Plasticity, Constraint, and Optimality in Reproductive Timing

Synchronous release of gametes or larvae by marine animals may be controlled by as many as four environmental cycles thereby enabling an array of reproductive timing patterns. Cohesive scenarios that account for the diversity of reproductive patterns are rare and tests of their adaptive significance have been even rarer. By exploiting plasticity in the timing of larval release, we isolated proximate factors regulating reproductive synchrony and provided evidence that predation ultimately may best explain the diversity of hatching patterns by brachyuran crabs. Tides entrain reproductive rhythms by many crabs, and therefore spatial and temporal variation in tides produces intraspecific variation in reproductive patterns. This was demonstrated by determining the timing of larval release of the same or sibling species of intertidal crabs in Pacific semidiurnal and Caribbean mixed semidiurnal tidal regimes on the two coasts of the Republic of Panama. The time of larval release varied during the year in the Caribbean, where entraining physical cycles exhibited complicated changes in phase, but not along the Pacific coast, where the phase relationships among physical cycles varied little year-round. Crabs timed larval release relative to the light-dark, tidal phase, and tidal amplitude cycles, but not the lunar cycle, suggesting that three rhythms determined when larvae were released. For each species we ranked these rhythms by the degree to which larval release kept phase with their entraining physical cycles. The species-specific hierarchies of rhythms we observed match those expected if the time of larval release minimizes predation on females, embryos, and newly hatched larvae. Such hierarchies enable crabs to track phase shifts of cycles in variable tidal environments and may enhance reproductive success across tidal regimes. However, larval release may be timed better in some tidal regimes than others due to differences in the phasing of environmental cycles. In some tidal regimes, larval release cannot be synchronized with all three physical cycles during the year, and hierarchies of rhythms may determine the timing and duration of breeding.