Unconventional lift-generating mechanisms in free-flying butterflies

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Abstract
Flying insects generate forces that are too large to be accounted for by conventional steady-state aerodynamics1,2. To investigate these mechanisms of force generation, we trained red admiral butterflies, Vanessa atalanta, to fly freely to and from artificial flowers in a wind tunnel, and used high-resolution, smoke-wire flow visualizations to obtain qualitative, high-speed digital images of the air flow around their wings. The images show that free-flying butterflies use a variety of unconventional aerodynamic mechanisms to generate force: wake capture3, two different types of leading-edge vortex3,4,5,6,7, active and inactive upstrokes8, in addition to the use of rotational mechanisms3 and the Weis–Fogh ‘clap-and-fling’ mechanism9,10,11,12. Free-flying butterflies often used different aerodynamic mechanisms in successive strokes. There seems to be no one ‘key’ to insect flight, instead insects rely on a wide array of aerodynamic mechanisms to take off, manoeuvre, maintain steady flight, and for landing.

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