Functional Microanatomy of the Feather-Bearing Integument: Implications for the Evolution of Birds and Avian Flight

A selective regime favoring a streamlining of body contours and surfaces is proposed as having been instrumental in driving the morphological and functional transformations of an unfeathered reptilian integument into a feather-bearing avian one. This hypothesis is consistent with a new, structurally and functionally coherent analysis of the microanatomy of the avian feather-bearing integument as a complex, integrated organ system that includes an intricate, hydraulic skeleto-muscular apparatus of the feathers, a dermo-subcutaneous muscle system of the integument, and a subcutaneous hydraulic skeletal system formed by fat bodies. Key elements of the evidence supporting the new hypothesis are (1) the presence of depressor feather muscles that are not needed as antagonists for the erector feather muscles, but can counteract external forces, such as air currents; (2) the fact that the highly intricate feather-bearing integument represents a machinery to move feathers or to stabilize them against external forces; (3) the crucial role of the coat of feathers in streamlining the body contours and surfaces of birds; (4) the aerodynamic role of feathers as pressure and turbulence sensors and as controllable temporary turbulators; and (5) the critical role that a streamlined body plays in avian flight and is likely to have played in the evolutionary transformations from ecologically and locomotorily versatile quadrupedal reptiles to volant bipedal birds without passing through parachuting or gliding stages. These transformations are likely to have occurred more than once. The ancestral birds were probably small, arboreal, hopping, and using flap-bounding, or intermittent bounding, flight.