A fundamental avian wing-stroke provides a new perspective on the evolution of flight

The evolution of avian flight remains one of biology's major controversies, with a long history of functional interpretations of fossil forms given as evidence for either an arboreal or cursorial origin of flight. Despite repeated emphasis on the 'wing-stroke' as a necessary avenue of investigation for addressing the evolution of flight, no empirical data exist on wing-stroke dynamics in an experimental evolutionary context. Here we present the first comparison of wing-stroke kinematics of the primary locomotor modes (descending flight and incline flap-running) that lead to level-flapping flight in juvenile ground birds throughout development. We offer results that are contrary both to popular perception and inferences from other studies. Starting shortly after hatching and continuing through adulthood, ground birds use a wing-stroke confined to a narrow range of less than 20 degrees , when referenced to gravity, that directs aerodynamic forces about 40 degrees above horizontal, permitting a 180 degrees range in the direction of travel. Based on our results, we put forth an ontogenetic-transitional wing hypothesis that posits that the incremental adaptive stages leading to the evolution of avian flight correspond behaviourally and morphologically to transitional stages observed in ontogenetic forms.