No Future Growth Enhancement Expected at the Northern Edge for European Beech due to Continued Water Limitation.

With ongoing global warming, increasing water deficits promote physiological stress on forest ecosystems with negative impacts on tree growth, vitality, and survival. How individual tree species will react to increased drought stress is therefore a key research question to address for carbon accounting and the development of climate change mitigation strategies. Recent tree-ring studies have shown that trees at higher latitudes will benefit from warmer temperatures, yet this is likely highly species-dependent and less well-known for more temperate tree species. Using a unique pan-European tree-ring network of 26,430 European beech (Fagus sylvatica L.) trees from 2118 sites, we applied a linear mixed-effects modeling framework to (i) explain variation in climate-dependent growth and (ii) project growth for the near future (2021-2050) across the entire distribution of beech. We modeled the spatial pattern of radial growth responses to annually varying climate as a function of mean climate conditions (mean annual temperature, mean annual climatic water balance, and continentality). Over the calibration period (1952-2011), the model yielded high regional explanatory power (R2 = 0.38-0.72). Considering a moderate climate change scenario (CMIP6 SSP2-4.5), beech growth is projected to decrease in the future across most of its distribution range. In particular, projected growth decreases by 12%-18% (interquartile range) in northwestern Central Europe and by 11%-21% in the Mediterranean region. In contrast, climate-driven growth increases are limited to around 13% of the current occurrence, where the historical mean annual temperature was below ~6°C. More specifically, the model predicts a 3%-24% growth increase in the high-elevation clusters of the Alps and Carpathian Arc. Notably, we find little potential for future growth increases (-10 to +2%) at the poleward leading edge in southern Scandinavia. Because in this region beech growth is found to be primarily water-limited, a northward shift in its distributional range will be constrained by water availability.

A critical thermal transition driving spring phenology of Northern Hemisphere conifers.

Despite growing interest in predicting plant phenological shifts, advanced spring phenology by global climate change remains debated. Evidence documenting either small or large advancement of spring phenology to rising temperature over the spatio-temporal scales implies a potential existence of a thermal threshold in the responses of forests to global warming. We collected a unique data set of xylem cell-wall-thickening onset dates in 20 coniferous species covering a broad mean annual temperature (MAT) gradient (-3.05 to 22.9°C) across the Northern Hemisphere (latitudes 23°-66° N). Along the MAT gradient, we identified a threshold temperature (using segmented regression) of 4.9 ± 1.1°C, above which the response of xylem phenology to rising temperatures significantly decline. This threshold separates the Northern Hemisphere conifers into cold and warm thermal niches, with MAT and spring forcing being the primary drivers for the onset dates (estimated by linear and Bayesian mixed-effect models), respectively. The identified thermal threshold should be integrated into the Earth-System-Models for a better understanding of spring phenology in response to global warming and an improved prediction of global climate-carbon feedbacks.

Photoperiod and temperature as dominant environmental drivers triggering secondary growth resumption in Northern Hemisphere conifers.

Wood formation consumes around 15% of the anthropogenic CO2 emissions per year and plays a critical role in long-term sequestration of carbon on Earth. However, the exogenous factors driving wood formation onset and the underlying cellular mechanisms are still poorly understood and quantified, and this hampers an effective assessment of terrestrial forest productivity and carbon budget under global warming. Here, we used an extensive collection of unique datasets of weekly xylem tissue formation (wood formation) from 21 coniferous species across the Northern Hemisphere (latitudes 23 to 67°N) to present a quantitative demonstration that the onset of wood formation in Northern Hemisphere conifers is primarily driven by photoperiod and mean annual temperature (MAT), and only secondarily by spring forcing, winter chilling, and moisture availability. Photoperiod interacts with MAT and plays the dominant role in regulating the onset of secondary meristem growth, contrary to its as-yet-unquantified role in affecting the springtime phenology of primary meristems. The unique relationships between exogenous factors and wood formation could help to predict how forest ecosystems respond and adapt to climate warming and could provide a better understanding of the feedback occurring between vegetation and climate that is mediated by phenology. Our study quantifies the role of major environmental drivers for incorporation into state-of-the-art Earth system models (ESMs), thereby providing an improved assessment of long-term and high-resolution observations of biogeochemical cycles across terrestrial biomes.

