Functional morphology of the praying mantis male genitalia (Insecta: Mantodea).

Male genitalia in praying mantids are highly complex, but we know little of how they function. I combined the micro-computed tomography of a copulating pair of the European mantis (Mantis religiosa) with public videos of copulation in various species of Mantodea and an analysis of literature. The function of each major element is reviewed. Copulation is divided into three phases: opening, anchoring and deposition. The opening is achieved by pulling the female subgenital plate with the male apical process. Multiple cases of female cooperation or resistance were observed and one case of coercion by the male. In species with the reduced apical process, female cooperation is mandatory. The male subgenital plate may participate in the opening as an integral part of the genitalia. After the opening, the conformation of the genitalia drastically changes, revealing activity of the genital papilla. Tight grasp on female genitalia is maintained solely by the clamp on the right phallomere, despite the overall complexity and predictions of sexual conflict theory. Other prominent elements show rhythmic motions, but their functions are not entirely clear and evidently involve spermatophore deposition, female stimulation or rival sperm removal. The opening and anchoring are similar in Mantodea and Blattodea, but achieved with non-homologous elements.