“Der fluglose Alk”: Johann Friedrich Naumann’s 1844 account of Pinguinus impennis (great auk)

The Atlantic auk assemblage includes four extant species, razorbill (Alca torda), dovekie (Alle alle), common murre (Uria aalge), and thick-billed murre (U. lomvia), and one recently extinct species, the flightless great auk (Pinguinus impennis). To determine the phylogenetic relationships among the species, a contiguous 4.2-kb region of the mitochondrial genome from the extant species was amplified using PCR. This region included one ribosomal RNA gene, four transfer RNA genes, two protein-coding genes, the control region, and intergenic spacers. Sets of PCR primers for amplifying the same region from great auk were designed from sequences of the extant species. The authenticity of the great auk sequence was ascertained by alternative amplifications, cloning, and separate analyses in an independent laboratory. Phylogenetic analyses of the entire assemblage, made possible by the great auk sequence, fully resolved the phylogenetic relationships and split it into two primary lineages, Uria versus Alle, Alca, and Pinguinus. A sister group relationship was identified between Alca and Pinguinus to the exclusion of ALLE: Phylogenetically, the flightless great auk originated late relative to other divergences within the assemblage. This suggests that three highly divergent species in terms of adaptive specializations, Alca, Alle, and Pinguinus, evolved from a single lineage in the Atlantic Ocean, in a process similar to the initial adaptive radiation of alcids in the Pacific Ocean.

Since the late 1600s it has been assumed that the Great AukPinguinus impenniswas similar to the Common GuillemotUria aalgeand Brünnich's GuillemotUria lomviain having a single, central brood patch. Through the examination of eight mounted museum specimens, we show that this is incorrect and that, like its closest relative the RazorbillAlca torda, the Great Auk had two lateral brood patches. We discuss how such misinformation persisted for so long. We also review the relationship between the number of brood patches and clutch size in the Alcidae. One implication of two brood patches is that the Great Auk would have incubated in a horizontal posture like the Razorbill, rather than in a semi‐upright posture like theUriaguillemots. Assuming that the Great Auk incubated like the Razorbill, it would probably have done so horizontally with its single egg pressed against one of the two lateral brood patches, positioned against the inside of one tarsus (and partially on the web of one foot), and with the wing on that side drooped to provide additional protection for the egg. Incubating in this way may have meant that the Great Auk's pyriform egg would have enabled it to use both level and sloping terrain, as in theUriaguillemots (but unlike the Razorbill). A horizontal incubation also has implications for estimates of their breeding density, which we estimate to have been around four pairs per square metre and, hence numbers on its largest known colony, Funk Island, Newfoundland (maximum 250 000 pairs).

We compared the shape and eggshell thickness of Great Auk Pinguinus impennis eggs with those of its closest relatives, the Razorbill Alca torda, Common Guillemot Uria aalge and Brünnich's Guillemot Uria lomvia, in order to gain additional insights into the breeding biology of the extinct Great Auk. The egg of the Great Auk was most similar in shape to that of Brünnich's Guillemot. The absolute thickness of the Great Auk eggshell was greater than that of the Common Guillemot and Razorbill egg, which is as expected given its greater size, but the relative shell thickness at the equator and pointed end (compared with the blunt end) was more similar to that of the Common Guillemot. On the basis of these and other results we suggest that Great Auk incubated in an upright posture in open habitat with little or no nest, where its pyriform egg shape provided stability and allowed safe manoeuvrability during incubation. On the basis of a recent phylogeny of the Alcidae, we speculate that a single brood patch, a pyriform egg and upright incubation posture, as in the Great Auk and the two Uria guillemots, is the ancestral state, and that the Razorbill – the Great Auk's closest relative – secondarily evolved two brood patches and an elliptical egg as adaptations for horizontal incubation, which provides flexibility in incubation site selection, allowing breeding in enclosed spaces such as crevices, burrows or under boulders, as well as on open ledges.

In dialogue with extinction studies scholarship and literary studies debates about methods of reading, this chapter explores how Walton Ford’s great auk paintings register species loss. In “Funk Island ~ or Good Conduct Well Chastised (1791)” (1998), Ford stages the death of the island’s great auk colony, once the largest breeding colony in North America. In “The Witch of St. Kilda ~ 1840” (2005), he portrays the capture of the last great auk in the British Isles. By attending to the content of both paintings, their relationship to each other, their art historical contexts, and the socioecological contexts of great auk exploitation and extinction, I argue that Ford employs and critiques the conventions of natural history illustration to present anthropogenic extinction as an explicitly colonialist and violent form of loss. In addition to parsing their critiques of natural history illustration, I attend to the ways in which the allusive and intertextual operations of both paintings produce reading practices that lure the viewer into assembling an archive of great auk extinction. Finally, this chapter suggests that reading great auk extinction stories with Walton Ford has important outcomes. First, Ford’s paintings, and the archive they evoke and constellate, proliferate a series of new dates for thinking a variety of Anthropocenes. These dates decenter more familiar Anthropocene narratives and highlight the exploitation of nonhuman creatures as fundamental to colonialisms. Second, by foregrounding the violence inherent in extinction, Ford’s paintings eclipse the effects of grief and melancholy that inform other cultural representations of extinction and plumb the existential horror that adheres to the irrevocable severing of genetic lines and socioecological relationships.

