A prehistoric form of great auk, Pinguinus alfrednewwni, is recorded from the Pliocene (5.2 to 1.64 million years ago) and other Pliocene flightless diving birds had wings even more penguin-like than those of the great auks. The oldest great auk remains, 75,000 years old, were found at Gibraltar. Great auks are depicted in late Stone Age cave paintings, 18,5000 years old, near Marseilles. Their remains have been found in Norwegian kitchen middens from 13,000 to 2,000 years ago. The great auk is linked to the discovery of North America by Icelandic Vikings who, noticing the seasonal appearance and departure of the birds, may have followed the huge rafts of migrating great auks and discovered Newfoundland to the west. Its closest living relative is the razorbill, which can fly and is a fifth of the size of the great auk. The great auk's similarity to southern hemisphere penguins is due to parallel evolution - they are quite unrelated.

The Icelanders called it "garefowl" (Anglicised spelling of geirfugl) meaning "spear-billed bird" while early navigators called it "penguin" either referring to its white head ("pen gwyn" is Welsh for white head) or "pen-winged" (referring to its pinioned wings) or simply because it was fat ("pinguis" in Latin). Auk comes from the Swedish "alka" and was used after the late 17th century. In French it is still called "grand pingouin" and in Spanish it is "pinguino grande" (both mean "big penguin").

It was the only flightless seabird in the northern hemisphere and its ungainliness on land resulted in early descriptions such as "it 'walked very awkwardly often tumbling over" and "stupid and tame". Captain Taverner, an English sailor, wrote of the great auk in the early 18th century: "the Penguin Islands [off the south-west coast of Newfoundland] are in the summer time covered with fowle of that name. They are as large as any Tame Goose; their wings are soe small that they can never fly, they get their food by Diving in the Sea".

In the water they were strong swimmers; according to one account: "Some of these birds were as large as geese, being black and white with a beak like a crow's. They are always on the water, not being able to fly in the air, inasmuch as they have only small wings about the size of half one's hand, with which however they move as quickly along the water as the other birds fly through the air."

The great auk (Alca impennis) was a white-breasted, black-backed, wedge-beaked diving bird. Though streamlined and excellent swimmers, they were flightless and ungainly on land. It was the strongest and swiftest of the northern swimming and diving birds, almost immune from marine predators and nested on uninhabited and almost inaccessible rocky sites in the rough North Atlantic. It was extremely populous, numbering tens of millions of birds. It was the original penguin; southern hemisphere penguins got their name due to their similarity to European auks and puffins.

It lived most of its life at sea and ranged across the north Atlantic. Apart from the short summer breeding season, it probably roosted at sea. It only came ashore to breed on flat, low-lying islands on which it was easy to beach, and where there was rich inshore feeding. Like modern rockhopper penguins, it might also have been able to scramble up cliffs in search of suitable nesting sites such as those at Bird Rocks (Magdalen Islands), Eldey and parts of St Kilda. Its natural enemies were Arctic wolves, Arctic foxes, polar bears and man, hence it chose small, uninhabited islands, away from the mainland.

Like penguins, great auks stood almost upright on land. In summer their plumage was blackish-brown on all the upper surfaces, with narrow white tips on the secondary wing plumage and large white spots on the face between the bill and the eye. Their throats and chins were dark, but their chests and bellies were white. The bill and feet were black, the webbed toes ending in short claws. The bill had a series of between 6 and 12 vertical grooves on both the upper and lower mandible towards the tip. Its eyes were dark, chestnut brown and the inside of its mouth appears to have been orange-yellow. In the winter, the white patches in front of the eyes reduced to a narrow white line, and the dark throat feathers were replaced by white plumage. It weighed 4 - 5 kg, with males and females differing slightly in size. It used its short (17 cm) but muscular wings for propulsion through the water and used its webbed feet as a rudder.