Tree growth response to drought partially explains regional-scale growth and mortality patterns in Iberian forests.

Tree-ring data has been widely used to inform about tree growth responses to drought at the individual scale, but less is known about how tree growth sensitivity to drought scales up driving changes in forest dynamics. Here, we related tree-ring growth chronologies and stand-level forest changes in basal area from two independent data sets to test if tree-ring responses to drought match stand forest dynamics (stand basal area growth, ingrowth, and mortality). We assessed if tree growth and changes in forest basal area covary as a function of spatial scale and tree taxa (gymnosperm or angiosperm). To this end, we compared a tree-ring network with stand data from the Spanish National Forest Inventory. We focused on the cumulative impact of drought on tree growth and demography in the period 1981-2005. Drought years were identified by the Standardized Precipitation Evapotranspiration Index, and their impacts on tree growth by quantifying tree-ring width reductions. We hypothesized that forests with greater drought impacts on tree growth will also show reduced stand basal area growth and ingrowth and enhanced mortality. This is expected to occur in forests dominated by gymnosperms on drought-prone regions. Cumulative growth reductions during dry years were higher in forests dominated by gymnosperms and presented a greater magnitude and spatial autocorrelation than for angiosperms. Cumulative drought-induced tree growth reductions and changes in forest basal area were related, but initial stand density and basal area were the main factors driving changes in basal area. In drought-prone gymnosperm forests, we observed that sites with greater growth reductions had lower stand basal area growth and greater mortality. Consequently, stand basal area, forest growth, and ingrowth in regions with large drought impacts was significantly lower than in regions less impacted by drought. Tree growth sensitivity to drought can be used as a predictor of gymnosperm demographic rates in terms of stand basal area growth and ingrowth at regional scales, but further studies may try to disentangle how initial stand density modulates such relationships. Drought-induced growth reductions and their cumulative impacts have strong potential to be used as early-warning indicators of regional forest vulnerability.

Chilling and forcing temperatures interact to predict the onset of wood formation in Northern Hemisphere conifers.

The phenology of wood formation is a critical process to consider for predicting how trees from the temperate and boreal zones may react to climate change. Compared to leaf phenology, however, the determinism of wood phenology is still poorly known. Here, we compared for the first time three alternative ecophysiological model classes (threshold models, heat-sum models and chilling-influenced heat-sum models) and an empirical model in their ability to predict the starting date of xylem cell enlargement in spring, for four major Northern Hemisphere conifers (Larix decidua, Pinus sylvestris, Picea abies and Picea mariana). We fitted models with Bayesian inference to wood phenological data collected for 220 site-years over Europe and Canada. The chilling-influenced heat-sum model received most support for all the four studied species, predicting validation data with a 7.7-day error, which is within one day of the observed data resolution. We conclude that both chilling and forcing temperatures determine the onset of wood formation in Northern Hemisphere conifers. Importantly, the chilling-influenced heat-sum model showed virtually no spatial bias whichever the species, despite the large environmental gradients considered. This suggests that the spring onset of wood formation is far less affected by local adaptation than by environmentally driven plasticity. In a context of climate change, we therefore expect rising winter-spring temperature to exert ambivalent effects on the spring onset of wood formation, tending to hasten it through the accumulation of forcing temperature, but imposing a higher forcing temperature requirement through the lower accumulation of chilling.

Forest resilience to drought varies across biomes.

Forecasted increase drought frequency and severity may drive worldwide declines in forest productivity. Species-level responses to a drier world are likely to be influenced by their functional traits. Here, we analyse forest resilience to drought using an extensive network of tree-ring width data and satellite imagery. We compiled proxies of forest growth and productivity (TRWi, absolutely dated ring-width indices; NDVI, Normalized Difference Vegetation Index) for 11 tree species and 502 forests in Spain corresponding to Mediterranean, temperate, and continental biomes. Four different components of forest resilience to drought were calculated based on TRWi and NDVI data before, during, and after four major droughts (1986, 1994-1995, 1999, and 2005), and pointed out that TRWi data were more sensitive metrics of forest resilience to drought than NDVI data. Resilience was related to both drought severity and forest composition. Evergreen gymnosperms dominating semi-arid Mediterranean forests showed the lowest resistance to drought, but higher recovery than deciduous angiosperms dominating humid temperate forests. Moreover, semi-arid gymnosperm forests presented a negative temporal trend in the resistance to drought, but this pattern was absent in continental and temperate forests. Although gymnosperms in dry Mediterranean forests showed a faster recovery after drought, their recovery potential could be constrained if droughts become more frequent. Conversely, angiosperms and gymnosperms inhabiting temperate and continental sites might have problems to recover after more intense droughts since they resist drought but are less able to recover afterwards.

