Penguin

or Dutch.[1]

For other uses, see Penguin (disambiguation).

Some dictionaries suggest a derivation from Welsh pen,

head and gwyn, white, including the Oxford English Dictionary, the American Heritage Dictionary,[5]
the Century Dictionary[5] and Merriam-Webster,[6] on
the basis that the name was originally applied to the
great auk, either because it was found on White Head Island (Welsh Pen Gwyn) in Newfoundland, or because it
had white circles around its eyes (though the head was
black). However, the Welsh word pen is also used to
mean front, foremost part or extremity and therefore white front is far more likely to be the sense in
which Welsh sailors used the term when referring to the
bird. Indeed the Welsh expression for bow or prow
of a ship is pen blaen.[7]

Penguins (order Sphenisciformes, family Spheniscidae) are a group of aquatic, ightless birds living almost exclusively in the Southern Hemisphere, especially
in Antarctica. Highly adapted for life in the water, penguins have countershaded dark and white plumage, and
their wings have evolved into ippers. Most penguins
feed on krill, sh, squid and other forms of sealife caught
while swimming underwater. They spend about half of
their lives on land and half in the oceans.

Although all penguin species are native to the Southern

Hemisphere, they are not found only in cold climates,
such as Antarctica. In fact, only a few species of penguin live so far south. Several species are found in the
temperate zone, and one species, the Galpagos penguin, An alternative etymology links the word to Latin pinguis,
lives near the equator.
which means fat. In Dutch, the alternative word for
penguin is fat-goose (vetgans see: Dutch wiki or dicThe largest living species is the emperor penguin (Aptenodytes forsteri): on average adults are about 1.1 m (3 ft tionaries under Pingun), and would indicate this bird received its name from its appearance.
7 in) tall and weigh 35 kg (77 lb) or more. The smallest penguin species is the little blue penguin (Eudyptula
minor), also known as the fairy penguin, which stands
around 40 cm (16 in) tall and weighs 1 kg (2.2 lb). Among 2 Systematics and evolution
extant penguins, larger penguins inhabit colder regions,
while smaller penguins are generally found in temper- 2.1 Living species and recent extinctions
ate or even tropical climates (see also Bergmanns rule).
Some prehistoric species attained enormous sizes, becoming as tall or as heavy as an adult human. These
were not restricted to Antarctic regions; on the contrary,
subantarctic regions harboured high diversity, and at least
one giant penguin occurred in a region not quite 2,000
km south of the equator 35 mya, in a climate decidedly
warmer than today.

Etymology

The word penguin rst appears in the 16th century as

a synonym for great auk.[1] When European explorers
discovered what are today known as penguins in the
Southern Hemisphere, they noticed their similar appearance to the great auk of the Northern Hemisphere, and
named them after this bird, although they are not closely
related.[2]

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

The number of extant penguin species is debated.

Depending on which authority is followed, penguin
biodiversity varies between 18 and 20 living species,
The etymology of the word penguin is still debated. The all in the subfamily Spheniscinae. Some sources conEnglish word is not apparently of French,[1] Breton[3] sider the white-ippered penguin a separate Eudyptula
or Spanish[4] origin (the latter two are attributed to the species, while others treat it as a subspecies of the
French word pingouin "auk"), but rst appears in English little penguin;[8][9] the actual situation seems to be more
1

SYSTEMATICS AND EVOLUTION

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

Eudyptula Little penguins

Little blue penguin, Eudyptula minor
White-ippered penguin, Eudyptula albosignata (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

Closeup of southern rockhopper penguin (Eudyptes chrysocome)

Waitaha penguin, Megadyptes waitaha (extinct)

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

penguin,

Eudyptes

Eastern rockhopper penguin, Eudyptes lholi

Northern rockhopper penguin, Eudyptes moseleyi
Royal penguin, Eudyptes schlegeli (disputed)
Macaroni penguin, Eudyptes chrysolophus
Chatham penguin, Eudyptes chathamensis (extinct)
Two king penguins and one gentoo penguin walking on a beach
on South Georgia, British overseas territory

complicated.[10] Similarly, it is still unclear whether the

royal penguin is merely a color morph of the macaroni
penguin. The status of the rockhopper penguins is also
unclear.
[11]

Updated after Marples (1962),

Acosta Hospitaleche
(2004),[12] and Ksepka et al. (2006).[13]

2.2 Fossil genera

Order Sphenisciformes
Basal and unresolved taxa (all fossil)
Waimanu basal (Middle-Late Paleocene)
Perudyptes (Middle Eocene of Atacama
Desert, Peru) basal?

