Penguin
Penguins | |
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Penguin species of different genera; from top-left, clockwise: Emperor penguin (Aptenodytes forsteri), snares penguin (Eudyptes robustus), little penguin (Eudyptula minor), yellow-eyed penguin (Megadyptes antipodes), gentoo penguin (Pygoscelis papua), African penguin (Spheniscus demersus) | |
Scientific classification ![]() | |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Aves |
Clade: | Austrodyptornithes |
Order: | Sphenisciformes Sharpe, 1891 |
Family: | Spheniscidae Bonaparte , 1831
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Modern genera | |
Aptenodytes | |
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Breeding range of penguins, all species (aqua); some species have wider seasonal migration ranges |
Penguins (
They spend roughly half of their lives on land and the other half in the sea. The largest living species is the
Etymology

The word penguin first appears in literature at the end of the 16th century.[8] 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.[9]
The etymology of the word penguin is still debated. The English word is not apparently of French,[8] Breton[10] or Spanish[11] origin (the latter two are attributed to the French word pingouin), but first appears in English or Dutch.[8]
Some dictionaries suggest a derivation from
An alternative etymology links the word to
Adult male penguins are called cocks, females are hens; a group of penguins on land is a waddle, and a group of penguins in the water is a raft.
Pinguinus
Since 1871, the Latin word Pinguinus has been used in scientific classification to name the genus of the great auk (Pinguinus impennis, meaning "penguin without flight feathers"),[16] which became extinct in the mid-19th century. As confirmed by a 2004 genetic study, the genus Pinguinus belongs in the family of the auks (Alcidae), within the order of the Charadriiformes.[17][18]
The birds currently known as penguins were discovered later and were so named by sailors because of their physical resemblance to the great auk. Despite this resemblance, however, they are not auks, and are not closely related to the great auk.
Systematics and evolution
Living and recently extinct species



The number of extinct penguin
Updated after Marples (1962),[22] Acosta Hospitaleche (2004),[23] and Ksepka et al. (2006).[3]
Subfamily Spheniscinae – modern penguins
Image | Genus | Living species |
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Aptenodytes Miller, JF, 1778 – great penguins |
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Pygoscelis Wagler, 1832 – brush-tailed penguins |
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Eudyptula Bonaparte, 1856 – little penguins |
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Spheniscus Brisson 1760 – banded penguins |
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Megadyptes Milne-Edwards, 1880 |
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Eudyptes Vieillot, 1816 – crested penguins |
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Fossil genera
Phylogeny of Spheniscidae[25] | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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Order Sphenisciformes[26]
- Basal and unresolved taxa (all fossil)
- Anthropodyptes (Middle Miocene)
- Arthrodytes (San Julian Late Eocene/Early Oligocene – Patagonia Early Miocene of Patagonia, Argentina)
- Aprosdokitos Hospitaleche, Reguero & Santillana 2017
- Crossvallia (Cross Valley Late Paleocene of Seymour Island, Antarctica)
- Ichthyopteryx Wiman 1905
- Kupoupou (Late Early-Middle Paleocene of Takatika Grit, New Zealand)
- Kaiika Fordyce & Tomas 2011 (Maxwell's penguin)
- Korora (Late Oligocene of S Canterbury, New Zealand)
- Inguza (Late Pliocene)
- Muriwaimanu (Late Paleocene of Canterbury, New Zealand)
- Nucleornis (Early Pliocene of Duinfontain, South Africa)
- Orthopteryx Wiman 1905
- Palaeoapterodytes (Late Oligocene/Early Miocene of Argentina)
- Pseudaptenodytes (Late Miocene/Early Pliocene)
- Sequiwaimanu (Late Paleocene)
- TasidyptesVan Tets & O'Connor 1983 nomen dubium (Hunter Island penguins)
- Tereingaornis (Middle Pliocene of New Zealand)
- Tonniornis (Late Eocene –? Early Oligocene of Seymour Island, Antarctica)
- Wimanornis (Late Eocene –? Early Oligocene of Seymour Island, Antarctica)
- SpheniscidaeA reconstruction of the ancient penguin Icadyptes
- Waimanu Jones, Ando & Fordyce 2006 (Middle-Late Paleocene)
- Kumimanu Mayr, 2017
- Delphinornis Wiman 1905 (Middle/Late Eocene? – Early Oligocene of Seymour Island, Antarctica)
- Marambiornis Myrcha et al. 2002 (Late Eocene –? Early Oligocene of Seymour Island, Antarctica)
- Mesetaornis Myrcha et al. 2002 (Late Eocene –? Early Oligocene of Seymour Island, Antarctica)
- Perudyptes Clarke et al. 2007 (Middle Eocene of Atacama Desert, Peru)
- Anthropornis Wiman 1905 (Middle Eocene? – Early Oligocene of Seymour Island, Antarctica)
- Palaeeudyptes Huxley 1859 (Middle/Late Eocene – Late Oligocene)
- Icadyptes Clarke et al. 2007 (Late Eocene of Atacama Desert, Peru)
- Pachydyptes Oliver 1930 (Late Eocene)
- Inkayacu Clarke et al. 2010 (Late Eocene of South America)
- Kairuku Ksepka et al. 2012 (Late Oligocene of E South Island, New Zealand)
- Paraptenodytes Ameghino 1891 (Early – Late Miocene/Early Pliocene)
- Archaeospheniscus Marples 1952 (Middle/Late Eocene – Late Oligocene)
- Duntroonornis Marples 1953 (Late Oligocene of Otago, New Zealand)
- Platydyptes Marples 1952 (Late Oligocene of New Zealand)[27]
- Dege Simpson 1979 (Early Pliocene of South Africa) – possibly Spheniscinae
- Marplesornis Simpson 1972 (Early Pliocene)
- Subfamily Palaeospheniscinae (slender-footed penguins) (fossil)
- Eretiscus Olson 1986 (Patagonia Early Miocene of Patagonia, Argentina)
- Palaeospheniscus Moreno & Mercerat 1891 (Early? – Late Miocene/Early Pliocene) – includes Chubutodyptes
- Subfamily Spheniscinae
- Spheniscidae gen. et sp. indet. CADIC P 21 (Leticia Middle Eocene of Punta Torcida, Argentina)[28]
- Spheniscidae gen. et sp. indet. (Late Oligocene/Early Miocene of Hakataramea, New Zealand)[29]
- Madrynornis (Puerto Madryn Late Miocene of Argentina)
The
Taxonomy
Some recent sources
Evolution

