In Tanyderidae, Blepharoceridae, and Cylindrotominae, the aedeagus of the male carries three seminal ducts which open independently. In the female, similarly, there are three spermathecae and their ducts lead to independent openings on the ninth sternum, spaced to correspond to those of the male. The aedeagal ducts are very narrow and arise from an ejaculatory pump supplied by the common duct of the testes; and during mating the aedeagal openings are closely applied to the spermathecal openings or inserted into the spermathecal ducts. The ejaculatory pump and closed system during mating occur in many, but not all, Nematocera and perhaps throughout the Brachycera, but the tripartite condition of the organs of sperm transfer is reduced to a bipartite condition in Phlebotominae and to a ’simple’ condition, with only a single opening of the aedeagus and spermathecal system, in nearly all other Diptera. The spermathecae themselves, however, remain three in number almost throughout the order, though their ducts unite before opening; and this pattern must have been derived from the tripartite pattern of the Tanyderidae, etc., which represents the basic condition in the order.
In the Culiciformia (Culicidae, Chironomidae, Simuliidae, etc.) on the other hand, there is no ejaculatory pump but its place is taken by a wide duct lined with folded cuticle and covered by a muscular coat. The aedeagus is not suited for close apposition to the spermathecal opening. During mating a larger quantity of sperm-containing material is deposited in the surrounding atrium, as a liquid (Culicidae) or often as a spermatophore. Sperm transfer can be completed rapidly, as the pair meet in flight in the swarm. One primitive family, the Dixidae, constitutes an exception to these generalizations.
The Bibionomorpha, except for Anisopus, were not studied.
A brief comment is made on the characteristic small size of the dipterous spermatheca and the fineness of the spermathecal ducts.