Bimodal and unimodal radial growth of Mediterranean oaks along a coast-inland gradient

Abstract

Wood formation during the growing season is shaped by the intra-annual variability of climatic conditions. In the Mediterranean, the cambial activity is seasonally constrained by winter low temperature and summer drought, resulting in bimodal growth patterns. Although bimodal growth is an ecologically important adaptation of woody species to seasonally dry environments, its variability across space and among species remains unclear. We combined direct monitoring of intra-annual radial growth using dendrometers and xylogenesis micro-sampling with indirect wood formation modeling to assess growth bimodality for two coexisting oak species across six sites in North-Eastern Spain. We focused on species with two contrasting life strategies, evergreen with diffuse-porous wood (Quercus ilex) and winter deciduous with ring-porous wood (Q. faginea/Q. pubescens). Study sites spanned from oceanic areas with high autumn precipitation to dry continental inland regions. We hypothesized that growth bimodality reflects the interaction between local precipitation patterns and species-specific traits. Our results revealed cambial activity in autumn after summer quiescence across the entire region and for both oak species. However, growth bimodality and the relative contribution of the autumn growth peak to the total annual growth was geographically structured and species-specific. Growth was unimodal under continental and Atlantic conditions. By contrast, growth bimodality was significant along the Mediterranean coast, where precipitation showed a bimodal pattern. Moreover, evergreen Q. ilex showed more growth bimodality compared with deciduous Q. faginea/Q. pubescens at Mediterranean sites. Different intra-annual distribution of radial growth could facilitate coexistence of both oak species through temporal niche partitioning. The growth bimodality helps to compensate for reduced growth in summer by autumn cambial resumption and, therefore, the differences in its intensity between sites and species may determine divergent responses of forest ecosystems to ongoing climate change.