Tree-ring-based drought reconstruction in the Iberian Range (east of Spain) since 1694.

Droughts are a recurrent phenomenon in the Mediterranean basin with negative consequences for society, economic activities, and natural systems. Nevertheless, the study of drought recurrence and severity in Spain has been limited so far due to the relatively short instrumental period. In this work, we present a reconstruction of the standardized precipitation index (SPI) for the Iberian Range. Growth variations and climatic signals within the network are assessed developing a correlation matrix and the data combined to a single chronology integrating 336 samples from 169 trees of five different pine species distributed throughout the province of Teruel. The new chronology, calibrated against regional instrumental climatic data, shows a high and stable correlation with the July SPI integrating moisture conditions over 12 months forming the basis for a 318-year drought reconstruction. The climate signal contained in this reconstruction is highly significant (p < 0.05) and spatially robust over the interior areas of Spain located above 1000 meters above sea level (masl). According to our SPI reconstruction, seven substantially dry and five wet periods are identified since the late seventeenth century considering ≥±1.76 standard deviations. Besides these, 36 drought and 28 pluvial years were identified. Some of these years, such as 1725, 1741, 1803, and 1879, are also revealed in other drought reconstructions in Romania and Turkey, suggesting that coherent larger-scale synoptic patterns drove these extreme deviations. Since regional drought deviations are also retained in historical documents, the tree-ring-based reconstruction presented here will allow us to cross-validate drought frequency and magnitude in a highly vulnerable region.

Do variations in leaf phenology affect radial growth variations in Fagus sylvatica?

We used a dendrochronological and leaf phenology network of European beech (Fagus sylvatica) in Slovenia, a transitional area between Mediterranean, Alpine and continental climatic regimes, for the period 1955-2007 to test whether year to year variations in leaf unfolding and canopy duration (i.e. time between leaf unfolding and colouring) influence radial growth (annual xylem production and tree ring widths) and if such influences are more pronounced at higher altitudes. We showed that variability in leaf phenology has no significant effect on variations in radial growth. The results are consistent in the entire region, irrespective of the climatic regime or altitude, although previous studies have shown that leaf phenology and tree ring variation depend on altitude. The lack of relationship between year to year variability in leaf phenology and radial growth may suggest that earlier leaf unfolding--as observed in a previous study--probably does not cause increased tree growth rates in beech in Slovenia.

SiCLIMA: High-resolution hydroclimate and temperature dataset for Aragón (northeast Spain).

A new high-resolution climatic gridded dataset was built for Aragón (northeast Spain) using a large collection of daily precipitation and temperature observations from more than 3000 weather stations. The grid covers, at the unprecedented spatial resolution of 0.25 km2, daily maximum and minimum temperatures and precipitation in the 1950-2020 period. The complex orography (from 70 to 3,400 m.a.s.l.) of the medium-sized region (∼48,000 km2) required a climate modelling method based on a spatially-dense weather monitoring network and local predictors. The 3-step workflow for grid creation consisted of: 1) a comprehensive quality control of raw observations, based on a spatial comparison with nearest data; 2) a climate reconstruction based on the creation of reference values, through multivariate linear regressions, for every day and location, based on the observed climate and terrain-based environmental variables; and 3) the prediction of precipitation and temperature values in a regular 500 × 500 m grid, based on the reconstructed data series. The resulting dataset improves the spatial representativity of climate and allows for a detailed analyses at landscape scale not only in climate studies but also in related disciplines such as hydrology or biogeography, amongst others.

Climate of the Pyrenees: Extremes indices and long-term trends.

