Does larval food availability ultimately select for seasonal reproduction in marine invertebrates with feeding larvae? A field test of Crisp’s Rule with the temperate sea star Pisaster ochraceus

Most organisms reproduce seasonally, including most species of marine invertebrates that live across wide ranges in habitat, depth and environmental conditions. We asked: What ultimately selects for seasonal reproduction in benthic marine invertebrates with planktotrophic larvae (characteristic of ≥70% of marine invertebrates)? We hypothesized that seasonal variation in food available for the larvae ultimately selects for timing of adult reproduction. Testing this hypothesis requires a whole life cycle perspective and approach. Using a known proximate cue, daylength, we shifted gametogenesis in the laboratory by six months in a seasonally breeding temperate sea star, the Ochre Star Pisaster ochraceus (Brandt 1835). We were then able to induce spawning, fertilize gametes and culture resulting embryos in vitro to produce feeding larvae six months out of phase with natural photoperiod. We field-reared these out-of-season larvae, similarly produced in-season conspecifics and similar larvae of an aseasonally breeding asteroid, the Bat Star Patiria miniata (Brandt 1835) in mesh-covered flow-through containers that were deployed in seasonally contrasting oceanographic conditions reflecting different productivity regimes and larval food availability in spring and fall. Larval development and survival were similar between seasons: for example, planktonic larval duration was 48 d to first metamorphosis in spring vs. 45 d in fall for P. ochraceus. Hence, temporal variation in available phytoplankton may not be an ultimate selection factor acting on these larvae to regulate timing of seasonal reproduction in the adults. Alternatively, in this and other species of marine invertebrates, selection acting on early benthic juveniles or other life stages may determine the timing of adult reproduction.