Great Auk-Pinguinus impennis

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Great Auk-Pinguinus impennis


Illustration of two great auks by John James Audubon (created in the 1800s), featured in his book The Birds of North America.

Temporal range: Middle Pleistocene (Chibanian) to Holocene (Meghalayan); ~400,000 years BP[1] to 1844-1852[2] CE

Scientific classification:

Life
Domain: Eukaryota
(unranked): Unikonta
(unranked): Opisthokonta
(unranked): Holozoa
(unranked): Filozoa
Kingdom: Animalia
Subkingdom: Eumetazoa
(unranked): Bilateria
Clade: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Clade: Olfactores
Clade: Craniata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Clade: Eugnathostomata
Clade: Teleostomi
Superclass: Tetrapoda
Clade: Reptiliomorpha
Clade: Amniota
Class: Reptilia or Clade: Sauropsida
Clade: Eureptilia
Clade: Romeriida
Clade: Diapsida
Clade: Neodiapsida
Clade: Archelosauria
Clade: Archosauromorpha
Clade: Archosauriformes
Clade: Crurotarsi
Clade: Archosauria
Clade: Avemetatarsalia
Clade: Ornithodira
Clade: Dinosauromorpha
Clade: Dinosauriformes
Clade: Dinosauria
Order: Saurischia
Clade: Eusaurischia
Clade: Theropoda
Clade: Neotheropoda
Clade: Averostra
Clade: Tetanurae
Clade: Orionides
Clade: Avetheropoda
Clade: Coelurosauria
Clade: Tyrannoraptora
Clade: Maniraptoriformes
Clade: Maniraptora
Clade: Pennaraptora
Clade: Paraves
Clade: Eumaniraptora
Clade: Averaptora
Clade: Avialae
Clade: Euavialae
Clade: Avebrevicauda
Clade: Pygostylia
Clade: Ornithothoraces
Clade: Euornithes
Clade: Orithuromorpha
Clade: Ornithurae
Clade: Neornithes
Infraclass: Neognathae
Clade: Neoaves
Clade: Aequorlitornithes
Order: Charadriiformes
Suborder: Lari
Clade: Pan-Alcidae?[3]
Family: Alcidae
Subfamily: Alcinae
Tribe: Alcini
Genus: †Pinguinus
Species: †P. impennis

The great auk is an extinct flightless shorebird (charadriiform) that lived in the North Atlantic from the Middle Pleistocene epoch (Chibanian)[1] to the mid-19th century.

Evolution:
The Atlantic auk assemblage can be split into two primary lineages: Uria (murres) and a clade containing razorbills (Alca; to which the great auk is most closely related), Alle (dovekies, or little auks), and Pinguinus (containing the great auk).[4]

The genus Pinguinus was present by the Pliocene epoch in the form of P. alfrednewtoni.[5] P. alfrednewtoni was probably not an ancestor to P. impennis, and was not only larger and more robust, but also even more specialized than the modern species in some respects. Pinguinus appears to have diverged into two lineages prior to the early Pliocene, possibly having eastern Atlantic and western Atlantic vicariants. When P. alfrednewtoni went extinct sometime after the early Pliocene, P. impennis appears to have replaced it.[6]

Fossil remains of P. impennis have been recovered from Chibanian (Middle Pleistocene)-aged deposits in Bermuda.[1] Hence, the modern species was certainly around by this point in time (~400,000 years ago[1]) and had established itself in the western Atlantic.

Description:

Great auk specimen (Bird No. 8; the Glasgow Auk) with a replica egg on exhibit in Kelvingrowe, Glasgow. © @ Mike Pennington->.

