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Penguins (
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
Sphenisciformes ,
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Spheniscidae ) are a group of aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: only one
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
, the
Galápagos penguin The Galápagos penguin (''Spheniscus mendiculus'') is a penguin endemic to the Galápagos Islands, Ecuador. It is the only penguin found north of the equator. Most inhabit Fernandina Island and the west coast of Isabela Island. The cool wat ...
, is found north of the
Equator The equator is a circle of latitude, about in circumference, that divides Earth into the Northern and Southern hemispheres. It is an imaginary line located at 0 degrees latitude, halfway between the North and South poles. The term can also ...
. Highly adapted for life in the water, penguins have
countershaded Countershading, or Thayer's law, is a method of camouflage in which an animal's coloration is darker on the top or upper side and lighter on the underside of the body. This pattern is found in many species of mammals, reptiles, birds, fish, an ...
dark and white plumage and flippers for swimming. Most penguins feed on krill,
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
, squid and other forms of
sea life Sea Life is a chain of commercial sea life-themed aquarium attractions. there are 53 Sea Life attractions (including standalone Sea Life centres, mini Sea Life features within resort theme parks, and Legoland submarine rides) around the world ...
which they catch with their bills and swallow it whole while swimming. A penguin has a spiny tongue and powerful jaws to grip slippery prey. They spend roughly half of their lives on land and the other half in the sea. The largest living species is the emperor penguin (''Aptenodytes forsteri''): on average, adults are about tall and weigh . The smallest penguin species is the little blue penguin (''Eudyptula minor''), also known as the fairy penguin, which stands around tall and weighs . Today, larger penguins generally inhabit colder regions, and smaller penguins inhabit regions with temperate or tropical climates. Some prehistoric penguin species were enormous: as tall or heavy as an adult human. There was a great diversity of species in subantarctic regions, and at least one giant species in a region around 2,000 km south of the
equator The equator is a circle of latitude, about in circumference, that divides Earth into the Northern and Southern hemispheres. It is an imaginary line located at 0 degrees latitude, halfway between the North and South poles. The term can also ...
35  mya, during the Late
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
, a climate decidedly warmer than today.


Etymology

The word ''penguin'' first appears in literature at the end of the 16th century. When European explorers discovered what are today known as penguins in the Southern Hemisphere, they noticed their similar appearance to the
great auk The great auk (''Pinguinus impennis'') is a species of flightless alcid that became extinct in the mid-19th century. It was the only modern species in the genus ''Pinguinus''. It is not closely related to the birds now known as penguins, wh ...
of the
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the Equator. For other planets in the Solar System, north is defined as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
, and named them after this bird, although they are not closely related. The etymology of the word ''penguin'' is still debated. The English word is not apparently of
French French (french: français(e), link=no) may refer to: * Something of, from, or related to France ** French language, which originated in France, and its various dialects and accents ** French people, a nation and ethnic group identified with Franc ...
, Breton or Spanishpingüino
. Diccionario de la lengua española. rae.es
origin (the latter two are attributed to the French word ), but first appears in English or Dutch. Some dictionaries suggest a derivation from
Welsh Welsh may refer to: Related to Wales * Welsh, referring or related to Wales * Welsh language, a Brittonic Celtic language spoken in Wales * Welsh people People * Welsh (surname) * Sometimes used as a synonym for the ancient Britons (Celtic peopl ...
, "head" and , "white", including the ''
Oxford English Dictionary The ''Oxford English Dictionary'' (''OED'') is the first and foundational historical dictionary of the English language, published by Oxford University Press (OUP). It traces the historical development of the English language, providing a c ...
'', the '' American Heritage Dictionary'', the '' Century Dictionary'' and '' Merriam-Webster'', on the basis that the name was originally applied to the great auk, either because it was found on
White Head Island White Head Island is an island located in the Bay of Fundy. It is off the east coast of Grand Manan Island, New Brunswick. In 2011 the island had a population of 162. White Head Island is governed as a local service district. Its economy is bas ...
( cy, Pen Gwyn) in Newfoundland, or because it had white circles around its eyes (though the head was black). An alternative etymology links the word to
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
, which means "fat" or "oil". Support for this etymology can be found in the alternative Germanic word for penguin, or "fat-goose", and the related Dutch word . 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 The great auk (''Pinguinus impennis'') is a species of flightless alcid that became extinct in the mid-19th century. It was the only modern species in the genus ''Pinguinus''. It is not closely related to the birds now known as penguins, wh ...
(''Pinguinus impennis'', meaning "penguin without flight feathers"), which became
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
in the mid-19th century. As confirmed by a 2004 genetic study, the genus ''Pinguinus'' belongs in the family of the
auk An auk or alcid is a bird of the family Alcidae in the order Charadriiformes. The alcid family includes the murres, guillemots, auklets, puffins, and murrelets. The word "auk" is derived from Icelandic ''álka'', from Old Norse ''alka'' (a ...
s (Alcidae), within the order of the
Charadriiformes Charadriiformes (, from ''Charadrius'', the type genus of family Charadriidae) is a diverse order of small to medium-large birds. It includes about 390 species and has members in all parts of the world. Most charadriiform birds live near water a ...
. 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. They do not belong in the genus ''Pinguinus'', and are not classified in the same family and order as the great auk. They were classified in 1831 by Charles Bonaparte in several distinct genera within the family Spheniscidae and order Sphenisciformes.


