The great auk (Pinguinus impennis), also known as the garefowl or penguin, is an extinct species of flightless alcid that first appeared around 400,000 years ago and was driven to extinction by human exploitation in the mid-19th century. It was the only modern species in the genus Pinguinus. It was not closely related to the penguins of the Southern Hemisphere, which were named for their resemblance to this species.
It bred on rocky, remote islands with easy access to the ocean and a plentiful food supply, a rarity in nature that provided only a few breeding sites for the great auks. During the non-breeding season, the auk foraged in the waters of the North Atlantic, ranging as far south as northern Spain and along the coastlines of Canada, Greenland, Iceland, the Faroe Islands, Norway, Ireland, and Great Britain.
The bird was about tall and weighed about , making it the largest alcid to survive into the modern era, and the second-largest member of the alcid family overall (the prehistoric Miomancalla was larger). It had a black back and a white belly. The black beak was heavy and hooked, with grooves on its surface. During summer, great auk plumage showed a white patch over each eye. During winter, the great auk lost these patches, instead developing a white band stretching between the eyes. The wings were only long, rendering the bird flightless. Instead, the great auk was a powerful swimmer, a trait that it used in hunting. Its favourite prey were fish, including Atlantic menhaden and capelin, and crustaceans. Although agile in the water, it was clumsy on land. Great auk pairs mated for life. They nested in extremely dense and social colonies, laying one egg on bare rock. The egg was white with variable brown marbling. Both parents participated in the incubation of the egg for around six weeks before the young hatched. The young left the nest site after two to three weeks, although the parents continued to care for it.
The great auk was an important part of many Native American cultures, both as a food source and as a symbolic item. Many Maritime Archaic people were buried with great auk bones. One burial discovered included a corpse covered by more than 200 great auk beaks, which are presumed to be the remnants of a cloak made of great auks' skins. Early European explorers to the Americas used the great auk as a convenient food source or as fishing bait, reducing its numbers. The bird's down was in high demand in Europe, a factor that largely eliminated the European populations by the mid-16th century. Around the same time, nations such as Great Britain began to realize that the great auk was disappearing, and it became the beneficiary of many early protection laws but despite them was still hunted.
Its growing rarity increased interest from European museums and private collectors in obtaining skins and eggs of the bird. On 3 June 1844, the last two confirmed specimens were killed on Eldey, off the coast of Iceland, ending the last known breeding attempt. Later reports of roaming individuals being seen or caught are unconfirmed. A report of one great auk in 1852 is considered by some to be the last sighting of a member of the species. The great auk is mentioned in several novels, and the scientific journal of the American Ornithological Society (now Ornithology) was named The Auk in honour of the bird until 2021.
Taxonomy and evolution
thumb|left|Fossil [[humerus of the Early Pliocene relative Pinguinus alfrednewtoni]]
Analysis of mtDNA sequences has confirmed morphological and biogeographical studies suggesting that the razorbill is the closest living relative of the great auk. The great auk also was related closely to the little auk or dovekie, which underwent a radically different evolution compared to Pinguinus. Due to its outward similarity to the razorbill (apart from flightlessness and size), the great auk often was placed in the genus Alca, following Linnaeus.
The oldest known fossil records of the modern great auk are from the Boxgrove Palaeolithic site of England and Lower Town Hill Formation of Bermuda, both of which are dated to the Middle Pleistocene at least 400,000 years BP. The Pliocene sister species, Pinguinus alfrednewtoni, and molecular evidence show that the three closely related genera diverged soon after their common ancestor, a bird probably similar to a stout Xantus's murrelet, had spread to the coasts of the Atlantic. Apparently, by that time, the murres, or Atlantic guillemots, already had split from the other Atlantic alcids. Razorbill-like birds were common in the Atlantic during the Pliocene, but the evolution of the little auk is sparsely documented. It is the only recorded British bird made extinct in historic times.
upright|thumbtime=0:00|thumb|alt=A large, stuffed bird with a black back, white belly, heavy bill, and white eye patch.|Turnaround video of Specimen No. 57 and a [[razorbill, Naturalis Biodiversity Center]]
The following cladogram shows the placement of the great auk among its closest relatives, based on a 2004 genetic study:
Pinguinus alfrednewtoni was a larger, and also flightless, member of the genus Pinguinus that lived during the Early Pliocene. Known from bones found in the Yorktown Formation of the Lee Creek Mine in North Carolina, it is believed to have split, along with the great auk, from a common ancestor. Pinguinus alfrednewtoni lived in the Western Atlantic, while the great auk lived in the Eastern Atlantic. After the former died out following the Pliocene, the great auk took over its territory.]]
The great auk was one of the 4,400 animal species formally described by Carl Linnaeus in his eighteenth-century work Systema Naturae, in which it was given the binomial Alca impennis. The name Alca is a Latin derivative of the Scandinavian word for razorbills and their relatives.
The species was not placed in its own scientific genus, Pinguinus, until 1791.
The Irish name for the great auk is , meaning "big seabird/auk". The Basque name is ', meaning "spearbill". Its early French name was apponatz, while modern French uses '. The Norse called the great auk geirfugl, which means "spearbird". This has led to an alternative English common name for the bird, garefowl or gairfowl. The Inuit name for the great auk was isarukitsok, which meant "little wing". It may be derived from the Welsh pen gwyn "white head", although the etymology is debated. When European explorers discovered what today are known as penguins in the Southern Hemisphere, they noticed their similar appearance to the great auk and named them after this bird, although biologically, they are not closely related. Whalers also lumped the northern and southern birds together under the common name "woggins".
