Kinematic Analysis of Symmetrical Flight Manoeuvres of Odonata

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Abstract
By analysis of slow-motion films of dragonflies and damselflies in free flight, released in front of a backdrop or startled during flight, the following flight parameters have been quantified for symmetrical manoeuvres: wingbeat frequency, relative durations of up-and downstroke, phase relationships of the beats of fore-and hindwings, stroke amplitude, mean stroke velocity, flight velocity, nondimensional flight velocity, advance ratio, acceleration, angle of attack and stroke plane. The wingbeat frequencies are higher in the smaller species and in those with relatively large wing loading. As a rule, Zygoptera have a wingbeat frequency only half that of Anisoptera. The stroke amplitude is almost always much larger in Zygoptera than in Anisoptera, which have a greater range of variation in this respect. Stroke velocity is higher in Anisoptera than in Zygoptera; it is also higher in the more elaborate flight manoeuvres than in others. The calculated stroke velocities resemble those actually measured. Anisoptera fly more rapidly than Zygoptera. With respect to the nondimensional flight velocities, it is notable that although the values for Anisoptera are higher than those for Zygoptera, they are exceeded by the Calopterygidae; the latter can fold their wings back during rapid forward flight and shoot away, as in the ‘ballistic’ flight of small songbirds. However, the advance ratio is higher in Anisoptera than in Calopterygidae. Anisoptera also perform better than Zygoptera with respect to acceleration. Three categories of phase relationships between the beats of the fore-and hindwings are established: counterstroking, phase-shifted stroking and parallel stroking. Counterstroking produces uniform flight, whereas the flight produced by phase-shifted and, in particular, parallel stroking is irregular. The angles of attack of the wings are shown to be associated with particular flight manoeuvres, as are the stroke planes. Flight manoeuvres are discussed without drawing detailed aerodynamic conclusions. The flight of Anisoptera is compared with that of Zygoptera.