2.2

Fossil genera

Spheniscidae gen. et sp. indet. CADIC P

21 (Leticia Middle Eocene of Punta Torcida,
Argentina)[14]
Delphinornis (Middle/Late Eocene? Early
Oligocene of Seymour Island, Antarctica)
Palaeeudyptinae, basal, new subfamily 1?
Archaeospheniscus (Middle/Late Eocene
Late Oligocene) Palaeeudyptinae? New subfamily 2?
Marambiornis (Late Eocene ?
Early
Oligocene of Seymour Island, Antarctica)
Palaeeudyptinae, basal, new subfamily 1?
Mesetaornis (Late Eocene ?
Early
Oligocene of Seymour Island, Antarctica) Palaeeudyptinae, basal, new subfamily
1?
Tonniornis (Late Eocene ? Early Oligocene
of Seymour Island, Antarctica)
Wimanornis (Late Eocene ? Early Oligocene
of Seymour Island, Antarctica)
Duntroonornis (Late Oligocene of Otago, New
A reconstruction of the ancient penguin Icadyptes
Zealand) possibly Spheniscinae
Korora (Late Oligocene of S Canterbury, New
Zealand)
Kairuku (Late Oligocene of E South Island,
New Zealand)

Subfamily Palaeeudyptinae Giant penguins (fossil)

Marplesornis (Early Pliocene) possibly

Spheniscinae

Crossvallia (Cross Valley Late Paleocene

of Seymour Island, Antarctica) tentatively assigned to this subfamily
Anthropornis (Middle Eocene? Early
Oligocene 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 belong in other subfamilies
Pachydyptes (Late Eocene)
Anthropodyptes (Middle Miocene) tentatively assigned to this subfamily

Nucleornis (Early Pliocene of Duinfontain,

South Africa) possibly Spheniscinae

Subfamily Paraptenodytinae Stout-footed

penguins (fossil)

Tereingaornis (Middle Pliocene of New

Zealand)

Arthrodytes
(San
Julian
Late
Eocene/Early Oligocene Patagonia
Early Miocene of Patagonia, Argentina)
Paraptenodytes
(Early

Late
Miocene/Early Pliocene)

Platydyptes (Late Oligocene of New Zealand)

possibly not monophyletic; Palaeeudyptinae,
Paraptenodytinae or new subfamily?[15]
Spheniscidae gen. et sp. indet. (Late
Oligocene/Early Miocene of Hakataramea,
New Zealand)[16]
Madrynornis (Puerto Madryn Late Miocene
of Argentina) possibly Spheniscinae
Pseudaptenodytes
Pliocene)

(Late

Miocene/Early

Dege (Early Pliocene of South Africa) possibly Spheniscinae

Inguza (Late Pliocene) probably Spheniscinae; formerly Spheniscus predemersus

Palaeoapterodytes (Late
Miocene of Argentina)

Family Spheniscidae

Oligocene/Early

Subfamily Palaeospheniscinae Slenderfooted penguins (fossil)

Eretiscus (Patagonia Early Miocene of
Patagonia, Argentina)

SYSTEMATICS AND EVOLUTION

Palaeospheniscus (Early?
Late that time less than 1,500 kilometres (930 mi) apart rather
Miocene/Early Pliocene) includes than the 4,000 kilometres (2,500 mi) of today. The most
Chubutodyptes
recent common ancestor of penguins and their sister clade
can be roughly dated to the CampanianMaastrichtian
The Early Oligocene genus Cruschedula was formerly boundary, around 7068 mya.[16][20][21] What can be said
thought to belong to Spheniscidae, however reexamina- as certainly as possible in the absence of direct (i.e., fostion of the holotype in 1943 resulted in the genus being sil) evidence is that, by the end of the Cretaceous, the penplaced in Accipitridae.[17] Further examination in 1980 guin lineage must have been evolutionarily well distinct,
though much less so morphologically; it is fairly likely that
resulted in placement as Aves incertae sedis.[18]
they were not yet entirely ightless at that time, as ightless birds have generally low resilience to the breakdown
2.3 Taxonomy
of trophic webs that follows the initial phase of mass extinctions because of their below-average dispersal capaSome recent sources[13][14] apply the phylogenetic taxon bilities (see also Flightless cormorant).
Spheniscidae to what here is referred to as Spheniscinae. Furthermore, they restrict the phylogenetic taxon
Sphenisciformes to ightless taxa, and establish the phy- 2.4.1 The basal fossils
logenetic taxon Pansphenisciformes as equivalent to the
Linnean taxon Sphenisciformes,[14] i.e., including any y- The oldest known fossil penguin species is Waimanu
ing basal proto-penguins to be discovered eventually. manneringi, which lived in the early Paleocene epoch
[20]
While they were
Given that neither the relationships of the penguin sub- of New Zealand, or about 62 mya.
families to each other nor the placement of the pen- not as well-adapted to aquatic life as modern penguins,
guins in the avian phylogeny is presently resolved, this is Waimanu were generally loon-like birds but already
[20]
confusing, so the established Linnean system is followed ightless, with short wings adapted for deep diving.
They swam on the surface using mainly their feet, but
here.
the wings were as opposed to most other diving birds
(both living and extinct) already adapting to underwater
2.4 Evolution
locomotion.[22]
Perudyptes from northern Peru was dated to 42 mya.
An unnamed fossil from Argentina proves that, by the
Bartonian (Middle Eocene), some 3938 mya,[23] primitive penguins had spread to South America and were in
the process of expanding into Atlantic waters.[14]
2.4.2 Palaeeudyptines

