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.

High risk, high gain? Trade-offs between growth and resistance to extreme events differ in northern red oak (Quercus rubra L.).

Information about the resistance and adaptive potential of tree species and provenances is needed to select suitable planting material in times of rapidly changing climate conditions. In this study, we evaluate growth responses to climatic fluctuations and extreme events for 12 provenances of northern red oak (Quercus rubra L.) that were tested across three trial sites with distinct environmental conditions in Germany. Six provenances each were sourced from the natural distribution in North America and from introduced stands in Germany. We collected increment cores of 16 trees per provenance and site. Dendroecological methods were used to compare provenance performance and establish climate-growth relationships to identify the main growth limiting factors. To evaluate the provenance response to extreme drought and frost events, three site-specific drought years were selected according to the Standardized Precipitation Evapotranspiration Index (SPEI) and 2010 as a year with an extreme late frost event. Resistance indices for these years were calculated and assessed in relation to overall growth performance. We observed a high variation in growth and in the climate sensitivity between sites depending on the prevailing climatic conditions, as well as a high intra-specific variation. Overall, summer drought and low temperatures in the early growing season appear to constrain the growth of red oak. The resistance of provenances within sites and extreme years showed considerable rank changes and interaction effects. We did not find a trade-off between growth and resistance to late frost, namely, fast growing provenances had a high frost hardiness. Further, there was no evidence for a trade-off between growth and drought hardiness. Still, responses to drought or late frost differ between provenances, pointing to dissimilar adaptive strategies. Provenances from introduced (i.e. German) stands represent suitable seed sources, as they combine a higher growth and frost hardiness compared to their North American counterparts. Drought hardiness was slightly higher in the slow-growing provenances. The results provide a better understanding of the variable adaptive strategies between provenances and help to select suitable planting material for adaptive forest management.

Early warning signals indicate climate change-induced stress in Norway spruce in the Eastern Carpathians.

Climate change is affecting forest ecosystems globally, in particular through warming as well as increases in the frequency and intensity of extreme events. Norway spruce (Picea abies (L.) Karst.) is one of the most important coniferous tree species in Europe. In recent extremely dry years in Central Europe, spruce suffered and large dieback has been observed. In parts of Eastern Europe, however, no large-scale decline in spruce has been reported so far, though anticipated changes in climate pose the question how the future of these forests may look like. To assess the current state of spruce forests in Eastern Europe, we established a tree-ring network consisting of 157 Norway spruce chronologies (from >3000 trees) of different ages distributed along elevational transects in the Eastern Carpathians, Romania. We evaluated early warning signals of climate change-induced stress, i.e. (1) growth decline, (2) increased sensitivity of tree growth (assessed over the statistics first-order autocorrelation and standard deviation), and (3) increased growth synchrony. A pronounced growth decline was observed over the last two decades, which was strongest in younger stands and at lower elevations. However, growth sensitivity and synchrony did not show consistent patterns, suggesting that forest decline may not be immediately imminent. Overall, our findings highlight an increased vulnerability of spruce in the Eastern Carpathians. With ongoing climate change, spruce dieback may be expected in this part of Europe as well.

Incorporating high-resolution climate, remote sensing and topographic data to map annual forest growth in central and eastern Europe.

To enhance our understanding of forest carbon sequestration, climate change mitigation and drought impact on forest ecosystems, the availability of high-resolution annual forest growth maps based on tree-ring width (TRW) would provide a significant advancement to the field. Site-specific characteristics, which can be approximated by high-resolution Earth observation by satellites (EOS), emerge as crucial drivers of forest growth, influencing how climate translates into tree growth. EOS provides information on surface reflectance related to forest characteristics and thus can potentially improve the accuracy of forest growth models based on TRW. Through the modelling of TRW using EOS, climate and topography data, we showed that species-specific models can explain up to 52 % of model variance (Quercus petraea), while combining different species results in relatively poor model performance (R2 = 13 %). The integration of EOS into models based solely on climate and elevation data improved the explained variance by 6 % on average. Leveraging these insights, we successfully generated a map of annual TRW for the year 2021. We employed the area of applicability (AOA) approach to delineate the range in which our models are deemed valid. The calculated AOA for the established forest-type models was 73 % of the study region, indicating robust spatial applicability. Notably, unreliable predictions predominantly occurred in the climate margins of our dataset. In conclusion, our large-scale assessment underscores the efficacy of combining climate, EOS and topographic data to develop robust models for mapping annual TRW. This research not only fills a critical void in the current understanding of forest growth dynamics but also highlights the potential of integrated data sources for comprehensive ecosystem assessments.

