Comparative and functional morphology of wing coupling structures in Trichoptera: Annulipalpia

Several orders of morphologically four-winged insects have evolved mechanisms that enforce a union between the mesothoracic and metathoracic wings (forewings and hindwings) during the wing beat cycle. Such mechanisms result in a morphologically tetrapterous insect flying as if it were functionally dipterous, and these mechanisms have been described for several insect orders. The caddisfly suborders Annulipalpia and Integripalpia (Trichoptera) each have evolved a wing coupling apparatus, with at least three systems having evolved within the suborder Annulipalpia. The comparative and inferred functional morphology of the putative wing coupling mechanisms is described for the annulipalpian families Hydropsychidae (subfamilies Macronematinae and Hydropsychinae), Polycentropodidae and Ecnomidae, and a novel form-functional complex putatively involved with at-rest forewing-forewing coupling is described for Hydropsychidae: Smicrideinae. It is proposed that the morphology of the wing coupling apparatuses of Hydropsychinae and Macronematinae are apomorphies for those clades.