The
The
Basal fossils
The oldest known
Palaeeudyptines
During the Late Eocene and the Early
Traditionally, most extinct species of penguins, giant or small, had been placed in the
But size plasticity seems to have been great at this initial stage of penguin
In any case, the gigantic penguins had disappeared by the end of the
Origin and systematics of modern penguins
Modern penguins constitute two undisputed clades and another two more basal genera with more ambiguous relationships.[32] To help resolve the evolution of this order, 19 high-coverage genomes that, together with two previously published genomes, encompass all extant penguin species have been sequenced.[38] The origin of the Spheniscinae lies probably in the latest Paleogene and, geographically, it must have been much the same as the general area in which the order evolved: the oceans between the Australia-New Zealand region and the Antarctic.[29] Presumably diverging from other penguins around 40 mya,[29] it seems that the Spheniscinae were for quite some time limited to their ancestral area, as the well-researched deposits of the Antarctic Peninsula and Patagonia have not yielded Paleogene fossils of the subfamily. Also, the earliest spheniscine lineages are those with the most southern distribution.
The genus Aptenodytes appears to be the basalmost divergence among living penguins.[3][39] They have bright yellow-orange neck, breast, and bill patches; incubate by placing their eggs on their feet, and when they hatch the chicks are almost naked. This genus has a distribution centred on the Antarctic coasts and barely extends to some Subantarctic islands today.
The
The Megadyptes–Eudyptes clade occurs at similar
Geography
The geographical and temporal pattern of spheniscine evolution corresponds closely to two episodes of
Later, an interspersed period of slight warming was ended by the
Relationship to other bird orders
Penguin ancestry beyond

What seems clear is that penguins belong to a clade of
Inside this group, penguin relationships are far less clear. Depending on the analysis and dataset, a close relationship to
A 2014 analysis of whole genomes of 48 representative bird species has concluded that penguins are the sister group of Procellariiformes,[44] from which they diverged about 60 million years ago (95% CI, 56.8-62.7).[45]
The distantly related puffins, which live in the North Pacific and North Atlantic, developed similar characteristics to survive in the Arctic and sub-Arctic environments. Like the penguins, puffins have a white chest, black back and short stubby wings providing excellent swimming ability in icy water. But, unlike penguins, puffins can fly, as flightless birds would not survive alongside land-based predators such as polar bears and foxes; there are no such predators in the Antarctic. Their similarities indicate that similar environments, although at great distances, can result in similar evolutionary developments, i.e. convergent evolution.[46]
Anatomy and physiology
Penguins are superbly adapted to aquatic life. Their wings have evolved to become flippers, useless for flight in the air. In the water, however, penguins are astonishingly agile. Penguins' swimming looks very similar to birds' flight in the air.[47] 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 camouflage – that is, they have black backs and wings with white fronts.[48] A predator looking up from below (such as an orca or a leopard seal) has difficulty distinguishing between a white penguin belly and the reflective water surface. The dark plumage on their backs camouflages them from above.
Gentoo penguins are the fastest underwater birds in the world. They are capable of reaching speeds up to 36 km (about 22 miles) per hour while searching for food or escaping from predators. They are also able to dive to depths of 170–200 meters (about 560–660 feet).[49] The small penguins do not usually dive deep; they catch their prey near the surface in dives that normally last only one or two minutes. Larger penguins can dive deep in case of need. Emperor penguins are the world's deepest-diving birds. They can dive to depths of approximately 550 meters (1,800 feet) while searching for food.[50]
Penguins either waddle on their feet or slide on their bellies across the snow while using their feet to propel and steer themselves, 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

