Species diversity in the soft-bottom intertidal zone: Biogenic structure, sediment, and macrofauna across mussel bed spatial scales

Mussels are important ecosystem engineers that have significant impacts on the ecology of the seafloor. In this study, we attempt to characterize and explain infaunal and epifaunal abundance patterns across scales in a spatially complex, intertidal, soft-bottom habitat: beds of the semi-infaunal mussel, Mytilus edulis. We used univariate and multivariate analyses in a nested sampling design at three locations in eastern Maine to show the relationships between environmental and macrofaunal assemblage variables. The results support the hypotheses that: 1) environmental and macrofaunal assemblage variables vary widely across spatial scales and exhibit maximum variance at different spatial scales; 2) abundance and diversity patterns of macrofauna are linked to spatial variation in environmental variables in the form of biogenic structure (live mussels and mussel shell hash), terrestrially-derived gravel, and, to a lesser degree, the proportion of silt-clay in the sediment; and 3) epifauna are generally positively correlated with hard-substrate features such as live mussels and gravel, and negatively correlated with the silt-clay content of the sediment, although the latter relationships were not strong. On the other hand, the results did not support the hypotheses that: 4) most infauna, specifically those with free-swimming larvae, are negatively correlated with live mussels and silt-clay proportion; instead, they were often positively correlated with live mussels and had weak, insignificant correlations with silt-clay; and 5) infauna without free-swimming larvae are positively correlated with live mussels and silt-clay proportion; instead, they showed some positive correlations with live mussels, but a mix of weak, insignificant correlations with silt-clay. Thus, mussel beds had a variety of scale-dependent relationships, both positive and negative, with each of the macrofaunal groups. The results show that biogenic structure in the form of live mussels and mussel shell hash is important in this system, as is terrestrial gravel, and that these variables are more important than silt-clay content for the macrofaunal assemblage. Mussel beds play an important role in regulating macrofaunal diversity in soft-bottom habitats, and their effects vary across spatial scales.