Major tree species of Central European forests differ in their proportion of positive, negative, and nonstationary growth trends.

Temperate forests are undergoing significant transformations due to the influence of climate change, including varying responses of different tree species to increasing temperature and drought severity. To comprehensively understand the full range of growth responses, representative datasets spanning extensive site and climatic gradients are essential. This study utilizes tree-ring data from 550 sites from the temperate forests of Czechia to assess growth trends of six dominant Central European tree species (European beech, Norway spruce, Scots pine, silver fir, sessile and pedunculate oak) over 1990-2014. By modeling mean growth series for each species and site, and employing principal component analysis, we identified the predominant growth trends. Over the study period, linear growth trends were evident across most sites (56% increasing, 32% decreasing, and 10% neutral). The proportion of sites with stationary positive trends increased from low toward high elevations, whereas the opposite was true for the stationary negative trends. Notably, within the middle range of their distribution (between 500 and 700 m a.s.l.), Norway spruce and European beech exhibited a mix of positive and negative growth trends. While Scots pine growth trends showed no clear elevation-based pattern, silver fir and oaks displayed consistent positive growth trends regardless of site elevation, indicating resilience to the ongoing warming. We demonstrate divergent growth trajectories across space and among species. These findings are particularly important as recent warming has triggered a gradual shift in the elevation range of optimal growth conditions for most tree species and has also led to a decoupling of growth trends between lowlands and mountain areas. As a result, further future shifts in the elevation range and changes in species diversity of European temperate forests can be expected.

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.

Dendrometers challenge the 'moon wood concept' by elucidating the absence of lunar cycles in tree stem radius oscillation.

Wood is a sustainable natural resource and an important global commodity. According to the 'moon wood theory', the properties of wood, including its growth and water content, are believed to oscillate with the lunar cycle. Despite contradicting our current understanding of plant functioning, this theory is commonly exploited for marketing wooden products. To examine the moon wood theory, we applied a wavelet power transformation to series of 2,000,000 hourly stem radius records from dendrometers. We separated the influence of 74 consecutive lunar cycles and meteorological conditions on the stem variation of 62 trees and six species. We show that the dynamics of stem radius consist of overlapping oscillations with periods of 1 day, 6 months, and 1 year. These oscillations in stem dimensions were tightly coupled to oscillations in the series of air temperature and vapour pressure deficit. By contrast, we revealed no imprint of the lunar cycle on the stem radius variation of any species. We call for scepticism towards the moon wood theory, at least as far as the stem water content and radial growth are concerned. We foresee that similar studies employing robust scientific approaches will be increasingly needed in the future to cope with misleading concepts.

Linkage between growth phenology and climate-growth responses along landscape gradients in boreal forests.

Boreal forests represent an important carbon sink and, therefore, significantly contribute to climate change mitigation. Tree-ring width series of boreal species reflect climate variation at the moment of tree-ring formation but also lagged climatic effects from dormancy preceding tree-ring formation and antecedent growing seasons. However, little is known about how the growth sensitivity to climate in specific intra-annual periods varies across the landscape. Here, we assessed growth responses to climate variation during the 45 months preceding the tree-ring formation for nine boreal stands of Picea glauca and Picea mariana distributed along the gradients of elevation and slope aspect. We combined process-based modeling of wood formation and remote sensing data to determine growth phenology at each site. Next, we classified intra-annual seasons with significant climate-growth correlations based on the timing of dormancy and growth periods. Both the phenology and the climate-growth relationships systematically shifted with elevation and, to a lower extent, also with slope orientation at the treeline. The mean duration of the growing season varied between 100 days at treelines above 900 m and 160 days at lowlands below 500 m. The growth at treelines was stimulated by temperature in the summer of the tree-ring formation year and two years before tree-ring formation. The period of significant climate-growth correlations during the current summer did not exceed three months in agreement with the local duration of the growing season. The growth of trees in lower elevations was instead stimulated by high temperature during the dormancy periods but restricted by high temperature in antecedent summer seasons. In conclusion, our study highlights the linkage between the timing of climate-growth sensitivity and growth phenology, primarily determined by proximity to the treeline. Consequently, accounting for landscape gradients in growth phenology is crucial for upscaling the climatic limits of boreal stands' growth as climate change progresses.

