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Penguins ( order Sphenisciformes , family Spheniscidae ) are a group of aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: only one species, the Galápagos penguin, is found north of the Equator. Highly adapted for life in the water, penguins have countershaded dark and white plumage and flippers for swimming. Most penguins feed on krill, fish, squid and other forms of sea life 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 35  mya, during the Late Eocene, 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 of the Northern Hemisphere, 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, 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 , "head" and , "white", including the ''Oxford English Dictionary'', 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 ( 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 , 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 (''Pinguinus impennis'', meaning "penguin without flight feathers"), 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.

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 is debated. Depending on which authority is followed, penguin biodiversity varies between 17 and 20 living species, all in the subfamily Spheniscinae. Some sources consider the white-flippered penguin 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 is a separate species or merely a color morph of the macaroni penguin. The status of the rockhopper penguins 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)
** '' 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)
** '' Tasidyptes'' Van 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)
* Spheniscidae
** ''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)
** '' 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)
*** 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'' 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 is presently resolved, this is confusing, so the established Linnean system is followed here.

Evolution

The evolutionary 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 somewhere in the general area of (southern) New Zealand and Byrd Land, Antarctica. Due to plate tectonics, 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 can be roughly dated to the Campanian–Maastrichtian 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 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 penguin species is ''Waimanu manneringi'', which lived in the early Paleocene epoch of New Zealand, or about 62 mya. While they were not as well-adapted to aquatic life as modern penguins, ''Waimanu'' 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 proves that, by the Bartonian (Middle Eocene), some 39–38 mya, primitive penguins had spread to South America and were in the process of expanding into Atlantic 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 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 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''). The oldest well-described giant penguin, the -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, fish-eating toothed whales, 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 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, 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'' 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 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 stage of the Early Miocene, roughly 20–15 mya.

The genera ''Spheniscus'' and ''Eudyptula'' contain species with a mostly Subantarctic distribution centred on South America; 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 (Late Oligocene), starting approximately 28 mya. While the two genera separated during this time, the present-day diversity is the result of a Pliocene

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