Environmental drivers of tree species richness in the southernmost portion of the Paranaense forests

Background The Rio de la Plata grassland region is dominated by temperate grasslands, with the scarce natural forests, influenced floristically by adjacent biogeographical provinces. Uruguay represents the southern limit for many tree species of the Paranaense Province, several of which inhabit the hillside forests. With many species shifting poleward due to climate change, we do not yet know how current environmental factors, particularly climatic ones, are linked to the tree diversity of this flora nowadays. The aim of this study is to understand the geographic pattern of tree richness in the hillside forests of Uruguay, evaluating the water-energy and the environmental heterogeneity hypotheses. The distribution of the hillside forest trees was obtained by compiling and updating the herbaria database and distribution maps of woody plants of Uruguay. The presence/absence of each species, and then the species richness, were georeferenced over a grid that covers Uruguay with 302 cells (660 km2). Over the same grid were compiled environmental variables associated with climate and environmental heterogeneity. The relationship between richness and environmental variables was studied by applying general linear models (GLM). As a strong autocorrelation was detected, a residuals auto-covariate term was incorporated into the GLM, to take into account the species richness spatial structure.Results The tree flora of the hillside forest was composed mainly by Paranaense species that show a latitudinal gradient, with two high richness cores, in the east and northeast of Uruguay. The final model including the environmental variables and the spatial term explained 84% of the variability of tree richness. Species richness showed a positive relationship with precipitation, forest cover, potential evapotranspiration and productivity, while a negative effect of temperature variation was found. The spatial component was the primary predictor, accounting for a 30% of spatial pattern of tree richness.Conclusions This study accounts for a large proportion of the environmental and spatial variations of the tree richness pattern of the Paranense flora in its southernmost portion. It brings support to both water-energy and environmental heterogeneity hypotheses, emphasizing the role of climate and its variation and the habitat availability on the hillside forest diversity.