Penguin tracks in the sand on Bruny Island, Tasmania

The evolutionary history of penguins is well-researched

and represents a showcase of evolutionary biogeography;
though as penguin bones of any one species vary much in
size and few good specimens are known, the alpha taxonomy of many prehistoric forms still leaves much to be
desired. Some seminal articles about penguin prehistory
have been published since 2005,[13][19][16][20] the evolution of the living genera can be considered resolved by
now.
The basal penguins lived around the time of the
CretaceousPaleogene extinction event somewhere in the
general area of (southern) New Zealand and Byrd Land,
Antarctica.[13] Due to plate tectonics, these areas were at

During the Late Eocene and the Early Oligocene (40

30 mya), some lineages of gigantic penguins existed.
Nordenskjoelds giant penguin was the tallest, growing
nearly 1.80 meters (5.9 feet) tall. The New Zealand giant penguin was probably the heaviest, weighing 80 kg or
more. Both were found on New Zealand, the former also
in the Antarctic farther eastwards.
Traditionally, most extinct species of penguins, giant or
small, had been placed in the paraphyletic subfamily
called Palaeeudyptinae. More recently, with new taxa
being discovered and placed in the phylogeny if possible, it is becoming accepted that there were at least two
major extinct lineages. One or two closely related ones
occurred in Patagonia, and at least one otherwhich is
or includes the paleeudyptines as recognized today occurred on most Antarctic and Subantarctic coasts.
But size plasticity seems to have been great at this initial
stage of penguin radiation: on Seymour Island, Antarctica, for example, around 10 known species of penguins
ranging in size from medium to huge apparently coexisted
some 35 mya during the Priabonian (Late Eocene).[24] It

2.4

Evolution

is not even known whether the gigantic palaeeudyptines

constitute a monophyletic lineage, or whether gigantism was evolved independently in a much restricted
Palaeeudyptinae and the Anthropornithinae whether
they were considered valid, or whether there was a wide
size range present in the Palaeeudyptinae as delimited as
usually done these days (i.e., including Anthropornis nordenskjoeldi).[13] The oldest well-described giant penguin,
the 5-foot (1.5 m)-tall Icadyptes salasi, actually occurred
as far north as northern Peru about 36 mya.
In any case, the gigantic penguins had disappeared by
the end of the Paleogene, around 25 mya. Their decline and disappearance coincided with the spread of the
Squalodontoidea and other primitive, sh-eating toothed
whales, which certainly competed with them for food,
and were ultimately more successful.[16] A new lineage,
the Paraptenodytes, which includes smaller but decidedly
stout-legged forms, had already arisen in southernmost
South America by that time. The early Neogene saw the
emergence 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 penguin
biodiversity of our time.

5
Bartonian,[26] but the range expansion and radiation that
led to the present-day diversity probably did not occur until much later; around the Burdigalian stage of the Early
Miocene, roughly 2015 mya.[16]
The genera Spheniscus and Eudyptula contain species
with a mostly Subantarctic distribution centered on South
America; some, however, range quite far northwards.
They all lack carotenoid coloration, and the former genus
has a conspicuous banded head pattern; they are unique
among living penguins by nesting in burrows. This group
probably radiated eastwards with the Antarctic Circumpolar Current out of the ancestral range of modern penguins throughout the Chattian (Late Oligocene), starting
approximately 28 mya.[16] While the two genera separated during this time, the present-day diversity is the
result of a Pliocene radiation, taking place some 42
mya.[16]

