Disentangling the climate-driven bimodal growth pattern in coastal and continental Mediterranean pine stands

Mediterranean climate promotes two distinct growth peaks separated by summer quiescence in trees. This bimodal pattern has been associated to favourable growing conditions during spring and autumn when mild temperatures and soil-water availability enhance cambial activity. Climatic models predict progressive warming and drying for the Mediterranean Basin, which could shorten or shift the spring and autumn growing seasons. We explored this idea by comparing two sites with different Mediterranean climate types (continental/dry and coastal/wet) and studied how climate drives the bimodal growth pattern in Aleppo pine (Pinus halepensis). Specifically we investigated the intra-annual changes in wood anatomy and the corresponding formation of density fluctuations (IADF). Trees on both sites were analyzed by dendrometer monitoring and by developing chronologies of wood anatomical traits. Radial-increment dynamics followed a similar bimodal pattern in both sites but coastal trees showed higher increments during the spring and autumn growth peaks, especially in autumn. The summer rest of cambium activity occurs almost one month earlier in the coastal than in the inland site. Lumen area and cell-wall thickness were significantly smaller in the continental site, while the increment rate of cell-wall thickness during an IADF event was much higher in the coastal pines. The accumulated soil moisture deficit was the main climatic constraint of tracheid enlargement in continental pines. Intra-annual density fluctuations were more frequent in the coastal trees where wood anatomy features recover to average values after such events, meanwhile inland trees presented a much lower recovery rate. Growth bimodality and the formation of density fluctuations were linked, but mild climate of the coastal site allows a longer growing season, which explains why trees in this area showed higher and more variable growth rates.