The lock-and-key mechanisms of the internal genitalia of the Noctuidae (Lepidoptera): How are they selected for?

In the Noctuidae, the owlet moths, the internal genitalia, i.e. the aedeagus and vesica (penis) in the males, and the bursa copulatrix in the females, together form a lock-and-key mechanism (LKM). The species-specific structures have their counterparts in the opposite sex. The internal LKM constitutes a specific reproductive isolation mechanism (lock-and-key hypothesis), which seem to be the rule in the ditrysian Lepidoptera, and also Occurs in the Carabidae (Coleoptera) and some other insects. In contrast, the external genitalia rarely have species-specific counterparts in the sexes. Several results indicate the presence of LKMs: In the Noctuidae, (1) heterospecific differences in the male vesica may prevent sperm transfer or lead to mechanical failure during copulation, (2) the more complicated the specific genitalia structures, the more aberrations may occur even in conspecific copulations, and (3) in many species pairs and groups, and in one large genus, Apamea, the structures in the opposite sexes show a strictly specific correspondence, but, (4) when there is precopulatory isolation due to differences in pheromone production or perception, the internal genitalia may be identical. Conversely, in the Colias butterflies (Pieridae), (5) frequent heterospecific hybridization is associated with the similarity of the internal genitalia. The LKMs seem to protect genomes against alien genes, supposedly selected for because of the lower fitness of specimens with an imprecise LKM and/or inferiority of hybrids. In the literature, the diversity of the noctuid genitalia has been ascribed to sexual selection, because the females were classified as polyandrous. Most species produce the main part of their eggs monandrously, and remate, if at all, in their old age, and are thus successively monandrous and polyandrous. The allopatric divergence in the structure of the internal genitalia of 39 Holarctic pairs of sister species of Noctuidae is suggested to be due to genetic drift. The insecure function of the female pheromones and external genitalia of males are illustrated with the aid of original photographs.