Systematics and evolution


Living and recently extinct species

The number of extinct penguin
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
is debated. Depending on which authority is followed, penguin
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...
varies between 17 and 20 living species, all in the
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus Genus ( plural genera ) is a taxonomic rank used in the biological classifica ...
Spheniscinae. Some sources consider the
white-flippered penguin The little penguin (''Eudyptula minor'') is a species of penguin from New Zealand. They are commonly known as little blue penguins or blue penguins owing to their slate-blue plumage and are also known by their Māori name . The Australian ...
a separate '' Eudyptula'' species, while others treat it as a subspecies of the little blue penguin; WilliamsDavis; Lloyd S. & Renner; M. (1995). '' Penguins''. London: T & A D Poyser. the actual situation seems to be more complicated. Similarly, it is still unclear whether the
royal penguin The royal penguin (''Eudyptes schlegeli'') is a species of penguin, which can be found on the sub-Antarctic Macquarie Island and adjacent islands. The International Union for Conservation of Nature (IUCN) classifies the royal penguin as near thr ...
is a separate species or merely a color morph of the macaroni penguin. The status of the
rockhopper penguin The rockhopper penguins are three closely related taxa of crested penguins that have been traditionally treated as a single species and are sometimes split into three species. Not all experts agree on the classification of these penguins. Some ...
s is also unclear. Updated after Marples (1962), Acosta Hospitaleche (2004), and Ksepka ''et al.'' (2006). Subfamily Spheniscinae – modern penguins


