What is a juvenile sea urchin? A comparative and phylogenetic survey of post-metamorphic juveniles

Abstract

Echinoid development progresses through embryonic and larval stages to metamorphosis and the adult form. Despite vast differences in embryos and larval forms, including bilaterally symmetric echinopluteus larvae, ovoid non-feeding larvae and brooded embryos, all metamorphose into juvenile sea urchins with pentaradial symmetry. The adult sea urchin body plan is initiated as the juvenile rudiment. The rudiment has been called the phylotypic stage for the class Echinoidea, a designation that implies little variation at this midpoint in development (e.g. Raff et al., 1991; Richardson, 1995; Raff, 1996). However, right at metamorphosis (upon eversion of the juvenile rudiment), variations in test symmetry, shape and number of spines, and number of skeletal plates, podia and pedicellariae are present in juveniles. This variation suggests either that there is no phylotypic stage or that such a stage occurs earlier in rudiment formation. To distinguish between these possibilities, I explored the patterns by which the juvenile rudiment is formed as well as the variation among juveniles approximately 1 day after metamorphosis in 19 echinoid taxa covering a broad taxonomic range including cidaroids, diadematids, irregular echinoids (spatangoids and clypeasteroids), arbaciids, temnopleurids, echinometrids and strongylocentrotids. Most of the material for analysis of juveniles was obtained by the author. Additional information was gathered from classical studies of metamorphosis. Data were collected on the number and shape of dorsal pedicellariae, juvenile and adult spines, primary and secondary podia, and juvenile test shape. When possible multiple individuals within a species were examined, revealing no or only minor trait variation. These data were mapped on a well-resolved phylogeny established from adult characters.