The great auk was somewhere around 78.6 cm long and weighed 5 kg. Its comparatively great size and small wings gave it a wing-loading comparable to those of medium-sized penguins (roughly 22 g*cm^-2). The bird’s flightlessness was evidently a result of extreme specialization for pursuit diving, just as with spheniscids.[7]

Plumage color depended on the season. The breeding (summer) plumage of this bird (as seen in the image above) is arguably the most recognizable plumage, and consists of a white patch over each eye. The non-breeding (winter?) plumage replaces the white patch with a white band, and a grey line from eye to ear. The summer moult had a chin and throat that was blackish-brown, and a bill with up to seven white grooves on the rostrum and twelve on the mandible. On the other hand, the winter moult had a white throat, and the bill had less white grooves.[8]

Generally, the head, neck, and back were glossy black; the underside was white; the feet and claws were black, the toe webbing was brownish-black; the inside of the mouth was yellow; and the iris was hazel or chestnut. Juveniles had necks that were mottled black and white, the white eye spot was absent, there was a grey line through the eye to below the ear, and the beak grooves were less prominent.[8]


Painting by John Gerrard Keulemans (made before 1903) of a great auk in summer (left) and winter (right) plumage.

The great auk was the most penguin-like of the auks. In fact, the term ‘penguin’ was originally applied to the great auk, before Southern Hemisphere Sphenisciformes were named after this bird.[8][9] Both this Northern Hemisphere alcid and the Southern Hemisphere Sphenisciformes were known as penguins until the former’s extinction.[9]

Diet:
Typical prey size for the great auk consisted of fish ranging from 70-190 mm SL (standard length, which is the total length of the fish minus the caudal fin), although fish as large as 240-320 mm SL are also known to have been preyed upon. The typical-sized prey included juvenile (1-2 year old) Brevoortia, Alosa, Mallotus, some Gadidae; subadults of Morone and Gadidae, and adult Gasterosteus. Almost all of these prey species could be found in the water column from near the bottom to near the surface. This suggests that great auks typically hunted in water less than ~18 meters deep and within 2 km of the shore.[10]

Swimming habits:
The wings of this bird propelled it underwater.[11] It was known to dive to depths of 250 feet or more, some even claiming 1 km. The great auk could perhaps hold its breath for 15 minutes.[12]

Reproduction:
Great auks were colonial breeders, forming vast colonies. Breeding season began in May and ended in mid-July. Females only laid one egg, which was incubated by both parents. The combined incubation and fledgling period was between 43 and 53 days, with the fledgling period being ~10 days. Eggs had a yellowish white to light ochre ground color, with varying patterns of black, brown, or greyish spots and lines, typically concentrated on the large end.[8]

Vocalizations:
The great auk’s calls were similar to the razorbill’s, except they were deeper and and louder. Also amongst its vocalizations were low croaking, gurgling noises, as well as a hoarse scream.[8]

Extinction:
“If you come for their Feathers you do not give yourself the trouble of killing them, but lay hold of one and pluck the best of the Feathers. You then turn the poor Penguin adrift, with his skin half naked and torn off, to perish at his leasure. This is not a very humane method but it is the common practize…

While you abide on this island you are in the constant practice of horrid cruelties for you not only skin them Alive, but you burn them Alive also to cook their Bodies with. You take a kettle with you into which you put a Penguin or two, you kindle a fire under it, and this fire is absolutely made of the unfortunate Penguins themselves. Their bodies being oily soon produce a Flame; there is no wood on the island.”
~Aaron Thomas, in an account on great auk feather hunting.[13]

The great auk always had natural predators, including polar bears (Ursus maritimus), orcas (Orcinus orca), and white-tailed eagles (Haliaeetus albicilla).[8] The Little Ice Age might have surrounded their best breeding islands with ice, allowing polar bears to access their colonies and reduce their populations via predation. They might also have reduced supplies of fish that the great auk depended on.[14]

However, it was really massive human exploitation that drove the great auk into extinction. Recent analysis shows no evidence that great auks were at risk of extinction prior to intensive human hunting beginning in the early 16th century. Even if great auks were under no threat by environmental change, human hunting alone would have been enough to drive it into extinction.[15]

The great auk was hunted all across its North Atlantic range for centuries, both for food and for its down. Although it was extremely hard to catch at sea, it was vulnerable when it emerged onto land to breed.[8]

