Abstract
Tree secondary growth is responsible for woody biomass accumulation and is a major component of carbon storage in forest ecosystems. Environmental constraints on secondary growth in Mediterranean ecosystems must, however, be described in more to details to better understand how they will be modified by climate change. This dissertation aims at studying the functional responses of Mediterranean holm oak (Quercus ilex) to seasonal and inter-annual climate variations through the study of carbon allocation to secondary growth. Different experimental approaches, at spatial scales ranging from tree rings to the ecosystem and at temporal scales from the day to several decades, were used to identify the main environmental constraints (water availability, temperature warming, competition) to secondary growth and carbon isotopic composition of tree rings. The phenology of stem growth shows evidence for a direct environmental control on annual growth by winter temperature and summer drought that is more limiting than the carbon supply from photosynthesis. Climate change from 1968 to 2013 resulted in earlier water limitation on secondary growth, which was compensated by earlier growth onset, due to warmer winter temperature, and higher water use efficiency, due to increased atmospheric CO2 concentration. Thinning reduced tree mortality and increased stem growth, so thinning management in old holm oak coppices could prepare the ecosystem to better withstand the increasing drought forecasted for the Mediterranean region.