The MegadyptesEudyptes clade occurs at similar

latitudes (though not as far north as the Galapagos
penguin), has its highest diversity in the New Zealand
region, and represents a westward dispersal. They are
characterized by hairy yellow ornamental head feathers;
their bills are at least partly red. These two genera
diverged apparently in the Middle Miocene (Langhian,
roughly 1514 mya), but again, the living species of
2.4.3 Origin and systematics of modern penguins
Eudyptes are the product of a later radiation, stretching
from about the late Tortonian (Late Miocene, 8 mya) to
Modern penguins constitute two undisputed clades and the end of the Pliocene.[16]
another two more basal genera with more ambiguous
The geographical and temporal pattern or spheniscine
relationships.[19] The origin of the Spheniscinae lies probevolution corresponds closely to two episodes of global
ably in the latest Paleogene, and geographically it must
cooling documented in the paleoclimatic record.[16] The
have been much the same as the general area in which
emergence of the Subantarctic lineage at the end of the
the order evolved: the oceans between the Australia-New
Bartonian corresponds with the onset of the slow period
Zealand region and the Antarctic.[16] Presumably divergof cooling that eventually led to the ice ages some 35 miling from other penguins around 40 mya,[16] it seems that
lion years later. With habitat on the Antarctic coasts dethe Spheniscinae were for quite some time limited to
clining, by the Priabonian more hospitable conditions for
their ancestral area, as the well-researched deposits of
most penguins existed in the Subantarctic regions rather
the Antarctic Peninsula and Patagonia have not yielded
than in Antarctica itself.[27] Notably, the cold Antarctic
Paleogene fossils of the subfamily. Also, the earliest
Circumpolar Current also started as a continuous circumspheniscine lineages are those with the most southern dispolar ow only around 30 mya, on the one hand forcing
tribution.
the Antarctic cooling, and on the other facilitating the
The genus Aptenodytes appears to be the basalmost di- eastward expansion of Spheniscus to South America and
vergence among living penguins[13][25] they have bright eventually beyond.[16] Despite this, there is no fossil evyellow-orange neck, breast, and bill patches; incubate by idence to support the idea of a crown radiation from the
placing their eggs on their feet, and when they hatch the Antarctic continent in the Paleogene.[27]
chicks are almost naked. This genus has a distribution
Later, an interspersed period of slight warming was ended
centered on the Antarctic coasts and barely extends to
by the Middle Miocene Climate Transition, a sharp drop
some Subantarctic islands today.
in global average temperature from 1412 mya, and simPygoscelis contains species with a fairly simple black- ilar abrupt cooling events followed at 8 mya and 4 mya;
and-white head pattern; their distribution is intermedi- by the end of the Tortonian, the Antarctic ice sheet was
ate, centered on Antarctic coasts but extending some- already much like today in volume and extent. The emerwhat northwards from there. In external morphology, gence of most of todays Subantarctic penguin species althese apparently still resemble the common ancestor of most certainly was caused by this sequence of Neogene
the Spheniscinae, as Aptenodytes ' autapomorphies are climate shifts.
in most cases fairly pronounced adaptations related to
that genus extreme habitat conditions. As the former
genus, Pygoscelis seems to have diverged during the

6
2.4.4

3 ANATOMY AND PHYSIOLOGY

Relationship to other bird orders

Penguin ancestry beyond Waimanu remains unknown

and not well-resolved by molecular or morphological
analyses. The latter tend to be confounded by the
strong adaptive autapomorphies of the Sphenisciformes;
a sometimes perceived fairly close relationship between
penguins and grebes is almost certainly an error based
on both groups strong diving adaptations, which are
homoplasies. On the other hand, dierent DNA sequence datasets do not agree in detail with each other either.
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 astonishingly

agile. Penguins swimming looks very similar to birds
ight in the air.[32] Within the smooth plumage a layer of
air is preserved, ensuring buoyancy. The air layer also
helps insulate the birds in cold waters. On land, penguins
use their tails and wings to maintain balance for their upright stance.
All penguins are countershaded for camouage that is,
they have black backs and wings with white fronts.[33]
Humboldt penguins in an aquarium. The penguin is an accomA predator looking up from below (such as an orca or a
plished swimmer, having ippers instead of wings.
leopard seal) has diculty distinguishing between a white
What seems clear is that penguins belong to a clade penguin belly and the reective water surface. The dark
of Neoaves (living birds except paleognaths and fowl) plumage on their backs camouages them from above.
that comprises what is sometimes called higher wa- Diving penguins reach 6 to 12 km/h (3.7 to 7.5 mph),
terbirds to distinguish them from the more ancient though there are reports of velocities of 27 km/h (17
waterfowl. This group contains such birds as storks, mph) (which are more realistic in the case of startled
rails, and the seabirds, with the possible exception of the ight). The small penguins do not usually dive deep; they
Charadriiformes.[28]
catch their prey near the surface in dives that normally last
Inside this group, penguin relationships are far less clear. only one or two minutes. Larger penguins can dive deep
Depending on the analysis and dataset, a close relation- in case of need. Dives of the large emperor penguin have
ship to Ciconiiformes[20] or to Procellariiformes[16] has been recorded reaching a depth of 565 m (1,854 ft) for
been suggested. Some think the penguin-like plotopterids up to 22 minutes.
(usually considered relatives of anhingas and cormorants)
may actually be a sister group of the penguins, and that
penguins may have ultimately shared a common ancestor
with the Pelecaniformes and consequently would have to
be included in that order, or that the plotopterids were
not as close to other pelecaniforms as generally assumed,
which would necessitate splitting the traditional Pelecaniformes in three.[29]

Penguins either waddle on their feet or slide on their bellies across the snow, a movement called tobogganing,
which conserves energy while moving quickly. They also
jump with both feet together if they want to move more
quickly 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 in
crowded colonies.[35] Their eyes are adapted for underwaA 2014 analysis of whole genomes of 48 representative ter vision, and are their primary means of locating prey
bird species has concluded that penguins are the sister and avoiding predators; in air it has been suggested that
group of Procellariiformes,[30] from which they diverged they are nearsighted, although research has not supported
about 60 million years ago (95% CI, 56.8-62.7).[31]
this hypothesis.[36]
Penguins have a thick layer of insulating feathers that
keeps them warm in water (heat loss in water is much
3 Anatomy and physiology
greater than in air). The emperor penguin has the largest
body mass of all penguins, which further reduces relative
Penguins are superbly adapted to aquatic life. Their surface area and heat loss. They also are able to control
vestigial wings have become ippers, useless for ight in blood ow to their extremities, reducing the amount of

Gentoo penguin swimming underwater at Nagasaki Penguin

Aquarium.

