Inverted sand dollars actively orient themselves in flow to increase likelihood of righting

The fact that sand dollars are often dislodged and inverted is an inescapable consequence of living at or slightly below the sediment–water interface. Once inverted, however, how do sand dollars effectively right themselves, given their small spines and stiff internal skeletons? Here, we examined the possibility that individuals of Mellita quinquiesperforata and Dendraster excentricus may take advantage of the interaction of their morphology and flow to increase the likelihood of righting. Based on flow tank observations, the critical velocity required to flip an inverted sand dollar varies with orientation and increases with test size. For both species, the critical velocity was lower when inverted sand dollars were oriented with the posterior margin facing directly downstream, compared with when the posterior margin was positioned in an upstream orientation. To test whether inverted sand dollars would actively rotate into a more advantageous position for flipping, we exposed inverted animals in three starting orientations – with their posterior edge directed upstream (the least favored position for flipping), perpendicular, and downstream to flow – to the minimum flow expected to induce flipping and compared their responses. Time‐lapse photography showed that regardless of initial orientation, within one hour, a majority of individuals of both species rotated into positions that were not statistically different from the downstream orientation (the most favored position for flipping). These results for D. excentricus were further confirmed in a field experiment. Taken together, these data suggest that inverted sand dollars are able to recognize flow direction and respond by modifying their orientation to maximize lift and drag for righting.