Fossil genera

Order Sphenisciformes * Basal and unresolved taxa (all
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
) ** ''
Anthropodyptes ''Anthropodyptes'' is a poorly known monotypic genus of extinct penguin. It contains the single species ''Anthropodyptes gilli'', known from a Middle Miocene humerus from Australia. The bone is somewhat similar to those found in members of ...
'' (Middle Miocene) ** '' Arthrodytes'' (San Julian Late Eocene/Early Oligocene – Patagonia Early Miocene of Patagonia, Argentina) ** '' Aprosdokitos'' Hospitaleche, Reguero & Santillana 2017 ** ''
Crossvallia ''Crossvallia'' is an extinct genus of penguins. It includes two species, ''C. unienwillia'' and ''?C. waiparensis''. Their anatomy suggests that the genus is closely related to the Anthropornithinae. Taxonomy Order Sphenisciformes * Family Sph ...
'' (Cross Valley Late Paleocene of Seymour Island, Antarctica) ** '' Ichthyopteryx'' Wiman 1905 ** ''
Kupoupou ''Kupoupou'' (meaning "diving bird" in Ta Rē Moriori) is an extinct genus of bird from the Paleocene-aged Takatika Grit of the Chatham Islands, New Zealand which is believed to be the oldest penguin genus known to date. The type species, ''K. ...
'' (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 ''Muriwaimanu'' is an extinct genus of early penguin. One species is known, ''Muriwaimanu tuatahi'', which was originally referred to '' Waimanu tuatahi'' in 2006 by Slack ''et al.''.Slack, K.E., Jones, C.M., Ando, T., Harrison G.L., Fordyce R.E ...
'' (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) ** '' Tasidyptes'' Van Tets & O'Connor 1983 nomen dubium (Hunter Island penguins) ** ''
Tereingaornis ''Tereingaornis moisleyi'', also referred to as Moisley's penguin, is a genus and species of extinct penguin from the Middle Pliocene of New Zealand. It was slightly smaller than the extant Fiordland crested penguin. It was described by Ron Sc ...
'' (Middle Pliocene of New Zealand) ** '' Tonniornis'' (Late Eocene –? Early Oligocene of Seymour Island, Antarctica) ** '' Wimanornis'' (Late Eocene –? Early Oligocene of Seymour Island, Antarctica) * Spheniscidae ** ''
Waimanu ''Waimanu'' is a genus of early penguin which lived during the Paleocene, soon after the Cretaceous–Paleogene extinction event, around 62–60 million years ago. It was about the size of an emperor penguin (1 metre). It is one of the most impor ...
'' 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 ''Anthropornis'' is a genus of giant penguin that lived 45-33 million years ago, during the Late Eocene and the earliest part of the Oligocene.Myrcha, A., Jadwiszczak, P., Tambussi, C.P., Noriega, J.I., Gazdzicki, A., Tatur, A., and Valle, R.A. ...
'' Wiman 1905 (Middle Eocene? – Early Oligocene of Seymour Island, Antarctica) ** '' Palaeeudyptes'' Huxley 1859 (Middle/Late Eocene – Late Oligocene) ** ''
Icadyptes ''Icadyptes'' is an extinct genus of giant penguins from the Late Eocene tropics of South America. Etymology The genus name is a combination of "''Ica''" for the Peruvian region where the type species was found and "dyptes" from the Gree ...
'' Clarke et al. 2007 (Late Eocene of Atacama Desert, Peru) ** ''
Pachydyptes ''Pachydyptes'' is an extinct genus of penguin. It contains the single species ''Pachydyptes ponderosus'', the New Zealand giant penguin. This taxon is known from a few bones from Late Eocene (37 to 34 MYA) rocks in the area of Otago Otag ...
'' Oliver 1930 (Late Eocene) ** ''
Inkayacu ''Inkayacu'' is a genus of extinct penguins. It lived in what is now Peru during the Late Eocene, around 36 million years ago. A nearly complete skeleton was discovered in 2008 and includes fossilized feathers, the first known in penguins. A stud ...
'' Clarke et al. 2010 (Late Eocene of South America) ** ''
Kairuku ''Kairuku'' is an extinct genus of penguin. It contains three species, ''K. grebneffi'', ''K. waitaki'' and ''K. waewaeroa''. This taxon is known from bones from 27 MYA (late Oligocene), from the Kokoamu Greensand Formation of New Zealand. I ...
'' 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) ** '' Dege'' Simpson 1979 (Early Pliocene of South Africa) – possibly Spheniscinae ** ''
Marplesornis ''Marplesornis novaezealandiae'', also referred to as Harris's penguin is a genus and species of extinct penguin from New Zealand. It was a relatively large penguin, about the same size as a king penguin. The age of the only known specimen is ...
'' Simpson 1972 (Early Pliocene) ** Subfamily Palaeospheniscinae (slender-footed penguins) (
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
) *** '' 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) *** Spheniscidae gen. et sp. indet. (Late Oligocene/Early Miocene of Hakataramea, New Zealand) *** '' Madrynornis'' (Puerto Madryn Late Miocene of Argentina) The Early Oligocene genus ''
Cruschedula ''Cruschedula'' is an enigmatic bird genus considered to be ''nomen dubium'' which consists of the single species ''Cruschedula revola''. Description The genus was first described by Florentino Ameghino in his 1899 volume ''Sinopsis geológi ...
'' was formerly thought to belong to Spheniscidae; however, re-examination of the holotype in 1943 resulted in the genus being placed in Accipitridae. Further examination in 1980 resulted in placement as Aves ''incertae sedis''.


Taxonomy

Some recent sources apply the phylogenetic taxon Spheniscidae to what here is referred to as Spheniscinae. Furthermore, they restrict the phylogenetic taxon Sphenisciformes to flightless taxa, and establish the phylogenetic taxon Pansphenisciformes as equivalent to the Linnean taxon Sphenisciformes, i.e., including any flying basal "proto-penguins" to be discovered eventually. Given that neither the relationships of the penguin subfamilies to each other nor the placement of the penguins in the avian
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological s ...
is presently resolved, this is confusing, so the established Linnean system is followed here.


Evolution

The
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary history of penguins is well-researched and represents a showcase of evolutionary biogeography. Although 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;Supplementary Material
the evolution of the living genera can be considered resolved by now. The basal penguins lived around the time of the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
somewhere in the general area of (southern) New Zealand and
Byrd Land Marie Byrd Land (MBL) is an unclaimed region of Antarctica. With an area of , it is the largest unclaimed territory on Earth. It was named after the wife of American naval officer Richard E. Byrd, who explored the region in the early 20th centur ...
, Antarctica. Due to
plate tectonics Plate tectonics (from the la, label= Late Latin, tectonicus, from the grc, τεκτονικός, lit=pertaining to building) is the generally accepted scientific theory that considers the Earth's lithosphere to comprise a number of larg ...
, these areas were at that time less than apart rather than the of today. The most recent common ancestor of penguins and their
sister clade In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
can be roughly dated to the Campanian
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the inte ...
boundary, around 70–68 mya. What can be said as certainly as possible in the absence of direct (i.e., fossil) evidence is that, by the end of the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
, the penguin lineage must have been evolutionarily well distinct, though much less so morphologically; it is fairly likely that they were not yet entirely flightless at that time, as flightless birds have generally low resilience to the breakdown of trophic webs that follows the initial phase of mass extinctions because of their below-average dispersal capabilities (''see also'' Flightless cormorant).