Being large and heavy makes a bird a more efficient diver. If necessary, it could dive down to 75 - 150 metres, holding its breath for up to 15 minutes, though most dives would have been shallow. Large size also means a slower metabolic rate that uses up air more slowly and allows a bird to stay under water for longer. The great auk's thick layer of subcutaneous fat provided insulation. Its body and short muscular wings were perfectly adapted to "flying through water". Every single feather had its own special muscle, enabling the great auk to flatten its feathers close to its body, trapping as little air as possible. This further reduced buoyancy. Underwater it would have been extremely fast and agile. Having a great gape, great auks could swallow fish whole; more energy efficient than returning to land to peck at their food. Off the coasts of Greenland they ate shorthorn sculpins and lumpsuckers. On Funk Island their diet included Atlantic menhaden up to 190 mm long, shad, capelin, stickleback and striped bass. They also ate crustaceans near the colonies. Outside of the breeding season, they were sighted as far as 550 km from land.

Its eyes may have been adapted to seeing better underwater than on land - it spent 90% of its time underwater. Large flat eyes would have maximised light intake in the murky depths and strong eye muscles might have allowed the lens of the eye to contract, allowing it to see well on both land and underwater. These latter adaptations are found in penguins, along with increased sensitivity to blue, green and near-UV light, and may well have existed in the great auk.

The great auk had a large, narrow bill that served as a formidable dagger. Life in densely populated breeding colonies tends to result in aggression. Around 1808, Fabricius wrote that a great auk, if caught, would "bite wildly all around" while Audobon in 1840 wrote that it would "bite everyone within reach of its powerful bill". Although Audobon's name is now associated with bird conservation, his own idea of preserving birds appeared to be preservation through taxidermy. Great auk "nests" may have been within one square meter of each other, not unlike the densely packed and aggressive blue-footed booby colonies in modern day Galapagos.

In 1718 a surveyor of Penguin Island off Newfoundland reported that French colonists of Placentia told him that "a Mann could not goe ashoar upon those Islands without Bootes, for otherwise they would spoil his Leggs, that they were Intirely covered with those fowles, soe close that a mann could not put his foot between them". An account from Iceland in 1824, 20 years before the last killing of great auks by humans, tells how a young man was pecked so severely by a cornered great auk that in spite of his calfskin jacket, the bird drew blood seriously.

The great auk's breeding habits were probably similar to those of its living relatives: an elaborate courtship ritual, mating on land and the parents taking turns to incubate the single egg. The couple, which possibly mated for life, might have reinforced their bond by mutual preening and beak-tapping. Although they were not described as very vocal, making only low groans and croaks, they probably called softly to one another. The egg was laid on bare rock and was elegantly patterned, wide at one end and narrow at the other so that it rolled in circles rather than rolling away. The adults probably stood upright to brood the egg and changing places would have been a painstaking affair. Incubation is estimated at 6 weeks.

The estimated weight of a newly hatched chick (based on the known weight of the egg) was 240 grammes. If the egg or chick was lost, it is likely that the breeding pair would try again if there was enough time remaining in the breeding season. The only description of young great auks comes from an 1808 account of a raid on the auk colony at the Geirfuglasker islets 40 km south-west of Cape Rejkjanes, south-west Iceland. British sailors spent several days on the island, clubbing the birds to death for food and for sport. Reports described downy grey chicks at sea, and chicks riding on their parents' backs. This means the chicks took to the water within a few days of hatching. This is based on the absence of chicks' remains and the lack of reports of chicks at great auk colonies. The diet of the chicks is unknown - they were too small to eat whole fish and there is no evidence that great auks could regurgitate food. They probably took a year to mature.

Since the birds only produced one egg each year and were vulnerable while brooding, the killing and the widescale theft of eggs from a breeding colony would have had a disastrous effect. Large colonies mean many beaks to defend against predators. Such birds would suffer from "under-crowding" if the numbers fell too low and might fail to breed at all. Isolated from a colony, the last pair of auks on Eldey Island in 1844 were too stressed to fight back. Without the backing of hundreds of other beaks, it made sense for them to flee rather than fight.

From the 10th century, great auk colonies in the Outer Hebrides and on islets off of Iceland were occasionally raided by sailors and fishermen as an alternative food source, though this small scale, local depredation had little effect on the great auk population. The records of great auks being hunted off the coast of Newfoundland are from 1497 when French fishing ships sailed after cod in the region. They took the "pingouins" and their eggs in such huge numbers, that they considered it unnecessary to stock their ships with food for the duration of their stay off the Grand Banks. In 1534, Jacques Cartier visited an "Island of Birds" off Newfoundland; his crew filled two boats with great auks in less than half an hour, and every ship salted down six barrelfuls of the birds. A Captain Mood recorded taking 100,000 eggs in a single day. The egg in the photos is preserved at Ipswich Museum, Ipswich, Suffolk, UK.