Penguins have a thick layer of insulating feathers that keeps them warm in water (heat loss in water is much greater than in air). The
The emperor penguin has the largest body mass of all penguins, which further reduces relative surface area and heat loss. They also are able to control blood flow to their extremities, reducing the amount of blood that gets cold, but still keeping the extremities from freezing. In the extreme cold of the Antarctic winter, the females are at sea fishing 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 centre of the heat pack.
Calculations of the heat loss and retention ability of marine endotherms [55] suggest that most extant penguins are too small to survive in such cold environments.[56] In 2007, Thomas and Fordyce wrote about the "heterothermic loophole" that penguins utilize in order to survive in Antarctica.[57] All extant penguins, even those that live in warmer climates, have a counter-current heat exchanger called the humeral plexus. The flippers 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 flippers. This system allows penguins to efficiently use their body heat and explains why such small animals can survive in the extreme cold.[58]
They can drink salt water because their supraorbital gland filters excess salt from the bloodstream.[59][60][61] The salt is excreted in a concentrated fluid from the nasal passages.
The great auk of the Northern Hemisphere, now extinct, was superficially similar to penguins, and the word penguin was originally used for that bird centuries ago. They are only distantly related to the penguins, but are an example of convergent evolution.[62]
Isabelline penguins
Perhaps one in 50,000 penguins (of most species) are born with brown rather than black plumage. These are called isabelline penguins. Isabellinism is different from albinism. Isabelline penguins tend to live shorter lives than normal penguins, as they are not well-camouflaged against the deep and are often passed over as mates.
Distribution and habitat
Although almost all penguin species are native to the Southern Hemisphere, they are not found only in cold climates, such as
Several authors have suggested that penguins are a good example of
Major populations of penguins are found in Angola, Antarctica, Argentina, Australia, Chile, Namibia, New Zealand, and South Africa.[70][71] Satellite images and photos released in 2018 show the population of 2 million in France's remote Ile aux Cochons has collapsed, with barely 200,000 remaining, according to a study published in Antarctic Science.[72]
Behaviour
Breeding
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 100 pairs for gentoo penguins to several hundred thousand in the case of king, macaroni and chinstrap penguins.[73] Living in colonies results in a high level of social interaction between birds, which has led to a large repertoire of visual as well as vocal displays in all penguin species.[74] Agonistic displays are those intended to confront or drive off, or alternately appease and avoid conflict with, other individuals.[74]
Penguins form monogamous pairs for a breeding season, though the rate the same pair recouples varies drastically. Most penguins lay two eggs in a clutch, although the two largest species, the emperor and the
Penguins generally only lay one brood; the exception is the little penguin, which can raise two or three broods in a season.[77]
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%.[75] The relatively thick shell forms between 10 and 16% of the weight of a penguin egg, presumably to reduce the effects of dehydration and to minimize the risk of breakage in an adverse nesting environment.[78] The yolk, too, is large and comprises 22–31% 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.[79]
When emperor penguin mothers lose a chick, they sometimes attempt to "steal" another mother's chick, usually unsuccessfully as other females in the vicinity assist the defending mother in keeping her chick.[80] In some species, such as emperor and king penguins, the chicks assemble in large groups called crèches.
Conservation status
The majority of living penguin species have declining populations. According to the IUCN Red List, their conservation statuses range from Least Concern through to Endangered.
Species | IUCN Red List Status | Trend | Mature Individuals | Last assessment |
---|---|---|---|---|
Emperor penguin, Aptenodytes forsteri | Near Threatened | Unknown | 2018[81] | |
King penguin, Aptenodytes patagonicus | Least Concern | Increasing | 2018[82] | |
Little penguin, Eudyptula minor | Least Concern | Stable | 469,760 | 2018[83] |
Southern rockhopper penguin, Eudyptes chrysocome | Vulnerable | Decreasing | 2,500,000 | 2018[84] |
Macaroni penguin, Eudyptes chrysolophus | Vulnerable | Decreasing | 2018[85] | |
Northern rockhopper penguin, Eudyptes moseleyi | Endangered | Decreasing | 480,600 | 2018[86] |
Fiordland penguin, Eudyptes pachyrynchus | Vulnerable | Decreasing | 2,500-9,999 | 2018[87] |
Snares penguin, Eudyptes robustus | Vulnerable | Stable | 63,000 | 2018[88] |
Royal penguin, Eudyptes schlegeli (disputed) | Near Threatened | Stable | 1,700,000 | 2018[89] |
Erect-crested penguin, Eudyptes sclateri | Endangered | Decreasing | 150,000 | 2016[90] |
Yellow-eyed penguin, Megadyptes antipodes | Endangered | Decreasing | 2,528-3,480 | 2018[91] |
Adélie penguin, Pygoscelis adeliae | Least Concern | Increasing | 7,580,000 | 2018[92] |
Chinstrap penguin, Pygoscelis antarctica | Least Concern | Decreasing | 8,000,000 | 2018[93] |
Gentoo penguin, Pygoscelis papua | Least Concern | Stable | 774,000 | 2018[94] |
African penguin, Spheniscus demersus | Endangered | Decreasing | 50,000 | 2018[95] |
Humboldt penguin, Spheniscus humboldti | Vulnerable | Decreasing | 32,000 | 2018[96] |
Magellanic penguin, Spheniscus magellanicus | Near Threatened | Decreasing | 2018[97] | |
Galápagos penguin , Spheniscus mendiculus
|
Endangered | Decreasing | 1,200 | 2018[98] |
Penguins and humans


