Bioengineers and their associated fauna respond differently to the effects of biogeography and upwelling

Temperature and primary production (often linked to nutrient supply) are two of the few factors influencing species diversity and abundances across mesoscale gradients, while at smaller scales the habitat complexity offered by bioengineers is important. Previous studies have illustrated effects of upwelling and biogeography on intertidal bioengineers, but it is not known if these processes influence assemblages associated with those bioengineers in a similar way. We examined the habitat structure offered by two species of mussels and their associated fauna in five regions across 3000 km and three biogeographic provinces of the South African coast, replicating upwelling and non-upwelling areas within each region. Upwelling and region influenced the structure of mussel beds (the density and size of mussels). In contrast, upwelling did not influence mesoscale differences in composition, abundance, and numbers of species of crustaceans, mollusks, or polychaetes in mussel beds. Regardless of trophic level or mode of reproduction, mussel bed fauna were influenced only by region. Regional differences were strongly influenced by biogeography. The associated fauna was, however, also strongly correlated with the structure of the habitats created by mussels. Our results support the importance of upwelling to a critical ecosystem engineer, but show that these effects do not extend directly to the assemblages of associated fauna, which are more influenced by regional-scale effects and biogeography. We suggest that mesoscale patterns in the associated fauna of this bioengineered habitat are driven by the direct effects of biogeography, combined with the influence of biogeography and upwelling on mussel bed structure.