Accuracy, realism and general applicability of European forest models.

Forest models are instrumental for understanding and projecting the impact of climate change on forests. A considerable number of forest models have been developed in the last decades. However, few systematic and comprehensive model comparisons have been performed in Europe that combine an evaluation of modelled carbon and water fluxes and forest structure. We evaluate 13 widely used, state-of-the-art, stand-scale forest models against field measurements of forest structure and eddy-covariance data of carbon and water fluxes over multiple decades across an environmental gradient at nine typical European forest stands. We test the models' performance in three dimensions: accuracy of local predictions (agreement of modelled and observed annual data), realism of environmental responses (agreement of modelled and observed responses of daily gross primary productivity to temperature, radiation and vapour pressure deficit) and general applicability (proportion of European tree species covered). We find that multiple models are available that excel according to our three dimensions of model performance. For the accuracy of local predictions, variables related to forest structure have lower random and systematic errors than annual carbon and water flux variables. Moreover, the multi-model ensemble mean provided overall more realistic daily productivity responses to environmental drivers across all sites than any single individual model. The general applicability of the models is high, as almost all models are currently able to cover Europe's common tree species. We show that forest models complement each other in their response to environmental drivers and that there are several cases in which individual models outperform the model ensemble. Our framework provides a first step to capturing essential differences between forest models that go beyond the most commonly used accuracy of predictions. Overall, this study provides a point of reference for future model work aimed at predicting climate impacts and supporting climate mitigation and adaptation measures in forests.

Jet stream position explains regional anomalies in European beech forest productivity and tree growth.

The mechanistic pathways connecting ocean-atmosphere variability and terrestrial productivity are well-established theoretically, but remain challenging to quantify empirically. Such quantification will greatly improve the assessment and prediction of changes in terrestrial carbon sequestration in response to dynamically induced climatic extremes. The jet stream latitude (JSL) over the North Atlantic-European domain provides a synthetic and robust physical framework that integrates climate variability not accounted for by atmospheric circulation patterns alone. Surface climate impacts of north-south summer JSL displacements are not uniform across Europe, but rather create a northwestern-southeastern dipole in forest productivity and radial-growth anomalies. Summer JSL variability over the eastern North Atlantic-European domain (5-40E) exerts the strongest impact on European beech, inducing anomalies of up to 30% in modelled gross primary productivity and 50% in radial tree growth. The net effects of JSL movements on terrestrial carbon fluxes depend on forest density, carbon stocks, and productivity imbalances across biogeographic regions.

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.

Species-specific effects of thermal stress on the expression of genetic variation across a diverse group of plant and animal taxa under experimental conditions.

Assessing the genetic adaptive potential of populations and species is essential for better understanding evolutionary processes. However, the expression of genetic variation may depend on environmental conditions, which may speed up or slow down evolutionary responses. Thus, the same selection pressure may lead to different responses. Against this background, we here investigate the effects of thermal stress on genetic variation, mainly under controlled laboratory conditions. We estimated additive genetic variance (VA), narrow-sense heritability (h2) and the coefficient of genetic variation (CVA) under both benign control and stressful thermal conditions. We included six species spanning a diverse range of plant and animal taxa, and a total of 25 morphological and life-history traits. Our results show that (1) thermal stress reduced fitness components, (2) the majority of traits showed significant genetic variation and that (3) thermal stress affected the expression of genetic variation (VA, h2 or CVA) in only one-third of the cases (25 of 75 analyses, mostly in one clonal species). Moreover, the effects were highly species-specific, with genetic variation increasing in 11 and decreasing in 14 cases under stress. Our results hence indicate that thermal stress does not generally affect the expression of genetic variation under laboratory conditions but, nevertheless, increases or decreases genetic variation in specific cases. Consequently, predicting the rate of genetic adaptation might not be generally complicated by environmental variation, but requires a careful case-by-case consideration.

Global assessment of relationships between climate and tree growth.