Isabelline Adlie penguin on Gourdin Island.

4 Distribution and habitat

blood that gets cold, but still keeping the extremities from
freezing. In the extreme cold of the Antarctic winter, the
females are at sea shing for food leaving the males to
brave the weather by themselves. They often huddle together to keep warm and rotate positions to make sure that
each penguin gets a turn in the center of the heat pack.
Calculations of the heat loss and retention ability of marine endotherms <ref name="Calculating just how small
a whale can be[37] suggest that most extant penguins are
too small to survive in such cold environments.[38] In
2007, Thomas and Fordyce wrote about the heterothermic loophole that penguins utilize in order to survive in
Antarctica.[39] All extant penguins, even those that live in
warmer climates, have a counter-current heat exchanger
called the humeral plexus. The ippers of penguins have
at least three branches of the axillary artery, which allows cold blood to be heated by blood that has already
been warmed and limits heat loss from the ippers. This
system allows penguins to eciently use their body heat
and explains why such small animals can survive in the
extreme cold.[40]

See also: List of Sphenisciformes by population

Although all penguin species are native to the Southern
Hemisphere, they are not found only in cold climates,
such as Antarctica. In fact, only a few species of penguin
actually live so far south. At least 10 species live in the
temperate zone; one, the Galpagos penguin, lives as far
north as the Galpagos Islands, but this is only made possible by the cold, rich waters of the Antarctic Humboldt
Current that ows around these islands.[45]
Several authors have suggested that penguins are a good
example of Bergmanns Rule[46][47] where larger bodied
populations live at higher latitudes than smaller bodied
populations. There is some disagreement about this, and
several other authors have noted that there are fossil penguin species that contradict this hypothesis and that ocean
currents and upwellings are likely to have had a greater
eect on species diversity than latitude alone.[48][49]

Major populations of penguins are found in Antarctica,

Argentina, Australia, Chile, New Zealand, and South
[50][51]
They can drink salt water because their supraorbital gland Africa.
lters excess salt from the bloodstream.[41][42][43] The salt
is excreted in a concentrated uid from the nasal passages.

The great auk of the Northern Hemisphere, now extinct, 5 Behaviour

was supercially similar to penguins, and the word penguin was originally used for that bird, centuries ago. They 5.1 Breeding
are only distantly related to the penguins, but are an example of convergent evolution.[44]
Penguins for the most part breed in large colonies, the
exceptions being the yellow-eyed and Fiordland species;
these colonies may range in size from as few as a 100
pairs for gentoo penguins, to several hundred thousand
in the case of king, macaroni and chinstrap penguins.[52]
3.1 Isabelline penguins
Living in colonies results in a high level of social interacof
Perhaps one in 50,000 penguins (of most species) are tion between birds, which has led to a large repertoire[53]
visual
as
well
as
vocal
displays
in
all
penguin
species.
born with brown rather than black plumage. These are
called isabelline penguins. Isabellinism is dierent from Agonistic displays are those intended to confront or drive
appease and avoid conict with, other
albinism. Isabelline penguins tend to live shorter lives o, or alternately
[53]
individuals.
than normal penguins, as they are not well-camouaged
against the deep, and are often passed over as mates.

Penguins form monogamous pairs for a breeding season,

PENGUINS AND HUMANS

Chinstrap penguins in Antarctica.

though the rate the same pair recouples varies drasti- A cook on the Endurance preparing a penguin for consumption
cally. Most penguins lay two eggs in a clutch, although
the two largest species, the emperor and the king penguins, lay only one.[54] With the exception of the emperor
penguin, where the male does it all, all penguins share the
incubation duties.[55] These incubation shifts can last days
and even weeks as one member of the pair feeds at sea.
Penguins generally only lay one brood; the exception is
the little penguin, which can raise two or three broods in
a season.[56]
Penguin eggs are smaller than any other bird species when
compared proportionally to the weight of the parent birds;
at 52 g (2 oz), the little penguin egg is 4.7% of its mothers weight, and the 450 g (1 lb) emperor penguin egg is
2.3%.[54] The relatively thick shell forms between 10 and
16% of the weight of a penguin egg, presumably to minimize the risk of breakage in an adverse nesting environment. The yolk, too, is large, and comprises 2231% of
the egg. Some yolk often remains when a chick is born,
and is thought to help sustain the chick if the parents are
delayed in returning with food.[57]

