Dispersal pathways, seed rains, and the dynamics of larval behavior.

Dispersing propagules (larvae, seeds, and spores) establish and maintain populations, which serve as templates for subsequent species interactions. Connectivity among demes derives, in large part, from connectivity between consecutive steps, release, transport, and settlement, in dispersal pathways. Concurrent measurements of individuals in each step are a necessary precursor to identifying governing mechanisms. Here we directly and definitively resolved the roles of physics and behavior in mediating dispersal pathways of an estuarine parasite between its intermediate hosts. Planktonic cercariae of Himasthla rhigedana, a parasitic flatworm, are functionally similar to lecithotrophic larvae of many free-living marine invertebrates. The combination of parasite life cycle characteristics and the relatively simple tidal flows in their habitat renders this system an effective model for dispersal studies. Simultaneous field measurements of larval release, transport, settlement, and the flow regime, together with mechanistic experiments, led to empirical understanding of host colonization. All dispersal steps were highly and significantly correlated over time and in space. This tight coupling resulted, unequivocally, from a suite of larval behaviors. Cercariae emerged from first intermediate host snails only during daytime flood tides, enhancing larval retention in the marsh. Daylight triggered downward swimming, and within seconds, cercariae overpowered turbulent mixing, landing in benthic habitat of second intermediate host snails and crabs. Larvae settled (encysted) on external regions of snails/crabs that, presumably, were most vulnerable to ingestion by definitive host shorebirds. In total, cercarial behaviors greatly foreshortened dispersal distances, magnified local parasite prevalence, and increased the likelihood of large-scale transmission by definitive hosts. Cracking open the black box of dispersal thus revealed mechanisms, connectivity, and ecological consequences of the larval stage.