Tree-ring records provide global high-resolution information on tree-species responses to global change, forest carbon and water dynamics, and past climate variability and extremes. The underlying assumption is a stationary (time-stable), quasi-linear relationship between tree growth and environment, which however conflicts with basic ecological and evolutionary theory. Indeed, our global assessment of the relevant tree-ring literature demonstrates non-stationarity in the majority of tested cases, not limited to specific proxies, environmental parameters, regions or species. Non-stationarity likely represents the general nature of the relationship between tree-growth proxies and environment. Studies assuming stationarity however score two times more citations influencing other fields of science and the science-policy interface. To reconcile ecological reality with the application of tree-ring proxies for climate or environmental estimates, we provide a clarification of the stationarity concept, propose a simple confidence framework for the re-evaluation of existing studies and recommend the use of a new statistical tool to detect non-stationarity in tree-ring proxies. Our contribution is meant to stimulate and facilitate discussion in light of our results to help increase confidence in tree-ring-based climate and environmental estimates for science, the public and policymakers.

Tree growth influenced by warming winter climate and summer moisture availability in northern temperate forests.

The role of future forests in global biogeochemical cycles will depend on how different tree species respond to climate. Interpreting the response of forest growth to climate change requires an understanding of the temporal and spatial patterns of seasonal climatic influences on the growth of common tree species. We constructed a new network of 310 tree-ring width chronologies from three common tree species (Quercus robur, Pinus sylvestris and Fagus sylvatica) collected for different ecological, management and climate purposes in the south Baltic Sea region at the border of three bioclimatic zones (temperate continental, oceanic, southern boreal). The major climate factors (temperature, precipitation, drought) affecting tree growth at monthly and seasonal scales were identified. Our analysis documents that 20th century Scots pine and deciduous species growth is generally controlled by different climate parameters, and that summer moisture availability is increasingly important for the growth of deciduous species examined. We report changes in the influence of winter climate variables over the last decades, where a decreasing influence of late winter temperature on deciduous tree growth and an increasing influence of winter temperature on Scots pine growth was found. By comparing climate-growth responses for the 1943-1972 and 1973-2002 periods and characterizing site-level growth response stability, a descriptive application of spatial segregation analysis distinguished sites with stable responses to dominant climate parameters (northeast of the study region), and sites that collectively showed unstable responses to winter climate (southeast of the study region). The findings presented here highlight the temporally unstable and nonuniform responses of tree growth to climate variability, and that there are geographical coherent regions where these changes are similar. Considering continued climate change in the future, our results provide important regional perspectives on recent broad-scale climate-growth relationships for trees across the temperate to boreal forest transition around the south Baltic Sea.

Removing the no-analogue bias in modern accelerated tree growth leads to stronger medieval drought.

In many parts of the world, especially in the temperate regions of Europe and North-America, accelerated tree growth rates have been observed over the last decades. This widespread phenomenon is presumably caused by a combination of factors like atmospheric fertilization or changes in forest structure and/or management. If not properly acknowledged in the calibration of tree-ring based climate reconstructions, considerable bias concerning amplitudes and trends of reconstructed climatic parameters might emerge or low frequency information is lost. Here we present a simple but effective, data-driven approach to remove the recent non-climatic growth increase in tree-ring data. Accounting for the no-analogue calibration problem, a new hydroclimatic reconstruction for northern-central Europe revealed considerably drier conditions during the medieval climate anomaly (MCA) compared with standard reconstruction methods and other existing reconstructions. This demonstrates the necessity to account for fertilization effects in modern tree-ring data from affected regions before calibrating reconstruction models, to avoid biased results.

Climatically controlled reproduction drives interannual growth variability in a temperate tree species.

Climatically controlled allocation to reproduction is a key mechanism by which climate influences tree growth and may explain lagged correlations between climate and growth. We used continent-wide datasets of tree-ring chronologies and annual reproductive effort in Fagus sylvatica from 1901 to 2015 to characterise relationships between climate, reproduction and growth. Results highlight that variable allocation to reproduction is a key factor for growth in this species, and that high reproductive effort ('mast years') is associated with stem growth reduction. Additionally, high reproductive effort is associated with previous summer temperature, creating lagged climate effects on growth. Consequently, understanding growth variability in forest ecosystems requires the incorporation of reproduction, which can be highly variable. Our results suggest that future response of growth dynamics to climate change in this species will be strongly influenced by the response of reproduction.

