Maintaining the avian wing aerofoil: Relationships between the number of primary and secondary flight feathers and under-lying skeletal size in birds

Bird wings vary in size and morphology in terms of both size and number of feathers and the underlying skeletal anatomy. The number of primary remiges does not seem to vary much between bird species but, by contrast, the number of secondary remiges is reported to range between 6 and 40 depending on bird size. Given that the primaries are attached to the manus, and the secondaries are attached to the ulna, it was predicted that as bone lengths increased with increasing size of the bird, then feather count would increase. Data were collected for 268 species from 25 different orders, and phylogenetically controlled analysis explored the allometry between feather count and bone size. The number of primaries was typically 10 or 11 and did not vary with manus size. By contrast, the number of secondaries increased with ulna length, but only in some orders. For example, in Gruiformes, the number of secondary feathers increased concomitantly with ulna length but despite a two orders of magnitude range in body mass, almost all species in the Passeriformes had nine secondary remiges. It is unclear why, for instance, species with an ulna length of 70 mm can have between 9 and 24 secondaries depending on their order. This variation in secondary feather number can be added to variation in relative wing bone lengths, flight feather lengths, flight feather mechanical properties, and flight feather vane densities as another potential mechanism of adaptation to flight requirements. The apparent constraint of wingspan is scaling as approximately body mass^1/3. Further research is needed to explore whether changes in secondary feather number relative to ulna length are accompanied by changes in feather vane width or the overlap of adjacent feathers and how this relates to wing aerodynamics.