The Great Auk (Pinguinus impennis) had two brood patches, not one: confirmation and implications

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.

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.

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 Great Auk Pinguinus impennis was a large, flightless alcid, endemic to the North Atlantic Ocean. It became extinct around 1844. Skeletal remains are used to document its (pre-)historic range. While these remains were considered rare from the southern North Sea, over the past five years 91 (sub-)fossil specimens have been recovered by citizen scientist fossil collectors from Dutch beaches that were nourished with sediments dredged from the bottom of the North Sea. Some of this material is now stored in museum collections. This paper lists the new remains and documents them through measurements and photographs. The material was recovered from fourteen new localities and one previously known locality in The Netherlands and has yielded four radiocarbon dates (1425–1300 BC till beyond 48,000 cal BP) which significantly increase the Great Auk's temporal range in this area. The sheer volume of remains alters our image of the Great Auk in the southern part of the North Sea from a rare bird to most likely a common or regular wintering bird over the past millennia.

The life, death and afterlife of one of the true icons of extinction, the Great Auk The great auk was a flightless, goose-sized bird superbly adapted for life at sea. Fat, flush with feathers and easy to capture, the birds were in trouble whenever sailors visited their once-remote breeding colonies. Places like Funk Island, off north-east Newfoundland, became scenes of unimaginable slaughter, with birds killed in their millions. By 1800 the auks of Funk Island were gone. A scramble by private collectors for specimens of the final few birds then began, a bloody, unthinking destruction of one of the world’s most extraordinary species. But their extinction in 1844 wasn’t the end of the great auk story, as the bird went on to have a remarkable afterlife; skins, eggs and skeletons became the focus for dozens of collectors in a story of pathological craving and unscrupulous dealings that goes on to this day. In a book rich with insight and packed with tales of birds and of people, Tim Birkhead reveals previously unimagined aspects of the bird’s life before humanity, its death on the killing shores of the North Atlantic, and the unrelenting subsequent quest for its remains. The great auk remains a symbol of human folly and the necessity of conservation. This book tells its story.

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.

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.

We investigated variation in the number and size of brood patches of female (n = 133) and male (n = 81) American Kestrels Falco sparverius in northcentral Saskatchewan, Canada. We analyzed brood patch size with respect to characteristics of individual birds such as sex, age, size, condition, and clutch volume. All incubating birds developed brood patches but females tended to have larger patches than males. The size of the brood patch was correlated significantly only with the mass of the female. Birds with large eggs or large clutches did not necessarily have large brood patches, and patch size was not associated with laying date or hatching success. In a comparison with other avian taxa, we found kestrels to be unusual in that their three discrete brood patches did not correspond to the modal clutch size of five eggs. Kestrels potentially have difficulty heating all their eggs simultaneously.

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.

Incubating birds transfer large amount of heat from the brood patch to the eggs during rewarming of cold eggs. If a vasoconstriction is present in the brood patch as in other parts of the body, it could possibly limit heat transfer to the eggs. To investigate this, heat transfer to water-circulated eggs was measured in incubating bantam hens (Gallus domesticus) and a black grouse hen (Lyrurus tetrix) during exposure to cold eggs. Egg temperature, egg surface temperature, heat production and cloacal temperature were also measured. At all levels of egg cooling, egg surface temperature and heat transfer to the eggs was stable throughout an exposure, except during resettling movements, which often changed egg surface temperature and the level of heart transfer. Egg surface temperature decreased linearly with egg temperature in both species, but was lower and more variable at low egg temperature in black grouse than in bantam hens. A higher proportion of the heat production was transferred to the eggs in the black grouse (corresponding to 109–118% of the increase above resting level) than previously reported in bantam hens. Clutch size did not affect this efficiency of heat transfer in black grouse. It is concluded that a vasoconstruction of the brood patch does not occur even under strong cold stress from the eggs. Heat transfer to the eggs is probably controlled more by behavioural adjustments than circulatory changes. An increase in brood patch blood flow probably occurs at relatively high egg temperature at the onset of egg rewarming. The efficiency of heat transfer, and thus the energetic cost of rewarming eggs, depends on the insulation of the bird and nest structure. The boreal/subarctic black grouse was able to reduce heat loss to the environment and transfer a higher proportion of its heat production to the eggs than the tropical bantam hen.

