Interactions between Climate and Nutrient Cycles on Forest Response to Global Change: The Role of Mixed Forests

Forest ecosystems are undergoing unprecedented changes in environmental conditions due to global change impacts. Modification of global biogeochemical cycles of carbon and nitrogen, and the subsequent climate change are affecting forest functions at different scales, from physiology and growth of individual trees to cycling of nutrients. This review summarizes the present knowledge regarding the impact of global change on forest functioning not only with respect to climate change, which is the focus of most studies, but also the influence of altered nitrogen cycle and the interactions among them. The carbon dioxide (CO₂) fertilization effect on tree growth is expected to be constrained by nutrient imbalances resulting from high N deposition rates and the counteractive effect of increasing water deficit, which interact in a complex way. At the community level, responses to global change are modified by species interactions that may lead to competition for resources and/or relaxation due to facilitation and resource partitioning processes. Thus, some species mixtures can be more resistant to drought than their respective pure forests, albeit it depends on environmental conditions and species’ functional traits. Climate change and nitrogen deposition have additional impacts on litterfall dynamics, and subsequent decomposition and nutrient mineralization processes. Elemental ratios (i.e., stoichiometry) are associated with important ecosystem traits, including trees’ adaptability to stress or decomposition rates. As stoichiometry of different ecosystem components are also influenced by global change, nutrient cycling in forests will be altered too. Therefore, a re-assessment of traditional forest management is needed in order to cope with global change. Proposed silvicultural systems emphasize the key role of diversity to assure multiple ecosystem services, and special attention has been paid to mixed-species forests. Finally, a summary of the patterns and underlying mechanisms governing the relationships between diversity and different ecosystems functions, such as productivity and stability, is provided.