Indirect closing of elytra by the prothorax in beetles (Coleoptera): general observations and exceptions.

Voluntary movements of the prothorax and the elytra in tethered flying beetles and manually induced movements of these parts in fresh dead beetles were recorded in 30 species representing 14 families. Participation of prothoracic elevation in the closing of the elytra was demonstrated in three ways. (i) The elevation was always simultaneous with elytral closing, in contrast to depression and elytral opening; a rare exception occurred in Lucanus cervus, whose elytra sometimes started to close before the cessation of wing strokes and the elevation of the prothorax. (ii) The manipulated elevation always induced closing of the spread elytra; the mechanical interaction between the hind edge of the pronotum and the roots of the elytra is a universal mechanism of closing the elytra in beetles. (iii) The prevention of pronoto-elytral contact in live beetles by the excision of the hind edge of the pronotum in front of the root prevented elytral closing after normal flight. Exceptions to this rule included some beetles that were able to close their elytra after such an excision: tiger beetles and diving beetles (seldomly) and rose chafers (always). This ability in Adephaga may be explained by attachments of the muscle actuating the 4th axillary plate, which differ from the attachments in Polyphaga. Cetoniinae open their elytra only by a small amount. It is proposed that their small direct adductors in combination with the elasticity of the sclerites are enough to achieve elytral closing without additional help from the prothorax.