Basal fossils

The oldest known
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
penguin species is ''Waimanu manneringi'', which lived in the early
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pala ...
epoch of New Zealand, or about 62 mya. While they were not as well-adapted to aquatic life as modern penguins, ''
Waimanu ''Waimanu'' is a genus of early penguin which lived during the Paleocene, soon after the Cretaceous–Paleogene extinction event, around 62–60 million years ago. It was about the size of an emperor penguin (1 metre). It is one of the most impor ...
'' were generally loon-like birds but already flightless, with short wings adapted for deep diving. They swam on the surface using mainly their feet, but the wings were – as opposed to most other diving birds (both living and extinct) – already adapting to underwater locomotion. '' Perudyptes'' from northern Peru was dated to 42 mya. An unnamed fossil from
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the List of South American countries by area, second-largest ...
proves that, by the Bartonian (Middle Eocene), some 39–38 mya, primitive penguins had spread to
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sou ...
and were in the process of expanding into
Atlantic The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the " Old World" of Africa, Europe ...
waters.


Palaeeudyptines

During the Late Eocene and the Early Oligocene (40–30 mya), some lineages of gigantic penguins existed. Nordenskjoeld's giant penguin was the tallest, growing nearly 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 In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus Genus ( plural genera ) is a taxonomic rank used in the biological classifica ...
called
Palaeeudyptinae Palaeeudyptinae, the giant penguins, is a paraphyletic subfamily of prehistoric penguins. It includes several genera of medium-sized to very large species, such as '' Icadyptes salasi'', '' Palaeeudyptes marplesi'', '' Anthropornis nordenskjoeldi'' ...
. More recently, with new taxa being discovered and placed in the
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological s ...
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 other—which 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). It 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 is usually done these days (i.e., including ''
Anthropornis nordenskjoeldi ''Anthropornis'' is a genus of giant penguin that lived 45-33 million years ago, during the Late Eocene and the earliest part of the Oligocene.Myrcha, A., Jadwiszczak, P., Tambussi, C.P., Noriega, J.I., Gazdzicki, A., Tatur, A., and Valle, R. ...
''). The oldest well-described giant penguin, the -tall ''
Icadyptes salasi ''Icadyptes'' is an extinct genus of giant penguins from the Late Eocene tropics of South America. Etymology The genus name is a combination of "''Ica''" for the Peruvian region where the type species was found and "dyptes" from the Gree ...
'', actually occurred as far north as northern
Peru , image_flag = Flag of Peru.svg , image_coat = Escudo nacional del Perú.svg , other_symbol = Great Seal of the State , other_symbol_type = National seal , national_motto = "Firm and Happy f ...
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, fish-eating
toothed whale The toothed whales (also called odontocetes, systematic name Odontoceti) are a parvorder of cetaceans that includes dolphins, porpoises, and all other whales possessing teeth, such as the beaked whales and sperm whales. Seventy-three species of t ...
s, which certainly competed with them for food, and were ultimately more successful. 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 Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...
of our time.


