LAUSR

Variation in rachis (central shaft) morphology in individual remiges (flight feathers) within and among species reflects adaptations to requirements imposed by aerodynamic forces, but the fine-scale variation of feather morphology across remiges is not well known. Here we describe how the shape of the rachis, expressed by the height/width ratio, changes along the longitudinal and lateral axis of the wing in four bird species with different flight styles: flapping-soaring (white storks), flapping-gliding (common buzzards), passerine-type (house sparrows) and continuous flapping (pygmy cormorants). Overall, in each wing feather, irrespective of species identity, rachis shape changed from circular to rectangular, from the base towards the feather tip. The ratio between the height and width of the calamus was similar across remiges in all species, whereas the ratio at the base, middle and tip of the rachis changed among flight feathers and species. In distal primaries of white storks and common buzzards, the ratio decreased along the feather shaft, indicating a depressed (wider than high) rachis cross section towards the feather tip, whereas the inner primaries and secondaries became compressed (higher than wide). In house sparrows, the rachis was compressed in each of the measurement points, except at the distal segment of the two outermost primary feathers. Finally, in pygmy cormorants, the width exceeds the height at each measurement point, except at the calamus. Our results may reflect the resistance of the rachis to in-plane and out-of-plane aerodynamic forces that vary across remiges and across study species. A link between rachis shape and resistance to bending from aerodynamic forces is further indicated by the change of the second moment of areas along the wing axes.