• Energy Research

As climate change increasingly affects forest ecosystems, detailed understanding of major effects is important to anticipate their consequences under future climate scenarios. The Mediterranean region is a prominent climate change hotspot, and evergreen cork oak (Quercus suber L.) woodlands are particularly climatically sensitive due to cork (bark) harvesting. Cork oak’s drought avoidance strategy is well-known and includes structural and physiological adaptations that maximise soil water uptake and transport and limit water use, potentially leading to reduced stem and cork growth. Trees’ responses to cope with water-limited conditions have been extensively described based on cork-rings width and, more recently, on cork-rings density, in dendroecological studies. However, so far, tree functional attributes and physiological strategies, namely photosynthetic metabolism adjustments affecting cork formation, have never been addressed and/or integrated on these previous cork-rings-based studies. In this study, we address the relation between carbon and oxygen stable isotopes of cork rings and precipitation and temperature, in two distinct locations of southwestern Portugal–the (wetter) Tagus basin peneplain and the (drier) Grândola mountains. We aimed at assessing whether the two climatic factors affect cork-ring isotopic composition under contrasting conditions of water availability, and, therefore, if carbon and oxygen signatures in cork can reflect tree functional (physiological and structural) responses to stressful conditions, which might be aggravated by climate change. Our results indicate differences between the study areas. At the drier site, the stronger statistically significant negative cork δ13C correlations were found with mean temperature, whereas strong positive cork δ18O correlations were fewer and found only with precipitation. Moreover, at the wetter site, cork rings are enriched in 18O and depleted in 13C, indicating, respectively, shallow groundwater as the water source for physiological processes related with biosynthesis of non-photosynthetic secondary tissues, such as suberin, and a weak stomatal regulation under high water availability, consistent with non-existent water availability constrains. In contrast, at the drier site, trees use water from deeper ground layers, depleted in 18O, and strongly regulate stomatal conductance under water stress, thus reducing photosynthetic carbon uptake and probably relying on stored carbon reserves for cork ring formation. These results suggest that although stable isotopes signatures in cork rings are not proxies for net growth, they may be (fairly) robust indicators of trees’ physiological and structural adjustments to climate and environmental changes in Mediterranean environments.

AbstractNorthern forests at the leading edge of their distributions may not show increased primary productivity under climate warming, being limited by climatic extremes such as drought. Looking beyond tree growth to underlying physiological mechanisms is fundamental for accurate predictions of forest responses to climate warming and drought stress. Within a 32-year genetic field trial, we analyze relative contributions of xylem plasticity and inferred stomatal response to drought tolerance in regional populations of a widespread conifer. Genetic adaptation leads to varying responses under drought. Trailing-edge tree populations produce fewer tracheids with thicker cell walls, characteristic of drought-tolerance. Stomatal response explains the moderate drought tolerance of tree populations in central areas of the species range. Growth loss of the northern population is linked to low stomatal responsiveness combined with the production of tracheids with thinner cell walls. Forests of the western boreal may therefore lack physiological adaptations necessary to tolerate drier conditions.

Ajut Marie Curie IF fellowship (No 659191); COST Action FP1106 STReESS

Aboveground carbon (C) sequestration in trees is important in global C dynamics, but reliable techniques for its modeling in highly productive and heterogeneous ecosystems are limited. We applied an extended dendrochronological approach to disentangle the functioning of drivers from the atmosphere (temperature, precipitation), the lithosphere (sedimentation rate), the hydrosphere (groundwater table, river water level fluctuation), the biosphere (tree characteristics), and the anthroposphere (dike construction). Carbon sequestration in aboveground biomass of riparian Quercus robur L. and Fraxinus excelsior L. was modeled (1) over time using boosted regression tree analysis (BRT) on cross-datable trees characterized by equal annual growth ring patterns and (2) across space using a subsequent classification and regression tree analysis (CART) on cross-datable and not cross-datable trees. While C sequestration of cross-datable Q. robur responded to precipitation and temperature, cross-datable F. excelsior also responded to a low Danube river water level. However, CART revealed that C sequestration over time is governed by tree height and parameters that vary over space (magnitude of fluctuation in the groundwater table, vertical distance to mean river water level, and longitudinal distance to upstream end of the study area). Thus, a uniform response to climatic drivers of aboveground C sequestration in Q. robur was only detectable in trees of an intermediate height class and in taller trees (>21.8m) on sites where the groundwater table fluctuated little (≤0.9m). The detection of climatic drivers and the river water level in F. excelsior depended on sites at lower altitudes above the mean river water level (≤2.7m) and along a less dynamic downstream section of the study area. Our approach indicates unexploited opportunities of understanding the interplay of different environmental drivers in aboveground C sequestration. Results may support species-specific and locally adapted forest management plans to increase carbon dioxide sequestration from the atmosphere in trees.

