

Future studies should better integrate disturbance in terms of temporal dynamics and dispersal rates in the EMS approach. We argue that sensitive species may survive in shallower water, which are more stable with regard to oxygen conditions and present greater habitat complexity, in contrast to deeper waters, which may experience periodic disturbance due to hypoxia. Species richness declined with depth (from 0.5 to 44.8 m). A quasi-nested clumped species loss pattern was also observed, which suggests nested habitat specialization. A quasi-Clementsian pattern was observed frequently, suggesting interdependent ecological relationships among species or similar response to an underlying environmental gradient across sites. We demonstrated several patterns associated with environmental variation and associated processes that could simultaneously assemble species to occur at the sites. We applied the elements of metacommunity structure (EMS) approach, allowing multiple hypothesis of variation in community structure to be tested. Metacommunity structure of soft-sediment benthic invertebrates was examined using a fine-grained (285 sites) data set collected during one summer across a large spatial extent (1700 km 2).

These systems are more open than many other systems and may thus exhibit relatively less discrete patterns in community structure across space.

The majority of studies in metacommunity ecology have focused on systems other than marine benthic ecosystems, thereby providing an impetus to broaden the focus of metacommunity research to comprise marine systems.