We utilized an extensive, multisource, cross-border dataset of daily meteorological observations from over 1500 stations in the Pyrenees, spanning from the mid-20th century to 2020, to examine the spatial and temporal climate patterns. Our focus was on 17 indices related to extreme precipitation and temperature events across the mountain range. The original data underwent rigorous quality control and homogenization processes, employing a comprehensive workflow that included spatial modeling based on environmental predictors. This process yielded two main outcomes: 1) a high-resolution gridded dataset (1 km2) of daily precipitation, maximum and minimum temperature from 1981 to 2020, allowing for a detailed analysis of spatial variations; and 2) an evaluation of long-term annual and seasonal trends from 1959 to 2020, using selection of high-quality data series that were homogenized to preserve their temporal structure and coherence. The findings revealed a clear elevation-related pattern in temperature indices (with the exception of tropical nights, which were predominantly observed on the Mediterranean side) and a distinct north-south latitudinal disparity in precipitation, turning longitudinal when focusing on extreme precipitation events. Overall, there was a notable and significant warming trend of 0.2 to 0.4 °C per decade, and a non-significant change of precipitation, with the exception of the southern and Mediterranean regions, where there was a notable decrease, approximately -3 % per decade, observed on an annual basis.

High preseason temperature variability drives convergence of xylem phenology in the Northern Hemisphere conifers.

Wood growth is key to understanding the feedback of forest ecosystems to the ongoing climate warming. An increase in spatial synchrony (i.e., coincident changes in distant populations) of spring phenology is one of the most prominent climate responses of forest trees. However, whether temperature variability contributes to an increase in the spatial synchrony of spring phenology and its underlying mechanisms remains largely unknown. Here, we analyzed an extensive dataset of xylem phenology observations of 20 conifer species from 75 sites over the Northern Hemisphere. Along the gradient of increase in temperature variability in the 75 sites, we observed a convergence in the onset of cell enlargement roughly toward the 5th of June, with a convergence in the onset of cell wall thickening toward the summer solstice. The increase in rainfall since the 5th of June is favorable for cell division and expansion, and as the most hours of sunlight are received around the summer solstice, it allows the optimization of carbon assimilation for cell wall thickening. Hence, the convergences can be considered as the result of matching xylem phenological activities to favorable conditions in regions with high temperature variability. Yet, forest trees relying on such consistent seasonal cues for xylem growth could constrain their ability to respond to climate warming, with consequences for the potential growing season length and, ultimately, forest productivity and survival in the future.

Identifying drivers of non-stationary climate-growth relationships of European beech.

The future performance of the widely abundant European beech (Fagus sylvatica L.) across its ecological amplitude is uncertain. Although beech is considered drought-sensitive and thus negatively affected by drought events, scientific evidence indicating increasing drought vulnerability under climate change on a cross-regional scale remains elusive. While evaluating changes in climate sensitivity of secondary growth offers a promising avenue, studies from productive, closed-canopy forests suffer from knowledge gaps, especially regarding the natural variability of climate sensitivity and how it relates to radial growth as an indicator of tree vitality. Since beech is sensitive to drought, we in this study use a drought index as a climate variable to account for the combined effects of temperature and water availability and explore how the drought sensitivity of secondary growth varies temporally in dependence on growth variability, growth trends, and climatic water availability across the species' ecological amplitude. Our results show that drought sensitivity is highly variable and non-stationary, though consistently higher at dry sites compared to moist sites. Increasing drought sensitivity can largely be explained by increasing climatic aridity, especially as it is exacerbated by climate change and trees' rank progression within forest communities, as (co-)dominant trees are more sensitive to extra-canopy climatic conditions than trees embedded in understories. However, during the driest periods of the 20th century, growth showed clear signs of being decoupled from climate. This may indicate fundamental changes in system behavior and be early-warning signals of decreasing drought tolerance. The multiple significant interaction terms in our model elucidate the complexity of European beech's drought sensitivity, which needs to be taken into consideration when assessing this species' response to climate change.

Partial asynchrony of coniferous forest carbon sources and sinks at the intra-annual time scale.

As major terrestrial carbon sinks, forests play an important role in mitigating climate change. The relationship between the seasonal uptake of carbon and its allocation to woody biomass remains poorly understood, leaving a significant gap in our capacity to predict carbon sequestration by forests. Here, we compare the intra-annual dynamics of carbon fluxes and wood formation across the Northern hemisphere, from carbon assimilation and the formation of non-structural carbon compounds to their incorporation in woody tissues. We show temporally coupled seasonal peaks of carbon assimilation (GPP) and wood cell differentiation, while the two processes are substantially decoupled during off-peak periods. Peaks of cambial activity occur substantially earlier compared to GPP, suggesting the buffer role of non-structural carbohydrates between the processes of carbon assimilation and allocation to wood. Our findings suggest that high-resolution seasonal data of ecosystem carbon fluxes, wood formation and the associated physiological processes may reduce uncertainties in carbon source-sink relationships at different spatial scales, from stand to ecosystem levels.