High resolution X-ray micro-computed tomography gives the ability to research objects in unprecedented detail in 3D without damaging them but applying these new techniques to specimens can be complex. In 2017 the Natural History Museum (NHM), London embarked on a ground-breaking project with University of Sheffield to compare extinct Great Auk Pinguinus impennis eggshell microstructure to that of their extant relatives to gain new insight into their breeding ecology. NHM has a ZEISS Xradia 520 Versa X-ray microscope capable of submicron X-ray imaging in 3D but using it required supporting and moving complete eggshells within the confined, potentially harsh, mechanised environment of the microscope without risk. Ensuring the correct position and orientation of each egg to image nine distinct areas on the eggshell was also a challenge. Collaboration with colleagues in the NHM Conservation and Imaging & Analysis Centres developed a bespoke solution to hold and protect the eggs during scanning. All six NHM Great Auk eggshells and the inside of the microscope were surface scanned using a handheld structured light scanner. Scan data produced 3D models from which accurate 3D printed plastic replicas were made of the three Great Auk eggs prioritised for research. Each replica was used to mould a two-part, custom-built, case for each egg constructed from conservation grade epoxy putty and lined with polyethylene foam. This provided close-fitting, durable cases which could be used for the 6-month duration of the project. Each case enclosed its matching Great Auk egg entirely and had the advantage of being rock-hard, electrically insulating and water, heat and chemical resistant. A system of three, interchangeable, tailor-made mounting brackets were designed that married with the cases and held them safely and precisely inside the microscope at the correct angles and positions for imaging. The structured light scan of the inside of the microscope was used to model the necessary rotational movements of the cases and brackets inside the scanner, ensuring that all movements had sufficient clearance to avoid risk of impact. This system successfully protected the fragile c. 200 year old eggs throughout 70 scanning sessions. This provides a methodology for high resolution X-ray micro-computed tomography imaging of any similarly sized, fragile, object.

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.

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.

ABSTRACTAlthough studies of osteological morphology, gross myology, myological histology, neuroanatomy, and wing‐scaling have all documented anatomical modifications associated with wing‐propelled diving, the osteohistological study of this highly derived method of locomotion has been limited to penguins. Herein we present the first osteohistological study of the derived forelimbs and hind limbs of wing‐propelled diving Pan‐Alcidae (Aves, Charadriiformes). In addition to detailing differences between wing‐propelled diving charadriiforms and nondiving charadriiforms, microstructural modifications to the humeri, ulnae and femora of extinct flightless pan‐alcids are contrasted with those of volant alcids. Histological thin‐sections of four species of pan‐alcids (Alca torda, †Alca grandis, †Pinguinus impennis, †Mancalla cedrosensis) and one outgroup charadriiform (Stercorarius longicaudus) were compared. The forelimb bones of wing‐propelled diving charadriiforms were found to have significantly thicker (∼22%) cortical bone walls. Additionally, as in penguins, the forelimbs of flightless pan‐alcids are found to be osteosclerotic. However, unlike the pattern documented in penguins that display thickened cortices in both forelimbs and hind limbs, the forelimb and hind limb elements of pan‐alcids display contrasting microstructural morphologies with thickened forelimb cortices and relatively thinner femoral cortices. Additionally, the identification of medullary bone in the sampled †Pinguinus impennis specimen suggests that further osteohistological investigation could provide an answer to longstanding questions regarding sexual dimorphism of Great Auks. Finally, these results suggest that it is possible to discern volant from flightless wing‐propelled divers from fragmentary fossil remains. Anat Rec, 297:188–199, 2014. © 2013 Wiley Periodicals, Inc.

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.

We used proteomic profiling to taxonomically classify extinct, alongside extant bird species using mass spectrometry on ancient bone-derived collagen chains COL1A1 and COL1A2. Proteins of Holocene and Late Pleistocene-aged bones from dodo (Raphus cucullatus) and great auk (Pinguinus impennis), as well as bones from chicken (Gallus gallus), rock dove (Columba livia), zebra finch (Taeniopygia guttata) and peregrine falcon (Falco peregrinus), of various ages ranging from the present to 1455 years old were analysed. HCl and guandine-HCL-based protein extractions from fresh bone materials yielded up to 60% coverage of collagens COL1A1 and COL1A2, and extractions from ancient materials yielded up to 46% coverage of collagens COL1A1 and COL1A2. Data were retrieved from multiple peptide sequences obtained from different specimens and multiple extractions. Upon alignment, and in line with the latest evolutionary insights, protein data obtained from great auk grouped with data from a recently sequenced razorbill (Alca torda) genome. Similarly, protein data obtained from bones of dodo and modern rock dove grouped in a single clade. Lastly, protein data obtained from chicken bones, both from ancient and fresh materials, grouped as a separate, basal clade. Our proteomic analyses enabled taxonomic classification of all ancient bones, thereby complementing phylogenetics based on DNA.