Ecological and methodological drivers of non-stationarity in tree growth response to climate.

Radial tree growth is sensitive to environmental conditions, making observed growth increments an important indicator of climate change effects on forest growth. However, unprecedented climate variability could lead to non-stationarity, that is, a decoupling of tree growth responses from climate over time, potentially inducing biases in climate reconstructions and forest growth projections. Little is known about whether and to what extent environmental conditions, species, and model type and resolution affect the occurrence and magnitude of non-stationarity. To systematically assess potential drivers of non-stationarity, we compiled tree-ring width chronologies of two conifer species, Picea abies and Pinus sylvestris, distributed across cold, dry, and mixed climates. We analyzed 147 sites across the Europe including the distribution margins of these species as well as moderate sites. We calibrated four numerical models (linear vs. non-linear, daily vs. monthly resolution) to simulate growth chronologies based on temperature and soil moisture data. Climate-growth models were tested in independent verification periods to quantify their non-stationarity, which was assessed based on bootstrapped transfer function stability tests. The degree of non-stationarity varied between species, site climatic conditions, and models. Chronologies of P. sylvestris showed stronger non-stationarity compared with Picea abies stands with a high degree of stationarity. Sites with mixed climatic signals were most affected by non-stationarity compared with sites sampled at cold and dry species distribution margins. Moreover, linear models with daily resolution exhibited greater non-stationarity compared with monthly-resolved non-linear models. We conclude that non-stationarity in climate-growth responses is a multifactorial phenomenon driven by the interaction of site climatic conditions, tree species, and methodological features of the modeling approach. Given the existence of multiple drivers and the frequent occurrence of non-stationarity, we recommend that temporal non-stationarity rather than stationarity should be considered as the baseline model of climate-growth response for temperate forests.

Increasing water-use efficiency mediates effects of atmospheric carbon, sulfur, and nitrogen on growth variability of central European conifers.

Climate controls forest biomass production through direct effects on cambial activity and indirectly through interactions with CO2, air pollution, and nutrient availability. The atmospheric concentration of CO2, sulfur and nitrogen deposition can also exert a significant indirect control on wood formation since these factors influence the stomatal regulation of transpiration and carbon uptake, that is, intrinsic water use efficiency (iWUE). Here we provide 120-year long tree-ring time series of iWUE, stem growth, climatic and combined sulfur and nitrogen (SN) deposition trends for two common tree species, Pinus sylvestris (PISY) and Picea abies (PCAB), at their lower and upper distribution margins in Central Europe. The main goals were to explain iWUE trends using theoretical scenarios including climatic and SN deposition data, and to assess the contribution of climate and iWUE to the observed growth trends. Our results showed that after a notable increase in iWUE between the 1950s and 1980s, this positive trend subsequently slowed down. The substantial rise of iWUE since the 1950s resulted from a combination of an accelerated increase in atmospheric CO2 concentrations (Ca) and a stable level of leaf intercellular CO2 (Ci). The offset of observed iWUE values above the trajectory of a constant Ci/Ca scenario was explained by trends in SN deposition (all sites) together with the variation of drought conditions (low-elevation sites only). Increasing iWUE over the 20th and 21st centuries improved tree growth at low-elevation drought-sensitive sites. In contrast, at high-elevation PCAB sites, growth was mainly stimulated by recent warming. We propose that SN pollution should be considered in order to explain the steep increase in iWUE of conifers in the 20th century throughout Central Europe and other regions with a significant SN deposition history.

