A genome-wide genetic association study reveals SNPs significantly associated with environmental variables and specific leaf area in European beech.

European beech is negatively affected by climate change and a further growth decline is predicted for large parts of its distribution range. Despite the importance of this species, little is known about its genetic adaptation and especially the genetic basis of its physiological traits. Here, we used genotyping by sequencing to identify SNPs in 43 German European beech populations growing under different environmental conditions. In total, 28 of these populations were located along a precipitation and temperature gradient in northern Germany, and single tree-based hydraulic and morphological traits were available. We obtained a set of 13,493 high-quality SNPs that were used for environmental and SNP-trait association analysis. In total, 22 SNPs were identified that were significantly associated with environmental variables or specific leaf area (SLA). Several SNPs were located in genes related to stress response. The majority of the significant SNPs were located in non-coding (intergenic and intronic) regions. These may be in linkage disequilibrium with the causative coding or regulatory regions. Our study gives insights into the genetic basis of abiotic adaptation in European beech, and provides genetic resources that can be used in future studies on this species. Besides clear patterns of local adaptation to environmental conditions of the investigated populations, the analyzed morphological and hydraulic traits explained most of the explainable genetic variation. Thus, they could successfully be altered in tree breeding programs, which may help to increase the adaptation of European beech to changing environmental conditions in the future.

Summer drought exposure, stand structure, and soil properties jointly control the growth of European beech along a steep precipitation gradient in northern Germany.

Increasing exposure to climate warming-related drought and heat threatens forest vitality in many regions on earth, with the trees' vulnerability likely depending on local climatic aridity, recent climate trends, edaphic conditions, and the drought acclimatization and adaptation of populations. Studies exploring tree species' vulnerability to climate change often have a local focus or model the species' entire distribution range, which hampers the separation of climatic and edaphic drivers of drought and heat vulnerability. We compared recent radial growth trends and the sensitivity of growth to drought and heat in central populations of a widespread and naturally dominant tree species in Europe, European beech (Fagus sylvatica), at 30 forest sites across a steep precipitation gradient (500-850 mm year-1 ) of short length to assess the species' adaptive potential. Size-standardized basal area increment remained more constant during the period of accelerated warming since the early 1980s in populations with >360 mm growing season precipitation (April-September), while growth trends were negative at sites with <360 mm. Climatic drought in June appeared as the most influential climatic factor affecting radial growth, with a stronger effect at drier sites. A decadal decrease in the climatic water balance of the summer was identified as the most important factor leading to growth decline, which is amplified by higher stem densities. Inter-annual growth variability has increased since the early 1980s, and variability is generally higher at drier and sandier sites. Similarly, within-population growth synchrony is higher at sandier sites and has increased with a decrease in the June climatic water balance. Our results caution against predicting the drought vulnerability of trees solely from climate projections, as soil properties emerged as an important modulating factor. We conclude that beech is facing recent growth decline at drier sites in the centre of its distribution range, driven by climate change-related climate aridification.

Soil water availability and branch age explain variability in xylem safety of European beech in Central Europe.

Xylem embolism resistance has been identified as a key trait with a causal relation to drought-induced tree mortality, but not much is known about its intra-specific trait variability (ITV) in dependence on environmental variation. We measured xylem safety and efficiency in 300 European beech (Fagus sylvatica L.) trees across 30 sites in Central Europe, covering a precipitation reduction from 886 to 522 mm year-1. A broad range of variables that might affect embolism resistance in mature trees, including climatic and soil water availability, competition, and branch age, were examined. The average P50 value varied by up to 1 MPa between sites. Neither climatic aridity nor structural variables had a significant influence on P50. However, P50 was less negative for trees with a higher soil water storage capacity, and positively related to branch age, while specific conductivity (Ks) was not significantly associated with either of these variables. The greatest part of the ITV for xylem safety and efficiency was attributed to random variability within populations. We conclude that the influence of site water availability on P50 and Ks is low in European beech, and that the high degree of within-population variability for P50, partly due to variation in branch age, hampers the identification of a clear environmental signal.

Branch xylem vascular adjustments in European beech in response to decreasing water availability across a precipitation gradient.

Crucial for the climate adaptation of trees is a xylem anatomical structure capable of adjusting to changing water regimes. Although species comparisons across climate zones have demonstrated anatomical change in response to altered water availability and tree height, less is known about the adaptability of tree vascular systems to increasing water deficits at the intraspecific level. Information on the between-population and within-population variability of xylem traits helps assessing a species' ability to cope with climate change. We investigated the variability of wood anatomical and related hydraulic traits in terminal branches of European beech (Fagus sylvatica L.) trees across a precipitation gradient (520-890 mm year-1) and examined the influence of climatic water balance (CWB), soil water capacity (AWC), neighborhood competition (CI), tree height and branch age on these traits. Furthermore, the relationship between xylem anatomical traits and embolism resistance (P50) was tested. Within-population trait variation was larger than between-population variation. Vessel diameter, lumen-to-sapwood area ratio and potential conductivity of terminal branches decreased with decreasing CWB, but these traits were not affected by AWC, whereas vessel density increased with an AWC decrease. In contrast, none of the studied anatomical traits were influenced by variation in tree height (21-34 m) or CI. Branch age was highly variable (2-22 years) despite equal diameter and position in the flow path, suggesting different growth trajectories in the past. Vessel diameter decreased, and vessel density increased, with increasing branch age, reflecting negative annual radial growth trends. Although vessel diameter was not related to P50, vessel grouping index and lumen-to-sapwood area ratio showed a weak, though highly significant, positive relationship to P50. We conclude that the xylem anatomy of terminal tree-top branches in European beech is modified in response to increasing climatic aridity and/or decreasing soil water availability, independent of a tree height effect.

FunAndes - A functional trait database of Andean plants.

We introduce the FunAndes database, a compilation of functional trait data for the Andean flora spanning six countries. FunAndes contains data on 24 traits across 2,694 taxa, for a total of 105,466 entries. The database features plant-morphological attributes including growth form, and leaf, stem, and wood traits measured at the species or individual level, together with geographic metadata (i.e., coordinates and elevation). FunAndes follows the field names, trait descriptions and units of measurement of the TRY database. It is currently available in open access in the FIGSHARE data repository, and will be part of TRY's next release. Open access trait data from Andean plants will contribute to ecological research in the region, the most species rich terrestrial biodiversity hotspot.