A shortening of the manus precedes the attenuation of other wing-bone elements in the evolution of flightlessness in birds.

Nudds, R. L. and Slove Davidson, J. 2010. A shortening of the manus precedes the attenuation of other wing-bone elements in the evolution of flightlessness in birds. — Acta Zoologica (Stockholm) 91: 115–122 This is the first study to present evidence for a general pattern of wing-bone attenuation during the early stages of the evolution of flightlessness. A comparative analysis using phylogenetic independent contrasts showed that in families that contain both flighted (volant) and flightless species, the volant species have shorter wings and total-arm (humerus + ulna + manus) lengths relative to their body masses than the species within their wholly volant sister families. A shortening of the manus may typify the early stages of the evolution of flightlessness, with the humerus and ulna attenuating later, perhaps because of their role in maintaining the position of the aerodynamically important alula. A shorter wing relative to body mass was not the result of the inverse (i.e. heavier body mass relative to wing length) because mean body masses of volant members of flightless families were similar to or lower than those of their wholly volant sister families. Despite finding a common trend in the wing morphologies of volant members of flightless families, it seems unlikely that a general model of selection pressures driving the evolution of flightlessness exists. At the very least, a dichotomy between those birds that have lost the ability to fly in order to gain the ability to swim and terrestrial forms, may persist. [ABSTRACT FROM AUTHOR]