Dominance of Fagus sylvatica in the Growing Stock and Its Relationship to Climate-An Analysis Using Modeled Stand-Level Climate Data.

In the future, climate change is expected to affect the spatial distribution of most tree species in Europe. The European beech (Fagus sylvatica), a drought-sensitive tree species, is currently distributed throughout Europe, where it is an ecologically and economically important species. In Slovenia, the European beech represents 33% of the growing stock, but such a proportion greatly varies across Europe. Whether such a variation is related to the climate environmental gradients or because of historical or management decisions is an as-yet unexplored question. For this study, we employed the Slovenian Forests Service inventory, where the proportion of beech in the forest stock has been monitored in 341,341 forest stands across the country. Modeled climate data from the SLOCLIM database, calculated for each of the stands, was also used to test the hypothesis that although beech forests have always been influenced by human activity, the dominance of beech trees in forest stands is at least partially dictated by the climate. The results showed the distribution of the main climate variables (annual precipitation, the share of summer and spring precipitation, and annual maximum and minimum temperatures) and how they affect the current dominance of beech trees at the stand level. Due to the large number and variability of forest stands studied, the results should be transferable to better understand and manage the climatic suitability and risks of Fagus sylvatica. The modeled data is publicly available in the web repository Zenodo.

Climate-change-driven growth decline of European beech forests.

The growth of past, present, and future forests was, is and will be affected by climate variability. This multifaceted relationship has been assessed in several regional studies, but spatially resolved, large-scale analyses are largely missing so far. Here we estimate recent changes in growth of 5800 beech trees (Fagus sylvatica L.) from 324 sites, representing the full geographic and climatic range of species. Future growth trends were predicted considering state-of-the-art climate scenarios. The validated models indicate growth declines across large region of the distribution in recent decades, and project severe future growth declines ranging from -20% to more than -50% by 2090, depending on the region and climate change scenario (i.e. CMIP6 SSP1-2.6 and SSP5-8.5). Forecasted forest productivity losses are most striking towards the southern distribution limit of Fagus sylvatica, in regions where persisting atmospheric high-pressure systems are expected to increase drought severity. The projected 21st century growth changes across Europe indicate serious ecological and economic consequences that require immediate forest adaptation.

Leaf δ18O of remaining trees is affected by thinning intensity in a semiarid pine forest.

Silvicultural thinning usually improves the water status of remaining trees in water-limited forests. We evaluated the usefulness of a dual stable isotope approach (δ¹³C, δ¹8O) for comparing the physiological performance of remaining trees between forest stands subjected to two different thinning intensities (moderate versus heavy) in a 60-year-old Pinus halepensis Mill. plantation in semiarid southeastern Spain. We measured bulk leaf δ¹³C and δ¹8O, foliar elemental concentrations, stem water content, stem water δ¹8O (δ¹8O(stem water)), tree ring widths and leaf gas exchange rates to assess the influence of forest stand density on tree performance. Remaining trees in low-density stands (heavily thinned) showed lower leaf δ¹8O, and higher stomatal conductance (g(s)), photosynthetic rate and radial growth than those in moderate-density stands (moderately thinned). By contrast, leaf δ¹³C, intrinsic water-use efficiency, foliar elemental concentrations and δ¹8O(stem water) were unaffected by stand density. Lower foliar δ¹8O in heavily thinned stands reflected higher g(s) of remaining trees due to decreased inter-tree competition for water, whereas higher photosynthetic rate was largely attributable to reduced stomatal limitation to CO2 uptake. The dual isotope approach provided insight into the early (12 months) effects of stand density manipulation on the physiological performance of remaining trees.

Evidence for the spatial segregation hypothesis: a test with nine-year survivorship data in a Mediterranean shrubland.