Origin and systematics of modern penguins

Modern penguins constitute two undisputed clades and another two more basal genera with more ambiguous relationships. To help resolve the evolution of this
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
, 19 high-coverage genomes that, together with two previously published genomes, encompass all extant penguin species have been sequenced. 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. Presumably diverging from other penguins around 40 mya, 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 The genus ''Aptenodytes'' contains two extant species of penguins collectively known as "the great penguins". Etymology The name "Aptenodytes" is a composite of Ancient Greek elements, "ἀ-πτηνο-δύτης" (without-wings-diver). Taxo ...
'' appears to be the basalmost divergence among living penguins. 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. '' Pygoscelis'' contains species with a fairly simple black-and-white head pattern; their distribution is intermediate, centred on Antarctic coasts but extending somewhat northwards from there. In external morphology, these apparently still resemble the common ancestor of the Spheniscinae, as ''Aptenodytes'' autapomorphies are, in most cases, fairly pronounced adaptations related to that genus' extreme
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
conditions. As the former genus, ''Pygoscelis'' seems to have diverged during the Bartonian, but the range expansion and radiation that led to the present-day diversity probably did not occur until much later; around the
Burdigalian The Burdigalian is, in the geologic timescale, an age or stage in the early Miocene. It spans the time between 20.43 ± 0.05 Ma and 15.97 ± 0.05 Ma (million years ago). Preceded by the Aquitanian, the Burdigalian was the first and longest w ...
stage of the Early
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
, roughly 20–15 mya. The genera ''
Spheniscus The banded penguins are penguins that belong to the genus ''Spheniscus''. There are four living species, all with similar banded plumage patterns. They are sometimes also known as "jack-ass penguins" due to their loud locator calls sounding simil ...
'' and '' Eudyptula'' contain species with a mostly Subantarctic distribution centred on
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sou ...
; some, however, range quite far northwards. They all lack carotenoid colouration 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 The Chattian is, in the geologic timescale, the younger of two ages or upper of two stages of the Oligocene Epoch/Series. It spans the time between . The Chattian is preceded by the Rupelian and is followed by the Aquitanian (the lowest stage ...
(Late Oligocene), starting approximately 28 mya. While the two genera separated during this time, the present-day diversity is the result of a
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58 The ''Megadyptes''–''Eudyptes'' clade occurs at similar
latitude In geography, latitude is a coordinate that specifies the north– south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from –90° at the south pole to 90° at the north ...
s (though not as far north as the
Galápagos penguin The Galápagos penguin (''Spheniscus mendiculus'') is a penguin endemic to the Galápagos Islands, Ecuador. It is the only penguin found north of the equator. Most inhabit Fernandina Island and the west coast of Isabela Island. The cool wat ...
), 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 The Langhian is, in the ICS geologic timescale, an age or stage in the middle Miocene Epoch/Series. It spans the time between 15.97 ± 0.05 Ma and 13.65 ± 0.05 Ma (million years ago) during the Middle Miocene.GeoWhen (2007) The Langhian was ...
, roughly 15–14 mya), but again, the living species of ''Eudyptes'' are the product of a later radiation, stretching from about the late Tortonian (Late Miocene, 8 mya) to the end of the Pliocene.


Geography

The geographical and temporal pattern of spheniscine evolution corresponds closely to two episodes of global cooling documented in the paleoclimatic record. The emergence of the Subantarctic lineage at the end of the Bartonian corresponds with the onset of the slow period of cooling that eventually led to the ice ages some 35 million years later. With habitat on the Antarctic coasts declining, by the Priabonian more hospitable conditions for most penguins existed in the Subantarctic regions rather than in Antarctica itself. Notably, the cold Antarctic Circumpolar Current also started as a continuous circumpolar flow only around 30 mya, on the one hand forcing the Antarctic cooling, and on the other facilitating the eastward expansion of ''
Spheniscus The banded penguins are penguins that belong to the genus ''Spheniscus''. There are four living species, all with similar banded plumage patterns. They are sometimes also known as "jack-ass penguins" due to their loud locator calls sounding simil ...
'' to South America and eventually beyond. Despite this, there is no fossil evidence to support the idea of crown radiation from the Antarctic continent in the Paleogene, although DNA study favors such a radiation. Later, an interspersed period of slight warming was ended by the
Middle Miocene Climate Transition The term Middle Miocene disruption, alternatively the Middle Miocene extinction or Middle Miocene extinction peak, refers to a wave of extinctions of terrestrial and aquatic life forms that occurred around the middle of the Miocene, roughly 14 mill ...
, a sharp drop in global average temperature from 14–12 mya, and similar abrupt cooling events followed at 8 mya and 4 mya; by the end of the Tortonian, the Antarctic ice sheet was already much like today in volume and extent. The emergence of most of today's Subantarctic penguin species almost certainly was caused by this sequence of Neogene climate shifts.


