Summary Body feathers ensure both waterproofing and insulation in waterbirds, but how natural variation in the morphological properties of these appendages relates to environmental constraints remains largely unexplored. Here, we test how habitat and thermal condition affect the morphology of body feathers, using a phylogenetic comparative analysis of five structural traits [i.e., total feather length, the lengths of the pennaceous (distal) and plumulaceous (proximal) sections, barb density, and pennaceous barbule density] from a sample of 194 European bird species. Body feather total length is shorter in aquatic than in terrestrial birds, and this difference between groups is due to the shorter plumulaceous feather section in aquatic birds. Indeed, a reduced plumulaceous section in feather length probably reflects the need to limit air trapped in the plumage to adjust the buoyancy of aquatic birds. In contrast, the high pennaceous barbule density of aquatic birds compared to their terrestrial counterparts reflects water resistance of the plumage in contact with water. Our results show that birds living in environments with low ambient temperature have long plumulaceous feather lengths, low barb density, and low pennaceous barbule density. Data also suggest that plumage probably has limited function in reducing the heat absorption of species living in hot environments. Our results have broad implications for understanding the suite of selection pressures driving the evolution of body feather functional morphology. It remains to be tested, however, how other feather traits, such as the density of plumage (feathers per unit area) and the relative number of different feather types, for example downy feathers, are distributed amongst birds with different water resistance and thermoinsulative needs. A http://onlinelibrary.wiley.com/doi/10.1111/1365-2435.12820/suppinfo is available for this article.

Studies modelling heat transfer of bird plumage design suggest that insulative properties can be attributed to the density and structure of the downy layer, whereas waterproofing is the result of the outer layer, comprised of contour feathers. In this study, we test how habitat and thermal condition affect feather mass and density of body feathers (contour, semiplume and downy feathers) measured on the ventral and dorsal sides of the body, using a phylogenetic comparative analysis of 152 bird species. Our results demonstrate that feather mass and the density of downy feathers are higher in species that inhabit colder environments, whereas total feather density is higher of species breeding under intermediate temperatures compared to the ones breeding under more extreme conditions. The density of contour feathers, depending on the body region, is either quadratically related or negatively correlated with minimum winter temperature. The density of contour and downy feathers, measured on both sides of the body, is higher in aquatic than in terrestrial birds. However, among the former, diving behaviour does not select for further increases in body feather mass or density. The results of this study provides key insights into how the plumage of birds is adapted to different environments and lifestyles and provides a basis for understanding the diverse range and the evolution of variation in these characteristics. A http://onlinelibrary.wiley.com/doi/10.1111/1365-2435.13019/suppinfo is available for this article.

Down feathers are the first feather types that appear in both the phylogenetic and the ontogenetic history of birds. Although it is widely acknowledged that the primary function of downy elements is insulation, little is known about the interspecific variability in the structural morphology of these feathers, and the environmental factors that have influenced their evolution. Here, we collected samples of down and afterfeathers from 156 bird species and measured key morphological characters that define the insulatory properties of the downy layer. We then tested if habitat and climatic conditions could explain the observed between-species variation in down feather structure. We show that habitat has a very strong and clearly defined effect on down feather morphology. Feather size, barbule length and nodus density all decreased from terrestrial toward aquatic birds, with riparian species exhibiting intermediate characters. Wintering climate, expressed as windchill (a combined measure of the ambient temperature and wind speed) had limited effects on down morphology, colder climate only being associated with higher nodus density in dorsal down feathers. Overall, an aquatic lifestyle selects for a denser plumulaceous layer, while the effect of harsh wintering conditions on downy structures appear limited. These results provide key evidence of adaptations to habitat at the level of the downy layer, both on the scale of macro-and micro-elements of the plumage. Moreover, they reveal characters of convergent evolution in the avian plumage and mammalian fur, that match the varying needs of insulation in terrestrial and aquatic modes of life.