A penguin encounters a human during Antarctic summer.

approach closer than about 3 meters (9.8 feet) at which

point they become nervous. This is also the distance that
Antarctic tourists are told to keep from penguins (tourists
When mothers lose a chick, they sometimes attempt to are not supposed to approach closer than 3 meters, but are
steal another mothers chick, usually unsuccessfully as not expected to withdraw if the penguins come closer).
other females in the vicinity assist the defending mother In June 2011, a penguin came ashore on New Zealands
in keeping her chick. In some species, such as emperor Peka Peka Beach, 3200 km o course on its journey to
penguins, young penguins assemble in large groups called Antarctica.[59] Nicknamed Happy Feet, after the movie of
crches.
the same name, it was suering from heat exhaustion and
had to undergo a number of operations to remove objects
like driftwood and sand from its stomach.[60] Happy Feet
was a media sensation, with extensive coverage on TV
6 Penguins and humans
and the web, including a live stream that had thousands
of views[61] and a visit from English actor Stephen Fry.[62]
Penguins seem to have no special fear of humans, and
have approached groups of explorers without hesitation. Once he had recovered, Happy Feet was released back
This is probably because penguins have no land predators into the water south of New Zealand.[63]
in Antarctica or the nearby oshore islands. Dogs preyed
upon penguins while they were allowed in Antarctica during the age of early human exploration as sled dogs, but 6.1 In popular culture
dogs are now banned from Antarctica.[58] Instead, adult
penguins are at risk at sea from predators such as sharks, Main article: Cultural depictions of penguins
the orca, and the leopard seal. Typically, penguins do not Penguins are popular around the world, primarily for

9
Several pro, minor, college and high school sport teams
have named themselves after the species, with the
Pittsburgh Penguins team in the National Hockey League
and the Youngstown State Penguins being the most recognizable.
The tendency of penguins to form large groups feeds the
stereotype that they all look exactly alike, a popular notion
exploited by cartoonists such as Gary Larson.
Penguins featured regularly in the cartoons of UK cartoonist Steve Bell in his strip in The Guardian Newspaper, particularly during and following the Falklands War,
and the well-known Opus the Penguin from the cartoons
of Berkeley Breathed, is also described as hailing from
the Falklands. Opus was a comical, existentialist penguin character in the cartoons Bloom County, Outland and
Opus. He was also the star in the Christmas show A Wish
for Wings That Work.

Tux the Linux kernel mascot

their unusually upright, waddling gait and (compared to

other birds) lack of fear of humans. Their striking blackand-white plumage is often likened to a white tie suit.
Mistakenly, some artists and writers have penguins based
at the North Pole. This is incorrect, as there are almost
no wild penguins in the Northern Hemisphere, except the
small group on the northernmost of the Galpagos. The
cartoon series Chilly Willy helped perpetuate this myth, as
the title penguin would interact with northern-hemisphere
species, such as polar bears and walruses.

In the mid-2000s, penguins became one of the most publicized species of animals that form lasting homosexual
couples. A childrens book, And Tango Makes Three, was
written about one such penguin family in the New York
Zoo.

7 References
[1] PINGOUIN : Etymologie de PINGOUIN. Centre National de Ressources Textuelles et Lexicales. Retrieved
January 25, 2010.
[2] Croord, Emily (1989). Gone Forever: The Great Auk.
New York: Crestwood House. p. 10. ISBN 0-89686459-6.

Penguins have been the subject of many books and lms, [3]
such as Happy Feet, Surfs Up and The Penguins of Madagascar, all CGI lms; March of the Penguins, a docu- [4]
mentary based on the migration process of the emperor
penguin; and a parody titled Farce of the Penguins. Mr. [5]
Poppers Penguins is a childrens book written by Richard
and Florence Atwater; it was named a Newbery Honor [6]
Book in 1939. Penguins have also found their way into
a number of cartoons and television dramas; perhaps the [7]
most notable of these is Pingu, created by Silvio Mazzola in 1986 and covering more than 100 short episodes.
At the end of 2009, Entertainment Weekly put it on its [8]
end-of-the-decade, best-of list, saying, Whether they
were walking (March of the Penguins), dancing (Happy [9]
Feet), or hanging ten (Surfs Up), these oddly adorable
birds took ight at the box oce all decade long. [64]
[10]
A video game called Pengo was released by Sega in 1982.
Set in Antarctica, the player controls a penguin character
who must navigate mazes of ice-cubes. The player is rewarded with cut-scenes of animated penguins marching, [11]
dancing, saluting and playing peekaboo. Several remakes
and enhanced editions have followed, most recently in
2012.

