Macrofaunal diversity and habitat structure in intertidal boulder fields

Low-midshore boulder fields in Europe are increasingly subject to degradation related to beach fishing for crabs and abalones. The aim of the study was to better understand the structure and species diversity of macrofaunal assemblages living in a low-midshore boulder field in order to define conservation strategies for this biotope. Sampling strategy involved different spatial scales (macro- and microstrata) relative to the complexity and heterogeneity of the habitat. Assemblages of species and the corresponding habitats were identified by multifactorial analysis and compared by ANOVAs. The results show a hierarchical organization of the macrofauna within the boulder field, corresponding to three spatial levels of habitat structure: (i) eight microhabitats at the lowest level of spatial organization, each defined by a specific assemblage (subcommunity); (ii) three habitats at a middle level combining these microhabitats, each associated with a specific community (open rock, protected rock and sediment); and (iii) three macrohabitats at the uppermost level (corresponding to the scale of the entire boulder field and including the main geomorphological features of the beach), each defined by a specific assemblage of species (boulders on boulders, boulders on bedrock, and boulders on sediment). Two microhabitats with particularly high species diversity were regarded as the most important ecological units of the field in terms of conservation of specific and functional biodiversity. Comparison of habitat/community parameters showed (i) that habitat heterogeneity was not an accurate indicator of faunal diversity, and (ii) that habitat complexity enhanced the species richness of the fauna, but only above a threshold value. This enhancement was due to semi-sheltered microhabitats, which were found only in the most complex areas of the boulder field. It is likely that this complexity affects species richness qualitatively more than by the diversity of microhabitats. In other words, a cross-scale effect is apparent in which high habitat complexity on the middle spatial scale creates microhabitats on the lowest spatial scale that are characterized by stable semi-sheltered environmental conditions conducive to a maximum of species.