The Tragic Tale of the Great Auk

This paper presents all available archaeozoological, ethnohistorical, and historical evidence on the Great Auk, Pinguinus impennis (L.), in Greenland. Substantial new contributions are: (1) the presentation of 132 Auk bones from seven archaeological sites, and (2) a translation of Otto Fabricius's original handwritten notes on the Great Auk in Greenland. A description of the Greenlandic Great Auk's zoogeography and biology is given as well as a description of the Inuit's Great Auk hunting methods and of their use of Great Auk products. Main conclusions are: (1) P. impennis probably occurred further north in Greenland c. 2000 BC than in historical times; (2) c. 1500 BC P. impennis possibly nested in West Greenland; (3) Between 1350 and 1800 AD Great Auks originating from breeding colonies near Newfoundland or Iceland wintered on the banks off West Greenland from Cape Farewell in the south to Maniitsoq in the north; (4) Great Auks were breeding in Southwest Greenland in small numbers in the 1760's and 1770'...

The great auk was once abundant and distributed across the North Atlantic. It is now extinct, having been heavily exploited for its eggs, meat, and feathers. We investigated the impact of human hunting on its demise by integrating genetic data, GPS-based ocean current data, and analyses of population viability. We sequenced complete mitochondrial genomes of 41 individuals from across the species’ geographic range and reconstructed population structure and population dynamics throughout the Holocene. Taken together, our data do not provide any evidence that great auks were at risk of extinction prior to the onset of intensive human hunting in the early 16th century. In addition, our population viability analyses reveal that even if the great auk had not been under threat by environmental change, human hunting alone could have been sufficient to cause its extinction. Our results emphasise the vulnerability of even abundant and widespread species to intense and localised exploitation.

The great auk was once abundant and distributed across the North Atlantic. It is now extinct, having been heavily exploited for its eggs, meat, and feathers. We investigated the impact of human hunting on its demise by integrating genetic data, GPS-based ocean current data, and analyses of population viability. We sequenced complete mitochondrial genomes of 41 individuals from across the species' geographic range and reconstructed population structure and population dynamics throughout the Holocene. Taken together, our data do not provide any evidence that great auks were at risk of extinction prior to the onset of intensive human hunting in the early 16th century. In addition, our population viability analyses reveal that even if the great auk had not been under threat by environmental change, human hunting alone could have been sufficient to cause its extinction. Our results emphasise the vulnerability of even abundant and widespread species to intense and localised exploitation.

We measured the bones of extinct great auks Pinguinis impennis that were killed during recent centuries on Funk Island off the northeast coast of Newfoundland. Comparisons of these measurements with those taken elsewhere suggest that great auks from Funk Island, which is situated in a Low Arctic oceanographic region, were larger than conspecifics from Boreal oceanographic regions. This finding is supported by extant alcid species that inhabit Boreal, Low Arctic or Boreal through High Arctic ocean regions and tend to increase in body size with increasing latitude (generally decreasing sea surface temperature). We suggest that paleoecological sea surface temperatures and food webs may have favored oceanographic-related variation in body sizes of great auks. The variances of the bone sizes of great auks from Funk Island were not less than those of a sample of great auk bones collected from Scandinavian archaeological sites that cover an extensive geographic range and that span seven millenia. This finding is inconsistent with a previously suggested latitudinal cline in body size among great auks in Scandinavia. Research techniques and studies that could address questions of great auk feeding ecology and population genetics are considered.

