Wing Movements Associated With Collision-Avoidance Manoeuvres During Flight in the Locust Locusta Migratoria

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Abstract
Flying locusts will try to avoid colliding with objects directly in their flight path. This study investigated the wing movements and behaviour patterns associated with collision avoidance. Tethered locusts were flown in a wind tunnel. Targets were transported at different speeds either directly towards the head of the animal or to one side of the midline but parallel to it. Changes in the form of the wingbeat for each of the wings were monitored using either a video camera or a high-speed ciné camera. Animals attempted to avoid an impending collision by making movements interpreted here as (a) increasing lift to fly over the object, (b) gliding and extending the forelegs to land on the object, and (c) steering to one side of the object. Steering was monitored by observation of abdominal movements. Steering to one side of an approaching target was reliably associated with an earlier and more pronounced pronation of the wings on the inside of the turn. Also, in the middle of the downstroke, the forewings were markedly asymmetrical. On the outside of the turn, the forewing was more elevated and separate from the hindwing. On the inside of the turn, the forewing was more depressed and often came down in conjunction with, or in advance of, the hindwing on that side. The forewing asymmetry correlated with the position of the target such that most attempted turns were in the direction that would take the animal around the closest edge. High-speed cinematography showed that the asymmetry was caused both by changes in the timing of the two wings and by changes in the angular ranges of the wingbeats. We propose that these changes in the form and timing of the wingbeats are likely to have swung the flight force vector around the long axis of the body to produce a banked turn around the closest edge of the object.