Burrowing Criteria and Burrowing Mode Adjustment in Bivalves to Varying Geoenvironmental Conditions in Intertidal Flats and Beaches

The response of bivalves to their abiotic environment has been widely studied in relation to hydroenvironmental conditions, sediment types and sediment grain sizes. However, the possible role of varying geoenvironmental conditions in their habitats remains poorly understood. Here, we show that the hardness of the surficial intertidal sediments varies by a factor of 20-50 due to suction development and suction-induced void state changes in the essentially saturated states of intertidal flats and beaches. We investigated the response of two species of bivalves, Ruditapes philippinarum and Donax semigranosus, in the laboratory by simulating such prevailing geoenvironmental conditions in the field. The experimental results demonstrate that the bivalve responses depended strongly on the varying geoenvironmental conditions. Notably, both bivalves consistently shifted their burrowing modes, reducing the burrowing angle and burial depth, in response to increasing hardness, to compensate for the excessive energy required for burrowing, as explained by a proposed conceptual model. This burrowing mode adjustment was accompanied by two burrowing criteria below or above which the bivalves accomplished vertical burrowing or failed to burrow, respectively. The suitable and fatal conditions differed markedly with species and shell lengths. The acute sensitivities of the observed bivalve responses to geoenvironmental changes revealed two distinctive mechanisms accounting for the adult-juvenile spatial distributions of Ruditapes philippinarum and the behavioral adaptation to a rapidly changing geoenvironment of Donax semigranosus. The present results may provide a rational basis by which to understand the ensuing, and to predict future, bivalve responses to geoenvironmental changes in intertidal zones.