Interspecific trait integration increases with environmental harshness: A case study along a metal toxicity gradient

Functional traits are commonly used to calculate a wide array of functional diversity indices to infer different mechanisms of community assembly and species coexistence. Recently, the degree of interspecific covariation between multiple functional traits has been suggested as a mechanism influencing both species distributions and abundances in communities. However, empirical assessments of this theory along environmental gradients are still scarce due to the lack of an appropriate method and of sufficiently strong environmental gradients. Here we compare interspecific trait integration (ITI) across plant communities along a marked gradient of copper toxicity in the soil, using new multivariate and bivariate indices. This was achieved using the range of the eigenvalues of a principal component analysis on the traits of the species in a local community (multivariate ITI index) and the correlations between traits in local communities (bivariate ITI index). We show that the plant metal tolerance strategy (i.e. leaf metal content) is relatively independent from leaf economics, while negatively correlated to plant size. In addition, our results indicate a weak support for the expected general patterns of trait syndromes, such as the 'leaf economics spectrum' or the 'leaf-height-seed', at the whole-community scale. This arises from an increase in multivariate trait integration along the soil copper gradient. The strongest trait integration is caused by an increase in the degree of association between certain traits on metal-rich soils. The multivariate trait integration explains species richness better than other commonly used functional diversity indices. Our study highlights the power of ITI, as well as its complementarity to other functional diversity indices, to investigate the variation in functional strategies and their drivers along environmental gradients. The increase in trait integration with soil metal toxicity in plant communities supports that highly constraining environments select increasingly coordinated sets of functional traits, in turn possibly driving the decrease in species richness. Further studies should assess the generality and underlying physiological mechanisms of such ecological patterns.