Environmental Versus Phylogenetic Controls on Leaf Nitrogen and Phosphorous Concentrations in Vascular Plants

Global patterns of leaf nitrogen (N) and phosphorus (P) stoichiometry have been interpreted as reflecting phenotypic plasticity in response to the environment, or as an overriding effect of the distribution of species growing in their biogeochemical niches. Here, we balance these contrasting views. We compile a global dataset of 36,413 paired observations of leaf N and P concentrations, taxonomy and 45 environmental covariates, covering 7,549 sites and 3,700 species, to investigate how species identity and environmental variables control variations in mass-based leaf N and P concentrations, and the N:P ratio. We find within-species variation contributes around half of the total variation, with 29%, 31%, and 22% of leaf N, P, and N:P variation, respectively, explained by environmental variables. Within-species plasticity along environmental gradients varies across species and is highest for leaf N:P and lowest for leaf N. We identified effects of environmental variables on within-species variation using random forest models, whereas effects were largely missed by widely used linear mixed-effect models. Our analysis demonstrates a substantial influence of the environment in driving plastic responses of leaf N, P, and N:P within species, which challenges reports of a fixed biogeochemical niche and the overriding importance of species distributions in shaping global patterns of leaf N and P.