Relationship to other bird orders

Penguin ancestry beyond ''
Waimanu ''Waimanu'' is a genus of early penguin which lived during the Paleocene, soon after the Cretaceous–Paleogene extinction event, around 62–60 million years ago. It was about the size of an emperor penguin (1 metre). It is one of the most impor ...
'' 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
grebe Grebes () are aquatic diving birds in the order Podicipediformes . Grebes are widely distributed freshwater birds, with some species also found in marine habitats during migration and winter. Some flightless species exist as well, most notably ...
s is almost certainly an error based on both groups' strong diving adaptations, which are homoplasies. On the other hand, different DNA sequence datasets do not agree in detail with each other either. What seems clear is that penguins belong to a clade of Neoaves (living birds except for paleognaths and fowl) that comprises what is sometimes called " higher waterbirds" to distinguish them from the more ancient
waterfowl Anseriformes is an order of birds also known as waterfowl that comprises about 180 living species of birds in three families: Anhimidae (three species of screamers), Anseranatidae (the magpie goose), and Anatidae, the largest family, which ...
. This group contains such birds as storks, rails, and the
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same envir ...
s, with the possible exception of the
Charadriiformes Charadriiformes (, from ''Charadrius'', the type genus of family Charadriidae) is a diverse order of small to medium-large birds. It includes about 390 species and has members in all parts of the world. Most charadriiform birds live near water a ...
. Inside this group, penguin relationships are far less clear. Depending on the analysis and dataset, a close relationship to Ciconiiformes or to
Procellariiformes Procellariiformes is an order of seabirds that comprises four families: the albatrosses, the petrels and shearwaters, and two families of storm petrels. Formerly called Tubinares and still called tubenoses in English, procellariiforms are oft ...
has been suggested. Some think the penguin-like plotopterids (usually considered relatives of cormorants and
anhinga The anhinga (; ''Anhinga anhinga''), sometimes called snakebird, darter, American darter, or water turkey, is a water bird of the warmer parts of the Americas. The word ''anhinga'' comes from ''a'ñinga'' in the Brazilian Tupi language and means ...
s) may actually be a sister group of the penguins and those 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 into three. A 2014 analysis of whole genomes of 48 representative bird species has concluded that penguins are the sister group of Procellariiformes, from which they diverged about 60 million years ago (95% CI, 56.8-62.7). 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 Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
.


Anatomy and physiology

Penguins are superbly adapted to aquatic life. Their
wing A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is e ...
s 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. 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
tail The tail is the section at the rear end of certain kinds of animals’ bodies; in general, the term refers to a distinct, flexible appendage to the torso. It is the part of the body that corresponds roughly to the sacrum and coccyx in mammal ...
s and wings to maintain balance for their upright stance. All penguins are
countershaded Countershading, or Thayer's law, is a method of camouflage in which an animal's coloration is darker on the top or upper side and lighter on the underside of the body. This pattern is found in many species of mammals, reptiles, birds, fish, an ...
for
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
– that is, they have black backs and
wing A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is e ...
s with white fronts. A predator looking up from below (such as an
orca The orca or killer whale (''Orcinus orca'') is a toothed whale belonging to the oceanic dolphin family, of which it is the largest member. It is the only extant species in the genus '' Orcinus'' and is recognizable by its black-and-white ...
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). 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. 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 hearing for birds; this is used by parents and chicks to locate one another in crowded
colonies In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
. Their eyes are adapted for underwater vision and are their primary means of locating prey and avoiding predators; in air it has been suggested that they are nearsighted, although research has not supported this hypothesis. 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 emperor penguin has a maximum feather density of about nine feathers per square centimeter which is actually much lower than other birds that live in antarctic environments. However, they have been identified as having at least four different ''types'' of feather: in addition to the traditional feather, the emperor has afterfeathers,
plumule A seedling is a young sporophyte developing out of a plant embryo from a seed. Seedling development starts with germination of the seed. A typical young seedling consists of three main parts: the radicle (embryonic root), the hypocotyl (embryo ...
s, and
filoplume Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and a premier e ...
s. The afterfeathers are downy plumes that attach directly to the main feathers and were once believed to account for the bird's ability to conserve heat when under water; the plumules are small down feathers that attach directly to the skin, and are much more dense in penguins than other birds; lastly the filoplumes are small (less than 1 cm long) naked shafts that end in a splay of fibers— filoplumes were believed to give flying birds a sense of where their plumage was and whether or not it needed preening, so their presence in penguins may seem inconsistent, but penguins also preen extensively. 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 suggest that most extant penguins are too small to survive in such cold environments. In 2007, Thomas and Fordyce wrote about the "heterothermic loophole" that penguins utilize in order to survive in Antarctica. 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. They can drink salt water because their
supraorbital gland The supraorbital gland is a type of lateral nasal gland found in some species of marine birds, particularly penguins, which removes sodium chloride from the bloodstream. The gland's function is similar to that of the kidneys, though it is much m ...
filters excess salt from the bloodstream. The salt is excreted in a concentrated fluid from the nasal passages. The
great auk The great auk (''Pinguinus impennis'') is a species of flightless alcid that became extinct in the mid-19th century. It was the only modern species in the genus ''Pinguinus''. It is not closely related to the birds now known as penguins, wh ...
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 Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
.