Description
upright|left|alt=A large bird with a black back, white belly, and white eye patch stands on a rock by the ocean, as a similar bird with a white stripe instead of an eyepatch swims.|thumb|Summer (standing) and winter (swimming) plumage, by [[John Gerrard Keulemans]]
Standing about tall and weighing approximately as adult birds, the flightless great auk was the second-largest member of both its family and the order Charadriiformes overall, surpassed only by the mancalline Miomancalla. It is, however, the largest species to survive into modern times. The great auks that lived farther north averaged larger in size than the more southerly members of the species. Males and females were similar in plumage, although there is evidence for differences in size, particularly in the bill and femur length. During winter the great auk moulted and lost this eye patch, which was replaced with a wide white band and a grey line of feathers that stretched from the eye to the ear. The bill was large at long and curved downward at the top; The legs were far back on the bird's body, which gave it powerful swimming and diving abilities.
Distribution and habitat
left|alt=A large, triangular rock rises from the misty waters, with more islands behind and northern gannets flying around it.|thumb|[[Stac an Armin, St. Kilda, Scotland, one locality where the great auk used to breed]]
The great auk was found in the cold North Atlantic coastal waters along the coasts of Canada, the Northeastern United States, Norway, Greenland, Iceland, the Faroe Islands, Great Britain, Ireland, France, and the Iberian Peninsula. and another from the Neolithic site of El Harhoura 2 in Morocco.
In the western Atlantic, Great auk bones have been found as far south as Florida, where it may have been present during some periods: approximately 1000 BC, AD 1000, and the 17th century. However, it has also been suggested that some of the bones discovered in Florida may be the result of aboriginal trading. For their nesting colonies the great auks required rocky islands with sloping shorelines that provided access to the sea. These were very limiting requirements and it is believed that the great auk never had more than 20 breeding colonies. The localities of only seven former breeding colonies are known: Papa Westray in the Orkney Islands, St. Kilda off Scotland, Grimsey Island, Eldey Island, Geirfuglasker near Iceland, Funk Island near Newfoundland, and the Bird Rocks (Rochers-aux-Oiseaux) in the Gulf of St. Lawrence. Records suggest that this species may have bred on Cape Cod in Massachusetts. After the chicks fledged, the great auk migrated north and south away from the breeding colonies and they tended to go southward during late autumn and winter.
Ecology and behaviour
left|alt=Two summer great auks, one swimming and facing right while another stands upon a rock looking left, are surrounded by steep, rocky cliffs.|thumb|Great Auks by [[John James Audubon, from The Birds of America (1827–1838)]]
The great auk was never observed and described by modern scientists during its existence and is only known from the accounts of laymen, such as sailors, so its behaviour is not well known and difficult to reconstruct. Much may be inferred from its close, living relative, the razorbill, as well as from remaining soft tissue. When they did run, it was awkwardly and with short steps in a straight line. Polar bears preyed on nesting colonies of the great auk. although after the breeding season, they had been sighted as far as from land. Other fish suggested as potential prey include lumpsuckers, shorthorn sculpins, cod, sand lance, as well as crustaceans. Great Auks began pairing in early and mid-May. They are believed to have mated for life (although some theorize that great auks could have mated outside their pair, a trait seen in the razorbill). It is believed that the variation in the egg streaks enabled the parents to recognize their egg among those in the vast colony.
Relationship with humans
upright|left|alt=A sketch of four bones of the great auk, all long. The first two on the left are shorter and hook and fatten at the end, while the third is straight. The fourth has a nub on both ends.|thumb|Illustration of two [[Humerus|humeri (1) and two tibiae (2), bones of the great auk uncovered by archaeologists in an ancient kitchen midden in Caithness]]
The great auk was a food source for Neanderthals more than 100,000 years ago, as evidenced by well-cleaned bones found by their campfires. Images believed to depict the great auk also were carved into the walls of the El Pendo Cave in Camargo, Spain, and Paglicci, Italy, more than 35,000 years ago, Nearly half of the bird bones found in graves at this site were of the great auk, suggesting that it had great cultural significance for the Maritime Archaic people. as well as from early 5th century Labrador, where the bird seems to have occurred only as stragglers. Early explorers, including Jacques Cartier, and numerous ships attempting to find gold on Baffin Island were not provisioned with food for the journey home, and therefore, used great auks as both a convenient food source and bait for fishing. Reportedly, some of the later vessels anchored next to a colony and ran out planks to the land. The sailors then herded hundreds of great auks onto the ships, where they were slaughtered. Great auk eggs were also a valued food source, as the eggs were three times the size of a murre's and had a large yolk. By the mid-16th century, the nesting colonies along the European side of the Atlantic were nearly all eliminated by humans killing this bird for its down, which was used to make pillows. Three men from St. Kilda caught a single "garefowl", noticing its little wings and the large white spot on its head. They tied it up and kept it alive for three days, until a large storm arose. Believing that the bird was a witch and was causing the storm, they then killed it by beating it with a stick.
upright|thumb|Specimen No. 3 in the [[Royal Belgian Institute of Natural Sciences, one of the two last birds killed on Eldey in 1844]]
The last colony of great auks lived on Geirfuglasker (the "Great Auk Rock") off Iceland. This islet was a volcanic rock surrounded by cliffs that made it inaccessible to humans, but in 1830, the islet submerged after a volcanic eruption, and the birds moved to the nearby island of Eldey, which was accessible from a single side. When the colony was discovered in 1835, nearly fifty birds were present. Museums, desiring the skins of the great auk for preservation and display, quickly began collecting birds from the colony.
Jón Brandsson and Sigurður Ísleifsson, the men who had killed the last birds, were interviewed by great auk specialist John Wolley, and Sigurður described the act as follows: