Independent effects of patch size and structural complexity on diversity of benthic macroinvertebrates.

Despite a long history of work on relationships between area and number of species, the details of mechanisms causing patterns have eluded ecologists. The general principle that the number of species increases with the area sampled is often attributed to a sampling artifact due to larger areas containing greater numbers of individuals. We manipulated the patch size and surface area of experimental mimics of macro-algae to test several models that can explain the relationship between abundance and species richness of assemblages colonizing different habitats. Our results show that patch size and structural complexity have independent effects on assemblages of macroinvertebrates. Regardless of their structural complexity, larger habitats were colonized by more species. Patch size did not have a significant effect on numbers of individuals, so the increased number of species in larger habitats was not simply a result of random placement associated with sampling increased number of individuals. Similarly, random placement alone could not explain differences in numbers of species among habitats with different structural complexity, contrary to suggestions that the relationship between number of species and surface area might also be a sampling artifact due to more complex habitats having larger areas and therefore sampling more individuals. Future progress would benefit from manipulating properties of habitat in conjunction with experimental manipulations of area.