Tympanal hearing in tachinid flies (Diptera, Tachinidae, Ormiini): the comparative morphology of an innovation.

Tympanal hearing organs have been reported only recently for Diptera. All the cases documented so far relate to parasitoid tachinid flies of the ormiine tribe. In the ormiine flies, the presence of tympanal hearing is functionally linked to their reproductive behavior. Indeed, female ormiine flies detect and localize their host, typically singing orthopterans, by hearing their calling songs. The three ormiine fly species investigated here at the comparative level share the key morphological features associated with tympanal hearing. The extent of these structural modifications becomes evident in the light of comparison with the closely related atympanate tachinid Myiopharus doryphorae. We document a series of eight characters that constitute specialized modifications of the ventral prothorax: (1) an inflation of the probasisternum, providing a rigid frame to span the large tympanal membranes; (2) an increased surface area of the prosternal membranes that constitute very thin, corrugated tympanal membranes; (3) a forked, broad presternum with tympanal pits to which the sensory organs directly attach; (4) several modifications of the tracheal system comprising the enlargement of the prosternal air sac, a supplementary tracheal tube to the prosternal air sac accompanied by a subpartioning of the spiracular atrium, and larger mesothoracic spiracles; (5) the presence of two scolopophorous chordotonal organs in the unpartitioned prosternal air sac; (6) stiff cuticular apodemes linking the chordotonal organs to the presternum; (7) reduction in size of the cervical sclerites; and (8) several structural modifications of the prosternal apophyses, creating new attachment sites for neck muscles. This comparative approach brings out differences and similarities of the homologous cuticular structures found on the ventral prothorax of both tympanate and atympanate tachinids. It is proposed that, given the degree of similarity between the ormiine hearing organs, the ormiine tribe is monophyletic, whereby all members of this tribe evolved from a common ancestor, an acoustic parasitoid of a singing orthopteran insect.