Detailed morphological investigation, mechanical testing and high-speed cinematography and stroboscopic examination of desert locusts, Schistocerca gregaria, in flight show that their hind wings are adapted to deform cyclically and automatically through the wing stroke and that the deformations are subtly dependent on the wings' structure: their shape, venation and vein design and the local properties of the membrane. The insects predominantly fly fast forwards, generating most force on the downstroke, and the hind wings generate extra lift by peeling apart at the beginning of the downstroke and by developing a cambered section during the stroke's translation phase through the 'umbrella effect' - an automatic consequence of the active extension of the wings' expanded posterior fan. Bending experiments indicate that most of the hind wing is more rigid to forces from below than from above and demonstrate that the membrane acts as a stressed skin to stiffen the structure.
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