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The organic carbon (Corg) stored in seagrass meadows is globally significant and could be relevant in strategies to mitigate increasing CO2 concentration in the atmosphere. Most of that stored Corg is in the soils that underlie the seagrasses. We explored how seagrass and soil characteristics vary among seagrass meadows across the geographic range of turtlegrass (Thalassia testudinum) with a goal of illuminating the processes controlling soil organic carbon (Corg) storage spanning 23° of latitude. Seagrass abundance (percent cover, biomass, and canopy height) varied by over an order of magnitude across sites, and we found high variability in soil characteristics, with Corg ranging from 0.08 to 12.59% dry weight. Seagrass abundance was a good predictor of the Corg stocks in surficial soils, and the relative importance of seagrass-derived soil Corg increased as abundance increased. These relationships suggest that first-order estimates of surficial soil Corg stocks can be made by measuring seagrass abundance and applying a linear transfer function. The relative availability of the nutrients N and P to support plant growth was also correlated with soil Corg stocks. Stocks were lower at N-limited sites than at P-limited ones, but the importance of seagrass-derived organic matter to soil Corg stocks was not a function of nutrient limitation status. This finding seemed at odds with our observation that labile standard substrates decomposed more slowly at N-limited than at P-limited sites, since even though decomposition rates were 55% lower at N-limited sites, less Corg was accumulating in the soils. The dependence of Corg stocks and decomposition rates on nutrient availability suggests that eutrophication is likely to exert a strong influence on carbon storage in seagrass meadows.

We studied the shapes of eggs from 955 extant bird species across the avian phylogeny, including 39 of 40 orders and 78% of 249 families. We show that the elongation component of egg shape (length relative to width) is largely the result of constraints imposed by the female's anatomy during egg formation, whereas asymmetry (pointedness) is mainly an adaptation to conditions during the incubation period. Thus, egg elongation is associated with the size of the egg in relation to both the size of the female's oviduct and her general body conformation and mode of locomotion correlated with pelvis shape. Egg asymmetry is related mainly to clutch size and the structure of the incubation site, factors that influence thermal efficiency during incubation and the risk of breakage. Importantly, general patterns across the avian phylogeny do not always reflect the trends within lower taxonomic levels. We argue that the analysis of avian egg shape is most profitably conducted within taxa where all species share similar life histories and ecologies, as there is no single factor that influences egg shape in the same way in all bird species.

Nearly 70 years ago, Preston published a pioneering study in which he provided formulae for the shapes of birds’ eggs. One of these formulae is universal in that it provides an almost perfect representation for all eggs, even pyriform ones, and is better than all other formulae published since. This essentially perfect representation of egg shape is obtained by estimating the parameters in Preston’s universal formula by least squares, using hundreds of measurements of the egg’s radii along its entire length. Preston’s universal formula can also be used to obtain an equation for avian egg shape that uses just 5 measurements (the length and 4 appropriately spaced diameters). The equation based solely on these 5 measurements provides an egg shape that is virtually indistinguishable from one based on hundreds of measurements. We demonstrate the usefulness of Preston’s formulations using digital photographs of eggs. Our perspective is that, despite a number of subsequent approaches, Preston’s original one has not been bettered and should be the standard for studying avian egg shape.

