Carefully controlled gas exchange across the eggshell is essential for the development of the avian embryo. Water vapour conductance (G(H2O)) across the shell, typically measured as mass loss during incubation, has been demonstrated to optimally ensure the healthy development of the embryo while avoiding desiccation. Accordingly, eggs exposed to sub-optimal gas exchange have reduced hatching success. We tested the association between eggshell G(H2O) and putative life-history correlates of adult birds, ecological nest parameters and physical characteristics of the egg itself to investigate how variation in G(H2O) has evolved to maintain optimal water loss across a diverse set of nest environments. We measured gas exchange through eggshell fragments in 151 British breeding bird species and fitted phylogenetically controlled, general linear models to test the relationship between G(H2O) and potential predictor parameters of each species. Of our 17 life-history traits, only two were retained in the final model: wet-incubating parent and nest type. Eggs of species where the parent habitually returned to the nest with wet plumage had significantly higher G(H2O) than those of parents that returned to the nest with dry plumage. Eggs of species nesting in ground burrows, cliffs and arboreal cups had significantly higher G(H2O) than those of species nesting on the ground in open nests or cups, in tree cavities and in shallow arboreal nests. Phylogenetic signal (measured as Pagel's λ) was intermediate in magnitude, suggesting that differences observed in the G(H2O) are dependent upon a combination of shared ancestry and species-specific life history and ecological traits. Although these data are correlational by nature, they are consistent with the hypothesis that parents constrained to return to the nest with wet plumage will increase the humidity of the nest environment, and the eggs of these species have evolved a higher G(H2O) to overcome this constraint and still achieve optimal water loss during incubation. We also suggest that eggs laid in cup nests and burrows may require a higher G(H2O) to overcome the increased humidity as a result from the confined nest microclimate lacking air movements through the nest. Taken together, these comparative data imply that species-specific levels of gas exchange across avian eggshells are variable and evolve in response to ecological and physical variation resulting from parental and nesting behaviours.

Department of Animal and Plant Sciences University of Sheffield Sheffield S10 2TN UK

Department of Psychology Hunter College and the Graduate Center of the City University of New York 695 Park Avenue New York NY 10065 USA

Department of Zoology and Laboratory of Ornithology Palacký University Olomouc CZ 771 46 Czech Republic

School of Earth and Environmental Sciences University of Adelaide SA 5005 Australia

Structure and Motion Laboratory The Royal Veterinary College University of London North Mymms Hatfield Herts AL9 7TA UK

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