Though the great auk was extensively killed for food and its eggs take, it was not until the birds came to the attention of the feather industry that they were headed for extinction. Around 1760, the supply of eider-down and feathers for feather beds was exhausted due to excessive hunting of breeding eider ducks and the destruction of their nesting grounds along the east coast of North America. The feather merchants sent out crews to the great auk nesting grounds and the birds were killed on an industrial scale. By 1810, Funk Island was the only west Atlantic 'rookery' left. The feather company crews returned each spring until they had killed every bird.

Funk Island first appeared on a chart in 1501 when cartographer, Pedro Renel called it the Y dos Ayes (Island of Birds) because of its large colonies of guillemots, razorbills and great auks. It was an inhospitable, barren rocky isle 0.5 miles long and 0.25 miles wide. There was no soil, and no plants, but the whole island was covered in millions of tons of dung, many feet thick, from millennia of nesting birds. During the June breeding season, as many as 100,000 pairs would have gathered there. Before Renel's chart, Vikings may have reached Funk Island and there is evidence that local aboriginal human populations hunted the great auk. As long ago as AD 500 Beothuk Indians canoed to Funk Island to collect its eggs; being 3 times the size of murre's eggs this was well worth the effort and danger. Though impossible to catch an auk in the water, even in a six-oared boat, on land it was easy to pick up if the hunter avoided its strong beak.

The island also acquired the name Isla de Pitigoen (Penguin Island) and the auks were so numerous that a number of men could slaughter enough great auks to fill a boat within half an hour. There are reports from the early 16th century of 4 or 5 tons of great auks being salted in barrels. In Tudor times, Richard Hakluyt (a collector and editor of voyagers' tales) reported Funk Island to be "so exceedingly full of birds that one would think they had been stowed there... in less than halfe an houre we filled two boats full of them, as if they had been stones, so that besides them that we did eat fresh, every ship did powder and salt five or sixe barrels full of them"

Funk Island became an important part of the cod fisheries and by 1536, Newfoundland's fisheries were so well known in Europe that a group of London merchants chartered and equipped two ships to take "thirty gentlemen to view its wonders and visit the island of the penguins." Those men also enjoyed "a great feast of great auks". In 1629, Captain Richard Whitbourne wrote in his "Discourse and Discovery of Newfoundland: "These Penguins are as big as geese... and they multiply so infinitely upon a certain flat island that men drive them from hence upon a board, into their boat by the hundred at that time, as if God had made the innocencie of so poor a creature to become such an admirable instrument for the sustenation of man". It is debatable whether great auks really could have been herded up a gangplank onto a boat as Whitbourne described.

The times and places the great auk came ashore were regular and predictable and sailors took advantage of this to stock up on fresh meat. In 1728 the mariner John Seller wrote "you may know when you are upon the Bank by the great quantity of fowls ... none are to be minded so much as the Pengwins [sic], for these never go without [beyond] the Bank, as the others do". This predictability led to its downfall when the feather industry hunted it. Around 1800, the great auk was found there in huge numbers; the island got its modern name from the stench ("funk") of the great auk industry.

By the 18th century, the Funk Island auks were slaughtered wholesale for their beautiful back feathers which were "soft and smooth as black silk". These were indispensable fashion accessories for trimming hats and dresses in Europe. Their oil was a valued fuel and according to the French fur trader Nicolas Denys, "The great auk (pennegoin) is another bird, variegated in white and black. It does not fly. it has only two stumps of wings with which it beats upon the water to aid in flying or diving. It is claimed that if it dives even to the bottom to seek its prey upon the Bank. it is found more than one hundred leagues from land, where, nevertheless, it comes to lay its eggs like the others. When they have had their young, they plunge into the water, and the young place themselves upon their backs and are carried like this as far as the Bank. There one sees some no larger than chickens, although they grow as large as geese. All those birds are [considered] good to eat by fishermen. As for myself I do not find them agreeable. They make fish oil. The fishermen collect them for this purpose. There are vessels which have made as much as ten or twelve puncheons of it [over 5000 litres]."

