Great Auk—the Extinct Penguin of the North

Table of Contents

“Today the original penguin is extinct.”

Robert Silverberg, 1967, p. 84

The great auk was a large, flightless seabird of the North Atlantic. Its population may once have numbered several million individuals. With its black-and-white plumage, upright posture, and short, flipper-like wings, it bore a striking resemblance to today’s penguins of the Southern Hemisphere. In fact, in historical usage it was the original penguin: the word initially referred to this North Atlantic bird and was only later applied to the similar-looking birds of the southern seas.

The name has been documented since the late 16th century. European sailors encountered great auks off Newfoundland, among other places, and called them penguins. When they encountered large, upright, flightless seabirds in the southern seas a little later, they applied the already familiar name to those animals. This shift in meaning occurred rapidly: according to the etymologist Anatoly Liberman (2011), the collections of travel accounts published by Richard Hakluyt in 1589 and 1600 contain early evidence that the word “penguin” was used for both the great auk and the birds of the Southern Hemisphere.

The scientific name Pinguinus impennis also preserves this naming history. Carl Linnaeus initially described the species as Alca impennis in 1758. Toward the end of the 18th century, Pierre Joseph Bonnaterre placed it in the separate genus Pinguinus. The species name impennis means “wingless” and refers to its greatly shortened wings, which were unsuitable for flight.

Where does the word “penguin” come from?

The origin of the word remains unclear. One frequently cited explanation derives it from the Welsh pen gwyn: pen means “head” and gwyn means “white.” The name may have referred to the large white patches in front of the great auk’s eyes. According to another hypothesis, pen gwyn originally referred to a rock or headland off Newfoundland that had been colored white by bird droppings. The place name was supposedly later applied to the birds living there. Both explanations are considered uncertain (Liberman, 2011).

The Russian zoologist Igor Akimushkin proposed another interpretation in 1972: English sailors may have referred to the great auk’s shortened wings as pin-wing or ping-wing. Liberman, however, considers this derivation an imaginative folk etymology. Nor is there evidence for a derivation proposed as early as the 16th century from the Latin pinguis, meaning “fat” or “well-fed.”

In Iceland and the Old Norse-speaking world, the great auk was known as Geirfugl or Geyrfugl. The Old Norse word combines geirr, meaning “spear” or “pike,” with fugl, meaning “bird.” It therefore means “spear bird” and probably refers to the large, pointed beak. The English name garefowl also derives from this term. According to Akimushkin, the bird appears under this name in Icelandic sagas and Eddic poems as well.

Penguin or auk?

The similarity between the northern and southern birds led to confusion for centuries. A linguistic distinction eventually developed in French. In 1760, Mathurin-Jacques Brisson used the term manchot for the flightless birds of the Southern Hemisphere. Georges-Louis Leclerc de Buffon adopted this distinction in 1783 and explained that the North Atlantic pingouins and the southern manchots belonged to different groups of birds. To this day, pingouin is the French term for the auks of the Northern Hemisphere, while the southern penguins are called manchots. Accordingly, the great auk is called the Grand Pingouin.

Great auks and penguins were not, in fact, closely related. The great auk belonged to the auk family, whereas penguins form a separate order of birds. Their similarity evolved independently through convergent evolution: both lineages produced streamlined, black-and-white diving birds whose shortened wings served primarily for underwater propulsion.

Great auk – fact sheet

alternative names	Giant auk, Garefowl, Grand Pingouin, Geirfugl, Geyrfugl, Gairfowl
scientific names	Pinguinus impennis, Alca impennis, Plautus impennis, Alca borealis, Penguin impennis, Pingouin impennis, Chenalopex impennis, Alca major, Mataeoptera impennis
original range	North Atlantic
date of extinction	last confirmed record: 1844; last credible sighting: 1852
causes of extinction	intensive hunting; collection of birds and eggs
IUCN status	extinct

Where the great auk lived and bred

The great auk is often portrayed as an Arctic bird. However, as early as 1861 the English ornithologist Alfred Newton rejected the idea that the species lived mainly among Arctic drift ice. He considered supposed records north of the Arctic Circle, particularly those from Spitsbergen, doubtful. Instead, the great auk was a bird of the temperate and subarctic waters of the North Atlantic.

In historical times it lived primarily in the cool, food-rich coastal and shelf waters between northeastern North America and northern Europe. In the western North Atlantic, its regular occurrence ranged from Labrador and Newfoundland across the Gulf of St. Lawrence to New England. Historic winter observations are known from Massachusetts Bay. In the east the species occurred near Greenland, Iceland, the Faroe Islands, the British Isles and on the Norwegian coast. Outside the breeding season, individual animals could have reached further south to the French Atlantic coast and the Iberian Peninsula.

This range was not uniformly populated. The great auk spent most of the year on the open sea and probably stayed mainly where cold ocean currents and productive fishing grounds provided sufficient food. Because it could not fly, it also completed its seasonal migrations by swimming.

Only a few islands were suitable for breeding

Probably between May and July, great auks came ashore to breed. Only a few islands and rocky outcrops were suitable. They had to be accessible from the water, have flat or gently sloping shores, and lie far enough from the mainland to protect the colonies from larger terrestrial predators. At the same time, sufficient food had to be available nearby.

The largest historically known breeding colony was on Funk Island, around 60 kilometers off the east coast of Newfoundland. Many thousands, possibly even tens of thousands, of pairs probably bred there. However, the colony’s exact size can no longer be determined. Other breeding sites in the western North Atlantic were probably located on the Rochers aux Oiseaux in the Gulf of St. Lawrence. The Penguin Islands off Newfoundland’s southern coast are also considered a former breeding area, although they are less well documented.

In the eastern North Atlantic, the Icelandic colonies are particularly well known. One of the last important breeding sites was the difficult-to-access rock of Geirfuglasker, southwest of Iceland. After it sank beneath the sea around 1830, the remaining great auks moved to the nearby island of Eldey. The last two confirmed breeding birds were killed there in 1844.

Earlier or smaller breeding populations probably also existed near St Kilda off Scotland’s west coast and on Papa Westray in the Orkney Islands. Other suspected colonies are poorly documented. Moreover, not all known breeding sites were occupied at the same time. Many disappeared long before the species’ final extinction. By the 18th and early 19th centuries, the last larger populations had become concentrated in a few refuges near Newfoundland and Iceland.

The prehistoric range

Archaeological and fossil finds show that the great auk was more widely distributed in earlier times. Bones have been found in Scandinavia, Denmark, the Netherlands, France, Spain, Portugal, and Italy, among other places. Records from the western Mediterranean are also known from the Pleistocene.

During the ice ages, cold marine waters extended farther south than they do today. Suitable living conditions may therefore have existed at times along the coasts of southern Europe. However, the finds do not prove that the great auk lived permanently, or even bred, at every site. People may also have transported killed birds, meat, or bones over long distances.

The prehistoric range reconstructed from bone finds is therefore larger than the known historical range. It shows where great auks have been recorded, but not where breeding colonies regularly existed.

The southernmost archaeological records in the western Atlantic come from Florida. In 1960, the paleontologist and ornithologist Pierce Brodkorb suggested that great auks may have reached the peninsula during particularly cool climatic phases in winter, around 1000 BC and AD 1000. This explanation is uncertain, however. The bones may also have reached Florida through Indigenous trade networks. Given the small number of finds and the long intervals between them, it remains unclear whether great auks repeatedly reached the region on their own.

An approximately 20,000-year-old cave painting from southern France may depict great auks. If this interpretation is correct, the people living there would have been familiar with the birds.

The southernmost known bone find comes from the El Harhoura 2 cave on the Moroccan Atlantic coast. It was discovered in a Neolithic layer. In 2010, Emilie Campmas and her research team discussed two possible explanations: either great auks occasionally reached North Africa in winter, or the bone was brought there by humans. A single find is therefore not enough to consider Morocco a regular part of the species’ range.

In historical times, the great auk was primarily a bird of the cool North Atlantic waters. Fossil and archaeological finds show, however, that during earlier climatic periods its range sometimes extended much farther south.

Relationships and evolutionary history

The great auk was the last known representative of the genus Pinguinus. The only other species known to date is Pinguinus alfrednewtoni. It lived about four million years ago, during the early Pliocene, on the Atlantic coast of North America. Fossil bones have been found in the Yorktown Formation at the Lee Creek Mine in North Carolina. Their dimensions fall within the range of size variation seen in the later great auk; the two species were already very similar in size and build (Olson, 1977).

Differences in the wing bones suggest that the wings of P. alfrednewtoni were less fully modified into rigid underwater-propulsion structures. Storrs L. Olson (1977) therefore considered the species to be a possible ancestor of the great auk. However, due to the incomplete fossil record, it cannot be determined whether it was actually its direct ancestor or a close branch of the same evolutionary line.

The great auk’s closest living relatives were identified using mitochondrial DNA. Truls Moum and Ulfur and Einar Árnason examined skin and feather samples from a great auk captured in Iceland in 1821. Their analysis, published in 2002, showed that the razorbill (Alca torda) is the great auk’s closest living relative. The little auk (Alle alle) forms the sister group to these two lineages, while the guillemots of the genus Uria diverged earlier.

Despite their close relationship, these species evolved very differently. The little auk remained a small seabird that feeds mainly on plankton. The razorbill evolved into a medium-sized diver that retained the ability to fly, while the great auk grew larger and eventually became completely flightless. Moum et al. (2002) interpreted these differences as distinct specializations within an originally Atlantic auk lineage.

The oldest fossils already attributed to the modern great auk, Pinguinus impennis, include finds from Boxgrove in southern England and from Bermuda. Both date to the Middle Pleistocene and are at least 400,000 years old. The Bermuda find consists of part of an upper bill and shows that great auks reached both sides of the Atlantic even then. It probably came from a bird that had traveled far south outside the breeding season and died there (Roberts et al., 1997; Olson, 2003).