-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

Although studies of osteological morphology, gross myology, myological histology, neuroanatomy, and wing-scaling have all documented anatomical modifications associated with wing-propelled diving, the osteohistological study of this highly derived method of locomotion has been limited to penguins. Herein we present the first osteohistological study of the derived forelimbs and hind limbs of wing-propelled diving Pan-Alcidae (Aves, Charadriiformes). In addition to detailing differences between wing-propelled diving charadriiforms and nondiving charadriiforms, microstructural modifications to the humeri, ulnae and femora of extinct flightless pan-alcids are contrasted with those of volant alcids. Histological thin-sections of four species of pan-alcids (Alca torda, †Alca grandis, †Pinguinus impennis, †Mancalla cedrosensis) and one outgroup charadriiform (Stercorarius longicaudus) were compared. The forelimb bones of wing-propelled diving charadriiforms were found to have significantly thicker (∼22%) cortical bone walls. Additionally, as in penguins, the forelimbs of flightless pan-alcids are found to be osteosclerotic. However, unlike the pattern documented in penguins that display thickened cortices in both forelimbs and hind limbs, the forelimb and hind limb elements of pan-alcids display contrasting microstructural morphologies with thickened forelimb cortices and relatively thinner femoral cortices. Additionally, the identification of medullary bone in the sampled †Pinguinus impennis specimen suggests that further osteohistological investigation could provide an answer to longstanding questions regarding sexual dimorphism of Great Auks. Finally, these results suggest that it is possible to discern volant from flightless wing-propelled divers from fragmentary fossil remains.

Although Charles and Emile Parzudaki were well-connected natural history dealers of nineteenth-century Paris, many aspects of their life and work remain unknown. The example of two letters from Emile Parzudaki to Robert Champley that accompanied three great auk ( Pinguinus impennis) eggs to the Zoologisches Forschungsmuseum Alexander Koenig in Bonn reveal new aspects of the Parzudaki enterprise, indicating at least some travelling and collecting activities of Charles Parzudaki beyond Europe.

A GREEABLY to the wish of the editor of NATURE that I should notice in its pages the lately-published volume whose title stands above, I undertake a responsibility of a kind which is for me as delicate as can be imposed upon anybody. It has long been no secret that for more than five-and-twenty years-since, indeed, the premature death, in 1859, of my friend and fellow-traveller, the late Mr. JOHN WOLLEY—I have had it in hand to prepare and eventually to produce a monograph of the presumably extinct species of bird, into the investigation of whose history he had thrown himself with all the energy of his character. During that time I am not conscious of having ever lost an opportunity of adding to my store of information on the subject, in doing which I was for several years assisted by the zeal of the late Mr. G. D. Rowley; and, though always having in view the ultimate publication of the monograph originally contemplated by Mr. Wolley, I never hesitated to supply any inquirer with the particulars for which he asked-as may be seen on reference to the publications of Dr. Victor Fatio1 and of Prof. Wilhelm Blasius2—both of whom I rejoice to think I was able in some measure to help. Nevertheless, each attempt to elucidate the natural history of the Garefowl only added to the number of still unanswered or unanswerable questions relating to it; and, amid numerous other occupations or duties, I have with difficulty been able to keep myself abreast of the ever-increasing contributions to the subject—many (I may say most) of them proving on investigation to have little or no foundation; and those which had the least, or none at all, generally giving the greatest trouble. The Great Auk, or Garefowl (Alca impennis, Linn.), its History, Archaeology, and Remains. By Symington Grieve, Edinburgh. 4to, pp. x. 141, and Appendix, pp. 58. (London: Jack, 1885.)

This article examines the historical context and afterlife of the great auk as a museum object. The focus is on the changing meaning of the bird as it travels across cultural boundaries, material forms and historical periods. It starts by focusing on how the bird was contextualized within Ole Worm’s cabinet in the 17th century and then moves on to more recent museum representations. By conducting interviews with museum staff and professionals and analysis of items stored and/or exhibited at the museum, the symbolic meanings and representative functions of the great auk are brought into light. As proposed in this paper, certain characteristics, values and meanings stem from the materiality of the objects but also from their travels and re-locations. The investigation reveals how the role of the great auk in relation to collective museum (and human) challenges such as emerging ecological limits and biodiversity loss has been developing.