Burrowing abilities and swash behavior of three crabs, Emerita analoga Stimpson, Blepharipoda occidentalis Randall, and Lepidopa californica Efford (Anomura, Hippoidea), of exposed sandy beaches.

To investigate factors related to the distribution of intertidal species, and specific predictions of the swash exclusion hypothesis for exposed sandy beaches, we compared the burrowing abilities and swash behavior of three species of anomuran crabs in the superfamily Hippoidea (Emerita analoga, Blepharipoda occidentalis and Lepidopa californica) which commonly inhabit the intertidal and shallow subtidal zones of beaches along the California coast. Burrowing times in the laboratory increased significantly with crab size for all species in five sediment grain sizes ranging from fine sand to gravel (0.15 to 3.24 mm). For each species, burrowing times differed significantly among sand grain sizes, ranging from 0.3 to 21.5 s. Burrowing times for the hippid crab, E. analoga, were relatively constant across sediment types, while those of the albuneid crabs, B. occidentalis and L. californica, were rapid in fine to medium sands, and much slower in coarser sediments. Our results indicate that E. analoga is a substrate generalist while L. californica and B. occidentalis are substrate sensitive. Pre-burrowing times and behavior, distance moved, and burrowing times differed among the species in the swash zone. Combined times of preburrowing and burrowing were shorter than the swash period (6 s) for most E. analoga individuals. Fifty percent of the individuals of L. californica reached the substrate and burrowed in the swash period, while no individuals of B. occidentalis burrowed in that time. Pre-burrowing behavior and time may be valuable in explaining spatial and temporal patterns in the distribution of hippoid crabs on California beaches. Our results support predictions of the swash exclusion hypothesis concerning the burrowing and locomotory abilities of sandy beach macrofauna. The substrate generalist characteristics, and unique orientation and swimming abilities of the hippid crab, E. analoga, in intertidal swash may help explain the success of this species and its congeners, and have important implications for understanding patterns of macrofauna community structure on exposed sandy beaches in California and other regions.