During the predictable nesting season, tens of thousands of birds were slaughtered at one end of the island, their eggs smashed or taken by collectors. At the other end of the island, hundreds of thousands more were unwilling to abandon their nests and remaining eggs. Their devotion to their single precious egg made them easy prey for the feather trade. Huge numbers of great auks were easily herded into stone enclosures, packed so tightly they could barely move. Men worked their way through the enclosures, swinging spiked clubs. Dead and stunned birds were thrown out of the enclosure and sued either as fuel for the fires or thrown into the boiling cauldrons on the fires. In the treeless land, the thick insulating layer of fat that protected the birds from arctic waters also served as fuel. In the cauldrons, the feathers were separated from the auks' bodies and scooped into sacks. The denuded, boiled bodies were raked from the cauldron and thrown down the cliff to lie rotting on the rocky seashore. The feather industry - and the fashion for feather trim - had not learned their lessons from the over-hunting of eider ducks.

Many witnesses described this slaughter and most have been recorded in Syminton Grieve's 19th century book "The Great Auk, or Garefowl, its history, archaeology and remains". In 1819, Rev L A Anspach suggested that the usefulness of great auks as aids to navigation was in danger of being lost to overexploitation. By the beginning of the 19th century, the great auk was extinct on Funk Island and Canada and only the ashpits remained as a reminder of the huge colonies that once nested there.

In 1785, an English adventurer and trapper George Cartwright, who had been living in Labrador since 1770, had written "A boat came in from Funk Island laden with birds, chiefly penguins ... it has been customary of late years for crews of men to live all summer on... that island, for the sole purpose of killing birds for the sake of their feathers... the destruction which they have made is incredible ... if a stop is not put to that practice the whole breed will be diminished to almost nothing, particularly the penguins: for this is now the only island they have left to breed upon."

Cartwright was describing islands in the vicinity of St Lawrence and the authorities paid heed to what he said. Ten earlier, Newfoundland's governing council had petitioned the British parliament to stop the wholesale slaughter of seabirds. Magistrates at St John's had introduced harsh penalties, including flogging, for those caught killing birds for their feathers or eggs though it was still permissible to take birds for fish bait. It was hard to police and regulate these rule and easy to make false claims about what use birds were being put to. In 1793, Chief Justice Reeves convicted three men from Greenspond for taking great auk eggs from Funk Island during the closed season. In 1786, a proclamation was finally issued banning the slaughter of great auks on Funk Island. It was already too late; the massive industrialised culling had already driven the great auk to the edge of extinction on the island.

In Europe, great auks continued to breed off the Iceland coast. A substantial auk settlement still existed on the Geirfuglasker ("Auk Rocks"), sea-lashed islets off the coast of Iceland. Ships trying to reach Geirfuglasker were liable to be wrecked on the dangerous rocks, but the determination of egg and specimen collectors meant even those dangerous islets were plundered, albeit not on the organised industrial scale of Funk Island. When the colony seemed in danger of extinction, it was left alone long enough for the population to recover sufficiently to make egg-collecting worthwhile again, though this was due more to dangerous sea conditions than to concerns over the great auk population. It was generally believed that nature would replenish herself and that there would always be great auk colonies elsewhere, so there were no restrictions on hunting or collecting eggs.

As the great auk died out in Newfoundland, its European population also declined. Its range was an arc across the North Atlantic from Canada to Greenland and Iceland, and across to Britain. Its bones of have also been found in Florida (bones were 3250 years old), New England, Labrador and coastal areas of Norway, Denmark, the Netherlands, Brittany, the Bristol Channel and even Gibraltar. These appear to date from a time when the climate in those latitudes was much colder than today.

In March 1830, the eruption of an underwater volcano caused a huge seaquake, obliterating some islets, including Geirfuglasker, and changing Iceland's coastline. Although damage was limited by the fact that it was not breeding season, the great auks nevertheless lost some of their safe breeding grounds. By 1840 they were becoming noticeably rarer and were forced to recolonize Eldey, much closer to the mainland and easier for men to reach. As numbers grew fewer, their value to collectors increased. Hunting for food also continued. Only about fifty great auks were killed in the years after 1830, indicating that numbers were critically low.