The great auk was not the only flightless auk in Earth’s history. The genera Mancalla, Miomancalla, and Praemancalla, among others, lived in the North Pacific. These Mancallinae belonged to a different and much older evolutionary lineage. Flightlessness evolved independently at least twice within the auk family: in the Atlantic lineage of Pinguinus and in the Pacific lineage of the Mancallinae (Smith, 2011; Smith & Clarke, 2015).

In the great auk, the wings were fully specialized for underwater propulsion—an adaptation that decisively shaped its body and way of life.

The great auk: size and weight

With a body length of about 70 to 85 centimeters, the great auk was considerably larger than any living auk and roughly the size of a small goose. Historical sources occasionally give a figure of around 90 centimeters. Such information is uncertain, however, because no living great auk was ever scientifically measured. Current estimates are based on historical descriptions and preserved skeletons and skins. Mounted specimens are only of limited use for measurements because their posture and body shape depend heavily on the taxidermist’s work.

An Atlantic puffin (Fratercula arctica) reaches a body length of only about 30 to 35 centimeters, and a razorbill about 38 to 43 centimeters. Even the common guillemot and Brünnich’s guillemot (Uria), among the largest living auks, remain much smaller at no more than about 50 centimeters and roughly one kilogram.

Within the wider auk family, however, the great auk may not have been the largest member. In 2015, N. Adam Smith estimated its weight at around 4.75 kilograms, compared with approximately 5.36 kilograms for the extinct Pacific species Miomancalla howardi. Since more recent studies indicate a substantially lower body mass for the great auk, however, the two species can be compared only to a limited extent on the basis of these older figures. What is certain is that M. howardi was also a very large, flightless auk and may even have been heavier than the great auk.

It is not known how much a great auk actually weighed. The long-held estimate of 4.5 to five kilograms goes back to the memory of a resident of the Faroe Islands. Decades later, he reported that a bird caught in 1808 weighed around nine Danish pounds. It is unknown whether the animal was actually weighed at the time or whether its weight was only estimated.

Robert Montgomerie and Tim Birkhead reassessed this estimate in 2025 using modern statistical methods. A comparison of preserved leg and thigh bones with those of living auks yielded an estimated body mass of around 3.44 kilograms. A second calculation, based on the volume of preserved eggs and the ratio of egg size to body weight in today’s auks, yielded around 3.68 kilograms.

Combining the two results, the researchers derived an average body mass of about 3.56 kilograms. The great auk may therefore have been considerably lighter than long assumed. A fresh egg probably weighed about 350 grams.

These estimates are not direct measurements either. Since no living auk species reaches the size of the great auk, a degree of uncertainty remains. Nevertheless, the new study suggests that the traditional image of a bird weighing almost five kilograms needs to be revised.

Plumage and possible sex differences

The great auk had the black-and-white coloration typical of many seabirds. Its back, head, and wings were dark, while its underparts were white. Its appearance changed over the course of the year, however. In summer it had large white patches in front of its eyes and a dark throat. In winter, the eye patches disappeared, while the throat and parts of the face turned white.

According to standard inventories, 74 of the 78 preserved skins show the great auk in summer plumage and only four in winter plumage. This marked imbalance is probably due primarily to the circumstances of their capture. During the breeding season, the birds gathered in large colonies on land and were easy to catch. In winter they lived widely scattered across the open North Atlantic and were much harder for hunters and collectors to reach.

The juvenile plumage is also known from only a few specimens. Young birds did not yet have the adults’ striking markings; no reliably identified chick specimen survives.

Males and females apparently did not differ in coloration. In 1986, however, Dieter Luther suggested there was slight sexual dimorphism: the sexes may have differed in body and bill size.

A 1988 morphometric study by Bradley C. Livezey offered some support for this hypothesis. Measurements of bills and femurs revealed two size classes. Because the remains examined generally cannot be assigned to a sex, it remains unclear whether the differences actually corresponded to males and females. If sexual dimorphism existed, it was probably only slight.

A life between sea and rocks

Much has been written about the extinction of the great auk, yet surprisingly little is known about its way of life. In 1984, Sven Axel Bengtson noted that no naturalist had ever reported encountering a great auk in its natural habitat. Our knowledge is therefore based primarily on accounts by sailors, fishermen, and bird hunters, as well as later comparisons with living auks. Historical observations and scientific reconstructions cannot always be clearly distinguished.

On land and in water

In 1858, John Wolley and Alfred Newton traveled to southwestern Iceland. They searched for any surviving great auks, examined bone finds, and interviewed people who had seen or captured the birds themselves. Newton published their accounts in 1861.

According to these accounts, great auks swam with their heads raised and their necks drawn in. When threatened, they did not try to flutter across the water’s surface but dived immediately. On the rocks, they stood more upright than razorbills and common guillemots and moved straight ahead in short steps. They occasionally made quiet croaking sounds and seemed to respond more strongly to noises than to visible movements. They did not defend their egg; captured birds could, however, bite fiercely with their powerful bills.

The reports show a bird that moved upright on land but had little mobility and tried to reach the water as quickly as possible when in danger. The supposedly weak response to visible movement is not evidence of poor vision. On their remote breeding islands, the birds may have had little experience with approaching terrestrial animals.

Its true strengths lay in the water. Like other auks, the great auk propelled itself with powerful strokes of its short wings, while its feet and body posture served for steering. Its wing bones were greatly shortened and had thick walls—adaptations for underwater propulsion. How deep or how long the great auk could dive was never measured. The literature often mentions depths of around 75 meters, but all information about its diving performance is based solely on comparisons with living species.

Outside the breeding season, great auks probably spent most of their lives on the open North Atlantic. Little is known about their migration routes, group sizes, or how long they remained in particular marine areas.

Isotope analyses by Keith A. Hobson and William A. Montevecchi from 1991 indicate that juveniles fed at a lower level of the food web than adult great auks. They probably received mainly krill-like crustaceans regurgitated by their parents. Adults fed primarily on fish from higher trophic levels.

Fish remains from archaeological layers on Funk Island confirm a fish-rich diet. The birds ate energy-rich schooling fish in particular, including capelin (Mallotus villosus) and members of the herring family. The great auk probably hunted mainly small to medium-sized fish in the productive waters surrounding its breeding colonies. The exact composition of its diet may have varied by region and season (Hobson & Montevecchi, 1991; Montevecchi & Kirk, 2020).

Breeding and rearing

Probably between May and July, great auks returned to breed on a few remote rocky islands. They nested close together, sometimes alongside razorbills, common guillemots, and other seabirds. Great auk eggs were elongated and distinctly pointed at one end; this shape probably made them particularly stable on the bare rock, where the great auk incubated its egg without a nest. Research by Tim Birkhead et al. (2020) also shows that the shell was especially strong where it rested on the ground.

Parental care was never scientifically observed. Both parents probably took part in incubating the egg and caring for the chick, as is common among related auk species. The incubation period is generally estimated at around 39 to 44 days.

Previous authors assumed that the young bird left the colony just a few days after hatching. This idea was probably based on a misinterpretation of historical accounts of St Kilda, which reported a brief presence of the great auk. However, since eggs were regularly collected there, the natural length of the breeding stay may have been underestimated. Comparisons with razorbills and guillemots tend to suggest that young great auks went into the sea after about 20 days and were presumably still cared for there by a parent (Birkhead, 2021). The fact that great auk pairs stayed together for life is plausible, but not proven.

It is difficult to reconstruct which natural predators preyed on the great auk. Adults were large and spent most of their lives in the open sea, so they were probably taken regularly only by larger marine predators. Eggs and young birds were probably more vulnerable at the breeding sites.

The great auk was not a clumsy animal but a highly specialized swimmer and diver. It became vulnerable where it had to reproduce: on a few flat islands that were easily accessible from the water. On land, the flightless bird had little chance of escaping humans. It was precisely this dependence on a small number of breeding sites that later became its decisive disadvantage.

Exploitation: From food bird to collector’s item

People hunted the great auk long before European sailors reached its large breeding colonies in the North Atlantic. Bones from prehistoric settlements and caves show that the birds were used for their meat, eggs, and fat for many millennia. Such finds come from Greenland, northeastern North America, Scandinavia, Great Britain, and the Bay of Biscay region. Remains from Gibraltar have even been linked to Neanderthals. It is no longer possible to determine precisely how this early hunting affected individual populations (Fisher & Lockley, 1954; Thomas et al., 2019).

A possible depiction of this early use can be found at Hjemmeluft near Alta in northern Norway. An approximately 6,000- to 5,000-year-old rock engraving there shows a person holding a large bird by the neck. In their 2026 study, Tim Birkhead and Robert Montgomerie consider it likely that the bird is a great auk. Its size relative to the person, powerful bill, webbed feet, and strikingly small, flipper-like wings support this interpretation.

The person may be deliberately holding the bird by the upper neck at arm’s length to avoid its bill. Historical accounts confirm that captured great auks could bite fiercely. The engraving may therefore record not only the bird’s appearance but also a method of capture. Four other bird images at Alta share similar features but are less distinct.

Archaeological finds support this interpretation. Great auk bones have been discovered at more than 50 sites along the Norwegian coast, most dating to between 6,000 and 2,000 years before the present. The nearest known sites lie about 150 kilometers northeast of Alta. It is unknown whether the birds bred in the region at the time or were merely caught there outside the breeding season.

Persecution after 1500

With European voyages across the North Atlantic, exploitation reached a new level from the 16th century onward. On the breeding islands, sailors could capture large numbers of the flightless birds in a short time. Akimushkin (1972) cites reports of more than a thousand great auks being killed in a single day, or of two boats being filled with them within half an hour.

