Seasonal variation in foliar properties in Mediterranean Pine forests of different post-fire age

Foliar properties play a crucial role in local and global biochemical cycles. Systematic variation in key leaf traits has been reported both between and within species. Intraspecific variation in leaf traits is controlled by micro-environmental conditions and follows seasonal patterns. In this study we examine the seasonal patterns of six foliar traits including leaf area (LA), leaf thickness (Lth), leaf mass per area (LMA), leaf dry matter content (LDMC), leaf area to sapwood area ratio (LA/SA) and branch wood density (WD) in addition to the associated parameters of the Michaelis-Menten light response curve (i.e. light saturated net photosynthetic rate (Asat), half saturation coefficient (Km) and dark respiration rate (Rd)). We measured on a monthly basis the foliar traits and developed light response curves in four Pinus brutia dominated stands along a post-fire chronosequence (15, 40, 70 and 90 years) from sunlit branches. Significant differences in the interannual trait variability were found between stands for LDMC, WD and Asat, with the highest variability identified in the younger plot. LA/SA, Rd and Km also showed strong interannual variability although not statistically different between plots. A mixed effect model analysis revealed high intraclass correlation coefficients for Km and Asat suggesting that net photosynthesis is following systematic seasonal patterns. Overall LA was higher and LDMC was lower in the oldest plot and WD was higher in the denser (40 years) plot. Interestingly gas exchange parameters did not show differences in their overall mean values. Across plots, Asat was strongly positively related to Km, and LMA was positively related to LDMC and Lth. LDMC was also positively related with Asat and negatively with Lth. A principal component analysis (PCA) revealed two major dimensions of intraspecific trait variability within our plots. The first PCA axis was positively related to Asat, Km, LDMC and LMA suggesting that regardless of the stand age needles are placed along a fast-slow carbon gain dimension with denser needles illustrating faster area-based photosynthesis. The second PCA axis was positively related to LA and Lth suggesting that bigger needles are also thicker. A subsequent permutational multivariate analysis of variance revealed that the centroids and the dispersion of the trait syndromes differed between stands, with the youngest plot illustrating higher trait dispersion and the oldest plot characterized by bigger and thicker needles. Thus, in older stands were competition for light is higher, needles are deployed to be bigger and thicker to optimize light capture, while in younger stands they are optimized along a leaf density - photosynthetic capacity spectrum depending on (more heterogeneous) microenvironmental conditions. Our findings illustrate that intraspecific variation can be attributed to either seasonal (abiotic) light availability or to (biotic) heterogeneity related to stand structure, and have important implications for local scale forest dynamics models.

«This research is co-financed by Greece and the European Union (European Social Fund- ESF) through the Operational Programme «Human Resources Development, Education and Lifelong Learning 2014-2020» in the context of the project “Carbon fluxes across a post-fire chronosequence in Pinus brutia Ten forests.” (MIS 5049513)».