AbstractTree rings provide an invaluable long‐term record for understanding how climate and other drivers shape tree growth and forest productivity. However, conventional tree‐ring analysis methods were not designed to simultaneously test effects of climate, tree size, and other drivers on individual growth. This has limited the potential to test ecologically relevant hypotheses on tree growth sensitivity to environmental drivers and their interactions with tree size. Here, we develop and apply a new method to simultaneously model nonlinear effects of primary climate drivers, reconstructed tree diameter at breast height (DBH), and calendar year in generalized least squares models that account for the temporal autocorrelation inherent to each individual tree's growth. We analyze data from 3811 trees representing 40 species at 10 globally distributed sites, showing that precipitation, temperature, DBH, and calendar year have additively, and often interactively, influenced annual growth over the past 120 years. Growth responses were predominantly positive to precipitation (usually over ≥3‐month seasonal windows) and negative to temperature (usually maximum temperature, over ≤3‐month seasonal windows), with concave‐down responses in 63% of relationships. Climate sensitivity commonly varied with DBH (45% of cases tested), with larger trees usually more sensitive. Trends in ring width at small DBH were linked to the light environment under which trees established, but basal area or biomass increments consistently reached maxima at intermediate DBH. Accounting for climate and DBH, growth rate declined over time for 92% of species in secondary or disturbed stands, whereas growth trends were mixed in older forests. These trends were largely attributable to stand dynamics as cohorts and stands age, which remain challenging to disentangle from global change drivers. By providing a parsimonious approach for characterizing multiple interacting drivers of tree growth, our method reveals a more complete picture of the factors influencing growth than has previously been possible.

Mixed forests of Quercus ilex L. and Pinus pinea L. are widely found throughout the Mediterranean Basin, being representative of two co-existing functional types: evergreen-sclerophyllous drought-resistant species and Mediterranean-adapted drought-avoidant conifers. Their contrasting physiological strategies to cope with water deficit influence all the processes regulating their growth such as wood formation, leading to peculiar tree-ring anatomical features such as intra-annual density fluctuations (IADFs). Intra-annual density fluctuations are abrupt changes in wood anatomical traits within a tree ring, appearing as latewood-like cells within earlywood or earlywood-like cells within latewood, and are frequently found in Mediterranean species as a response to seasonal climate changes. In this study, we characterized the anatomical traits and composition of carbon and oxygen stable isotopes in IADFs occurring in tree rings of Q. ilex and P. pinea trees co-existing at a same site in Southern Italy, in order to link their xylem hydraulic properties with the related physiological mechanisms. The relationships between IADF occurrence and seasonal mean temperature and total precipitation were investigated, with the aim of assessing whether they can be used as indicators of species-specific responses to intra-annual climate fluctuations. Results show that IADF period of formation is during autumn months for both species. The influence of climate on IADF occurrence was found to be an indicator of species-specific response to climate: an increased stomatal conductance associated to the formation of a wood safer against embolism was found in Q. ilex, while a tighter stomatal control associated to a more efficient wood with regard to hydraulic conductivity occurred in P. pinea. Moreover, the assessment of the influence of climate on IADF occurrence indicates that, with rising temperatures, Q. ilex would form fewer IADFs compared with P. pinea. Other study cases are desirable to assess the suggested forecasts and to link the plasticity of the species to form IADFs with their effective adaptive capability to compete for resources, and to explain how it may influence future population development.

Summary Elevated CO2 increases intrinsic water use efficiency (WUEi) of forests, but the magnitude of this effect and its interaction with climate is still poorly understood. We combined tree ring analysis with isotope measurements at three Free Air CO2 Enrichment (FACE, POP‐EUROFACE, in Italy; Duke FACE in North Carolina and ORNL in Tennessee, USA) sites, to cover the entire life of the trees. We used δ13C to assess carbon isotope discrimination and changes in water‐use efficiency, while direct CO2 effects on stomatal conductance were explored using δ18O as a proxy. Across all the sites, elevated CO2 increased 13C‐derived water‐use efficiency on average by 73% for Liquidambar styraciflua, 77% for Pinus taeda and 75% for Populus sp., but through different ecophysiological mechanisms. Our findings provide a robust means of predicting water‐use efficiency responses from a variety of tree species exposed to variable environmental conditions over time, and species‐specific relationships that can help modelling elevated CO2 and climate impacts on forest productivity, carbon and water balances.