IMAGINATION has long had a large share in the accounts given of the Gare-fowl or Great Auk, notwithstanding the efforts of those who have tried to set forth nothing but the truth on the subject, yet I do not call to mind meeting with so “many inventions” regarding it as have appeared in the newspapers within the last week, on the occasion of the recent sale of a specimen of the egg of that bird. I should occupy too much space were I to dwell upon them; but I would ask for the admission of a few lines in which to state what is known exactly of the origin of that specimen, which I well remember in the collection of the late Mr. Yarrell. He told me, as he told others of his friends, that he bought it in Paris; and, to the best of my belief, not many years after the peace of 1815. In a little curiosity-shop of mean appearance, he saw a number of eggs hanging on a string; he recognised one of them as an egg of Alca impennis, and asking their price was told that they were one franc apiece, except the large one, which from its size was worth two francs. He paid the money and walked away with the egg in his hat. That is the whole story on which so imposing an edifice has been built, and the only “variant” of it deserving of consideration is to the effect that the price of the big egg was five instead of two francs. I may add that this-simple story was published by the late owner of the egg, the Baron Louis d'Hamonville, in the Bulletin of the French Zoological Society for 1891 (tome xvi. p. 34).

IN your last issue (p. 412), I observe a letter from Prof. Newton, in which he gives his version of the history of the egg of this extinct bird, which was recently sold by auction for £315. There is no doubt that the egg was brought to this, country by Yarrell, who purchased it in France some time before 1838, in which year it was figured by Hewitson in his well-known work on birds‘ eggs. But the question is, where-abouts in France did he find it? Prof. Newton, who well remembers it in the collection of Yarrell, says: “He told me, as he told others of his friends, that he bought it in Paris, in a little curiosity shop of mean appearance,” and that he paid two francs for it, He adds that the only “variant” of this story deserving of consideration, is to the effect that the price was five instead of two francs. If this were the only “variant,” it would not be worth further discussion. But there is a very different story told of it in Mr. Symington Grieve's important work on “The Great Auk, its history, archaeology, and remains,” published in 1885. At p. 105 of this volume, Mr. Grieve writes of this very egg:—

In many incubating birds, heat transfer from parent to egg is facilitated by the brood patch, an area of ventral abdominal skin that becomes highly vascularised, swells and loses its down feathers around the time of laying. Only the female develops a brood patch in most passerine species, but males of some species can incubate and maintain the eggs at similar temperatures to females even without a brood patch. Here we used a novel application of infrared thermography to examine sex differences in parental care from a physiological perspective. Using incubating male and female zebra finches (Taeniopygia guttata), a species in which the male lacks a brood patch, we measured the surface temperature of the ventral plumage overlying the abdomen and a reference area that does not contact the eggs (thorax) twice per pair. In half of the pairs, clutch size was experimentally enlarged between the two sets of measurements to increase incubation demand. We found that the temperature differential between abdomen and thorax plumage was greater in females than in males, and that abdomen plumage was warmer after clutch enlargement than before in females but not in males. These findings are consistent with morphological sex differences in brood patch development and suggest that male and female zebra finches differ in the way they regulate abdomen versus general body surface temperature in response to variation in incubation demand.

1. In birds, local competition for food between pairs during the nestling phase may affect nestling growth and survival. A decrease in clutch size with an increase in breeding density could be an adaptive response to this competition. To investigate whether breeding density causally affected the clutch size of great tits (Parus major), we manipulated breeding density in three out of eight study plots by increasing nest-box densities. We expected clutch size in these plots to be reduced compared to that in control plots. 2. We analysed both the effects of variation in annual mean density (between-year comparisons) and experimental density (within-year comparison between plots) on clutch size variation, the occurrence of second broods and nestling growth. We examined within-female variation in clutch size to determine whether individual responses explain the variation over years. 3. Over the 11 years, population breeding density increased (from 0.33 to 0.50 pairs ha(-1)) while clutch size and the occurrence of second broods decreased (respectively from 10.0 to 8.5 eggs and from 0.39 to 0.05), consistent with a negative density-dependent effect for the whole population. Nestling growth showed a declining but nonsignificant trend over years. 4. The decline in population clutch size over years was primarily explained by changes occurring within individuals rather than selective disappearance of individuals laying large clutches. 5. Within years, breeding density differed significantly between manipulated plots (0.16 pairs ha(-1) vs. 0.77 pairs ha(-1)) but clutch size, occurrence of second broods and nestling growth were not affected by the experimental treatment, resulting in a discrepancy between the effects of experimental and annual variation in density on reproduction. 6. We discuss two hypotheses that could explain this discrepancy: (i) the decline in breeding performance over time was not due to density, but resulted from other, unknown factors. (ii) Density did cause the decline in breeding performance, but this was not due to local competition in the nestling phase. Instead, we suggest that competition acting in a different phase (e.g. before egg laying or after fledgling) was responsible for the density effect on clutch size among years.

Effects of artificial eggs on prolactin secretion, steroid levels, brood patch development, incubation onset and clutch size in the yellow-eyed penguin ( Megadyptes antipodes)