Seabirds are the most threatened of any living group of birds, continuing a larger pattern of elevated Holocene bird extinctions on islands and coastlines. The Great Auk (Charadriiformes: Pinguinus impennis) was found on both coasts of the Atlantic during the Holocene until its last sighting on Iceland in 1844. Far more is known about the population structure and genetic diversity of NE Atlantic populations, and the latest surviving populations were documented from the British Isles in 1834. While sightings from Canada suggest Great Auks disappeared by 1800, no systematic evaluation of extinction timing has been conducted for this coast. Determining extinction timing of the Great Auk in Maine allows a comparison to be made to populations in other areas of the Atlantic Ocean, and raises the question: was the Maine population’s fate different due to regional, cultural, or other factors? There is a single eye-witness record in the late 17th century at “Black Point”, now Scarborough, Maine. To address this gap, we compiled a radiocarbon dataset on associated material from Maine archaeological shell middens. These 91 dates from 13 sites situate the Great Auk in Maine from about 180 to 4,555 years before present. The majority of these dates are from charcoal samples, but also include shells, ceramics, and bone, and cultural contexts span the Middle and Late Ceramic Periods. To account for differences in stratigraphic control and sampling material, we assigned quality scores, and used these scores to run a sensitivity analysis in extinction timing with the GRIWM model. Disentangling the spatiotemporal dynamics of the Great Auk extinction in Maine is useful in determining how to conserve current species in decline and modern insular seabirds in Maine, such as the puffin. Future study will include new radiocarbon dating of bones as well as isotopic and morphometric analysis to unfold more chapters of the Maine Great Auk’s narrative.

The story of the killing of the ‘last’ great auk (Pinguinus impennis) in Britain, apparently put to death as a witch at Stac an Armin in the St Kilda archipelago c. 1840, is well known. However, other accounts claim that an auk was killed on the main island, Hirta, having been condemned to death by the celebrated men’s ‘parliament’. The historical veracity of three differing stories, which recount discreditable deeds in a deeply Christian community, is evaluated; it seems that fewest difficulties are raised if two great auks were killed, one on Hirta and the other on Stac an Armin. It is argued that this kind of avicide was a ‘ritual’ killing, to be understood in its historical context. The auk-killing probably took place in the mid to late 1840s, after the St Kilda minister had departed in the wake of the Disruption of 1843 - a particularly unsettling time within this small island community. A possible sighting of a pair of great auks on Soay (St Kilda) in 1890 is also briefly discussed.

Stable isotopic ratios of animal tissues are related to those of their foods and can be used in palaeoecological reconstructions, including those of extinct animals. Nitrogen isotopic analyses of marine organisms from coastal Newfoundland and Georges Bank were used to construct a model predicting collagen δ15N values for seabirds feeding at various trophic levels (TL). This model was tested by measuring bone collagen δ15N values of extant alcids from the northwest Atlantic and high Arctic. Isotopic analysis of bone collagen of the extinct great auk (Pinguinus impennis), the last flightless seabird in the northern hemisphere, indicate that this species occupied a trophic continuum from TL3 (crustacean diets) to TL5 (diets of piscivorous fish). We suggest that (a) great auk chicks and juveniles occupied lower trophic levels and probably consumed euphausiids, and (b) great auks fed offspring via regurgitation, as do dovekies (Alle alle), the only extant fully planktivorous alcid in the Atlantic, and unrelated penguins of the southern hemisphere.

One hundred and seventy-three years ago, the last two Great Auks, Pinguinus impennis, ever reliably seen were killed. Their internal organs can be found in the collections of the Natural History Museum of Denmark, but the location of their skins has remained a mystery. In 1999, Great Auk expert Errol Fuller proposed a list of five potential candidate skins in museums around the world. Here we take a palaeogenomic approach to test which—if any—of Fuller’s candidate skins likely belong to either of the two birds. Using mitochondrial genomes from the five candidate birds (housed in museums in Bremen, Brussels, Kiel, Los Angeles, and Oldenburg) and the organs of the last two known individuals, we partially solve the mystery that has been on Great Auk scholars’ minds for generations and make new suggestions as to the whereabouts of the still-missing skin from these two birds.

In this final contribution on the identification of the birds painted in the Palaeolithic Grotte Cosquer as Great Auks, it is noticed that the birds need to be seen within a Palaeolithic hunter–gatherer's view of the world, which is not the same as that of a modern natural historian or taxonomist.

