Co-evolution of the mating position and male genitalia in insects: a case study of a hangingfly.

Hangingflies are unique for the male providing a nuptial gift to the female during mating and taking a face-to-face hanging copulation with the female. Their male genitalia are peculiar for an extremely elongated penisfilum, a pair of well-developed epandrial lobes (9th tergum), and a pair of degenerated gonostyli. However, the co-evolution of their face-to-face copulation behavior and the male genitalia has rarely been studied hitherto. In this paper the mating behavior of the hangingfly Bittacus planus Cheng, 1949 was observed under laboratory conditions, and the morphology of the male and female external genitalia was investigated using light and scanning electron microscopy. The male provides an insect prey as a nuptial gift to the female in courtship and mating process, and commits a face-to-face copulation. During copulation, the male abdomen twists temporarily about 180° to accommodate their face-to-face mating position. The aedeagal complex has an extremely elongated penisfilum, corresponding to the elongated spermathecal duct of the female. The well-developed epandrial lobes serve as claspers to grasp the female subgenital plate during copulation, replacing the function of gonostyli, which are greatly reduced in Bittacidae. The modified proctiger assists the penisfilum to stretch and to enter into the female spermathecal duct. The possible reasons why this species might mate face-to-face are briefly discussed.