Aquatic turning performance by the whirligig beetle: constraints on maneuverability by a rigid biological system.

Turning performance is constrained by morphology, where the flexibility of the body and the mobility and position of the control surfaces determine the level of performance. The use of paddling appendages in conjunction with the rigid bodies of aquatic arthropods could potentially limit their turning performance. Whirligig beetles (Coleoptera: Gyrinidae) are rigid-bodied, but these aquatic insects can swim rapidly in circular patterns. Turning performance of swimming whirligig beetles (Dineutes horni) was assessed by videotaping beetles in a small (115 mm diameter) arena at 500 frames s(-1) and 1000 frames s(-1). Curved trajectories were executed as continuous powered turns. Asymmetrical paddling of the outboard legs was used to power the turn. Turns were produced also by abduction of the inboard elytra and vectored thrust generated from sculling of the wing at 47.14 Hz. The abducted elytra increased drag and acted as a pivot. Swimming speeds varied from 0.06 m s(-1) to 0.55 m s(-1) (4.7-44.5 L s(-1)). Relative minimum radius was 24% of body length. Maximum rate of turn was 4428 degrees s(-1) with maximum centripetal acceleration of 2.86 g. Turning radius was weakly associated with swimming velocity, although minimum values of the radius showed no correlation with velocity. Turning rate was also related indirectly to radius and directly to centripetal acceleration. Compared to vertebrates with flexible bodies, the relative turning radius of whirligig beetles is constrained by a rigid body and use of drag-based propulsive mechanisms. However, these mechanisms permit continuous turning, and the size of the beetle permits higher turn rates with lower centripetal accelerations.