A current focus of ecology is the investigation of spatial effects on population and community dynamics; however, spatiotemporal theory remains largely untested by empirical observations or experimental studies. For example, the segregation hypothesis predicts that intraspecific aggregation should increase the importance of intraspecific competition relative to interspecific competition, thereby enhancing local coexistence in plant communities. We applied recent methods of point pattern analysis to analyze a unique long-term data set on fully mapped seedling emergence and subsequent survival in a Mediterranean gorse shrubland after experimental fires and simulated torrential rainfall events. Our overall aim was to test if the observed spatial patterns were consistent with the segregation hypothesis during the entire community dynamics from early seedling emergence to the establishment of a mature community, i.e., we explored if the observed initial segregation did indeed prevent interspecific competition from becoming dominant. We used random labeling as the null model and specific test statistics to evaluate different biological effects of the spatial interactions that determine mortality. We found that mortality was clearly not random. Comparison of the probability of mortality in dependence on the distance to conspecific and to heterospecific plants showed that mortality was controlled almost entirely by intraspecific interactions, which is consistent with the segregation hypothesis. Dead plants were aggregated and segregated from surviving plants, indicating two-sided scramble competition. Spatial interactions were density dependent and changed their sign over the course of time from positive to negative when plants grew to maturity. The simulated torrential rainfall events and subsequent erosion caused nonspecific mortality of seedlings but did not reduce the prevalence of intraspecific competition. Our results provide support for the hypothesis that the spatial distribution of plants may profoundly affect competition and can be an important determinant in the coexistence of species and biodiversity.

Precipitation is not limiting for xylem formation dynamics and vessel development in European beech from two temperate forest sites.

We investigated the dynamics of xylem differentiation processes and vessel characteristics in Fagus sylvatica L. to evaluate the plasticity of xylem structures under different environmental conditions. In 2008-10, analyses were performed on microcores collected weekly from two temperate sites: Menina planina (1200 m above sea level (a.s.l.)) and Panska reka (400 m a.s.l.). The duration between the onset and end of major cell differentiation steps and vessel characteristics (i.e., density, VD; mean diameter, MVD; mean area, MVA; and theoretic conductivity area, TCA) were analysed in the first and last quarters of the xylem rings, also in respect of local weather conditions (precipitation, temperature). Although the onset, duration and end of xylem formation phases differed between the two sites, the time spans between the successive wood formation phases were similar. Significant differences in MVD, MVA and TCA values were found between the first and last quarters of xylem increment, regardless of the site and year. Vessel density, on the other hand, depended on xylem-ring width and differed significantly between the sites, being about 30% higher at the high elevation site, in beech trees with 54% narrower xylem rings. Vessel density in the first quarter of the xylem ring showed a positive correlation with the onset of cell expansion, whereas a negative correlation of VD with the cessation of cell production was found in the last quarter of xylem increment. This may be explained by year-to-year differences in the timing of cambial reactivation and leaf development, which effect hormonal regulation of radial growth. No significant linkage between intra-annual weather conditions and conduit characteristics was found. It can thus be presumed that precipitation is not a limiting factor for xylem growth and cell differentiation in beech at the two temperate study sites and sites across Europe with similar weather conditions.

Soil moisture and its role in growth-climate relationships across an aridity gradient in semiarid Pinus halepensis forests.

In Mediterranean areas with limited availability of water, an accurate knowledge of growth response to hydrological variables could contribute to improving management and stability of forest resources. The main goal of this study is to assess the temporal dynamic of soil moisture to better understand the water-growth relationship of Pinus halepensis forests in semiarid areas. The estimates of modelled soil moisture and measured tree growth were used at four sites dominated by afforested Pinus halepensis Mill. in south-eastern Spain with 300 to 609mm mean annual precipitation. Firstly, dendrochronological samples were extracted and the widths of annual tree rings were measured to compute basal area increments (BAI). Secondly, soil moisture was estimated over 20 hydrological years (1992-2012) by means of the HYDROBAL ecohydrological model. Finally, the tree growth was linked, to mean monthly and seasonal temperature, precipitation and soil moisture. Results depict the effect of soil moisture on growth (BAI) and explain 69-73% of the variance in semiarid forests, but only 51% in the subhumid forests. This highlights the fact that that soil moisture is a suitable and promising variable to explain growth variations of afforested Pinus halepensis in semiarid conditions and useful for guiding adaptation plans to respond pro-actively to water-related global challenges.