Different roles for geography, energy and environment in determining three facets of freshwater molluscan beta diversity at broad spatial scales

Current understanding of different facets of beta diversity and their underlying determinants remains limited at broad scales in the freshwater realm. We examined the geographical patterns and spatial congruence of three beta diversity facets of freshwater molluscs across all of China, and evaluated the relative importance of environmental and spatial factors underlying the observed patterns. Taxonomic (β-TD), functional (β-FD) and phylogenetic (β-PD) beta diversity were calculated for 212 drainage basins belonging to 10 hydrographic regions using compiled occurrence data of 313 molluscan species. Geographical patterns of the three diversity facets were visualized on maps and pairwise spatial congruence among them was evaluated using regression on distance matrices. Variation partitioning and multivariate regression trees were used to assess the relative importance of different factors underlying beta diversity patterns. Beta diversity maps revealed that geographical patterns of β-TD and β-PD showed strong spatial clustering and were well matched with hydrographic regions' boundaries, while β-FD showed only moderate spatial aggregation. The three facets were only moderately congruent, with over 60% of the variation in one facet remaining unexplained by any other facet. Remarkably, all diversity facets were best explained by the spatial factors with considerable unique effects. Environmental filtering associated with energy gradients also made a large contribution, while habitat availability only explained minor fractions of the variation in beta diversity. At the national scale, β-TD and β-PD were more strongly related to spatial processes, whereas β-FD was more strongly associated with energy gradients. Our results suggested that, for freshwater organisms with low dispersal capacity, dispersal processes may override environmental filtering in driving geographical diversity patterns. However, different ecological drivers were important for each diversity facet. Importantly, rather weak spatial congruence among the different diversity facets stresses the need to incorporate functional and phylogenetic facets into the development of conservation planning.