Penguins have no special fear of humans and will often approach groups of people. This is probably because penguins have no land predators in Antarctica or the nearby offshore islands. They are preyed upon by other birds like skuas, especially in eggs and as fledglings. Other birds like petrels, sheathbills, and gulls also eat the chicks. Dogs preyed upon penguins while they were allowed in Antarctica during the age of early human exploration as sled dogs, but dogs have long since been banned from Antarctica.[99] Instead, adult penguins are at risk at sea from predators such as sharks, orcas, and leopard seals. Typically, penguins do not approach closer than around 9 feet (2.7 meters), at which point they appear to become nervous.[100]
In June 2011, an emperor penguin came ashore on New Zealand's Peka Peka Beach, 3,200 kilometres (2,000 mi) off course on its journey to Antarctica.[101] Nicknamed Happy Feet, after the film of the same name, it was suffering from heat exhaustion and had to undergo a number of operations to remove objects like driftwood and sand from its stomach.[102] Happy Feet was a media sensation, with extensive coverage on TV and the web, including a live stream that had thousands of views[103] and a visit from English actor Stephen Fry.[104] Once he had recovered, Happy Feet was released back into the water south of New Zealand.[105]
In popular culture

Penguins are widely considered endearing for their unusually upright, waddling gait, swimming ability and (compared to other birds) lack of fear of humans. Their black-and-white plumage is often likened to a white tie suit. Some writers and artists have penguins based at the North Pole, but there are no wild penguins in the Arctic. The cartoon series Chilly Willy helped perpetuate this myth, as the title penguin would interact with Arctic or sub-Arctic species, such as polar bears and walruses.
Penguins have been the subject of many books and films, such as
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, dancing, saluting and playing peekaboo. Several remakes and enhanced editions have followed, most recently in 2012. Penguins are also sometimes depicted in music.[107]
In 1941, DC Comics introduced the avian-themed character of the Penguin as a supervillain adversary of the superhero Batman (Detective Comics #58). He became one of the most enduring enemies in Batman's rogues gallery. In the 60s Batman TV series, as played by Burgess Meredith, he was one of the most popular characters, and in Tim Burton's reimagining of the character in the 1992 film Batman Returns, he employed an actual army of penguins (mostly African penguins and king penguins).
Several pro, minor, college and high school sport teams in the United States have named themselves after the species, including the Pittsburgh Penguins team in the National Hockey League and the Youngstown State Penguins in college athletics.
Penguins featured regularly in the cartoons of U.K. cartoonist Steve Bell in his strip in The Guardian newspaper, particularly during and following the Falklands War. 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 animated Christmas TV special A Wish for Wings That Work.
In the mid-2000s, penguins became one of the most publicized species of animals that form lasting
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{{cite web}}
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Bibliography
- Williams; Tony D. (1995). The Penguins – Spheniscidae. Oxford: ISBN 978-0-19-854667-2.
External links
Library resources about Penguin |
- Two new fossil penguin species found in Peru. news.nationalgeographic.com
- Information about penguins at pinguins.info
- Integrated Taxonomic Information System (archived 17 February 2006)
- Penguin information on 70South (archived 15 March 2006)
- Penguin research projects on the web
- Penguin videos and photos on the Internet Bird Collection (archived 27 December 2015)
- Penguin World
- Penguins in Encyclopedia of New Zealand(archived 5 September 2008)
- Seaworld Penguin Information (archived 17 October 2013)
- "Lessons in a Land of Wind and Ice" from National Wildlife Magazine 1/15/2010
- Curious Penguins
- Live 24/7 camera inside a penguin habitat