Distinct growth phenology but similar daily stem dynamics in three co-occurring broadleaved tree species.

Dendrometers offer a useful tool for long-term, high-resolution monitoring of tree responses to environmental fluctuations and climate change. Here, we analyze a 4-year dendrometer dataset (2014-17) on European beech (Fagus sylvatica L.), common hornbeam (Carpinus betulus L.) and pedunculate oak (Quercus robur L.), co-occuring in a mixed broadleaved forest in northeastern Germany. In our analyses, we focus both on seasonal growth dynamics as well as on the environmental forcing of daily stem-size variations. Over the study period with contrasting weather conditions, we observed species- and year-specific differences in growth phenology (i.e., growth onset, cessation and duration). Oak was characterized by early growth onset and long growth duration in all years as compared with beech and hornbeam. The analysis on the environmental forcing of daily stem dynamics revealed, however, highly similar responses for the studied species, with current-day vapor pressure deficit and sunshine duration negatively, and relative humidity and precipitation positively affecting stem size. When considering lagged effects, environmental conditions often oppositely affected stem-size changes. No consistent seasonality in environmental responses was detected, though specific weather conditions were found to affect temporal patterns in individual years. We suggest that the high similarity in environmental forcing observed between tree species can be explained by daily stem-size changes mainly reflecting tree water status rather than tree growth. Our results stress that correcting dendrometer series for reversible stem hydrological changes is of utmost importance to better quantify tree growth from dendrometers in future.

Species distribution models predict temporal but not spatial variation in forest growth.

Bioclimate envelope models have been widely used to illustrate the discrepancy between current species distributions and their potential habitat under climate change. However, the realism and correct interpretation of such projections has been the subject of considerable discussion. Here, we investigate whether climate suitability predictions correlate to tree growth, measured in permanent inventory plots and inferred from tree-ring records. We use the ensemble classifier RandomForest and species occurrence data from ~200,000 inventory plots to build species distribution models for four important European forestry species: Norway spruce, Scots pine, European beech, and pedunculate oak. We then correlate climate-based habitat suitability with volume measurements from ~50-year-old stands, available from ~11,000 inventory plots. Secondly, habitat projections based on annual historical climate are compared with ring width from ~300 tree-ring chronologies. Our working hypothesis is that habitat suitability projections from species distribution models should to some degree be associated with temporal or spatial variation in these growth records. We find that the habitat projections are uncorrelated with spatial growth records (inventory plot data), but they do predict interannual variation in tree-ring width, with an average correlation of .22. Correlation coefficients for individual chronologies range from values as high as .82 or as low as -.31. We conclude that tree responses to projected climate change are highly site-specific and that local suitability of a species for reforestation is difficult to predict. That said, projected increase or decrease in climatic suitability may be interpreted as an average expectation of increased or reduced growth over larger geographic scales.

Tuning the Voices of a Choir: Detecting Ecological Gradients in Time-Series Populations.

This paper introduces a new approach-the Principal Component Gradient Analysis (PCGA)-to detect ecological gradients in time-series populations, i.e. several time-series originating from different individuals of a population. Detection of ecological gradients is of particular importance when dealing with time-series from heterogeneous populations which express differing trends. PCGA makes use of polar coordinates of loadings from the first two axes obtained by principal component analysis (PCA) to define groups of similar trends. Based on the mean inter-series correlation (rbar) the gain of increasing a common underlying signal by PCGA groups is quantified using Monte Carlo Simulations. In terms of validation PCGA is compared to three other existing approaches. Focusing on dendrochronological examples, PCGA is shown to correctly determine population gradients and in particular cases to be advantageous over other considered methods. Furthermore, PCGA groups in each example allowed for enhancing the strength of a common underlying signal and comparably well as hierarchical cluster analysis. Our results indicate that PCGA potentially allows for a better understanding of mechanisms causing time-series population gradients as well as objectively enhancing the performance of climate transfer functions in dendroclimatology. While our examples highlight the relevance of PCGA to the field of dendrochronology, we believe that also other disciplines working with data of comparable structure may benefit from PCGA.

Can We Use Tree Rings of Black Alder to Reconstruct Lake Levels? A Case Study for the Mecklenburg Lake District, Northeastern Germany.