Great auks provided meat for ships’ crews, were salted, or were used as fishing bait. People also used their eggs, fat, feathers, and down. Because the birds nested close together on a few islands and could not fly away on land, they were easy to herd together. According to Luther (1986), the great auk’s size made it a particularly rewarding target for sailors, fishermen, bird catchers, and sealers.

A manuscript drawing made by Guðni Sigurðsson in 1770 shows how such hunting expeditions were conducted. It depicts bird catchers on Geirfuglasker and shows how their boats were anchored by the stern to prevent them from striking the rocks in the heavy surf (Petersen, 1995).

From the late 18th century onward, great auks on Funk Island were killed en masse, especially for their feathers. Fuller (2000) describes the process vividly:

“The birds were also slaughtered for their feathers. On Funk Island, the (…) animals were rounded up, beaten to death or immobile, and then thrown into cauldrons of boiling water, which caused their feathers to come off; the fires under these cauldrons were stoked with the fat and oil of the auks, which had already suffered a similar fate.”

On St Kilda, too, almost every part was used. James Fisher and R. M. Lockley (1954) mention an account from around 1682 stating that the birds’ fat was placed in a bird’s stomach, dried by the fireplace, and then sold on the mainland as a painkiller. The author of the original account, however, had probably never visited St Kilda.

The rarity increased the value

As the great auk became increasingly rare in the early 19th century, the purpose of its persecution also changed. Museums, dealers, and private collectors now sought skins, skeletons, and eggs in growing numbers. Only a few scientific collections possessed specimens of the bird, so prices rose sharply as it became rarer.

The large eggs were especially coveted. Their coloration and markings varied from specimen to specimen, allowing collectors to own several without regarding them as mere duplicates (Fuller, 2000). According to Hume (2017), incubated eggs were sometimes discarded and only fresh specimens retained. Because a pair of great auks laid only one egg per year, every egg collected meant the loss of an entire year’s brood.

Akimushkin (1972) summarized this final phase as follows:

“Collectors completed the tragedy; the great auk became a rarity. Museums and enthusiasts paid large sums for eggs and skins. […] A great auk skin could fetch almost a hundred crowns. That was more than the fat and feathers from an entire sloop-load of birds.”

Collecting did not initiate the centuries-long decline, but it became a decisive factor in the final phase. Just when the species would have needed special protection, its rarity and scientific value made every great auk still within reach a coveted object. The last documented birds were no longer killed for food or feathers but were obtained specifically for zoological collections.

The demise of the last breeding colonies

The great auk did not decline everywhere at the same time. Archaeological finds indicate that populations in the eastern North Atlantic declined early—possibly due to prolonged human exploitation, but perhaps also under the influence of environmental change. Many easily accessible breeding sites probably disappeared before the species’ history was documented in greater detail. Individual colonies nevertheless survived on remote islands off Scotland, the Faroe Islands, and Iceland into the 18th and 19th centuries.

The end of Funk Island

Funk Island was probably home to the largest historically known breeding colony of the great auk. Its exact size is unknown; the often-cited figure of around 10,000 breeding pairs is based on a later estimate. Accounts by early European sailors leave no doubt, however, that an extraordinary number of birds lived there.

In 1534, Jacques Cartier’s crews filled several boats and barrels with great auks in a short time. In the centuries that followed, the birds were regularly used as food and fishing bait or killed for their fat and feathers (Fisher & Lockley, 1954).

According to David Day (1981), hunting on Funk Island intensified again in the late 18th century. After populations of eiders, which were also hunted for their down, declined, feather collectors increasingly turned to great auks. In a travel account published in 1792, the English officer and trader George Cartwright warned that the colony was in danger of disappearing:

“In recent years it has become common for several crews of men to live on Funk Island throughout the summer and kill birds solely for their feathers. The destruction they have caused is unimaginable. If this practice is not stopped soon, the entire population will be almost completely wiped out—especially the penguins, since this is now the only island where they still breed. All the other islands lie so close to the coast of Newfoundland that they are constantly being plundered.”

As late as 1794, the English sailor Aaron Thomas witnessed the mass and extremely cruel killing of birds on Funk Island. A few years later, the colony was gone; by 1800, the great auk was considered extinct on Funk Island (Luther, 1986). Its largest known breeding population had thus collapsed within just a few decades.

The Last Birds of the British Isles

The breeding sites on the European side of the Atlantic also gradually disappeared. On Papa Westray in the Orkney Islands, a single pair was still breeding in 1812. The female was killed with a stone while on her egg. The male remained at the breeding site and was shot the following year by the collector William Bullock. The islanders called the pair the “King and Queen of the Auks” (Fisher & Lockley, 1954).

In May 1834, a great auk was captured at the entrance to Ireland’s Waterford Harbour and kept in captivity for several months before it died. The last reliable British record probably dates to 1840: islanders captured a live great auk on Stac an Armin and killed it three days later (Fuller, 2000).

Geirfuglasker—the Last Major Refuge

Several Icelandic rocks bore the name Geirfuglasker. A small colony on the rock of that name near the Westman Islands probably died out around 1800. The larger and much harder-to-reach rock southwest of the Reykjanes Peninsula, however, remained occupied for several more decades (Fisher & Lockley, 1954).

Geirfuglasker was surrounded by heavy surf and could be reached only at considerable risk. Older accounts mention large numbers of birds and eggs being collected, but the size of the colony is uncertain. A later estimate suggested only about 300 to 1,500 breeding pairs (Petersen, 1995).

The British crew of the Salamine caused particularly heavy losses in 1808. According to Newton (1861), the men spent an entire day killing great auks and trampling their eggs and chicks. In 1813, another expedition brought at least 24 great auks to Reykjavík; additional birds were salted or taken to the Faroe Islands.

When the Danish ornithologist Frederik Faber, the natural-history enthusiast Count Frederik Christian Raben, and the botanist A. M. Mørch visited Geirfuglasker in 1821, they found no great auks there. Raben managed to land at great risk and narrowly escaped drowning. The colony was not yet completely extinct, however: in 1828, at least one adult bird was captured there for the Copenhagen museum (Fisher & Lockley, 1954).

Around 1830, Geirfuglasker disappeared beneath the sea. This is often linked to earthquakes or submarine volcanic activity, although the exact events are not clearly documented. In the same year, great auks were found on the rock of Eldey, closer to the coast (Fisher & Lockley, 1954).

The last years on Eldey

Eldey offered the birds less protection than Geirfuglasker. At one point a gently sloping rock slab led from the water to the breeding area, making it relatively easy for catchers to land there. As early as 1830, around 20 or 21 great auks were caught for traders on two trips. The following year, 24 more animals were added, and in 1834 at least nine birds and several eggs. Skins and eggs were still collected from Eldey in 1840 and 1841 (Fisher & Lockley, 1954).

Newton later estimated that approximately 60 great auks had been captured on Eldey between 1830 and 1844. Adding together every individual account known to him produced a total of as many as 87 birds. Because some reports may overlap or be based on uncertain memories, however, he did not consider this figure a reliable count. A large proportion of the skins and eggs that entered European collections at the time probably came from Eldey.

The killing of the last two great auks

In early June 1844, a party of fourteen set out for Eldey. The expedition had been initiated by the merchant Carl F. Siemsen, who wanted to obtain great auks for zoological collections. Because of the heavy surf, only three men could set foot on the rock: Sigurdur Isleifsson, Jón Brandsson, and Ketill Ketilsson.

Among common guillemots and razorbills, they discovered two great auks and an egg. The birds did not attempt to attack the intruders but ran away beneath the cliff. One was driven into a corner and the other trapped near the edge of the rock. Ketill found an already damaged egg on a rock ledge and put it back down.

Alfred Newton summarized the participants’ statements in his 1861 report:

“The great auks did not show the slightest inclination to repel the intruders, but immediately ran under the high cliff (…). They uttered no cry of alarm, and moved with their short steps almost as quickly as a man could walk. Jon, with outstretched arms, drove one into a corner, where he soon caught it. Sigur and Ketil pursued the second, and the former seized it close to the edge of the rock (…). Ketil returned (…) to the (…) ledge, (…) and saw an egg lying on the lava plate, which he knew was from the great auks. He picked it up, but when he found it was broken, he put it back down (…) The birds were strangled and thrown into the boat (…).”

The two birds were then sold for 80 rigsbankdaler to the dealer Christian Hansen. They were skinned in Reykjavík. Parts of their internal organs later entered the zoological collection in Copenhagen, while the skins were sold on separately.

Newton’s report, published in 1861, is the principal contemporary source for these events. It was based primarily on the notes of John Wolley, who, together with Newton, interviewed numerous people in Iceland in 1858—including twelve of the fourteen participants in the final expedition. The accounts were therefore recorded only 14 years after the event, although some were conveyed through interpreters. Details may consequently have been affected by lapses in memory, translation, and differing descriptions.

In 1858, Newton and Wolley still hoped to find surviving birds on Eldey or the more distant Geirfugladrangur. The dangerous surf prevented them from landing, however. Later searches produced no evidence. At Bæjarsker they also found bones from at least eight great auks, some bearing cut marks. The finds documented the birds’ former use by people but offered no evidence of their continued survival.

Since the voyage of 1844, no living great auk has been unequivocally identified. Whether the two birds killed on Eldey were actually the last representatives of their species remains an open question.

People as a decisive factor

Human persecution of the great auk has been well documented since the 16th century. For a long time, however, it remained unclear whether intensive hunting alone caused its extinction or whether its populations had already been weakened by climatic changes or other natural factors.

