1. Flight Laboratory, Division of Biological Sciences, University of Montana, 32 Campus Drive, Missoula, Montana 59812, USA

Correspondence to: Kenneth P. Dial¹ Correspondence and requests for materials should be addressed to K.P.D. (Email: kdial@mso.umt.edu).

Abstract

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^{1, 2, 3, 4}, 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 (Fig. 1). We offer results that are contrary both to popular perception and inferences from other studies^{5, 6, 7}. Starting shortly after hatching and continuing through adulthood, ground birds use a wing-stroke confined to a narrow range of less than 20°, when referenced to gravity, that directs aerodynamic forces about 40° above horizontal, permitting a 180° 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.

Figure 1: Locomotor development during ontogeny in the chukar partridge from hatching to adulthood.

Our data suggest a default or basal wing-stroke is used by young and adults and may exist in all birds (Supplementary Videos). The fundamental wing-stroke described herein is used days after hatching and during all ages and over multiple behaviours (that is, flap-running, descending and level flight) and is the foundation of our new ontogenetic-transitional wing hypothesis. At hatching, chicks can ascend inclines as steep as 60° by crawling on all four limbs. From day 8 through adulthood, birds use a consistently orientated stroke-plane angle over all substrate inclines during wing-assisted incline running (red arcs) as well as during descending and level flight (blue arcs). Estimated force orientations from this conserved wing-stroke are limited to a narrow wedge (see Fig. 3b).

High resolution image and legend (289K)

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