Life history diversity and evolution in the Asterinidae.

Asterinid sea stars have the greatest range of life histories known for the Asteroidea. Larval form in these sea stars has been modified in association with selection for planktonic, benthic, or intergonadal developmental habitats. Life history data are available for 31 species and molecular data for 28 of these. These data were used to assess life history evolution and relationships among asterinid clades. Lecithotrophy is prevalent in Asterinidae, with at least 6 independent origins of this developmental mode. Morphological differences in the attachment complex of brachiolaria larvae were evident among species with planktonic lecithotrophy. Some features are clade specific while others are variable within clades. Benthic brachiolariae are similar in Aquilonastra and Parvulastra with tripod-shaped larvae, while the bilobed sole-shaped larvae of Asterina species appear unique to this genus. Multiple transitions and pathways have been involved in the evolution of lecithotropy in the Asterinidae. Although several genera have a species with a planktonic feeding larva in a basal phylogenetic position, relative to species with planktonic or benthic lecithotrophy, there is little evidence for the expected life history transformation series from planktonic feeding, to planktonic non-feeding, to benthic non-feeding development. Intragonadal development, a life history pattern unique to the Asterinidae, arose three times through ancestors with benthic or pelagic lecithotrophy. Evolution of lecithotrophy appears more prevalent in the Asterinidae than other asteroid families. As diverse modes of development are discerned in cryptic species complexes, new insights into life history evolution in the Asterinidae are being generated.