Archaeological finds indicate changes, at least regionally. Great auk bones are becoming rarer in younger finds in England, Scotland and Scandinavia, which could indicate a decline or shift in populations in the eastern North Atlantic. It cannot be determined from the bone finds whether hunting, environmental changes or both factors were responsible (Thomas et al. 2019).

Newton (1861) did not consider natural changes the principal cause of the decline. Only one important breeding site was lost through natural processes, whereas the birds at the remaining colonies had been deliberately persecuted over a long period. Bengtson (1984) was more cautious: because data on population trends, hunting pressure, and possible environmental changes were lacking, the effects of people and climate could hardly be separated.

A study published in 2019 and led by the biologist Jessica E. Thomas therefore investigated whether the great auk population had already declined before intensive European hunting began in the early 16th century. The research team combined genetic analyses with data on ocean currents and population modeling. To do so, they examined the complete mitochondrial genomes of 41 great auks from almost the entire former range. The remains were between approximately 170 and 15,000 years old.

The genetic data provided no evidence of a long-term population decline. Mitochondrial diversity remained high until the recent past: only two of the 41 birds examined had the same haplotype in the DNA section that could be compared across all samples. There was no evidence of declining diversity in the younger specimens either. A small, isolated remnant population persisting for many generations would normally be expected to show far greater genetic impoverishment.

Nor was there a clear geographic separation among populations. Birds from different parts of the North Atlantic formed no clearly distinct populations. This suggests that genetic exchange occurred between regions, at least over longer periods. Experiments with GPS-tracked drifters showed possible connections between former breeding areas near St Kilda, Iceland, and Funk Island. The researchers emphasized, however, that present-day ocean currents provide only speculative clues to the birds’ former migration routes.

The reconstruction of population history revealed a largely constant effective population size within the temporal resolution of the data. A longer-term decline immediately before the start of intensive hunting was not evident. However, the shallow temporal depth of the mitochondrial lineages suggests a much older population bottleneck in the late Pleistocene, possibly related to climatic fluctuations. Because of the great uncertainty, this bottleneck cannot be assigned to a specific event. However, the high number of unique haplotypes and the presumably large subsequent total population suggest that the great auk had recovered from it.

Would hunting alone have been enough?

In a population viability analysis, the team also examined whether human hunting could wipe out even a very large population within a few centuries. The models assumed total populations of two or six million birds. At the same time, deliberately favorable assumptions were made about the species’ survival, including a high reproductive rate and low natural mortality.

Under these especially conservative conditions, the population could withstand the annual removal of up to 9% of its initial numbers. If 10.5% of the birds and 5% of the potential eggs were removed each year, however, the species became extinct in every simulation within 350 years. For an initial population of two million birds, this would correspond to 210,000 birds killed and around 25,700 eggs collected each year. Even without egg collection, hunting pressure posed a substantial risk of extinction. Under more realistic assumptions based on the razorbill, considerably lower harvests already created a serious threat.

These figures do not reconstruct the actual numbers taken. They merely show the scale that would theoretically have been sufficient. Historical accounts, however, make hunting pressure of this magnitude seem plausible. From the late 15th century onward, around 300 to 400 European ships visited the fishing grounds off Newfoundland each year. One contemporary account describes the crews of two ships catching about 1,000 great auks within half an hour. If each of 400 ships had hunted at a comparable rate for only half an hour per year, around 200,000 birds would already have been killed. Additional losses resulted from nets, fishing lines, and egg collection.

What genetics cannot show

The study does not prove that environmental changes had no influence at all on distribution or individual populations. A regional decline in the eastern North Atlantic may also have occurred without leaving a discernible genetic trace. In addition, only maternally inherited mitochondrial DNA was examined. It represents a single genetic marker; nuclear DNA might have provided finer resolution of population structure. The sample of 41 specimens was also comparatively small for a population-genetic study.

Importantly, a rapid collapse can remain almost invisible in the genetic data. For a long-lived species, there may be too few generations between the start of a rapid decline and extinction for genetic diversity to decrease significantly. The analyses therefore cannot determine exactly when the final collapse began or whether it was preceded by several regional declines.

Nevertheless, an overall picture emerges: before intensive European persecution began, the great auk was not a species that had long been genetically impoverished and was already heading inevitably toward extinction. Its population was probably large and widespread, with populations in different regions well connected to one another. The modeling also shows that the documented human exploitation alone may have been sufficient to wipe out several million great auks within a few centuries.

The fate of the great auk shows that even common and widespread species are not safe from extinction. If a slowly reproducing species is exploited over a long period at a small number of densely populated breeding sites, its population can collapse without any preceding natural environmental crisis (Thomas et al., 2019).

After 1844: Reports of Surviving Great Auks

The killing of the pair on Eldey in 1844 is considered the last confirmed record of the great auk. It is unlikely, however, that these two birds were truly the last of their species. Possible great auks were reported several times in the following decades, but none of these reports could be confirmed beyond doubt. Errol Fuller wrote in 2000:

“It is unrealistic to assume that this pair were actually the last of their species. Undoubtedly there were still a few specimens left here and there. But shortly after 1844, somewhere in the deep waters of the North Atlantic or (…) on a windswept reef (…) the last great auk died.”

The Norwegian report from 1848

One of the earliest reports comes from northern Norway. In 1855, Lorenz Brodtkorb of Vardø told John Wolley that in April 1848 he had seen four unusual birds in a strait between Vardø and the islands of Hornø and Reinøya. After he shot one, the others did not fly away but continued swimming. The dead bird had a black back, dark head and neck, a white patch by the eye, a narrow pointed bill, and strikingly small wings (Shuker, 2019).

Newton was generally skeptical of records far north of the Arctic Circle. More recent archaeological finds show, however, that great auks reached as far north as Norway, at least in prehistoric times. Birkhead and Montgomerie (2026) even consider former breeding in the region possible: Alta still lies at the edge of the boreal marine zone, and numerous great auk bones have been discovered at sites in northern Norway. They therefore state explicitly that the archaeological evidence and the rock engraving lend some credibility to the Vardø report. Doubts center mainly on the bird’s description, particularly its narrow, pointed bill.

Brodtkorb left the carcass on the beach; the tide washed it away the next day. His observation can therefore no longer be verified. The narrow, pointed bill he described argues against identification as a great auk and is more consistent with a common loon or guillemot. The great auk had a large, powerful bill compressed from side to side. Newton therefore considered the identification doubtful as early as 1861. Wolley was also unable to find a matching bird at the stated location in 1855.

The possible sighting of 1852

The report by the Scottish ornithologist Henry Maurice Drummond-Hay is generally considered the most credible later sighting. In December 1852, he observed an unusual diving bird at close range along the outer edge of the Newfoundland Banks. Newton (1861) reported:

“Drummond-Hay has reported to me that in December 1852, as he was passing along the outer edge of the Newfoundland Banks, he saw what he was convinced was a great auk. At first he thought it was a common loon; but then he recognized the large beak and the white spots, which left him in no doubt. The bird dived at a distance of thirty or forty yards [27–37 m] from the steamer.”

Drummond-Hay was an experienced ornithologist and later became a co-founder and the first president of the British Ornithologists’ Union. The observation site was also within the species’ former range. The report cannot be confirmed, however: Drummond-Hay was apparently the only observer, and no specimen, drawing, or more detailed description survives.

The fact that the encounter took place in December also requires explanation. At that time of year, the great auk wore its winter plumage, which lacked the large white patches in front of the eyes. By the “white spots,” Drummond-Hay may have meant the white throat or underparts of the otherwise dark bird. Despite this uncertainty, his sighting is often treated as the last credible indication that the species still survived. BirdLife International and the IUCN also list 1852 as the year of the last sighting of a living great auk.

Later, increasingly uncertain reports

Newton also heard of a carcass supposedly found in Trinity Bay on Newfoundland’s east coast in 1853. His attempts to obtain further information were unsuccessful, however.

Another story from West Greenland became known in 1869. The American polar explorer Isaac Israel Hayes learned there that a great auk had been caught on one of the Walvis Islands two years earlier. The Greenlandic hunter did not recognize the bird as a special rarity and, as Hayes wrote, “immediately set about eating it.” (Shuker, 2019). Because neither the bird nor a contemporary description survives, the report cannot be verified.

Further reports also came from the British Isles. In 1880, the Reverend G. C. Green wrote about an elderly boatman on the Scottish island of Arran. The man described a black-and-white bird the size of a goose or turkey, with very short wings, and identified the great auk in an illustration. He even claimed to have shot three such birds. After his death, however, his sons could show Green only a common loon, which they believed to be the sought-after “Arran auk” (Shuker, 2019).

Two further stories from East Greenland are difficult to assess. The Danish ornithologist Otto Helms recorded them in his 1926 survey of the birds of Angmagssalik. In 1884/85, Gustav Holm heard of a man whose grandfather had once caught a very large, flightless bird with small wings. When this supposedly happened remained unclear. In 1896, Johan Petersen recorded a more recent report: about six years earlier, a Greenlander was said to have pursued a common-loon-sized bird in Angmagssalik Fjord that could not fly and escaped in rough seas. Apart from its size and flightlessness, however, no distinguishing features were described.

Reports from the 20th century

Supposed great auks continued to be reported even in the 20th century. In 1971, the nature writer J. Wentworth Day recounted a sighting made by his acquaintance Edward Valpy in 1927. Valpy saw a large, flightless bird in the Lofoten Islands slide from a rock into the water. A local boatbuilder reportedly said that the animal had been known there for a long time (Shuker, 2019).

In 1929, the naturalist H. A. A. Dombrain also reported an unknown bird beneath a quay in the Lofoten Islands. Two men who saw it independently selected the great auk as the closest match from several pictures (Fuller, 1999; Shuker, 2019). Both reports became known only later, however, and provided no verifiable evidence.

