Evaluating the correlation between area, environmental heterogeneity, and species richness using terrestrial isopods (Oniscidea) from the Pontine Islands (West Mediterranean)

Area and environmental heterogeneity influence species richness in islands. Whether area or environmental heterogeneity is more relevant in determining species richness is a central issue in island biogeography. Several models have been proposed, addressing the issue, and they can be reconducted to three main hypotheses developed to explain the species-area relationship: (1) the area-per se hypothesis (known also as the extinction-colonisation equilibrium), (2) the random placement (passive sampling), and the (3) environmental heterogeneity (habitat diversity). In this paper, considering also the possible influence of geographic distance on island species richness, we explore the correlation between area, environmental heterogeneity, and species richness by using faunistic data of Oniscidea inhabiting the Pontine Islands, a group of five small volcanic islands and several islets in the Tyrrhenian Sea, located about 60 km from the Italian mainland. We found that the colonisation of large Pontine Islands may occur via processes independent of geographic distance which could instead be an important factor at a much smaller scale. Such processes may be driven by a combination of anthropogenic influences and natural events. Even in very small-size island systems, environmental heterogeneity mostly contributes to species richness. Environmental heterogeneity could influence the taxocenosis structure and, ultimately, the number of species of Oniscidea via direct and indirect effects, these last mediated by area which may or may not have a direct effect on species richness.