In this study, we explore the potential to reconstruct lake-level (and groundwater) fluctuations from tree-ring chronologies of black alder (Alnus glutinosa L.) for three study lakes in the Mecklenburg Lake District, northeastern Germany. As gauging records for lakes in this region are generally short, long-term reconstructions of lake-level fluctuations could provide valuable information on past hydrological conditions, which, in turn, are useful to assess dynamics of climate and landscape evolution. We selected black alder as our study species as alder typically thrives as riparian vegetation along lakeshores. For the study lakes, we tested whether a regional signal in lake-level fluctuations and in the growth of alder exists that could be used for long-term regional hydrological reconstructions, but found that local (i.e. site-specific) signals in lake level and tree-ring chronologies prevailed. Hence, we built lake/groundwater-level reconstruction models for the three study lakes individually. Two sets of models were considered based on (1) local tree-ring series of black alder, and (2) site-specific Standardized Precipitation Evapotranspiration Indices (SPEI). Although the SPEI-based models performed statistically well, we critically reflect on the reliability of these reconstructions, as SPEI cannot account for human influence. Tree-ring based reconstruction models, on the other hand, performed poor. Combined, our results suggest that, for our study area, long-term regional reconstructions of lake-level fluctuations that consider both recent and ancient (e.g., archaeological) wood of black alder seem extremely challenging, if not impossible.

Peatland pines as a proxy for water table fluctuations: disentangling tree growth, hydrology and possible human influence.

Dendrochronological investigations of Scots pine (Pinus sylvestris L.) growing on Männikjärve peatland in central Estonia showed that annual tree growth of peatland pines can be used as a proxy for past variations of water table levels. Reconstruction of past water table levels can help us to better understand the dynamics of various ecological processes in peatlands, e.g. the formation of vegetation patterns or carbon and nitrogen cycling. Männikjärve bog has one of the longest water table records in the boreal zone, continuously monitored since 1956. Common uncertainties encountered while working with peatland trees (e.g. narrow, missing and wedging rings) were in our case exacerbated with difficulties related to the instability of the relationship between tree growth and peatland environment. We hypothesized that the instable relationship was mainly due to a significant change of the limiting factor, i.e. the rise of the water table level due to human activity. To test our hypothesis we had to use several novel methods of tree-ring chronology analysis as well as to test explicitly whether undetected missing rings biased our results. Since the hypothesis that the instable relationship between tree growth and environment was caused by a change in limiting factor could not be rejected, we proceeded to find possible significant changes of past water table levels using structural analysis of the tree-ring chronologies. Our main conclusions were that peatland pines can be proxies to water table levels and that there were several shifting periods of high and low water table levels in the past 200 years.

Climate change and European forests: what do we know, what are the uncertainties, and what are the implications for forest management?

The knowledge about potential climate change impacts on forests is continuously expanding and some changes in growth, drought induced mortality and species distribution have been observed. However despite a significant body of research, a knowledge and communication gap exists between scientists and non-scientists as to how climate change impact scenarios can be interpreted and what they imply for European forests. It is still challenging to advise forest decision makers on how best to plan for climate change as many uncertainties and unknowns remain and it is difficult to communicate these to practitioners and other decision makers while retaining emphasis on the importance of planning for adaptation. In this paper, recent developments in climate change observations and projections, observed and projected impacts on European forests and the associated uncertainties are reviewed and synthesised with a view to understanding the implications for forest management. Current impact assessments with simulation models contain several simplifications, which explain the discrepancy between results of many simulation studies and the rapidly increasing body of evidence about already observed changes in forest productivity and species distribution. In simulation models uncertainties tend to cascade onto one another; from estimating what future societies will be like and general circulation models (GCMs) at the global level, down to forest models and forest management at the local level. Individual climate change impact studies should not be uncritically used for decision-making without reflection on possible shortcomings in system understanding, model accuracy and other assumptions made. It is important for decision makers in forest management to realise that they have to take long-lasting management decisions while uncertainty about climate change impacts are still large. We discuss how to communicate about uncertainty - which is imperative for decision making - without diluting the overall message. Considering the range of possible trends and uncertainties in adaptive forest management requires expert knowledge and enhanced efforts for providing science-based decision support.