Geographic mosaic of mutually exclusive dominance of obligate commensals in symbiotic communities associated with a burrowing echiuran worm

Species richness and community composition of symbionts associated with the burrowing echiuran worm, Ochetostoma erythrogrammon Leuckart & Rüppell, 1828, were quantitatively surveyed on subtropical intertidal flats in the Ryukyu Archipelago, Japan. Overall, we recorded seven species of burrow associates, including at least six obligate commensals. According to symbiont community composition, the study sites were largely divided into two groups; one was characterized by the dominance of a snapping shrimp, Alpheus barbatus Coutière, 1897, and the other by the dominance of a scale worm, Lepidonotus sp. Furthermore, a granulometric analysis showed that the sediment characteristics differed significantly between shrimp- and scale worm-dominant sites in terms of median diameter and inclusive graphic standard deviation. These results suggest that these symbionts have different habitat requirements, resulting in the mutually exclusive dominance pattern. Our findings indicate that habitat heterogeneity is important to the evaluation and conservation of the symbiotic diversity in intertidal flats.