Evolution of the middle leg basal articulations in flies (Diptera)

The morphology of the coxa and trochanter was studied in 205 species from 68 fly families to compare these structures with respect to ability to fly in a streamlined posture, with the middle legs pointing forward and pressed to the thorax. Only Brachycera are able to attain this posture. The forward turn of the coxa at this position is hindered by the junction of the coxa with the pleuron. Recovery of mobility is gained in two ways. (1) By reduction of the contact zone between coxa and pleurite, as in Asiloidea, Bombyloidea, and Empidoidea. Within these flies, the streamlined posture was recorded in Bombyliidae and in a robber-fly, Laphria flava . Others fly with their middle legs straddled laterally or trailing backwards. (2) Longitudinal splitting of the coxa into three coxites provides intracoxal mobility in most Tabanoidea and Cyclorrhapha. The hind and medial coxites rotate about the front coxite and change the coxo-trochanteral axis, thus compensating for restricted protraction. Separation of the hind coxite appears in primitive Tabanoidea, and a separate middle coxite was found in several families among the Nematocera. The streamlined posture was recorded in horse-flies, stratiomyids, and in many Cyclorrhapha except Micropezidae and Hippoboscidae. There is morphological evidence for a possible secondary fusion of coxites at least in Dolichopodidae and Opetidae as well as for the origin of Cyclorrhapha from a miniature ancestor.