Macroclimatic structuring of spatial phylogenetic turnover in liverworts

Phylogenetic turnover has emerged as a powerful tool to identify the mechanisms by which biological communities assemble. When significantly structured along environmental gradients, phylogenetic turnover evidences phylogenetic niche conservatism, a critical principle explaining patterns of species distributions at different spatio-temporal scales. Here, we quantify the contribution of geographic or macroclimatic drivers to explain patterns of phylogenetic turnover in an entire phylum of land plants, namely liverworts. We further determine whether climatic niche conservatism has constrained the distribution of liverworts in the course of their evolutionary history. Two datasets, one insular, focused on 60 archipelagos and including 2346 species, and the second global, including 6334 species in 451 oceanic and continental Operational Geographic Units (OGUs) worldwide, were assembled. Phylogenetic turnover among OGUs was quantified through πst statistics. πst-throughtime profiles were generated at 1 myr intervals along the phylogenetic time-scale and used to compute the correlation between πst, current geographic distance and macroclimatic variation with Mantel tests based on Moran spectral randomization to control for spatial autocorrelation. The contribution of macroclimatic variation to phylogenetic turnover was about four-times higher than that of geographic distance, a pattern that was consistently observed in island and global geographic settings, and with datasets including or excluding species-poor OGUs. The correlation between phylogenetic turnover and geographic distance rapidly decayed at increasing phylogenetic depth, whereas the relationship with macroclimatic variation remained constant until 100 mya. Our analyses reveal that changes in the phylogenetic composition among liverwort floras across the globe are primarily shaped by macroclimatic variation. They demonstrate the relevance of macroclimatic niche conservatism for the assembly of liverwort floras over very large spatial and evolutionary time scales, which may explain why such a pervasive biodiversity pattern as the increase of species richness towards the tropics also applies to organisms with high dispersal capacities.