The population on the eastern Atlantic was smaller (likely because these auks were exploited for longer), but even so it was quite populous on relatively inaccessible breeding colonies (e.g. Funk Island) up to the 16th century. By the mid-16th century, the European population had drastically declined.[8]

There were concerns over the decline of the species. The first attempt at protecting them came in 1753, when Newfoundland petitioned Great Britain to cease the slaughter of great auks; it was unsuccessful. Penalties against killing adults or stealing eggs did exist, but it was still permitted to kill birds for use as fish bait. Eventually in 1794, Britain finally banned killing the birds, but by this point the great auk was no longer a breeding species in Newfoundland. The Funk Island great auks were driven to extinction sometime between 1785 and 1800.[8]

Despite the new ban on killing adult great auks, the birds were still slaughtered. Eggs were also sought after in a wasteful manner; eggers only took eggs without embryos, while discarding those containing an embryo. By the beginning of the 19th century, the population was reduced to a few hundred individuals. What is widely considered to be the last authentic account of a British great auk was in July of 1840, on Stac-an-Armin, St. Kilda, Scotland.[8][13] This bird was found on a rocky ledge and caught in its sleep. Its legs were tied up and it was kept alive for three days after its capture. After a storm began after some noise it had made, the men who captured the bird thought it was a witch and killed it.[13]

The species’ last Icelandic stronghold, and one of the last locations in which it lived, was a small island called Geirfuglasker. This island had a tremendous advantage over any other in that, due to the dangerous currents and volcanic activity, it was almost impossible for sailors to land on it. However, Geirfuglasker was submerged by a volcanic explosion in 1830, forcing the great auks on the island to take refuge on the island of Eldey.[16]

Although Eldey was difficult to land on, it was not impossible. After 1830, Icelanders have proven capable of regularly landing on Eldey and catching birds.[16]

On June 3, 1844, a group of fishermen landed on Eldey with the intent to collect more great auk specimens. One breeding pair was found incubating an egg. The adults were strangled to death (their skins were preserved and their carcasses were preserved in spirits), while the egg was apparently smashed. This account is generally accepted as the last genuine record of a great auk[8], although there is a supposed sighting from December 1852 that has been accepted by the IUCN.[2][8] The species was never seen again.[8]


A stuffed great auk specimen (the "Rivoli Auk", specimen No. 77) on display at the Academy of Natural Sciences of Drexel University. Image source->


References:

[1] Olson, S. L. (2003). A fossil of the Great Auk Pinguinus impennis from Middle Pleistocene deposits in Bermuda. Atlantic Seabirds, 5(2), 81-84.
[2] www.iucnredlist.org/species/22694856/93472944

[3] Smith, N. A. (2011). Taxonomic revision and phylogenetic analysis of the flightless Mancallinae (Aves, Pan-Alcidae). ZooKeys, (91), 1.
[4] 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.
[5] Olson, S. L. (1977). A great auk, Pinguinis [sic], from the Pliocene of North Carolina (Aves: Alcidae). Proceedings of the biological Society of Washington.
[6] Olson, S. L., & Rasmussen, P. C. (2001). Miocene and Pliocene birds from the Lee Creek Mine, North Carolina. Smithsonian Contributions to Paleobiology, 90, 233-365.
[7] Livezey, B. C. (1988). [url=https://sora.unm.edu/sites/default/files/journals/auk/v105n04/p0681-p0698.pdfMorphometrics of flightlessness in the Alcidae[/url]. The Auk, 105(4), 681-698.
[8] Hume, J. P., & Walters, M. (2012). Extinct birds (Vol. 217). A&C Black.
[9] Crofford, E. (1989). The Great Auk. New York: Crestwood House. p. 10.
[10] Olson, S. L., Swift, C. C., & Mokhiber, C. (1979). An attempt to determine the prey of the Great Auk (Pinguinus impennis). The Auk, 790-792.
[11] Morris, F. O. (1857). A history of British birds (Vol. 6). John C. Nimmo.
[12] Cokinos, C. (2009). Hope is the thing with feathers: a personal chronicle of vanished birds. Penguin. p. 311.
[13] Gaskell, J. (2000). Who killed the great auk?. Oxford University Press on Demand.
[14] Ehrlich, P., Dobkin, D. S., & Wheye, D. (1988). Birder’s handbook. Simon and Schuster. p. 201.

