Courtship, copulation, and sperm trading in the sea slug, Navanax inermis (Opisthobranchia: Cephalaspidea)
Navanax inermis are internally fertilizing, simultaneous hermaphrodites that normally mate in pairs. Copulations usually occur in bouts with active alternation of sexual roles and function. Over the study, individuals copulated equally often and for equal periods of time as each sex. The sequence of behaviors involved in courtship, copulation, and role switching is described. While the sequence of behaviors associated with each sexual role is quite stereotyped, the temporal relationship between the behaviors of the two partners is variable. A mating bout is initiated when one individual assumes the male role, finding and following a conspecific mucous trail, and then courts and copulates as a male. Each individual initiated approximately one-half of the copulatory bouts in which it was involved; size played no role in determining which individual initiated the bout. Our observations suggest that in N. inermis, individuals are always willing to be female, while willingness to be male varies over time. The mating system of N. inermis is based on sperm trading. Individuals apparently act as males, donating sperm, to obtain sperm, and are more willing to be fertilized than to fertilize, in violation of Bateman's principle. In N. inermis, females control fertilization and therefore copulation involves greater risks for males than females. The sperm-trading system is "enforced" by the maintenance of intromission until the partner shows courtship behaviour.