Further reports from the Lofoten Islands in the late 1930s and the 1940s may have resulted from an unusual introduction experiment. In 1936, nine king penguins arrived in Norway aboard the SS Neptune. The birds had been provided by Lars Christensen, an entrepreneur involved in Antarctic whaling; the initiative was organized by the Norwegian Society for the Conservation of Nature. Two pairs were released near Røst in the Lofoten Islands, and two more pairs and a juvenile near Gjesvær in Finnmark. In subsequent years, macaroni and African penguins were also released near Røst. Penguins were occasionally seen along the northern Norwegian coast until 1949. The released birds cannot explain the reports from 1927 and 1929, but they may account for some of the later alleged great auk sightings (Norwegian Polar Institute, 2011).

The extent to which the great auk had become the subject of legends and hopes was still evident in the mid-1980s. Members of a Scottish yacht club traveled to Papa Westray to investigate alleged sightings. The search, known as “Operation Aukfinder,” turned out to be a promotion for a whisky brand (Shuker, 2019).

When did the great auk actually die out?

None of the reports after 1844 produced a photograph, a positively identifiable carcass, bones, or other verifiable evidence. Many reports were recorded years or decades later, were based on oral stories, or contained few unique features. Confusion with common loons, guillemots, and later even released penguins was possible.

It is nonetheless plausible that individual great auks survived in other parts of the North Atlantic after 1844. The death of the last two known breeding birds did not necessarily mean that every other individual disappeared at the same moment. Survival until the end of the 19th century or even into the 20th, however, would have required a small breeding population to remain undiscovered for decades without leaving any clear trace.

The two birds killed on Eldey in 1844 therefore remain the last confirmed record. Drummond-Hay’s 1852 sighting is regarded as the last credible but unconfirmed indication of a surviving great auk.

Were there different forms of the great auk?

The great auk was once widespread across large parts of the North Atlantic. Since bone finds from different regions show certain differences in size, it has been repeatedly suggested that geographically separate populations or even subspecies may have existed.

Anne Karin Hufthammer conducted one of the first more extensive studies in her unpublished 1982 thesis. Comparing Scandinavian bones, she initially suggested that the birds may have become smaller from south to north.

Gary P. Burness and William A. Montevecchi reached a different conclusion in 1992. The great auks from Funk Island that they studied were larger on average than birds from boreal regions, including Scandinavian sites. They could not, however, confirm a consistent north-south division within Scandinavia. They therefore interpreted the differences not as evidence of western and eastern Atlantic subspecies but as probable oceanographically driven variation in size. The cold, low-Arctic marine environment around Funk Island may have favored larger bodies through its temperatures and feeding conditions.

Differences in average body size are not sufficient to distinguish subspecies. Environmental conditions, food availability, age, sex, and individual variation can also affect an animal’s dimensions.

A more extensive study by Anne Karin and Karl Ove Hufthammer confirmed in 2023 that variation in size cannot be reduced to a simple geographic rule. The researchers assessed 477 great auk bones from 53 Norwegian sites and 55 time periods. Most of the finds were between around 6,000 and 2,000 years old. Body size showed no consistent change over time or along a north-south or east-west gradient. Differences were generally weak and discernible only in individual bone measurements.

In her 2018 study of humeri from different populations, Jessica Thomas likewise found no significant population-specific differences in size. The bones therefore could not be reliably assigned to a western or eastern form.

Genetics provides additional evidence. In 2019, Thomas et al. examined the complete mitochondrial genomes of 41 great auks from different parts of the former range, including Funk Island, Iceland, Great Britain, Scandinavia, the Netherlands, and Spain. The genetic lineages showed no clear geographic division. Instead, the researchers found high mitochondrial diversity and evidence of long-term exchange between regions of the North Atlantic.

Great auks may have varied in size from region to region. Birds from Funk Island in particular appear to have been larger on average than many specimens from boreal regions. Such differences, however, can be explained by environmental conditions and natural variation within a species. Neither bone measurements nor the mitochondrial DNA examined to date provide convincing evidence of permanently separate populations or subspecies. The great auk is therefore classified as a species with no recognized subspecies.

Great auks in human care

Individual great auks were kept alive temporarily. Although no specimen is known to have been exhibited in a modern public zoo, some captured birds were kept privately for weeks or months. In the 17th century, a great auk was probably even held in the Royal Menagerie at Versailles. Scientific interest, courtly curiosity, and later the increasing value of the rare birds mattered more than any attempt to care for them permanently or breed them.

Ole Worm and the riddle of the white neck ring

The earliest well-documented case concerns the Danish physician and naturalist Ole Worm. In the 17th century, he received a young live great auk from the Faroe Islands and kept it in Copenhagen for several months. According to Henry Wemyss Feilden (1872), Worm cared for the bird personally, presumably in connection with his private natural-history collection, which also included two mounted great auks. Worm’s bird was therefore one of the few individuals that could be observed alive over an extended period.

The Faroese chronicler Lucas Debes also apparently knew the great auk from personal experience in the 17th century. The account by Fisher and Lockley (1954) does not make clear, however, whether he kept a bird himself or merely examined a captured one.

An illustration in Ole Worm’s posthumously published collection catalog, the 1655 Museum Wormianum, probably depicts the bird that was kept alive. Worm had sent the manuscript to the Elzevier printing house himself; after his death in 1654, his son Villum Worm oversaw its publication. It is not known who drew the great auk or prepared the image for printing.

The white neck ring, which was not part of the great auk’s natural plumage, makes the image particularly puzzling. Citing Errol Fuller’s The Great Auk (1999), Petra Tjitske Kalshoven (2018) adopts the interpretation that Worm’s bird wore a collar. The illustrator may have rendered it as a white band of plumage, which later artists then copied as a supposed natural feature.

In 2016, Arturo Valledor de Lozoya et al. proposed a second possibility. In Museum Wormianum, the great auk was mistakenly called Anser Magellanicus seu Pinguinus, associating it with the Magellanic penguin (Spheniscus magellanicus). Carolus Clusius had previously described a corresponding southern bird with white feathers on its neck. This may explain why the feature was added or emphasized when Worm’s illustration was made.

Clusius also knew the actual great auk, however. In his work Exoticorum libri decem, published in 1605, he had it depicted under the name Mergus americanus. This earlier woodcut shows no white neck ring. Clusius probably received the source image from the French collector Jacques Plateau.

It is no longer possible to determine today whether Worm’s illustration actually shows a collar or whether the white ring was the result of confusion with a southern penguin. It also remains unknown whether the feature was already included in a template created during Worm’s lifetime or was only added when the book was being prepared for printing.

More early depictions

For a long time, Worm’s illustration was considered the only known depiction probably based on a living great auk. Another possible example is Daniel King’s drawing from 1652. It shows a large black-and-white bird on a rocky outcrop on the Calf of Man. Kenneth Williamson identified it as a great auk in 1939. Fisher and Lockley (1954) called the image the earliest British depiction of the species, but emphasized the lack of contemporary written evidence for great auks on the Calf of Man. It is therefore unknown whether King actually observed a living great auk there.

Worm’s image nevertheless has a special status. It is the earliest depiction that can be linked directly to a great auk known to have been kept alive, because Worm himself reported on the bird in his possession. Clusius’s older woodcut, by contrast, was based on an image sent to him, while it is uncertain whether Daniel King used a living bird as his model.

A great auk in Versailles?

Another great auk was probably kept in the Royal Menagerie at Versailles between 1666 and 1670. Valledor de Lozoya et al. (2016) examined two color illustrations and an engraving attributed to Nicolas Robert. Robert worked for Louis XIV, depicting plants and animals from the royal collections.

One of the illustrations belongs to the so-called Vélins du Roi, the royal collection of natural-history images. Another appears in an album of birds from the Versailles menagerie and the collection of Gaston d’Orléans. An engraving from a collection of rare birds at the Versailles menagerie shows great auks in several different postures. This need not mean that several birds were kept at the same time; the same individual may have been depicted in different positions.

No written documents are known concerning the origin, arrival, or keeping of a great auk at Versailles. Given the context in which the images were created and their independent compositions, however, Valledor de Lozoya and his co-authors consider it probable that Robert was able to observe a living great auk. The menagerie was not a public zoo but a courtly animal collection intended for display and entertainment, while also permitting rare animals to be observed and visually documented.

The following depiction, probably created between 1666 and 1670, is attributed to Nicolas Robert:

The depiction shows a great auk swimming in a natural environment. In the background a smaller bird appears standing, another diving. The large furrowed beak, the white spot in front of the eye and the severely shortened wings are clearly reproduced. Unlike Worm’s great auk, the animal does not wear an unnatural white neck ring.

The single picture does not prove that a great auk actually lived in Versailles. However, together with the other depictions attributed to Robert and their close connection to the royal menagerie, it provides strong evidence that a live bird was kept and portrayed there in the 1660s.

Captive great auks in the 19th century

In 1821, a great auk was captured on the remote Scottish island of St Kilda. The naturalist John Fleming was given the opportunity to examine the living bird. According to Fuller (2000), Fleming tried to feed it fresh fish and fatten it up. He also allowed it to swim in the sea, tying a cord to one foot to prevent its escape. This restraint proved inadequate: the great auk escaped during one of its daily swims. Bengtson (1984) also cites Worm’s and Fleming’s birds as examples of great auks occasionally being kept alive.

Alfred Newton (1861) reported another case from Iceland. A total of 24 great auks were captured during an expedition to Eldey in 1831. One reached Keflavík alive but died or was killed there a short time later. Apparently, there had been no plan to keep it alive for long.