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 Brunnich'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 Brunnich'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 studied the ground colors and maculations of 161 Common Murre (Uria aalge) eggs laid by 43 females in 3 small breeding groups on the cliffs of Skomer Island, Wales, in 2016–2018. Both the colors and maculations varied much more among than within females, providing quantitative evidence for the egg traits that might facilitate the parents' ability to identify their own eggs on the crowded breeding ledges where the density is typically ∼20 eggs m–2. Ground colors had a trimodal distribution of hue values (whitish to pale brown, pale blue, or vivid blue-green) and maculations ranged from none to complex squiggles and blotches. The eggs laid by each female in different years were similar to one another, and replacement eggs laid by females within years were also more similar to their first egg than to other eggs in the same breeding group. Egg appearance did not differ among the 3 breeding groups that we studied. Our findings thus support anecdotal observations that, within and between years, female Common Murres lay eggs that have similar ground colors and maculations. We do not, however, find evidence that there is much difference among the eggs laid in different parts of a colony. LAY SUMMARY For more than a century, naturalists have marveled at the diversity of colors and markings on the eggs of Common Murres (Uria aalge) breeding in dense coastal colonies, where they lay and incubate a single egg. Largely anecdotal observations have suggested that each female lays distinctive eggs, and that eggs laid on the same cliff ledge tend to be more similar than expected if pairs are distributed randomly. We measured the appearance of 161 eggs laid by 43 Common Murres over a 3-year period on Skomer Island, Wales. Each female consistently laid eggs within a narrow range of whitish to pale brown (11 females) or pale blue to vivid blue-green ground color (32 females) and similar markings both within- and between breeding seasons. Despite offspring tending to return to breed near their parents, eggs laid within a few meters of one another were no more similar to each other than they were to eggs laid on cliff ledges 200 m away.

Edward Walter Wade (1864–1937), author of The Birds of Bempton Cliffs (1903, 1907), is almost unknown. He worked as a clerk for the family timber company in Hull and in his spare time visited Bempton Cliffs, where he climbed (routinely without a rope) to observe the seabirds and to collect eggs. He was an active member of the Hull Scientific and Field Naturalists' Club and became one of their Vice-Presidents and President. Between 1902 and 1920 he was a member of the British Ornithologists' Union. Wade's published comments on guillemots and their pyriform-shaped eggs, at Bempton, were unusually insightful, in part because his observations were made at close range (a consequence of his climbing ability), and partly because he thought carefully about what he saw. Wade was among the first to point out that the spinning-like-a-top explanation for the guillemot's pyriform egg shape was incorrect based on his own observations. An appendix lists Wade's 69 articles, mostly about birds in Yorkshire, published between 1901 and 1930.

The adaptive significance of avian egg shape is a long-standing problem in biology. For many years, it was widely believed that the pyriform shape of the Common Murre (Uria aalge) egg allowed it to either “spin like a top” or “roll in an arc,” thereby reducing its risk of rolling off the breeding ledge. There is no evidence in support of either mechanism. Two recent alternative hypotheses suggest that a pyriform egg confers mechanical strength and minimizes the risk of dirt contamination of the blunt end. We present a new hypothesis: that the Common Murre egg's pyriform shape confers stability on the breeding ledge, thus reducing the chance that it will begin to roll. We tested this hypothesis by measuring the stability of Common Murre and Razorbill (Alca torda) eggs of different shapes on slopes of 20°, 30°, and 40° above the horizontal. Common Murre eggs were more stable, and easier to stabilize, than the more elliptical Razorbill eggs. Within Common Murre eggs, more pyriform eggs were more stable. From a fitness perspective, the stability of the Common Murre egg on a slope seems likely to confer an advantage and thus may be a strong force of natural selection favoring the pyriform shape.

A recent broad comparative study suggested that factors during egg formation – in particular ‘flight efficiency’, which explained only 4% of the interspecific variation – are the main forces of selection on the evolution of egg shape in birds. As an alternative, we tested whether selection during the incubation period might also influence egg shape in two taxa with a wide range of egg shapes, the alcids (Alcidae) and the penguins (Spheniscidae). To do this, we analysed data from 30 species of these two distantly related but ecologically similar bird families with egg shapes ranging from nearly spherical to the most pyriform eggs found in birds. The shape of pyriform eggs, in particular, has previously proven difficult to quantify. Using three egg‐shape indices – pointedness, polar‐asymmetry and elongation – that accurately describe the shapes of all birds’ eggs, we examined the effects of egg size, chick developmental mode, clutch size and incubation site on egg shape. Linear models that include only these factors explained 70–85% of the variation in these egg‐shape indices, with incubation site consistently explaining > 60% of the variation in shape. The five species of alcids and penguins that produce the most pyriform eggs all incubate in an upright posture on flat or sloping substrates, whereas species that incubate in a cup nest have more spherical eggs. We suggest that breeding sites and incubation posture influence the ability of parents to manipulate egg position, and thus selection acting during incubation may influence egg‐shape variation across birds as a whole.

