Measurements of area and the (island) species-area relationship:: new directions for an old pattern

The species-area relationship is one of the strongest empirical generalizations in geographical ecology, yet controversy persists about some important questions concerning its causality and application. Here, using more accurate measures of island surface size for five different island systems, we show that increasing the accuracy of the estimation of area has negligible impact on the fit and form of the species-area relationship, even though our analyses included some of the most topographically diverse island groups in the world. In addition, we show that the inclusion of general measurements of environmental heterogeneity (in the form of the so-called choros model), can substantially improve the descriptive power of models of island species number. We suggest that quantification of other variables, apart from area, that are also critical for the establishment of biodiversity and at the same time have high explanatory power (such as island age, distance, productivity, energy, and environmental heterogeneity), is necessary if we are to build up a more predictive science of species richness variation across island systems.