In between breeding systems: neither dioecy nor androdioecy explains sexual polymorphism in functionally dioecious worms

Related species share genetic and developmental backgrounds. Therefore, separate-sex species that share recent common ancestors with hermaphroditic species may have hidden genetic variation for sex determination that causes some level of lability of expression of gender. Worms of the polychaete species Ophryotrocha labronica have separate, dimorphic sexes and their ancestor was hermaphroditic. Ophryotrocha labronica has a worldwide distribution and populations may differ in the degree of gender specialization. We analyzed the extent to which O. labronica had fixed or labile expression of gender. We found that there were up to four different sexual phenotypes, namely, pure males, males with oocytes, pure females, and females with sperm; the relative frequency of these sexual phenotypes varied in three geographically-distant populations. These sexual morphs had either male or female morphology. However, populations differed in the extent to which worms were sexually dimorphic. In the less dioecious-like population (in which pure males and females were virtually absent, all worms had both oocytes and sperm and sexual dimorphism was relatively weak), males with oocytes had slightly plastic female allocation that depended on mating opportunities-a clearly hermaphroditic trait. Males with oocytes and females with sperm were not functional hermaphrodites. They only used one type of gametes to reproduce and in this respect they probably differed from many cases of inconstancy of gender described in the literature. We consider these populations as novel examples of intermediate states between androdioecy and dioecy. This study contributes to our understanding of breeding systems as continuous gradients rather than as distinct clear-cut alternatives.