Surface-Skimming Stoneflies: A Possible Intermediate Stage in Insect Flight Evolution

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Abstract
Insect wings appear to have evolved from gills used by aquatic forms for ventilation and swimming, yet the nature of intermediate stages remains a mystery. Here a form of nonflying aerodynamic locomotion used by aquatic insects is described, called surface skimming, in which thrust is provided by wing flapping while continuous contact with the water removes the need for total aerodynamic weight support. Stoneflies surface skim with wing areas and muscle power output severely reduced, which indicates that surface skimming could have been an effective form of locomotion for ancestral aquatic insects with small protowings and low muscle power output.