Only 8 breeding locations are known. The largest, at Funk Island, was wiped out by the feather industry. There were at least three nesting islands off Newfoundland during the 15th and 16th centuries, as well as other islands off the tip of Nova Scotia and in Massachusetts Bay. There was a colony in the Magdalen Islands (Bird Rocks) in the Gulf of the St Lawrence River and great auks possibly also bred off Cape Breton as well. In addition to the Icelandic colonies at Eldey and Geirfuglasker (Auk Rocks), there were colonies at St Kilda, the Faeroes, Papa Westray in Orkney, and possibly one at the Calf of Man. There was also a colony on Penguin Island, off Cape La Hune on the south-west coast of Newfoundland. During the breeding season, the birds hunted relatively close to their colonies, ranging about 20 km.

Great auks seem to have deserted Bird Rocks soon after 1700; French navigator Jacques Cartier had captured several "apponatz" (great auks) there in 1534 during his expedition on behalf of the French king. They had gone from St Kilda by 1760. The last one killed in the Faeroes was in 1808, although there were reports of a few there until the 1840s. In the Faeroes, great auks were apparently rounded up at sea and driven ashore. The last pair on Papa Westray were killed in 1812. The war with Napoleon also affected the great auk population; in 1813, British blockades cut off supplies to Danish ports. The governor of the Faeroe Islands sent a gunboat to Iceland to obtain food; this included slaughtering great auks on Geirfuglasker. By 1860, the great auk was officially extinct.

Despite sailors' descriptions, the total population was probably not as huge as they believed. Even the largest colonies covered only modest areas of land. After breeding, the birds may have migrated both southward and northward with their young. Northward-bound migrating great auks were observed in August 1771 by George Cartwright as they moved in and out of the Gulf of St Lawrence. Great auks leaving the Iceland colonies in July may have travelled 20 km per day to reach Greenland by September. Dense breeding colonies and migrations of large numbers of birds may have given the illusion of a larger population than really existed.

With hunting for the feather industry banned on Funk Island, the great auk's final demise was due to naturalists who competed to collect and categorise animal and plant specimens and bird eggs for their collections. In America, Audubon reported that they were regularly sighted off the coast of Massachusetts in the winter months during the 18th century; he also had a specimen in his collection. Museums and zoos paid high prices for rare creatures. A stuffed great auk was an essential exhibit for any respectable museum. They were also essential exhibits in zoos, but with little knowledge of how to care for them, zoos found their great auks short-lived attractions and were obliged to replace them frequently.

A 19th century auction catalogues shows the high prices paid once serious egg collecting began. In 1832, a great auk's egg fetched £15 15s 6d - almost twice the average annual income for a skilled worker (£9 10s). In 1894, an egg sold for £315; the average annual income was £83. In 1898 a great auk's skin together with an egg fetched £630 (equivalent to £50,000 today). In the early 1970s, the remains of a great auk were sold to an American collector for $30,000. In 1971, Iceland's Natural History Museum paid £9,000 for a mounted specimen auctioned at Sotheby's; the museum was prepared to pay up to £20,000.

In the 19th century, Symington Grieve commented "To most people interested in natural history, it would seem that much more interest attached to skins, skeletons, or individual bones of the bird rather than its eggs. The last teach little regarding the habits and structure of the bird compared with others. Yet the prices obtained for eggs are about as high as those obtained for skins, and quite out of all proportion higher than any price obtained for skeletons or bones." Great auk's eggs were status symbols and one major collector owned 9 such eggs. On his death, there was a rumour that his family planned to sell off the eggs, causing a short-lived, but panicky fluctuation in the highly specialised bird's egg market. To end speculation, the family had to publish an open letter in newspapers stating that they had no intention of selling the eggs.

An article in the satirical "Punch" magazine of 24th March 1888 described the sale of a great auk's egg: "A golden egg again - another great auk's egg has just turned up, been put up for aukshun, and knocked down again, without being smashed, fortunately, frail a curiosity as it was to come under the hammer. Mr Stevens of King Street, Covent Garden, sold a very fine egg of the great auk for £225..."

