Genital coupling, morphology and evolution of male holding structures in Cicadinae (Hemiptera: Cicadidae)

Abstract

Male and female genitalia include some of the most complex and morphologically diverse structures in Metazoa. Ornamentations in genitalia have been studied in several groups, and a variety of functional roles have been proposed. Although complex features of the genitalia have been observed in internal genitalia in cicadas, their functions have not yet been elucidated. These ornamentations, together with precopulatory sexual selection, make cicadas good models for evolutionary studies on genital coupling. We explore the structural interaction of male and female genitalia in Guyalna bonaerensis (Berg) (Cicadinae) and the morphology of male ornamentations in Cicadinae generally. We group these ornamentations into two traits according to their inferred function: anchoring or gripping. We analyse the theca and vesica of 24 species and perform ancestral trait reconstruction under maximum likelihood and stochastic mapping on a Bayesian tree. Ornamentations of the male vesica and the female seminal ampoule possibly ensure male attachment by working as an active lock to avoid the premature termination of intercourse. These ornamentations emerged independently in different lineages in Cicadinae, reinforcing the suggestion that they are important adaptations to achieve complete copulation. Our results foster questions for the field of sexual selection and associated mechanisms shaping the evolution of male and female genitalia.