There is evidence that N concentration in hardwood leaf litter is reduced when plants are raised in an elevated CO2 atmosphere. Reductions in the N concentration of leaf litter have been found for tree species raised under elevated CO2, with reduction in N concentration ranging from ca. 50% for sweet chestnut (Castanea sativa) to 19% for sycamore (Acer platanoides). However, the effects of elevated CO2 on the chemical composition of litter has been investigated only for a limited number of species. There is also little information on the effects of increased CO2 on the quality of root tissues. If we consider, for example, two important European forest ecosystem types, the dominant species investigated for chemical changes are just a few. Thus, there are whole terrestrial ecosystems in which not a single species has been investigated, meaning that the observed effects of a raised CO2 level on plant litter actually has a large error source. Few reports present data on the effects of elevated CO2 on litter nutrients other than N, which limits our ability to predict the effects of elevated CO2 on litter quality and thus on its decomposability. In litter decomposition three separate steps are seen: (i) the initial stages, (ii) the later stages, and (iii) the final stages. The concept of "substrate quality," translated into chemical composition, will thus change between early stages of decomposition and later ones, with a balanced proportion of nutrients (e.g., N, P, S) being required in the early decomposition phase. In the later stages decomposition rates are ruled by lignin degradation and that process is regulated by the availability of certain nutrients (e.g., N, Mn), which act as signals to the lignin-degrading soil microflora. In the final stages the decomposition comes to a stop or may reach an extremely low decomposition rate, so low that asymptotic decomposition values may be estimated and negatively related to N concentrations. Studies on the effects of changes in chemical composition on the decomposability of litter have mainly been made during the early decomposition stages and they generally report decreased litter quality (e.g., increased C/N ratio), resulting in lower decomposition rates for litter raised under elevated CO2 as compared with control litter. No reports are found relating chemical changes induced by elevated CO2 to litter mass-loss rates in late stages. By most definitions, at these stages litter has turned into humus, and many studies demonstrated that a raising of the N level may suppress humus decomposition rate. It is thus reasonable to speculate that a decrease in N levels in humus would accelerate decomposition and allow it to proceed further. There are no experimental data on the long-term effect of elevated CO2 levels, and a decrease in the storage of humus and nutrients could be predicted, at least in temperate and boreal forest systems. Future works on the effects of elevated CO2 on litter quality need to include studies of a larger number of nutrients and chemical components, and to cover different stages of decomposition. Additionally, the response of plant litter quality to elevated CO2 needs to be investigated under field conditions and at the community level, where possible shifts in community composition (i.e., C3 versus C4 ; N2 fixers versus nonfixers) predicted under elevated CO2 are taken into account.Key words: climate change, substrate quality, carbon dioxide, plant litter, chemical composition, decomposition.
Leaf traits as indicators of resource-use strategy in floras with succulent species