The hunting of the great auk ( Pinguinus impennis), which led to its extinction in the mid-nineteenth century, is well documented. However, the discovery of archives providing new details on this species is a rare event. A manuscript dealing with seabirds and their ‘fishing’, written in 1720–1722 by François Le Masson du Parc (1671–1741), an attaché for the Normandy maritime administration, was acquired in 2019 from an auction house. This unpublished and unstudied manuscript comprises the sixth and final volume of the ‘Histoire des pesches’. As part of a national policy to regulate French maritime fisheries, Le Masson du Parc completed the description of marine resources and fisheries in his ‘Histoire des pesches’ which was abundantly illustrated by Pierre Le Chevalier (1688–after 1763). However, this monumental work was never published and the manuscripts were dispersed after the author's death. The section recently purchased provides valuable information on the spatiotemporal evolution of many populations of North Atlantic seabirds, including the great auk, in a context of anthropogenic pressures. This species is named using several hitherto unknown Latin and vernacular names of great importance for the exploration or reinterpretation of archives. It is mentioned as common and one of the most popular birds consumed by cod fishermen on the banks of Newfoundland and that it was caught using baited hooks from boats. The text is accompanied by two illustrations, including a life drawing of three great auks, which are among the oldest known illustrations of the species. These 300-year-old archives constitute a valuable testimony for the historical ecology of this iconic, extinct bird.

-Common Murres (Uria aalge) exhibit an unusual molt sequence. Primary molt begins at a focus between P4 and P7 and progresses in two rapid concurrent waves, proximally to P1, and distally to P10. The only other birds known to have a similar molt sequence are caracaras and falcons (Falconidae), parrots (Psittaciformes), and Pied Kingfishers (Ceryle rudis). Great Auks (Pinguinus impennis) also appear to have followed the same primary-molt sequence. Phylogenies for the Alcidae indicate that Great Auks, Common Murres, Thickbilled Murres (Uria lomvia), Razorbills (Alca torda), and Dovekies (Alle alle) share a common ancestor and are more closely related to one another than to other alcids. This suggests that the unusual sequence of primary molt in Common Murres is a shared-derived character that occurs in the other four species in their clade but has been overlooked. Adult male Common Murres have significantly shorter secondaries and longer primaries, on average, than do adult females, resulting in a slightly higher aspect ratio in males. Secondary molt begins when primary molt is more than one-third completed. Secondaries are replaced rapidly but sequentially (not synchronously or simultaneously); molt appears to proceed from two foci, proximally from S1 to S4, and both proximally and distally from S8, but more data are needed to clarify this point. Rectrix molt begins when primary molt is two-thirds completed. Rectrix loss and replacement occur rapidly, possibly synchronously, and in no apparent order. Adults molt about two weeks later than nonbreeding subadults. We found no differences in the timing of molt between the sexes in adults or subadults. Duration of flight-feather molt can vary from less than 25 days to more than 80 days, possibly reflecting interyear variation in prey abundance. Received 12 February 1997, accepted 9 December 1997. RELATIVE TO MOST OTHER TOPICS in ornithology, virtually all aspects of the molting process are poorly documented and understood (Pyle et al. 1987, Jenni and Winkler 1994). This is especially true in seabirds because most species, including all species that become flightless during molt, undergo molt at sea during the nonbreeding season (Palmer 1962, Glutz von Blotzheim and Bauer 1982, Cramp 1977, 1983, Warham 1996). Among alcids, molt has been better studied in Common Murres (Uria aalge) than in many other species (Verwey 1922, 1924, Salomonsen 1944, Stresemann and Stresemann 1966, Birkhead and Taylor 1977), but many aspects of their molt remain poorly known (contra Harris and Wanless 1990). As an extension of studies on seabird entanglement in gill nets in Puget Sound (Thompson et al. 1998), we studied molt in Common 4E-mail: thompcwt@dfw.wa.gov Murres. The phenology of breeding in Common Murres differs by two months or more among geographic areas. Thus, flight-feather molt scores in postbreeding adults might be useful for identifying breeding location (e.g. Oregon vs. Washington); this is important for determining the demographic effect of mortality caused by gill nets or other anthropogenic activities on different breeding populations of Common Murres. MATERIALS AND METHODS In 1993, commercial gill net fisheries occurred for summer sockeye salmon (Onchorhynchus nerka) in northern Puget Sound and for fall chum salmon (0. keta) in Hood Canal and central Puget Sound, Washington (Pierce et al. 1994). To evaluate various modified gill net designs, test fisheries were done in 1993 by Washington Sea Grant, and in 1993 and 1996 by Washington Sea Grant and Washington Department of Fish and Wildlife. Entangled seabirds were collected: (1) from commercial fisheries in 1993 from 1

