Piecing Together the Extinct Great Auk

Reviewed by: The Tragic Tale of the Great Auk Elizabeth Bush Thornhill, Jan The Tragic Tale of the Great Auk; written and illus. by Jan Thornhill. Groundwood, 2016 44p ISBN 978-1-55498-865-5 $18.95 R* Gr. 3-6 The short of it is that the great auk, a flightless seabird, is extinct. The long of it, however, is that the story of this extinction is one of considerable complexity, with plot threads involving evolution, geology, human history, and several strokes of plain bad luck. Thornbill’s approach to this historical event is so advanced it’s simple: she tells the tale. No chapter headings, no sidebars, no highlighted vocabulary, no literary gimmicks. She just tells the tale. And what a tale it is. Evolution favored the great auk with a body wonderfully adapted to hunting fish in the cold, open sea. With short wings, and legs set unusually far back, it was a speedy, agile swimmer. The trade-off, though, was the inability to fly, and since birds cannot lay eggs in mid ocean, auks were forced ashore to mate, lay their eggs, and rear their fledglings in an environment in which their only mobility was an awkward waddle. Fortunately the great auk always managed to find nesting grounds on storm-wracked North Atlantic coasts, on uninhabitable islands, or in any number of craggy, cliff-protected spots that foiled predators or at least limited the predators’ threat to their overall numbers. Even early humans, who are known to have developed a taste for great auk, couldn’t make a dent in their thriving populations. Once human hunters took to the sea in ever more efficient boats, though, auk colonies began to diminish and even disappear, losing the race to find protected nesting grounds. Was this the beginning of the tragic end? Or was it when the last excellent rocky island refuge sank in a volcanic eruption? Or when legal efforts to protect the endangered birds against hunters proved too little, too late? Or when the great auk’s rarity made it a target for collectors, who hustled the very last survivors to their demise in the nineteenth century ? Thornhill engages readers with well-placed questions that anticipate their curiosity. “So why didn’t The Wobble [an New England nickname for the Great Auk] avoid land entirely?” “How, then, did it successfully raise its young for millennia?” “So is that all that’s left of the Great Auk? A few sad taxidermy displays and blown eggs?” She also exploits the natural momentum of the auks’ history: as the tale shifts chronological scale from evolutionary time to the historical time, the pacing accelerates and the details become more plentiful and sharply focused. Humans appear on the scene, and we learn about the tempting richness of auk egg omelettes and pancakes; the usefulness of a fatty auk carcass as a substitute for firewood; the execution of the last known auk in the British Isles for witchcraft; the skyrocketing value of rare auk eggs that drove a collector to crush one specimen in order to raise the value of another; Iceland’s 1971 purchase of a stuffed auk and its [End Page 111] celebratory arrival: “A bird that never flew in life flew into Iceland strapped into its very own seat on an airplane.” Digital artwork, which makes clever use of fine white lines detailing foamy ocean and ghostly images of the decimated species, could easily pass for mixed-media compositions. In keeping with the great auk’s very long backstory and relatively short but deadly connection with mankind, most spreads feature auks in the wild rather than alongside their human predators. A notable trio, though, delivers serious chills: a museum gallery featuring a pair of taxidermied auks; the stuffed auk on its way to Iceland, seen through an airplane window; and a view of the booted legs of the men who strangled the last remaining auks, which now dangle at the hunters’ sides. Yes, Nature dealt the great auks some weak hands, but they beat the odds for countless millennia, until humans finally drove them from the table. Lists of resources, references, extinct species, and great auk names in thirteen...

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.

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.

