Animal aloft: the origins of aerial behavior and flight

Diverse taxa of animals exhibit remarkable aerial capacities, including jumping, mid-air righting, parachuting, gliding, landing, controlled maneuvers, and flapping flight. The origin of flapping wings in hexapods and in 3 separate lineages of vertebrates (pterosaurs, bats, and birds) greatly facilitated subsequent diversification of lineages, but both the paleobiological context and the possible selective pressures for the evolution of wings remain contentious. Larvae of various arboreal hemimetabolous insects, as well as many adult canopy ants, demonstrate the capacity for directed aerial descent in the absence of wings. Aerial control in the ancestrally wingless archaeognathans suggests that flight behavior preceded the origins of wings in hexapods. In evolutionary terms, the use of winglets and partial wings to effect aerial righting and maneuvers could select for enhanced appendicular motions, and ultimately lead to powered flight. Flight behaviors that involve neither flapping nor wings are likely to be much more widespread than is currently recognized. Further characterization of the sensory and biomechanical mechanisms used by these aerially capable taxa can potentially assist in reconstruction of ancestral winged morphologies and facilitate our understanding of the origins of flight.