IGonzalo, Hank2011, Cam0028, Glevum, Ornithodiez, Penguin6617, Mjd15, Justincheng12345-bot, Soulparadox, JYBot, Dexbot,Palacesblowlittle, Twiter is the best, Rotlink, Exenola, Zalunardo8, Ryenocerous, Shaztastic, Sillycap, Ccevol2014, Ldowd16, NGCBen-jamin and Anonymous: 787

10.2 Images File:135_-_Cap_Virgenes_-_Manchot_de_Magellan_-_Janvier_2010.JPG Source: http://upload.wikimedia.org/wikipedia/

commons/3/3b/135_-_Cap_Virgenes_-_Manchot_de_Magellan_-_Janvier_2010.JPG License: CC BY-SA 3.0 Contributors: Own workOriginal artist: Martin St-Amant (S23678)

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EtymologySystematics and evolutionLiving species and recent extinctionsFossil generaTaxonomyEvolutionThe basal fossilsPalaeeudyptinesOrigin and systematics of modern penguinsRelationship to other bird orders

Anatomy and physiologyIsabelline penguins

Distribution and habitatBehaviourBreeding

Penguins and humansIn popular culture

ReferencesBibliographyExternal linksText and image sources, contributors, and licensesTextImagesContent license