Scaling of avian primary feather length.

The evolution of the avian wing has long fascinated biologists, yet almost no work includes the length of primary feathers in consideration of overall wing length variation. Here we show that the length of the longest primary feather (f(prim)) contributing to overall wing length scales with negative allometry against total arm (ta = humerus+ulna+manus). The scaling exponent varied slightly, although not significantly so, depending on whether a species level analysis was used or phylogeny was controlled for using independent contrasts: f(prim) is proportional to ta(0.78-0.82). The scaling exponent was not significantly different from that predicted (0.86) by earlier work. It appears that there is a general trend for the primary feathers of birds to contribute proportionally less, and ta proportionally more, to overall wingspan as this dimension increases. Wingspan in birds is constrained close to mass (M(1/3)) because of optimisation for lift production, which limits opportunities for exterior morphological change. Within the wing, variations in underlying bone and feather lengths nevertheless may, in altering the joint positions, permit a range of different flight styles by facilitating variation in upstroke kinematics.