Species introductions and the phylogenetic and functional structure of California's grasses

Many species assemblages represent a nonrandom subset of a larger species pool. When an assemblage tends to contain close evolutionary relatives or species with similar functional traits, it can be described as phylogenetically or functionally clustered. Clustering is often interpreted as evidence for filtering by some combination of environmental and biotic factors. At sufficiently large spatial extents, however, biogeographic barriers can also lead to strong clustering. Here, we suggest that the breakdown of biogeographic barriers associated with human introductions of exotic species can be used as an unintentional experiment to assess their importance in driving phylogenetic and functional structure. An important role of biogeographic barriers would be revealed by a breakdown in clustering, particularly phylogenetic clustering, following species introductions. On the other hand, a role of filtering can be supported by similar patterns of clustering in the native and exotic assemblages along environmental gradients. We test these predictions using the grasses of California, a diverse group including many introduced species. Native grass assemblages in the state are highly clustered with respect to the global grass species pool, both phylogenetically and functionally. Within the state, variation in the strength of clustering is well explained by climatic variables, suggesting an important role for environmental-biotic filtering. Further, subregions within the state with highly clustered native assemblages also contain highly clustered exotic assemblages. Contrary to expectation, though, the introduction of exotic species led to even more strongly clustered assemblages. We conclude that biogeographic barriers have generally not excluded the major grass lineages (e.g., tribes) from the state and likely act only on finer taxonomic scales (for example, excluding particular genera). Our approach should prove broadly applicable and contribute to improved understanding of broad-scale patterns of assemblage structure.