Better documented is a female great auk captured off County Waterford, Ireland, in May 1834. The young, weakened bird first came into the possession of Francis Davis and then of Jacob Goff of Horetown. According to Waterford Treasures, it survived for approximately four months in human care. It was reportedly fed potatoes, milk, and trout (Fisher & Lockley, 1954). The bird is considered the last confirmed great auk in Ireland.

Unlike those of most other great auks kept alive, this bird’s subsequent fate is known. After its death, the physician and ornithologist Robert Burkitt prepared the body. Around ten years later, the specimen was given to the Zoological Museum of Trinity College Dublin, where it remains today.

Not every instance of captivity arose from scientific curiosity. Fuller (2000) reports a great auk found sleeping on the rock of Stac an Armin near St Kilda in July 1840. The islanders took it to a hut and held it there for three days. During a storm, they began to fear that the bird might be a witch, and killed it with stones. The incident is considered the last reliable record of the great auk in Great Britain.

No great auk in a modern zoo

Although the first modern zoological gardens existed when the last great auks were still alive, no specimen is known to have been kept permanently in such a facility. Most of the animals captured were killed immediately, preserved as specimens and sold to museums, dealers or private collectors. There was neither a coordinated conservation attempt nor any effort to establish a conservation-breeding program.

In this context, Alfred Newton’s final words from 1861 are revealing. At that time, he still considered it possible that individual great auks had survived on a rock near Iceland that was difficult to access. At the same time, he recognized that their rediscovery would probably also mean their final end:

“Whether the great auk is already extinct or still exists in an unknown location, it is clear that its extinction, if it has not already occurred, must immediately follow its rediscovery. I must therefore implore all those connected with this matter to do their utmost to ensure that such a rediscovery is put to the best possible use. (…) Surely no more suitable place could be found to house the very last great auks than the gardens of the Zoological Society of London (…).”

No such arrangement ever came about. The few great auks verifiably kept alive by people remained isolated cases. None became part of a breeding attempt, and none could help prevent the species’ disappearance.

The great auk in museums and collections

As it became rarer, the great auk turned into a coveted collector’s item in the 19th century. Museums and private individuals sought skins, skeletons, and eggs; the last birds were deliberately captured so they could be mounted and sold. Collecting contributed to the species’ final disappearance, but it also ensured that an unusually large number of remains survived. Compared with the spectacled cormorant, which became extinct at about the same time, the great auk is now documented by numerous specimens, eggs, and bones.

The high market value also encouraged forgeries and incorrectly assembled specimens. Akimushkin (1972) gives the example of a supposed great auk in the Darmstadt State Museum. Closer examination revealed that only the skull came from a great auk, while the plumage had been taken from flying auks. A similar specimen in Shrewsbury Museum is said to have been assembled from parts of several bird species.

Identification of the last great auk pair

The remains of the last reliably documented pair of great auks are especially important. After the two birds were killed on Eldey in June 1844, a pharmacist had them skinned in Reykjavík and sent the skins and preserved organs to Denmark. Their hearts, eyes, and esophagi remain in the Natural History Museum of Denmark in Copenhagen. The skins, however, were resold and changed hands several times. For more than 170 years, no one knew which collections they had entered. Fuller (1999) called the search for them an “ornithological mystery”.

In 2017, Jessica E. Thomas and her team compared the mitochondrial genomes of the preserved organs with five specimens considered possible candidates on the basis of their provenance. The male’s DNA matched that of a mounted great auk in the Royal Belgian Institute of Natural Sciences in Brussels. This identified the skin of the last reliably documented male. None of the specimens examined at the time, however, matched the female.

Historical documents later showed that the provenances of two specimens from the former collection of the English ornithologist George Dawson Rowley had apparently been confused. Both birds were acquired in 1934 by the ornithologist and collector Vivian Hewitt and, after his collection was dispersed, went to Los Angeles and Cincinnati in 1974. Because photographs and sales documents were initially assigned incorrectly, the research team examined the Los Angeles specimen in 2017. Newly evaluated auction catalogs, letters, and ownership records, however, indicated that the female’s sought-after skin was in Cincinnati.

This hypothesis was confirmed by a study published in 2025 by Emily L. Cavill, Jessica E. Thomas, and other researchers. A small tissue sample was taken from the underside of one foot of the specimen at the Cincinnati Museum Center’s Museum of Natural History & Science. The mitochondrial genome reconstructed from it matched the DNA from the female’s heart preserved in Copenhagen exactly. The sequence differed from those of the other great auks used for comparison at no fewer than eleven base positions.

Mitochondrial DNA does not always distinguish individual animals unambiguously, because close relatives can share the same haplotype. Mitochondrial diversity in the great auk was extremely high, however, and none of the other published mitochondrial genomes matched the female’s heart. Together with the reconstructed ownership history, this indicates that the Cincinnati specimen is indeed the skin of the female killed on Eldey in 1844 (Cavill et al., 2025).

The specimens in Brussels and Cincinnati are now considered to be the skins of the last reliably observed pair of great auks. The associated organs continue to be stored in Copenhagen.

Remains preserved and lost

Standard inventories today list 78 preserved skins or mounted specimens, 75 eggs, and 24 complete skeletons. There are also numerous individual bones, skulls, and subfossil remains. The exact total can vary slightly depending on the counting method and the condition of individual objects; more recent publications therefore also refer to around 80 preserved specimens. Almost every known item is held in a museum or scientific collection, with only a few remaining in private ownership (Fuller, 1999; Luther, 1986).

Several specimens were lost in the 20th century. A skin at the natural history museum in Mainz was destroyed during the Second World War. Another specimen burned in a 1978 fire at the Museu Bocage in Lisbon. A specimen formerly held in Szczecin has been considered lost since the Second World War. A complete skeleton at the Zoological Institute in Kiel was also destroyed in 1944 or 1945.

Almost all of the preserved skins show the great auk in summer plumage. Specimens in winter or juvenile plumage, on the other hand, are extremely rare and were not always classified consistently in older inventories.

The juvenile plumage is also known only incompletely. Some specimens of birds that were not yet fully grown survive, including in Newcastle upon Tyne and Prague, but no definite chick specimen has been preserved. According to historical descriptions, newly hatched chicks initially had dark gray down (Luther, 1986).

The following overview presents a selection of well-known great auk specimens, eggs, and skeletons in Germany, elsewhere in Europe, North America, and Australia (after Fuller, 1999; Luther, 1986).

List of Great Auk museum holdings (selection)

Germany

• Berlin — Museum für Naturkunde (MfN): 1 mounted specimen in summer plumage (probably collected off the coast of Reykjanes in the summer of 1830)
• Bonn — Museum Koenig (ZFMK): 1 mounted specimen in summer plumage, 1 complete skeleton, and 3 eggs (damaged, probably from Newfoundland)
• Braunschweig — Staatliches Naturhistorisches Museum (SNHM): 1 mounted female specimen in summer plumage, 1 incomplete skull, other bones, and 3 eggs
• Bremen — Übersee-Museum (ÜMB): 1 mounted specimen in summer plumage (probably from Eldey, collected in 1840 or 1841)
• Darmstadt — Hessisches Landesmuseum (HLMD): 1 skull
• Dresden — Museum für Tierkunde (MTD): 1 mounted female specimen in summer plumage, 1 skeleton or several bones, and 1 egg
• Düsseldorf — Aquazoo Löbbecke Museum: 1 egg (severely damaged)
• Erlangen — Zoologische Sammlung der FAU (ZSFAU): 1 egg
• Frankfurt am Main — Senckenberg Natural History Museum (SMF): 1 mounted specimen in summer plumage and 1 complete skeleton
• Gotha — Museum der Natur (MNG): 1 mounted specimen in summer plumage
• Göttingen — Zoological Museum (ZMG): 1 skeleton, probably from Funk Island
• Hanover — Lower Saxony State Museum (NLMH): 1 mounted specimen in summer plumage, collected in Iceland in 1830
• Kiel — Zoological Museum (ZMK): 1 mounted specimen in summer plumage (probably from Iceland)
• Köthen — Naumann Museum (NMK): 1 mounted specimen in summer plumage
• Leipzig — Zoological Teaching and Study Collection (ZLSL): 1 mounted specimen in summer plumage
• Munich — Museum Mensch und Natur (MMN): 2 mounted specimens in summer plumage (at least 1 male)
• Nuremberg — Natural History Museum Nuremberg (NHMN): 1 egg
• Oldenburg — State Museum of Nature and Man (LNM): 1 mounted specimen in summer plumage (from Iceland, 1840/41) and 1 egg
• Stuttgart — State Museum of Natural History (SMNS): 1 mounted specimen in summer plumage (from Iceland, purchased in 1867)
• Wittenberg — Museum of Natural History and Ethnology “Julius Riemer” (closed since 2011): 1 skeleton (assembled from the bones of different individuals)