[15] Thomas, J. E., Carvalho, G. R., Haile, J., Rawlence, N. J., Martin, M. D., Ho, S. Y., ... & Castruita, J. A. S. (2019). Demographic reconstruction from ancient DNA supports rapid extinction of the great auk. Elife, 8, e47509.
[16] Fuller, E. (2003). The Great Auk: the extinction of the original penguin (Vol. 3). Bunker Hill Publishing, Inc.

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I have a book that mentions great auk aggression and an account of one attacking a person.

"This mighty hunter was equipped with a prodigious weapon. The great auk had a large, narrow bill that it could use when threatened as a formidable dagger. The ornithological historian Jeremy Gaskell has unearthed contemporary accounts that give ample proof of the great auk's aggression and willingness to defend itself. Aggression within densely populated breeding colonies is likely, and early observers mention that if caught, the bird would 'bite everyone within reach of its powerful bill' (Audubon, c. 1840) or 'bite wildly all around' (Fabricius, 1808)" (Gill & West, 186).

"So severe underpopulation would cause stress, demoralization and a tendency to flee rather than fight. But an account Gaskell discovered from Iceland in 1824, only twenty years before the raid on Eldey, tells how a young man was pecked so severely by a cornered great auk that in spite of the calfskin jacket he was wearing the bird was able to draw blood seriously" (Gill & West, 190).

This-> is the book in question. Unfortunately, no preview is available (I was only able to share these excerpts because I own a copy of the book).

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Picture of a skeleton.

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Having recently revamped the OP of this profile, I have decided to celebrate by adding in more content.

These are the organs of the last two great auk specimens, now housed at the Zoological Museum of Copenhagen. The spirit jar on the far right appears to contain the eyeballs, while the second to the left contains the hearts.


Image source->

A genetic analysis of these preserved organs and a list of candidate skin specimens compiled by Errol Fuller shows that, while the male organs match the skin of an auk specimen in Brussels (RBINS 5355) (MK135), the female organs don't match any in the aforementioned list. A specimen from the Cincinnati Museum of Natural History and Science may be a potential candidate for being the last female killed on Eldey (Thomas et al., 2017).

Below is a screen capture I took from reference [16] in the OP above. It is a mummified specimen found on Funk Island, the likes of which I never even knew we had until today.


And below are a few shots of one of the only preserved non-breeding plumage specimens, this one from Copenhagen. This seems to be sometimes said to be the only preserved non-breeding plumage specimen, but according to Wikipedia's article, there are four specimens that are non-summer plumage, two of which are immature.

Image taken from here->



Image from here->

As mentioned in the OP above, great auks were hard to catch in the sea, but vulnerable during breeding season. This is most likely why almost all preserved skins are of the summer breeding plumage.

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This is an Irish specimen. It was caught and died in 1834, and was the last specimen seen alive in Ireland. It was described as a young female, and one of the only two immature skin specimens.


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Here it is on the website of the Zoological Museum of Dublin's collection.

collections.ed.ac.uk/coimbra/record/97383

And just for the sake of it, here's a nice illustration depicting the last two great auks on Eldey, before they were killed.

© @ Michael Rothman->

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This paper (Chávez-Hoffmeister, 2020->) has a bit of information on the great auk. According to the principal component analysis in Figure 3, it falls within the fish/cephalopod dietary preference amongst pursuit diving birds, which seems consistent with the remains of prey eaten by the great auk. Admittedly, though, the fish/cephalopod diet morphospace is quite large.

This book, Bird Recognition-> (by James Fisher, 1947) is the last identification guide that includes the great auk being formally described and figured. I remember reading a while back that the bird continued to be featured in field guides long after its actual extinction (and that searches and bounties for specimens continued long afterwards as well), although it appears by this time that people were catching on that the auk was extinct.