Differential elemental stoichiometry of two Mediterranean evergreen woody plants over a geochemically heterogeneous area

Leaf nutrient composition and stoichiometry reflect complex interactions of the plant with its environment and are useful traits to explore ecological processes and relationships. In the present study, the foliar elemental compositions of two common Mediterranean woody species, the evergreen broad-leaved Quercus ilex and the coniferous Pinus pinaster growing in an area of Central Italy known for geochemical and geothermal anomalies, were investigated. To assess the site-specific and age-dependent pattern of foliar composition and stoichiometry, macronutrients (C, N, P, K, Mg, S) and trace elements (Al, As, Ba, Cd, Cr, Cu, Fe, Hg, Ni, Pb, Sb, V, Zn) were determined in leaves and needles of three different ages (6-, 12- and 24-month-old) collected from metalliferous (geothermal, mining) and rural areas. Leaves of Q. ilex showed comparatively high concentrations of micronutrients (i.e., Cu, Fe and Zn), while needles of P. pinaster accumulated significantly high concentrations of potentially toxic elements (i.e., As, Pb and S). No significant trend was found in elemental concentrations in relation to the age of leaves and needles. Multi-element stoichiometry of P. pinaster was driven by the geochemical heterogeneity of the sites, suggesting plastic adaptation at the sites with the most selective edaphoclimatic conditions (i.e., patches with nutrient poor and metalliferous soils). On the other hand, the content of both nutrients and potentially toxic elements in Q. ilex leaves varied little across the study area, reflecting stoichiometric stability; this is consistent with the ecophysiological features of Q. ilex as a late-successional species with a dominant role in the ecosystems of the Mediterranean area. Our findings demonstrate the value of foliar stoichiometric traits for understanding plant adaptation in a heterogeneous environment and also the consequences of biotic interactions during succession.