Europe

• Abbeville, France — Le Beffroi — Musée Boucher-de-Perthes-Manessier: 1 mounted specimen in summer plumage
• Aberdeen, Scotland — University of Aberdeen Zoology Museum (ABDUZ): 1 egg
• Amiens, France — Musée de Picardie: 1 mounted specimen in summer plumage
• Angers, France — Muséum des Sciences Naturelles d’Angers (MSN): 1 egg (severely damaged)
• Autun, France — Muséum d’histoire naturelle Jacques de La Comble (MHNA): 1 mounted specimen in summer plumage
• Birmingham, England — Birmingham Museum and Art Gallery (BMAG): 1 mounted specimen in summer plumage
• Wrocław, Poland — Muzeum Przyrodnicze Uniwersytetu Wrocławskiego (MPUWr): 2 mounted specimens in summer plumage
• Bristol, England — Bristol Museum & Art Gallery (BRSMG): 2 eggs
• Brussels, Belgium — Royal Belgian Institute of Natural Sciences (RBINS): 1 mounted male specimen in summer plumage—one of the last two known great auks, collected in 1844 (together with the specimen in Cincinnati)
• Cambridge, England — University Museum of Zoology (UMZC): 1 skeleton, 1 mounted specimen in summer plumage, and 8 eggs
• Cardiff, Wales — Amgueddfa Cymru — National Museum Wales (NMW): 1 mounted specimen in summer plumage and 1 egg
• The Hague, Netherlands — Museon (MUS): 1 mounted specimen in summer plumage
• Dublin, Ireland — Trinity College Zoological Museum (TCZM): 1 mounted specimen in molt—the bird captured off Waterford in 1834
• Edinburgh, Scotland — National Museum of Scotland (NMS): 1 mounted specimen in summer plumage (acquired in 1895), 1 skeleton, and 10 eggs
• Florence, Italy — Museo di Storia Naturale dell’Università di Firenze — La Specola (MZUF): 1 mounted specimen in summer plumage
• Glasgow, Scotland — Kelvingrove Museum (KAGM): 1 mounted specimen in summer plumage
• Graz, Austria — Universalmuseum Johanneum (UMJ): 1 mounted specimen in summer plumage
• Gothenburg, Sweden — Naturhistoriska Museum (NHMG): 1 skeleton
• Helsinki, Finland — Finnish Museum of Natural History, University of Helsinki (FMNH): 1 egg
• Inverness, Scotland — Inverness Museum & Art Gallery (IMAG): 2 eggs
• Copenhagen, Denmark — Statens Naturhistoriske Museum (SNM): 2 mounted specimens (one in summer plumage and one in winter plumage), the eyes, hearts, and esophagi of the last two known great auks from 1844, preserved in ethanol, and 1 egg
• Lausanne, Switzerland — Naturéum — Département de Zoologie (NDZL): 1 mounted specimen in summer plumage
• Le Havre, France — Muséum d’histoire naturelle du Havre (MHNH): 1 mounted specimen in summer plumage
• Leiden, Netherlands — Naturalis Biodiversity Center (NBC): 2 mounted specimens in summer plumage, 2 eggs, 1 feather, 1 skeleton, and various bones (1 specimen and 1 egg were previously held at the Zoölogisch Museum Amsterdam)
• Lille, France — Musée d’histoire naturelle de Lille (MHNL, Lille): 1 mounted specimen in summer plumage
• Liverpool, England — World Museum, National Museums Liverpool (NML-VZ): 1 egg formerly owned by the 13th Earl of Derby, whose collection passed to Liverpool after his death in 1851
• London, England — Hunterian Museum (HM): 1 skeleton
• London, England — Natural History Museum (BMNH): several mounted specimens or skins
• Lund, Sweden — Biological Museum, Lund University (MZLU): 1 mounted specimen in summer plumage
• Moscow, Russia — State Darwin Museum (SDM): 1 mounted specimen in summer plumage (formerly owned by Wilhelm Blasius, director of the Natural History Museum in Braunschweig)
• Nantes, France — Muséum d’histoire naturelle de Nantes (MHNNT): 1 mounted specimen in summer plumage
• Neuchâtel, Switzerland — Muséum d’histoire naturelle de Neuchâtel (MHNN): 1 mounted specimen in summer plumage
• Newcastle upon Tyne, England — Great North Museum: Hancock (NEWHM): 2 mounted specimens, 1 in summer plumage and 1 in juvenile plumage (old summer plumage or winter plumage), and 1 egg
• Norwich, England — Norwich Castle Museum & Art Gallery (NWHCM): 1 mounted specimen in summer plumage, possibly collected on Geirfuglasker before 1822, and 2 eggs
• Oslo, Norway — Natural History Museum, University of Oslo (NHMO) — 1 mounted specimen in summer plumage
• Oxford, England — Oxford University Museum of Natural History (OUMNH): 1 egg
• Paris — Muséum national d’Histoire naturelle (MNHN): 1 mounted specimen in summer plumage, several preserved skins, 1 skeleton, and 3 eggs
• Pisa, Italy — Museo di Storia Naturale (MSNP): 1 mounted specimen in summer plumage
• Prague, Czechia — National Museum (NM): 1 mounted juvenile specimen in winter plumage and 1 mounted specimen in summer plumage
• Reykjavík, Iceland — Icelandic Institute of Natural History (IINH), Garðabær, Iceland: 1 mounted specimen in summer plumage (purchased for Iceland at Sotheby’s in 1971) and 1 egg
• Rome, Italy — Museo Civico di Zoologia (MCZR): 1 mounted specimen in summer plumage
• Rouen, France — Muséum d’histoire naturelle de Rouen (MHNR): 1 mounted specimen in summer plumage
• Scarborough, England — Scarborough Museums and Galleries (SCARB): 1 egg
• Shrewsbury, England — Shropshire Museums (SM): 1 mounted specimen in summer plumage (probably made from a combination of skins from other seabirds)
• Spalding, England — Spalding Gentlemen’s Society Museum (SGS): 1 egg
• Stockholm, Sweden — Naturhistoriska riksmuseet (NRM): 1 mounted specimen in summer plumage
• Strasbourg, France — Zoological Museum, Johann Hermann Collection (ZMUS): 1 mounted specimen in summer plumage
• St. Petersburg, Russia — Zoological Museum of the Zoological Institute of the Russian Academy of Sciences (ZIN RAS): 1 mounted specimen in summer plumage
• Tring, England — Natural History Museum (NHMUK): 4 specimens in summer plumage (some possibly study skins rather than mounts), 1 skeleton, and 6 eggs (including former Tomkinson egg no. 53)
• Turin, Italy — Museo Regionale di Scienze Naturali (MRSN; MZUT collection): 1 mounted specimen in summer plumage
• Vienna, Austria — Natural History Museum Vienna (NHMW): 1 mounted specimen in summer plumage
• Worksop, England — Osberton Hall near Worksop, Nottinghamshire, Foljambe private collection: 1 mounted specimen in summer plumage and possibly 1 egg
• York, England — Yorkshire Museum (YORYM): 2 mounted specimens in summer plumage

USA, Canada & Australia

• Andover, Massachusetts, USA — Phillips Academy — Peabody Institute of Archaeology: 1 mounted specimen in summer plumage
• Cambridge, Massachusetts, USA — Museum of Comparative Zoology, Harvard University (MCZ): 2 mounted specimens in summer plumage, 1 skeleton, and 10 eggs
• Cincinnati, Ohio, USA — Cincinnati Museum Center (CMC): 1 mounted female specimen—one of the last two known great auks, collected in 1844 (together with the specimen in Brussels)—and 1 egg
• Los Angeles, California, USA — Natural History Museum of Los Angeles County (NHMLAC): at least 1 mounted specimen in summer plumage
• New Haven, Connecticut, USA — Yale Peabody Museum (YPM): 1 skeleton
• New York, USA — American Museum of Natural History (AMNH): 3 mounted specimens or skins in summer plumage, 1 skeleton, and 1 egg
• Ottawa, Canada — Canadian Museum of Nature (CMN): 1 skeleton
• Toronto, Canada — Royal Ontario Museum (ROM): 1 mounted specimen in summer plumage, collected on Eldey—the only surviving great auk skin in Canada—and 1 skeleton
• Philadelphia, Pennsylvania, USA — The Academy of Natural Sciences of Drexel University (ANSP): 1 mounted specimen in summer plumage, 1 skeleton, 1 skull, and 1 egg
• St. John’s, Newfoundland — The Rooms Provincial Museum: 1 skeleton
• Sydney, New South Wales, Australia — Australian Museum (AM): 1 skeleton
• Victoria, British Columbia, Canada — Royal BC Museum (RBCM): 1 skeleton
• Washington, D.C., USA — Smithsonian National Museum of Natural History (USNM): 1 mounted specimen in summer plumage, 1 skeleton, and 1 egg

Can the Great Auk be brought back?

Strictly speaking, the great auk cannot be “cloned.” The transfer of a cell nucleus used to clone mammals cannot be applied in the same way because of birds’ particular reproductive biology. Moreover, no living great auk cells have survived, only DNA from museum specimens in varying states of degradation.

The organization Revive & Restore has presented the great auk as a possible candidate for so-called de-extinction. As early as 2015, 22 experts discussed its basic feasibility. In the Ocean Genomics Horizon Scan, published in 2019, the organization then outlined a possible research pathway. The principal living species under consideration is the razorbill, the great auk’s closest living relative. Its genome would have to be compared with the most complete possible reconstruction of the great auk genome and then modified in a targeted manner. The aim would be to produce offspring with key traits of the extinct species—such as its body size, shortened wings, and inability to fly.

The greatest difficulty lies in passing such genetic changes to the next generation. The proposed method would involve modifying primordial germ cells, which later develop into eggs and sperm. These cells could be transferred into the embryo of another bird species. The resulting bird would outwardly belong to the recipient species, but it would carry the modified germ cells in its reproductive organs and could produce corresponding offspring.

Revive & Restore proposed developing the necessary procedures first in the more easily kept buttonquails of the genus Turnix, using sterile domestic chickens as possible recipients. Only later would corresponding methods be applied to razorbill germ cells. The organization itself described the cultivation and transfer of germ cells from wild bird species as one of the greatest technical hurdles. The concept therefore describes not an imminent reintroduction project but a multistage research program whose implementation still faces numerous unresolved problems (Novak, 2019).

Even if the project succeeded, it would not create a genetically identical great auk. The result would instead be a modified descendant of the razorbill possessing as many physical and perhaps ecological traits of the extinct bird as possible. Revive & Restore therefore calls such animals ecological surrogates, or “proxies,” for the extinct species. Genetic modification alone could not guarantee that they would develop the behavior, reproduction, and way of life of great auks.

