A Third Type of Defensive Behavior in the Tenebrionid Beetle Zophobas atratus Pupae

Pupae of the tenebrionid beetle Zophobas atratus Fabricius (Coleoptera: Tenebrionidae) exhibit two types of reflex abdominal motions in response to tactile stimulation: circular rotation and lateral bending to close pinching devices (gin-traps). In the present study, the pupa exhibited novel, sequential abdominal movements at 0.3-2.2 sec after the onset of mechanical stimulation. The most effective stimulation was gentle, double brushing on the ventral surface of an abdominal segment (sternite). The sequential abdominal movements consisted of the following three types of discrete elementary motions (100-350 ms in duration): rapid vibration of 30-40 Hz, circular rotation (or swing), and small wiggling movements. A sequence of abdominal movements generally started with a few bouts of vibration, but the number and order of subsequent motions varied considerably among different sessions and conditions. A restrained pupa often showed a prolonged sequence of many motions, including several rotations, whereas an unrestrained pupa often shortened the sequence by skipping a few rotations after the displacement of its whole body induced by the first abdominal rotation. Stimulation of two types of mechanosensitive sensilla, the hair sensilla (touch sensors) and campaniform sensilla (strain sensors), seemed to be necessary to initiate the defensive response. In natural environments, crawling of a small predator (or parasitoid) on the surface of the abdomen or repeated attacks of a large predator may induce this defensive response in the pupae.