Morphology-flow interactions lead to stage-selective vertical transport of larval sand dollars in shear flow

Many larvae and other plankton have complex and variable morphologies of unknown functional significance. We experimentally and theoretically investigated the functional consequences of the complex morphologies of larval sand dollars, Dendraster excentricus (Eschscholtz), for hydrodynamic interactions between swimming and turbulent water motion. Vertical shearing flows (horizontal gradients of vertical flow) tilt organisms with simple geometries (e.g. spheres, ellipsoids), causing these organisms to move horizontally towards downwelling water and compromising their abilities to swim upwards. A biomechanical model of corresponding hydrodynamic interactions between turbulence-induced shear and the morphologically complex four-, six- and eight-armed stages of sand dollar larvae suggests that the movements of larval morphologies differ quantitatively and qualitatively across stages and shear intensities: at shear levels typical of calm conditions in estuarine and coastal environments, all modeled larval stages moved upward. However, at higher shears, modeled four- and eight-armed larvae moved towards downwelling, whereas six-armed larvae moved towards upwelling. We also experimentally quantified larval movement by tracking larvae swimming in low-intensity shear while simultaneously mapping the surrounding flow fields. Four- and eight-armed larvae moved into downwelling water, but six-armed larvae did not. Both the model and experiments suggest that stage-dependent changes to larval morphology lead to differences in larval movement: four- and eight-armed stages are more prone than the six-armed stage to moving into downwelling water. Our results suggest a mechanism by which differences can arise in the vertical distribution among larval stages. The ability to mitigate or exploit hydrodynamic interactions with shear is a functional consequence that potentially shapes larval evolution and development.