The shape of species abundance distributions across spatial scales

Species abundance distributions (SADs) describe community structure and are a key component of biodiversity theory and research. Although different distributions have been proposed to represent SADs at different scales, a systematic empirical assessment of how SAD shape varies across wide scale gradients is lacking. Here, we examined 11 empirical large-scale datasets for a wide range of taxa and used maximum likelihood methods to compare the fit of the logseries, lognormal, and multimodal (i.e., with multiple modes of abundance) models to SADs across a scale gradient spanning several orders of magnitude. Overall, there was a higher prevalence of multimodality for larger spatial extents, whereas the logseries was exclusively selected as best fit for smaller areas. For many communities the shape of the SAD at the largest spatial extent (either lognormal or multimodal) was conserved across the scale gradient, despite steep declines in area and taxonomic diversity sampled. Additionally, SAD shape was affected by species richness, but we did not detect a systematic effect of the total number of individuals. Our results reveal clear departures from the predictions of two major macroecological theories of biodiversity for SAD shape. Specifically, neither the Neutral Theory of Biodiversity (NTB) nor the Maximum Entropy Theory of Ecology (METE) are able to accommodate the variability in SAD shape we encountered. This is highlighted by the inadequacy of the logseries distribution at larger scales, contrary to predictions of the NTB, and by departures from METE expectation across scales. Importantly, neither theory accounts for multiple modes in SADs. We suggest our results are underpinned by both inter- and intraspecific spatial aggregation patterns, highlighting the importance of spatial distributions as determinants of biodiversity patterns. Critical developments for macroecological biodiversity theories remain in incorporating the effect of spatial scale, ecological heterogeneity and spatial aggregation patterns in determining SAD shape