Harper, Douglas. penguin. Online Etymology Dictionary.

pingino. Diccionario de la lengua espaola. rae.es
penguin. Wordnik.com. Retrieved October 7, 2014.
Penguin Denition. Merriam-webster.com. August
31, 2012. Retrieved September 8, 2013.
Pen - Denition. Prifysgol Cymru. Retrieved February
18, 2015.
Williams
Davis; Lloyd S. & Renner; M. (1995). Penguins. London:
T & A D Poyser. ISBN 0-7136-6550-5
Banks, Jonathan C.; Mitchell, Anthony D.; Waas, Joseph
R. & Paterson, Adrian M. (2002). An unexpected pattern of molecular divergence within the blue penguin (Eudyptula minor) complex (PDF). Notornis 49 (1): 2938.
Marples, B. J. (1962): Observations on the history of penguins. In: Leeper, G. W. (ed.), The evolution of living
organisms. Melbourne, Melbourne University Press. pp.
408416.

10

[12] (Spanish) Acosta Hospitaleche, Carolina (2004) Los

pinginos (Aves, Sphenisciformes) fsiles de Patagonia.
Sistemtica, biogeografa y evolucin. Doctoral thesis, Department of Natural Sciences and Museum, Universidad
Nacional de La Plata. La Plata, Argentina.
[13] Ksepka, D. T. B., Sara; Giannini, Norberto P; (2006).
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Cladistics 22 (5): 412441.
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[14] Clarke, Julia A.; Olivero, Eduardo B. & Puerta, Pablo
(2003). Description of the earliest fossil penguin
from South America and rst Paleogene vertebrate locality of Tierra Del Fuego, Argentina (PDF). American Museum novitates 3423: 118. doi:10.1206/00030082(2003)423<0001:dotefp>2.0.co;2.
[15] Platydyptes novaezealandiae; holotype. Collections Online. Museum of New Zealand Te Papa Tongarewa. Retrieved July 16, 2010.
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Proceedings of the Royal Society B: Biological Sciences
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[17] Simpson, G.G. (1946). Fossil penguins (PDF). Bulletin
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[18] Olson, S.L. (1985). Faunal Turnover in South American Fossil Avifaunas: The Insuciencies of the Fossil Record (PDF). Evolution 39 (5): 11741177.
doi:10.2307/2408747. JSTOR 2408747.
[19] Bertelli, S.; Giannini, N. P. (2005). A phylogeny of extant penguins (Aves: Sphenisciformes) combining morphology 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 Evolution. Molecular Biology and Evolution 23 (6): 1144
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[21] The exact divergence dates according to Baker et al.
(2006) mentioned in this section are not as precisely resolved as it appears to be due to uncertainties of the
molecular clock used.
[22] Elliott, K. H.; Ricklefs, R. E.; Gaston, A. J.; Hatch, S.
A.; Speakman, J. R.; Davoren, G. K. (2013). High ight
costs, but low dive costs, in auks support the biomechanical hypothesis for ightlessness in penguins. Proceedings 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 Seymour
Island, Antarctica: taxonomy (PDF). Polish Polar Research 27 (1): 362.

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[64] Geier, Thom; Jensen, Je; Jordan, Tina; Lyons, Margaret;

Markovitz, Adam; Nashawaty, Chris; Pastorek, Whitney; Rice, Lynette; Rottenberg, Josh; Schwartz, Missy;
Slezak, Michael; Snierson, Dan; Stack, Tim; Stroup,
Kate; Tucker, Ken; Vary, Adam B.; Vozick-Levinson, Simon; Ward, Kate, The 100 Greatest Movies, TV Shows,
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8 Bibliography
Williams; Tony D. (1995). The Penguins Spheniscidae. Oxford: Oxford University Press. ISBN 019-854667-X.

12

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 Collection
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

EXTERNAL LINKS

13

10
10.1

Text and image sources, contributors, and licenses

Text

Penguin Source: http://en.wikipedia.org/wiki/Penguin?oldid=661948277 Contributors: Derek Ross, Vicki Rosenzweig, Bryan Derksen,
Malcolm Farmer, Magnus~enwiki, DanKeshet, Danny, XJaM, Kchishol1970, TeunSpaans, Cprompt, Norm, Menchi, Tannin, Ixfd64,
Jedimike, Arpingstone, Paul A, Tengai~enwiki, Pcb21, Egil, Ahoerstemeier, Stan Shebs, Jimfbleak, Muriel Gottrop~enwiki, Theresa
knott, JWSchmidt, Kingturtle, , Error, Kils, Nikai, DaMatriX, Kaihsu, GCarty, Rl, Ideyal, Timwi, Nohat, Dysprosia, Jay,
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14