The body mass and egg mass of the Great Auk Pinguinus impennis were never measured before the bird was driven to extinction in 1844. Previous studies conducted before 1990 used data from related species to estimate the mass of an adult bird at 4500–5000 g, and the fresh mass of its egg as 327–372 g. In the present study, we use a larger dataset of measurements from extant alcids, and statistical methods that control for the effects of phylogeny, to provide new estimates for those traits. The presumed body mass of the Great Auk was initially derived from a hearsay report from the 19th century, and then supported by subsequent comparative analyses based on skeletal measurements. Our new best estimates from currently available data show that the Great Auk's body mass was probably closer to 3560 g and its fresh egg mass was about 350 g. This new body mass estimate is the average of predictions from independent regressions of body mass on (1) tibiotarsus and femur lengths (3441 g) and (2) egg volume (3681 g). We calculated the Great Auk's fresh egg mass from a regression of fresh egg mass on egg volume in the extant alcids. Providing more accurate estimates of the body and egg mass of Great Auk can inform speculation about the developmental mode, ecology and life history of this iconic, extinct species.

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.

The collection is extremely strong in diversity of island species throughout the world. Also the collection contains significant historical collections of importance: James Henry Fleming, James A. Munro, Hoyes Lloyd, part of the Haverschmidt collection from the Guyanas, R. G. Lanning, H.B. Haugh collection of birds eggs from Southern Ontario. The Fleming collection, once considered the most comprehensive private collection of birds in North America, contains many unique collections from Africa, Europe, India, China, and Island Archipelagos of the world. Representative collections of the magnificent Birds of Paradise (Passeriformes; Paradisaeidae) and Bower Birds (Passeriformes; Ptilonorhynchidae) (from the Fleming Collection) has been described as within the top 10 in world by researchers. Sub-collections of the J.H. Fleming, J.A. Munro collections are considered historically significant to the Ornithological community. Other significant collections are The New World Sparrows (Passeriformes: Emberizidae); the North American Wood Warblers (Passeriformes: Parulidae); Shorebirds, Gulls and Auks (Charadriiformes) and the Chicken-like birds (Galliformes) of North America, New World Vireos and Allies (Passeriformes: Vireonidae). The study skin collection contains one of the largest collections in the world of extinct birds including 132 Passenger Pigeon specimens (skins) and New Zealand Huias famed for their sexually dimorphic bills. 1 specimen of the extinct Labrador Duck (1 of 54 mounted skins in world). 1 specimen of the extinct Great Auk (1 of 78 mounted skins in world), miscellaneous skeletal parts of Great Auk specimens, 14 specimens of the extinct Carolina Parakeet and many other species. The skin collection contains historically significant holdings of the highly endangered Hawaiian Honeycreepers (Passeriformes: Fringillidae Drepanididae) from the Fleming collection, 1840s through to 1913, and the famous Darwin’s finches. The frozen tissue collection contains the largest collection of blood and DNA from endangered kiwi populations in New Zealand, as well as bone shavings and DNA from the 14 extinct species of giant moas. Extensive series of New World Owls. Extensive series of New World Flycatchers. Large series of New World Sparrows.

Extinctions, Specific

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.

The evolutionary significance of mass extinctions

Dickerson Coquina Pit (DCP) is a sand and coquina mine located in Saint Lucie County, Florida, 16 km inland from the Atlantic Ocean. Collections were made from a middle Pleistocene (late Irvingtonian Land Mammal Age) vertebrate fossil bed in the Okeechobee Formation at DCP from 2002 until the pit was flooded in 2008. Vertebrate fossils from DCP also were discovered in fill material used in beach replenishment on North Hutchinson Island, St. Lucie County. The avifauna of DCP is based on 65 fossil elements representing 12 orders, 16 families, and 26 species, with habitat preferences ranging from woodlands and prairies to fresh water and estuarine wetlands to the open ocean. Extinct or extralocal species include a transitional Wild Turkey Meleagris cf. M. gallopavo, transitional Short-tailed Albatross Phoebastria cf. P. albatrus, a large stork Ciconia maltha, a large crane Grus sp., Great Auk Pinguinus impennis, and Carolina Parakeet Conuropsis carolinensis. The taphonomy and paleoecology of the Dickerson Coquina Pit generally resemble those of other shell beds in Central and South Florida, such as the Irvingtonian Leisey Shell Pit of Hillsborough County. The avifauna from DCP is distinctive, however, in featuring Florida’s first Pleistocene records of four marine species (Short-tailed Albatross, Northern Gannet Morus bassanus, Great Cormorant Phalacrocorax carbo, and Great Auk) and the Carolina Parakeet.

