Vortex Formation in the Tethered Flight of the Peacock Butterfly Inachis IO L. (Lepidoptera, Nymphalidae) and Some Aspects of Insect Flight Evolution

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Abstract
High-speed filming has been used to investigate the performance of the peacock butterfly Inachis io while flying in a wind-tunnel. The wake of the butterfly in ‘feeding’ flight is a system of discrete pairs of vortex rings: in each pair the vortex rings are coupled at right angles. The flow distribution around the butterfly and the dynamics of the vortex rings suggest that useful force is produced continuously throughout the wingbeat. The butterfly's flapping flight can be divided into three successive stages: during the downstroke, force generation can be explained by quasi-steady aerofoil action; during the upstroke and supination, by unsteady aerofoil action; and during pronation, by a jet mechanism. The study of airflow around the peacock butterfly throws light on the evolutionary changes in the pattern of interaction between insect wings and the air. At the first stage of the evolution of insect flight, documented in a subimago of the mayfly Heptagenia sulphurea and some other primitive insects, flapping wings generate a system of coupled vortex rings; the aerodynamic force, being perpendicular to the stroke plane, coincides with the direction of the longitudinal body axis. At the second stage, this force is directed forwards and upwards relative to the body axis; the vortex wake is the same as that at the first stage. From this point, two paths of evolution are possible. The first leads to the vortex pattern recorded in the peacock butterfly. The second is typically found in higher orders, where the narrow and relatively short wings flap with lower amplitude and higher frequency, leaving in their wake two chains of uncoupled small vortex rings.