There are also practical questions: the animals would have to be bred in sufficient numbers, prepared for life in the open sea and settled in suitable, permanently protected breeding places. The ability to find food, choose mates and raise young birds is not determined exclusively by individual genes. An externally similar bird would therefore not necessarily be a complete replacement for the lost species.

The great auk cannot truly be brought back. Perhaps one day a bird could be created that resembles it in appearance and ecology—but it could not replace the irretrievably lost original. The fate of the great auk is therefore a reminder that preserving a species is incomparably easier than trying to recreate it after extinction.

---

Sources

• Akimushkin, I. (1972). Threatened with extinction? Animal tragedies caused by humans. F.A. Brockhaus.
• Bengtson, S.-A. (1984). Breeding Ecology and Extinction of the Great Auk (Pinguinus impennis): Anecdotal Evidence and Conjectures. The Auk, 101(1). 1–12. https://doi.org/10.1093/auk/101.1.1
• BirdLife International. 2021. Pinguinus impennis. The IUCN Red List of Threatened Species 2021: e.T22694856A205919631. https://dx.doi.org/10.2305/IUCN.UK.2021-3.RLTS.T22694856A205919631.en
• Birkhead, T., Russel, D., Garbout, A., et al. (2020). New insights from old eggs—the shape and thickness of Great Auk Pinguinus impennis eggs. Ibis, 162(4). 1345–1354. https://doi.org/10.1111/ibi.12820
• Birkhead, T. R. (2021): The chick-rearing period of the Great Auk: a mystery solved. British Birds, 114(1), pp. 27–30.
  https://britishbirds.co.uk/journal/article/chick-rearing-period-great-auk-mystery-solved
• Birkhead, T. R., & Montgomerie, R. (2026). A Neolithic rock engraving apparently showing a Great Auk being captured. Ibis, 168(1). 377–382. https://doi.org/10.1111/ibi.13452
• Boran, E. (n.d.). Clusius on penguins. Birds at the Edward Worth Library. https://birds.edwardworthlibrary.ie/flightless-birds/clusius-on-penguins/
• Brodkorb, P. (1960). Great auk and common murre from a Florida midden. The Auk, 77. 342–343. https://doi.org/10.2307/4082490
• Burness, G. P., & Montevecchi, W. A. (1992). Oceanographic-related variation in the bone sizes of extinct Great Auks. Polar Biology, 11. 545–551. https://doi.org/10.1007/BF00237947
• Campmas, E., Laroulandie, V., Michel, P., et al. (2010). A great auk (Pinguinus impennis) in North Africa: Discovery of a bone remain in a Neolithic layer of El Harhoura 2 Cave (Temara, Morocco). In W. Prummel, J. T. Zeiler, & D. C. Brinkhuizen (ed.), Birds in archaeology: Proceedings of the 6th Meeting of the ICAZ Bird Working Group in Groningen, 23–27 August 2008 (Groningen Archaeological Studies, Vol. 12, pp. 233–240). Barkhuis Publishing & Groningen University Library.
• Cartwright, G. (1792). A journal of transactions and events during a residence of nearly sixteen years on the coast of Labrador (Vol. 3). Allin and Ridge.
• Cavill, E.L., Hernández Alonso, G., Imhoff, E., et al. (2025). The hunt for the last great auks: Ancient DNA resolves a 180-year-old mystery. Zoological Journal of the Linnean Society, 205(1), zlaf097. https://doi.org/10.1093/zoolinnean/zlaf097
• Cokinos, C. (2000). Hope is the thing with feathers: A personal chronicle of vanished birds. New York: J.P. Tarcher & Putnam.
• Day, D. (1981). The doomsday book of animals: A natural history of vanished species. Viking Press.
• De Lozoya, A. V., García, D. G., Parish, J. (2016). A great auk for the Sun King. Archives of Natural History, 43(1). 41–56. https://doi.org/10.3366/anh.2016.0345
• Feilden, H.W. (1872). The birds of The Faroe Islands. The Zoologist, 2(7).
• Fisher, J., & Lockley, R. M. (1954). Sea-Birds: An introduction to the natural history of the sea-birds of the North Atlantic. The Riverside Press.
• Fuller, E. (1999). The Great Auk.
• Fuller, E. (2000). Extinct birds. Oxford University Press.
• Helms, O. (1926). The birds of Angmagssalik. Meddelelser om Grønland, 58, 207–274.
• Hobson, K.A., & Montevecchi, W.A. (1991). Stable isotope determination of the trophic status of the Great Auk. Oecologia, 87. 528–531. https://doi.org/10.1007/bf00320416
• Hufthammer, A. K., & Hufthammer, K. O. (2023). The Great Auk in Norway. From common to locally extinct. International Journal of Osteoarchaeology, 33. https://doi.org/10.1002/oa.3161
• Hume, J.P. (2017). Extinct birds. Bloomsbury Natural History.
• Kalshoven, P. T. (2018). Piecing Together the Extinct Great Auk: Techniques and Charms of Contiguity. Environmental Humanities, 10(1). 150–170. https://doi.org/10.1215/22011919-4385507
• Liberman, A. (2011, June 8). Penguin. OUPblog. https://blog.oup.com/2011/06/penguin/
• Linnaeus, C. (1758). Systema naturae per regna tria naturae (10th ed., Vol. 1). Laurentii Salvii.
• Livezey, B.C. (1988). Morphometrics of flightlessness in the Alcidae. The Auk, 105. 681–698. https://doi.org/10.1093/auk/105.4.681
• Luther, D. (1986). The extinct birds of the world. A. Ziemsen Verlag.
• Montevecchi, W. A., & Kirk, D. A. (2020). Great Auk (Pinguinus impennis), version 1.0. In Birds of the World (A.F. Poole and F.B. Gill, Editors). Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi.org/10.2173/bow.greauk.01
• Montgomerie, R.D., & Birkhead, T.R. (2025). Estimating the mass of the great auk (Pinguinus impennis) and its egg. Ibis, 167(1). 237–247. https://doi.org/10.1111/ibi.13350
• Moum, T., Arnason, U., & Arnason, E. (2002). Mitochondrial DNA Sequence Evolution and Phylogeny of the Atlantic Alcidae, Including the Extinct Great Auk (Pinguinus impennis). Molecular Biology and Evolution, 19(9). 1434–1439. https://doi.org/10.1093/oxfordjournals.molbev.a004206
• Newton, A. (1861). Abstract of Mr. J. Wolley’s Researches in Iceland respecting the Gare-fowl or Great Auk (Alca impennis, Linn.). Ibis, 2. 374–399.
• Novak, B. (2019). De-extinction of the great auk. Revive & Restore. https://ocean.reviverestore.org/big-ideas8/de-extinction/
• Norwegian Polar Institute (2011, December 5): Did you know that penguins used to live in Lofoten? South Pole 1911–2011. https://sorpolen2011.npolar.no/en/did-you-know/2011-12-05-penguins-used-to-live-in-lofoten.html
• Olson, S.L. (1977). A great auk, Pinguinus, from the Pliocene of North Carolina (Aves: Alcidae). Proceedings of the Biological Society of Washington, 90(3), 690–697.
• Olson, S.L. (2003). A fossil of the Great Auk (Pinguinus impennis) from Middle Pleistocene deposits on Bermuda. Atlantic Seabirds, 5(2), 81–84.
• Roberts, M. B., Parfitt, S. A., Pope, M. I., Wenban-Smith, F. F., Macphail, R. I., Locker, A., & Stewart, J. R. (1997). Boxgrove, West Sussex: Rescue excavations of a Lower Palaeolithic landsurface (Boxgrove Project B, 1989–91). Proceedings of the Prehistoric Society, 63, 303–358. https://doi.org/10.1017/S0079497X00002474
• Petersen, Æ. (1995). Some aspects of the history of the great auk in Iceland. Náttúrufræðingurinn, 65(1–2), 53–66.
• Shuker, K. (2019). Late Survival of the Lost Auk? Garefowl, Great Auks, and a Paradox of Penguins.
  https://karlshuker.blogspot.com/2019/03/late-survival-of-lost-auk-garefowl.html
• Silverberg, R. (1967). The auk, the dodo, and the oryx: Vanished and vanishing creatures. Thomas Y Crowell.
• Smith, N.A. (2011). Taxonomic revision and phylogenetic analysis of the flightless Mancallinae (Aves, Pan-Alcidae). ZooKeys, 91, 1–116. https://doi.org/10.3897/zookeys.91.709
• Smith, N.A., & Clarke, J.A. (2015). Systematics and evolution of the Pan-Alcidae (Aves, Charadriiformes). Journal of Avian Biology, 46(2), 125–140. https://doi.org/10.1111/jav.00487
• Smith, N.A. (2015). Evolution of body mass in the Pan-Alcidae (Aves, Charadriiformes): The effects of combining neontological and paleontological data. Paleobiology, 1(1). 1–19. https://doi.org/10.1017/pab.2015.24
• Thomas, J.E., Carvalho, G.R., Haile, J., et al. (2017). An “aukward” tale: A genetic approach to discover the whereabouts of the last great auks. Genes, 8(6), 164. https://doi.org/10.3390/genes8060164
• Thomas, J.E. (2018). Evolution & extinction of the great auk: A palaeogenomic approach [Doctoral dissertation, Bangor University and University of Copenhagen]. Bangor University Research Portal.
• Thomas, J.E., Carvalho, G.R., Haile, J., et al. (2019). Demographic reconstruction from ancient DNA supports rapid extinction of the great auk. eLife, 8, e47509. https://doi.org/10.7554/eLife.47509
• Waterford Treasures. (n.d.). Waterford and the last Great Auk of Ireland. https://waterfordtreasures.com/waterford-and-the-last-great-auk-of-ireland/