This paper describes two watercolours and one engraving of the great auk (Pinguinus impennis) executed by the French painter Nicolas Robert probably between 1666 and 1670. Despite their interest as early images of this extinct species and the first ones rendered in colour, they have not been mentioned in the literature. The images suggest that the bird was kept alive in the menagerie at the palace in Versailles, where Robert portrayed it for Louis XIV; Robert's paintings are collectively known as “Les vélins du Roi”.

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.)

In this paper, we modify dynamic occupancy models developed for detection-nondetection data to allow for the dependence of local vital rates on neighborhood occupancy, where neighborhood is defined very flexibly. Such dependence of occupancy dynamics on the status of a relevant neighborhood is pervasive, yet frequently ignored. Our framework permits joint inference about the importance of neighborhood effects and habitat covariates in determining colonization and extinction rates. Our specific motivation is the recent expansion of the Barred Owl (Strix varia) in western Oregon, USA, over the period 1990-2010. Because the focal period was one of dramatic range expansion and local population increase, the use of models that incorporate regional occupancy (sources of colonists) as determinants of dynamic rate parameters is especially appropriate. We began our analysis of 21 years of Barred Owl presence/nondetection data in the Tyee Density Study Area (TDSA) by testing a suite of six models that varied only in the covariates included in the modeling of detection probability. We then tested whether models that used regional occupancy as a covariate for colonization and extinction outperformed models with constant or year-specific colonization or extinction rates. Finally we tested whether habitat covariates improved the AIC of our models, focusing on which habitat covariates performed best, and whether the signs of habitat effects are consistent with a priori hypotheses. We conclude that all covariates used to model detection probability lead to improved AIC, that regional occupancy influences colonization and extinction rates, and that habitat plays an important role in determining extinction and colonization rates. As occupancy increases from low levels toward equilibrium, colonization increases and extinction decreases, presumably because there are more and more dispersing juveniles. While both rates are affected, colonization increases more than extinction decreases. Colonization is higher and extinction is lower in survey polygons with more riparian forest. The effects of riparian forest on extinction rates are greater than on colonization rates. Model results have implications for management of the invading Barred Owl, both through habitat alteration and removal.

Extinction in the Anthropocene.

Communities in fragmented landscapes are often assumed to be structured by species extinction due to habitat loss, which has led to extensive use of the species-area relationship (SAR) in fragmentation studies. However, the use of the SAR presupposes that habitat loss leads species to extinction but does not allow for extinction to be offset by colonization of disturbed-habitat specialists. Moreover, the use of SAR assumes that species richness is a good proxy of community changes in fragmented landscapes. Here, we assessed how communities dwelling in fragmented landscapes are influenced by habitat loss at multiple scales; then we estimated the ability of models ruled by SAR and by species turnover in successfully predicting changes in community composition, and asked whether species richness is indeed an informative community metric. To address these issues, we used a data set consisting of 140 bird species sampled in 65 patches, from six landscapes with different proportions of forest cover in the Atlantic Forest of Brazil. We compared empirical patterns against simulations of over 8 million communities structured by different magnitudes of the power-law SAR and with species-specific rules to assign species to sites. Empirical results showed that, while bird community composition was strongly influenced by habitat loss at the patch and landscape scale, species richness remained largely unaffected. Modeling results revealed that the compositional changes observed in the Atlantic Forest bird metacommunity were only matched by models with either unrealistic magnitudes of the SAR or by models ruled by species turnover, akin to what would be observed along natural gradients. We show that, in the presence of such compositional turnover, species richness is poorly correlated with species extinction, and z values of the SAR strongly underestimate the effects of habitat loss. We suggest that the observed compositional changes are driven by each species reaching its individual extinction threshold: either a threshold of forest cover for species that disappear with habitat loss, or of matrix cover for species that benefit from habitat loss.