The management of fluid and wave resistances by whirligig beetles.

Whirligig beetles (Coleoptera: Gyrinidae) are semi-aquatic insects with a morphology and propulsion system highly adapted to their life at the air-water interface. When swimming on the water surface, beetles are subject to both fluid resistance and wave resistance. The purpose of this study was to analyse swimming speed, leg kinematics and the capillarity waves produced by whirligig beetles on the water surface in a simple environment. Whirligig beetles of the species Gyrinus substriatus were filmed in a large container, with a high-speed camera. Resistance forces were also estimated. These beetles used three types of leg kinematics, differing in the sequence of leg strokes: two for swimming at low speed and one for swimming at high speed. Four main speed patterns were produced by different combinations of these types of leg kinematics, and the minimum speed for the production of surface waves (23 cm s(-1)) corresponded to an upper limit when beetles used low-speed leg kinematics. Each type of leg kinematics produced characteristic capillarity waves, even if the beetles moved at a speed below 23 cm s(-1). Our results indicate that whirligig beetles use low- and high-speed leg kinematics to avoid maximum drag and swim at speed corresponding to low resistances.