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
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest cont ...
. In fact, only a few species of penguin actually live so far south. Several species live in the temperate zone; one, the
Galápagos penguin The Galápagos penguin (''Spheniscus mendiculus'') is a penguin endemic to the Galápagos Islands, Ecuador. It is the only penguin found north of the equator. Most inhabit Fernandina Island and the west coast of Isabela Island. The cool wat ...
, lives as far north as the
Galápagos Islands The Galápagos Islands ( Spanish: , , ) are an archipelago of volcanic islands. They are distributed on each side of the equator in the Pacific Ocean, surrounding the centre of the Western Hemisphere, and are part of the Republic of Ecuad ...
, but this is only made possible by the cold, rich waters of the Antarctic Humboldt Current that flows around these islands. Also, though the climate of the Arctic and Antarctic regions is similar, there are no penguins found in the Arctic. Several authors have suggested that penguins are a good example of Bergmann's Rule 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 effect on species diversity than latitude alone. Major populations of penguins are found in
Angola , national_anthem = "Angola Avante"() , image_map = , map_caption = , capital = Luanda , religion = , religion_year = 2020 , religion_ref = , coordinat ...
,
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest cont ...
,
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the List of South American countries by area, second-largest ...
,
Australia Australia, officially the Commonwealth of Australia, is a sovereign country comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands. With an area of , Australia is the largest country by ...
,
Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the eas ...
,
Namibia Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and ea ...
,
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island coun ...
, and
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...
. Satellite images and photos released in 2018 show the population of 2 million in
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
's remote Ile aux Cochons has collapsed, with barely 200,000 remaining, according to a study published in Antarctic Science.


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. Williams, p. 17 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. Williams, p. 57 ''Agonistic'' displays are those intended to confront or drive off, or alternately appease and avoid conflict with, other individuals. 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
king penguin The king penguin (''Aptenodytes patagonicus'') is the second largest species of penguin, smaller, but somewhat similar in appearance to the emperor penguin. There are two subspecies: ''A. p. patagonicus'' and ''A. p. halli''; ''patagonicus'' ...
s, lay only one. Williams, p. 23 With the exception of the emperor penguin, where the male does it all, all penguins share the incubation duties. 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. Penguin eggs are smaller than any other bird species when compared proportionally to the weight of the parent birds; at , the little penguin egg is 4.7% of its mothers' weight, and the emperor penguin egg is 2.3%. 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. 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. Williams, p. 24 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. 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.


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 Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest cont ...
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. Instead, adult penguins are at risk at sea from predators such as
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachi ...
s,
orca The orca or killer whale (''Orcinus orca'') is a toothed whale belonging to the oceanic dolphin family, of which it is the largest member. It is the only extant species in the genus '' Orcinus'' and is recognizable by its black-and-white ...
s, and leopard seals. Typically, penguins do not approach closer than around 9 feet (3 meters), at which point they appear to become nervous. In June 2011, an emperor penguin came ashore on New Zealand's Peka Peka Beach, off course on its journey to Antarctica. 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. Happy Feet was a media sensation, with extensive coverage on TV and the web, including a live stream that had thousands of views and a visit from English actor Stephen Fry. Once he had recovered, Happy Feet was released back into the water south of New Zealand.