Présence du grand pingouin,Pinguinus impennis (aves, charadriiformes) dans le Pléistocène du Portugal

The paper considers the assemblage of bird and fish bones from a Romano-British settlement on the Isle of Portland, on the southern coast of England. Compared with contemporary sites, the assemblage includes an unusually large number of fish bones from a wide range of marine species, including large cod, other Gadidae, several species of seabream, scad and bass. The bird assemblage includes bones of a butchered great auk. This provides the first evidence that this extinct species was nesting off the shores of central southern England and being exploited for food in this period. Other seabirds identified included razorbill, great northern diver and gannet. The species represented are discussed in relation to other Romano-British sites, particularly the Roman town of Dorchester, situated 15 km away. Many of the species have been discovered on only a few contemporary sites and the presence of the seabream in particular indicates that seawater temperatures may have been warmer than until very recently. Possible cultural changes in diet and food procurement in the Roman period are also considered.

A total of 477 bones of the great auk (Pinguinus impennis) from 53 localities and 55 periods in Norway are studied. All but two, are archaeological sites from the Holocene, mainly from 6000–2000 cal years bp. The two non‐anthropogenic sites date to the Ice Age: probably 36,000–34,500 and 14,690–12,890 years bp. The bones are mainly unburned and well preserved but fractured. Except for the open‐air sites in northern Norway, the bones are mainly from rock‐shelters and caves. In combining archive data, chronological information, and morphometrical studies, we suggest the great auk disappeared from the most southern part of Norway (and Sweden and Denmark) prior to 4000 years bp: a decline in distribution 2000 years ago: It became absent from the Norwegian coast 1000 years ago. Data suggest that it was distributed on the coast and in the fjord systems in winter and early spring. The presence of bones of juveniles/subadults indicates that it was also distributed in northern Norway in the autumn. To evaluate possible size differences, in time and space, nine bone elements have been measured according to standard recommendations. Multiple imputation was used to handle missing data before any statistical analysis. Analyses indicate that bones from Nordland are larger than from the rest of the country, while bones from the northernmost sites are smaller. At some localities, size differences, especially in total length of the bones, are found. It has not been verified if this is due to individual variation or sexual differences. The great auk became extinct in the 19th century. The study supports the theory that human predation at breeding sites was the main cause of its extinction.

Data collected from skin specimens of the 23 Recent species of Alcidae, skeletal material for Recent and fossil alcids, and published data on body mass and wing area were used to describe the morphometric characteristics of flightlessness in the Great Auk (Pinguinus impennis) and the fossil mancalline auks. A regression equation confirmed a body-mass estimate (5 kg) for P. impennis (Bédard 1969). The size and relatively small wings produced wing-loading of roughly 22 g· cm-2, comparable to those of medium-sized penguins. Multivariate analysis of external measurements underscored the uniquely large size, relatively short wings, and moderately deep bill of Pinguinus compared to other Recent alcids. Analysis of skeletal measurements revealed that the genera of flightless Alcidae (Pinguinus, Mancalla, Praemancalla, and Alcodes) were characterized by relatively short distal wing elements and dorsoventral flattening of all major wing elements, in combination with relatively large core and pelvic dimensions. These differences were most pronounced in Mancalla, moderately developed in Praemancalla, and smallest in Pinguinus. Estimated body mass (1-4 kg) for selected fossil mancallines exceeded the largest flighted alcids (Uria) but was less than for Pinguinus. Pinguinus was a comparatively large piscivore sharing many morphological features with the Razorbill (Alca torda) and murres (Uria spp.). Its flightlessness evidently was a consequence of extreme specialization for pursuit diving, convergent with that of the Spheniscidae. Loss of flight imposed significant requirements on breeding sites and foraging habitats of the Great Auk and presumably the mancallines, and rendered Pinguinus exceptionally vulnerable to human exploitation.