Evo-Devo in Ophiuroids: The Switch from Planktotrophy to Lecithotrophy in Ophionereis

Abstract

Understanding the evolution of development (evo-devo) in the Ophiuroidea and the pathways in the switch from a feeding to a nonfeeding larva is complicated by the variability in the phenotype of the metamorphic larva, being a reduced yolky ophiopluteus in some species (type I development) and a vitellaria larva in others (type II development). We investigated evo-devo in the family Ophionereididae, a group dominated by lecithotrophic development through a vitellaria larva. We reared the planktotrophic larvae of Ophionereis fasciata to settlement to determine the metamorphic phenotype. Counter to expectations, O. fasciata did not exhibit type II metamorphosis through a vitellaria, although it did exhibit transient vitellaria-like features. Resorption of the larval arms in the same interradial positions where the ciliary bands form in vitellariae gave them a fleeting vitellaria-like appearance. Development of O. fasciata exhibits heterochronic features in early formation of the skeletal primordium of the third pair (postoral) of larval arms and in the presettlement juvenile early appearance of the juvenile terminal arm plates on external view in parallel with larval arm resorption. Development of the fourth pair (posterodorsal) of larval arms, the last pair to be formed, is plastic, with 44% of larvae exhibiting partial arm growth. Heterochronic traits in development, as seen in O. fasciata, may have facilitated evolution of a lecithotrophic mode of development in Ophionereis. Comparison of the ophiopluteus of O. fasciata and the vestigial pluteus of O. schayeri provided insights into the simplification of larval form from the ancestral (feeding larva) state in Ophionereis. The diverse metamorphic phenotypes in ophiuroids indicate that type I and type II development may not be completely divergent lines of evo-devo and point to selective pressure in the pelagic-benthic transition in the evolution of ophiuroid development.