The last stand of the great auk is well documented. On 3 June 1844 a group of fishermen, led by Vilhjalmar Hakonarsson, set out from Keflavik to inhospitable Eldey [Fire] Island, off the extreme south-western tip of Iceland in search of great auks. They were working on commission from Carl Siemsen, a dealer in Reykjavik, who had a purchaser lined up. The men tracked down only one pair of birds, which ran from them, and caught and strangled them, taking care not to damage the feathers or break the skin. The single egg, which would have been valuable if whole, was smashed in the melee so they left it. According to 19th century historian Symington Grieve:

"As the men clambered up they saw two garefowl sitting among numberless other rock-birds (murres and razorbills) and at once gave chase. The garefowl showed not the slightest disposition to repel the invaders, but immediately ran along under the high cliff, their heads erect, their wings somewhat extended. The uttered no cry of alarm, and moved, with their short steps, about as quickly as a man could walk. Jon Brandsson, with outstretched arms, drove one into a corner where he soon had it fast. Siguror Islefsson and Ketil Ketilsson pursued the second, and the former seized it close to the edge of the rock, here risen to a precipice some fathoms high, the water being directly below it. Ketil then returned to the sloping shelf whence the birds had started, and saw an egg lying on the lava slab, which he knew to be a garefowl's. He took it up, but finding it broken put it down again. Whether there was another egg is uncertain. All this took much less time than it takes to tell."

Hakonarsson returned to Eldey in 1846 and 1860 in the hope of finding more great auks, but without success. After the 1844 expedition, there was a reliable sighting in 1852, but with too few breeding individuals, the great auk was effectively extinct.

There was a general refusal to believe the birds rare, let alone extinct. Despite the difficulty reported in obtaining specimens (indicating the bird's rarity), collectors and hunters believed continued hunting would do no real harm since nature would always replenish the numbers. They believed there were undiscovered great auk colonies further north. In 1857, 13 years after the killings on Eldey, Johannes Iapetus Steenstrup, Professor of Zoology at Copenhagen University published his definitive account of the great auk's range. Despite exhaustive research, he found no trace of surviving colonies in the Arctic regions or elsewhere. Despite his report, ornithologists continued to seek the great auk in the vain hope that a colony might be found or a bird sighted. As late as 1891, a Scottish newspaper reported an alleged sighting of great auks on St Kilda. As late as 1945, it was still listed as alive in some bird guides.

The entry for Great Auk in Harmsworth natural History (1910): The typical members (Alca) of the family are characterised by the large size of the compressed beak, marked in front by oblique grooves, and feathered at its base close up to the slit-like nostrils, which are almost concealed by a dense, velvety feathering, completely filling the pits in which they are situated. The wings are more or less short, and the tail is graduated, with its component feathers pointed. On account of its extinction during the nineteenth century, as well as from being the largest representative of the family, and the only bird in the Northern Hemisphere incapable of flight, the great auk, or garefowl (A. impennis), is a species of great interest. In common with many other northern sea-birds, it was formerly known as the penguin - a name now transferred to the well-known birds of the Southern Hemisphere; and in size may be roughly compared to a goose, its total length being about 32 inches. It is especially characterised by the rudimentary condition of the wings, which, owing to the reduction in the length of the ulna and bones of the digits, were quite useless in flight; while it is further distinguished by the beak being equal in length to the head, and furnished with numerous grooves on its lower as well as on its upper half. In colour the plumage of the head, neck, and back is black, but the under parts, as well as a characteristic spot in front of the eye, are white. As already mentioned, the tail is peculiar in carrying sixteen feathers.

As in the allied species, the eggs are relatively large in proportion to the size of the bird, often measuring just over 5 inches in length; and they have also the same elongated form, with one end much larger than the other. They have a creamy-white ground colour, marked with black or brown streaks and blotches, with underlying grey patches. It is of interest to note in this connection that about seventy-two great auk’s eggs are known to exist, and of these fifty-one are, it is believed, in Great Britain. A very fine specimen which came into the market some years ago realised 315 guineas.

All that remains of the great auk now are 78 mounted skins (most from Eldey Island), 24 complete skeletons, 2 collections of preserved viscera, and around 75 eggs. The skinned corpses of Eldey island's last great auks are preserved in spirits at the Royal Museum in Copenhagen. In 1971, Iceland's Natural History Museum paid £9,000 for a stuffed great auk and had been willing to bid up to £20,000, if necessary, to acquire the specimen. It was ironic that the country where the great auk had finally been wiped out was willing to pay so much to bring the great auk home. Elsewhere, there are hopes that preserved specimens and eggs might yield enough DNA for the great auk to be a candidate for cloning attempts.

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