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 The North Pole, also known as the Geographic North Pole or Terrestrial North Pole, is the point in the Northern Hemisphere where the Earth's axis of rotation meets its surface. It is called the True North Pole to distinguish from the Ma ...
, but there are no wild penguins in the
Arctic The Arctic ( or ) is a polar regions of Earth, polar region located at the northernmost part of Earth. The Arctic consists of the Arctic Ocean, adjacent seas, and parts of Canada (Yukon, Northwest Territories, Nunavut), Danish Realm (Greenla ...
. 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 '' Happy Feet'', '' Surf's Up'' and '' Penguins of Madagascar'', all CGI films; '' March of the Penguins'', a documentary based on the migration process of the emperor penguin; and ''
Farce of the Penguins ''Farce of the Penguins'' is a 2007 American direct-to-video parody nature documentary written and directed by ''Full House'' star Bob Saget. It is a parody of the French feature-length nature documentary '' March of the Penguins'' (2005), directe ...
'', a parody of the documentary. '' Mr. Popper's Penguins'' is a children's book written by Richard and Florence Atwater; it was named a Newbery Honor Book in 1939. Penguins have also appeared in a number of cartoons and television dramas, including '' Pingu'', created by Silvio Mazzola in 1986 and covering more than 100 short episodes. At the end of 2009, ''
Entertainment Weekly ''Entertainment Weekly'' (sometimes abbreviated as ''EW'') is an American digital-only entertainment magazine based in New York City, published by Dotdash Meredith, that covers film, television, music, Broadway theatre, books, and popular ...
'' put it on its end-of-the-decade "best-of" list, saying, "Whether they were walking (''March of the Penguins''), dancing (''Happy Feet''), or hanging ten (''Surf's Up''), these oddly adorable birds took flight at the box office all decade long." 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.Shuker, R. (2012). ''Understanding popular music culture''. Routledge. In 1941,
DC Comics DC Comics, Inc. ( doing business as DC) is an American comic book publisher and the flagship unit of DC Entertainment, a subsidiary of Warner Bros. Discovery. DC Comics is one of the largest and oldest American comic book companies, with the ...
introduced the avian-themed character of the Penguin as a
supervillain A supervillain or supercriminal is a variant of the villainous stock character that is commonly found in American comic books, usually possessing superhuman abilities. A supervillain is the antithesis of a superhero. Supervillains are of ...
adversary of the
superhero A superhero or superheroine is a stock character that typically possesses ''superpowers'', abilities beyond those of ordinary people, and fits the role of the hero, typically using his or her powers to help the world become a better place, ...
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 penguin The African penguin (''Spheniscus demersus''), also known as Cape penguin or South African penguin, is a species of penguin confined to southern African waters. Like all extant penguins, it is flightless, with a streamlined body and wings stiffe ...
s and
king penguin The king penguin (''Aptenodytes patagonicus'') is the second largest species of penguin, smaller, but somewhat similar in appearance to the emperor penguin. There are two subspecies: ''A. p. patagonicus'' and ''A. p. halli''; ''patagonicus'' ...
s). Several pro, minor, college and high school sport teams in the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country Continental United States, primarily located in North America. It consists of 50 U.S. state, states, a Washington, D.C., ...
have named themselves after the species, including the
Pittsburgh Penguins The Pittsburgh Penguins (colloquially known as the Pens) are a professional ice hockey team based in Pittsburgh. They compete in the National Hockey League (NHL) as a member of the Metropolitan Division of the Eastern Conference, and have playe ...
team in the
National Hockey League The National Hockey League (NHL; french: Ligue nationale de hockey—LNH, ) is a professional ice hockey sports league, league in North America comprising 32 teams—25 in the United States and 7 in Canada. It is considered to be the top ranke ...
and the
Youngstown State Penguins The Youngstown State Penguins are the athletic teams of Youngstown State University of Youngstown, Ohio. The university is a member of the National Collegiate Athletic Association's (NCAA) Division I, and the Penguins compete in football as m ...
in
college athletics College athletics encompasses non-professional, collegiate and university-level competitive sports and games. World University Games The first World University Games were held in 1923. There were originally called the ''Union Nationale des ...
. Penguins featured regularly in the cartoons of U.K. cartoonist Steve Bell in his strip in ''
The Guardian ''The Guardian'' is a British daily newspaper. It was founded in 1821 as ''The Manchester Guardian'', and changed its name in 1959. Along with its sister papers '' The Observer'' and '' The Guardian Weekly'', ''The Guardian'' is part of the ...
'' newspaper, particularly during and following the
Falklands War The Falklands War ( es, link=no, Guerra de las Malvinas) was a ten-week undeclared war between Argentina and the United Kingdom in 1982 over two British dependent territories in the South Atlantic: the Falkland Islands and its territorial ...
. 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 ''Bloom County'' is an American comic strip by Berkeley Breathed which originally ran from December 8, 1980, until August 6, 1989. It examined events in politics and culture through the viewpoint of a fanciful small town in Middle America, whe ...
'', '' Outland'' and '' Opus''. He was also the star in the animated Christmas TV special ''
A Wish for Wings That Work ''A Wish for Wings That Work: An Opus Christmas Story'' is a children's book by Berkeley Breathed that was published in 1991. It was made into an animated television special that same year. The book and special feature characters from Breathed' ...
''. In the mid-2000s, penguins became one of the most publicized species of animals that form lasting homosexual couples. A children's book, ''
And Tango Makes Three ''And Tango Makes Three'' is a children's book written by Peter Parnell and Justin Richardson and illustrated by Henry Cole which was published in 2005. The book tells the story of two male penguins, Roy and Silo, who create a family together. Wi ...
'', was written about one such penguin family in the New York Zoo.


References


Bibliography

*


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 the Encyclopedia of New Zealand
Seaworld Penguin Information

"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
{{Authority control Articles containing video clips Flightless birds Seabirds Danian first appearances Taxa named by Charles Lucien Bonaparte Extant Danian first appearances