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TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

isdabomb, Crouchingtigershiningdragon, Mehmet Karatay, Mishappy, Hawkeye2010, SieBot, Kkgirl94, Pallab1234, Miettarox, Hansonton,
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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 work
Original artist: Martin St-Amant (S23678)
File:Antarctic_adelie_penguins_(js)_21.jpg Source:
http://upload.wikimedia.org/wikipedia/commons/f/fd/Antarctic_adelie_
penguins_%28js%29_21.jpg License: CC BY 3.0 Contributors: Own work Original artist: Jerzy Strzelecki
File:AntarcticaSummer.jpg Source: http://upload.wikimedia.org/wikipedia/commons/2/2a/AntarcticaSummer.jpg License: CC-BYSA-3.0 Contributors: http://en.wikipedia.org/wiki/Image%3AAntarcticaSummer.jpg Original artist: Roux
File:CSIRO_ScienceImage_11266_Penguin_tracks.jpg Source:
http://upload.wikimedia.org/wikipedia/commons/b/b1/CSIRO_
ScienceImage_11266_Penguin_tracks.jpg License: CC BY 3.0 Contributors: http://www.scienceimage.csiro.au/image/11266 Original
artist: Liese Coulter, CSIRO
File:CharlesGreenwithPenguin.png Source: http://upload.wikimedia.org/wikipedia/commons/c/cb/CharlesGreenwithPenguin.png License: Public domain Contributors: Scanned from The Endurance by Caroline Alexander ISBN 074754123X. Original artist: Frank Hurley
File:Chinstrap_Penguin.jpg Source: http://upload.wikimedia.org/wikipedia/commons/c/cd/Chinstrap_Penguin.jpg License: CC BY 2.0
Contributors: Chinstrap Penguin Original artist: Gilad Rom from Israel
File:Commons-logo.svg Source: http://upload.wikimedia.org/wikipedia/en/4/4a/Commons-logo.svg License: ? Contributors: ? Original
artist: ?
File:Falkland_Islands_Penguins_88.jpg Source: http://upload.wikimedia.org/wikipedia/commons/f/ff/Falkland_Islands_Penguins_
88.jpg License: CC BY 2.0 Contributors: ickr.com Original artist: Ben Tubby
File:Icadyptes_BW.jpg Source: http://upload.wikimedia.org/wikipedia/commons/3/37/Icadyptes_BW.jpg License: CC BY 3.0 Contributors: Own work Original artist: Nobu Tamura (http://spinops.blogspot.com)
File:Orcas_and_penguins.jpg Source: http://upload.wikimedia.org/wikipedia/commons/d/d9/Orcas_and_penguins.jpg License: CC
BY-SA 3.0 Contributors: Transferred from en.wikipedia Original artist: Brocken Inaglory
File:Penguins_on_Gourdin_Island.jpg Source: http://upload.wikimedia.org/wikipedia/commons/0/05/Penguins_on_Gourdin_Island.
jpg License: Public domain Contributors: Transferred from en.wikipedia Original artist: Peyre at en.wikipedia
File:Penguins_walking_-Moltke_Harbour,_South_Georgia,_British_overseas_territory,_UK-8.jpg
Source:
http:
//upload.wikimedia.org/wikipedia/commons/9/9e/Penguins_walking_-Moltke_Harbour%2C_South_Georgia%2C_British_overseas_
territory%2C_UK-8.jpg License: CC BY-SA 2.0 Contributors: Two King Penguins and a Gentoo Penguin walk along the beach Original
artist: Liam Quinn from Canada
File:Pygoscelis_antarctica_trying_to_get_to_iceberg.wmv.OGG Source:
http://upload.wikimedia.org/wikipedia/commons/9/9c/
Pygoscelis_antarctica_trying_to_get_to_iceberg.wmv.OGG License: CC BY-SA 3.0 Contributors: Own work Original artist: Brocken
Inaglory
File:Pygoscelis_papua_-Nagasaki_Penguin_Aquarium_-swimming_underwater-8a.jpg Source:
http://upload.wikimedia.org/
wikipedia/commons/d/db/Pygoscelis_papua_-Nagasaki_Penguin_Aquarium_-swimming_underwater-8a.jpg License: CC BY-SA 2.0
Contributors: originally posted to Flickr as Penguin can y Original artist: Ken FUNAKOSHI
File:Red_Pencil_Icon.png Source: http://upload.wikimedia.org/wikipedia/commons/7/74/Red_Pencil_Icon.png License: CC0 Contributors: Own work Original artist: Peter coxhead
File:Spheniscus_humboldti_-swimming_-aquarium-8a.jpg
Source:
http://upload.wikimedia.org/wikipedia/commons/c/c4/
Spheniscus_humboldti_-swimming_-aquarium-8a.jpg License: CC BY 2.0 Contributors: originally posted to Flickr as swimming?
Flying? penguin Original artist: su neko
File:Tux.svg Source: http://upload.wikimedia.org/wikipedia/commons/3/35/Tux.svg License: Attribution Contributors: [1] Original artist:
Larry Ewing, Simon Budig, Anja Gerwinski
File:Wikispecies-logo.svg Source: http://upload.wikimedia.org/wikipedia/commons/d/df/Wikispecies-logo.svg License: CC BY-SA 3.0
Contributors: Image:Wikispecies-logo.jpg Original artist: (of code) cs:User:-xfi-

10.3

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Content license

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