The adaptive significance of avian egg shape in birds is poorly understood. The pyriform (pear-like) shape of the Common Guillemot's Uria aalge egg has long been considered to be an adaptation to prevent eggs rolling off the bare cliff ledges on which this species breeds. Rolling was thought to be prevented either by the egg spinning like a top, which is not the case, or by rolling in an arc, which it does but with little influence on whether the egg will fall from a ledge. We therefore sought alternative explanations for the pyriform shape of the Common Guillemot's egg. This species breeds in extremely dense colonies, which makes their eggs vulnerable to mechanical damage from conspecifics, and to contamination by debris such as faeces and soil. We present evidence consistent with both these possible explanations. First, the pyriform shape of Common Guillemot eggs means that a higher proportion of the eggshell lies in contact with the substrate and this may minimize the effect of impacts. Resistance to impacts may be further enhanced because their eggshells are especially thick where they are in contact with the substrate. Secondly, Common Guillemot eggs are often heavily contaminated with faecal material and other debris during incubation. Most contamination is on the pointed end of the egg where it is in contact with the substrate; the pyriform shape thus keeps the blunt end of the egg, which has the highest porosity, relatively free of contamination, which in turn may facilitate both gas exchange during incubation and the hatching process, because the chick emerges from the blunt end of the egg.

Birds are arguably the most evolutionarily successful extant vertebrate taxon, in part because of their ability to reproduce in virtually all terrestrial habitats. Common guillemots, Uria aalge, incubate their single egg in an unusual and harsh environment; on exposed cliff ledges, without a nest, and in close proximity to conspecifics. As a consequence, the surface of guillemot eggshells is frequently contaminated with faeces, dirt, water and other detritus, which may impede gas exchange or facilitate microbial infection of the developing embryo. Despite this, guillemot chicks survive incubation and hatch from eggs heavily covered with debris. To establish how guillemot eggs cope with external debris, we tested three hypotheses: (1) contamination by debris does not reduce gas exchange efficacy of the eggshell to a degree that may impede normal embryo development; (2) the guillemot eggshell surface is self-cleaning; and, (3) shell accessory material (SAM) prevents debris from blocking pores, allowing relatively unrestricted gas diffusion across the eggshell. We show that (1) natural debris reduces the conductance of gases across the guillemot eggshell by blocking gas exchange pores. Despite this problem, we find (2) no evidence that guillemot eggshells are self-cleaning, but instead show that (3) the presence of SAM on the eggshell surface largely prevents pore blockages from occurring. Our results demonstrate that SAM is a crucial feature of the eggshell surface in a species whose eggs are frequently in contact with debris, acting to minimise pore blockages and thus ensure a sufficient rate of gas diffusion for embryo development.

Birds are arguably the most evolutionarily successful extant vertebrate taxon, in part because of their ability to reproduce in virtually all terrestrial habitats. Common guillemots

Describing the range of avian egg shapes quantitatively has long been recognized as difficult. A variety of approaches has been adopted, some of which aim to capture the shape accurately and some to provide intelligible indices of shape. The objectives here are to show that a (four‐parameter) method proposed by Preston (1953, The Auk, 70, 160) is the best option for quantifying egg shape, to provide and document an R program for applying this method to suitable photographs of eggs, to illustrate that intelligible shape indices can be derived from the summary this method provides, to review shape indices that have been proposed, and to report on the errors introduced using photographs of eggs at rest rather than horizontal.