Limitation by vapour pressure deficit shapes different intra-annual growth patterns of diffuse- and ring-porous temperate broadleaves.

Understanding the effects of temperature and moisture on radial growth is vital for assessing the impacts of climate change on carbon and water cycles. However, studies observing growth at sub-daily temporal scales remain scarce. We analysed sub-daily growth dynamics and its climatic drivers recorded by point dendrometers for 35 trees of three temperate broadleaved species during the years 2015-2020. We isolated irreversible growth driven by cambial activity from the dendrometer records. Next, we compared the intra-annual growth patterns among species and delimited their climatic optima. The growth of all species peaked at air temperatures between 12 and 16°C and vapour pressure deficit (VPD) below 0.1 kPa. Acer pseudoplatanus and Fagus sylvatica, both diffuse-porous, sustained growth under suboptimal VPD. Ring-porous Quercus robur experienced a steep decline of growth rates with reduced air humidity. This resulted in multiple irregular growth peaks of Q. robur during the year. By contrast, the growth patterns of the diffuse-porous species were always right-skewed unimodal with a peak in June between day of the year 150-170. Intra-annual growth patterns are shaped more by VPD than temperature. The different sensitivity of radial growth to VPD is responsible for unimodal growth patterns in both diffuse-porous species and multimodal growth pattern in Q. robur.

Trends in climatically driven extreme growth reductions of Picea abies and Pinus sylvestris in Central Europe.

Extreme tree growth reductions represent events of abrupt forest productivity decline and carbon sequestration reduction. An increase in their magnitude can represent an early warning signal of impending tree mortality. Yet the long-term trends in extreme growth reductions remain largely unknown. We analyzed the trends in the proportion of trees exhibiting extreme growth reductions in two Central-European conifer species-Pinus sylvestris (PISY) and Picea abies (PCAB)-between 1901 and 2018. We used a novel approach for extreme growth reduction quantification by relating their size to their mean recurrence interval. Twenty-eight sites throughout Czechia and Slovakia with 1120 ring width series representing high- and low-elevation forests were inspected for extreme growth reductions with recurrence intervals of 15 and 50 years along with their link to climatic drivers. Our results show the greatest growth reductions at low-elevation PCAB sites, indicating high vulnerability of PCAB to drought. The proportions of trees exhibiting extreme growth reductions increased over time at low-elevation PCAB, decreased recently following an abrupt increase in the 1970-1980s at high-elevation PCAB, and showed nonsignificant trends in high- and low-elevation PISY. Climatic drivers of extreme growth reductions, however, shifted over time for all site categories as the proportion of low-temperature-induced extreme growth reductions declined since the 1990s, whereas events caused by drought consistently increased in frequency during the same period. We observed higher growth volatility at the lower range of distribution compared with the upper range margin of PISY and PCAB. This will undoubtedly considerably impact tree growth and vitality as temperatures and incidence of drought in Central Europe are expected to further increase with ongoing climate change.

Forward Modeling Reveals Multidecadal Trends in Cambial Kinetics and Phenology at Treeline.

Significant alterations of cambial activity might be expected due to climate warming, leading to growing season extension and higher growth rates especially in cold-limited forests. However, assessment of climate-change-driven trends in intra-annual wood formation suffers from the lack of direct observations with a timespan exceeding a few years. We used the Vaganov-Shashkin process-based model to: (i) simulate daily resolved numbers of cambial and differentiating cells; and (ii) develop chronologies of the onset and termination of specific phases of cambial phenology during 1961-2017. We also determined the dominant climatic factor limiting cambial activity for each day. To asses intra-annual model validity, we used 8 years of direct xylogenesis monitoring from the treeline region of the Krkonose Mts. (Czechia). The model exhibits high validity in case of spring phenological phases and a seasonal dynamics of tracheid production, but its precision declines for estimates of autumn phenological phases and growing season duration. The simulations reveal an increasing trend in the number of tracheids produced by cambium each year by 0.42 cells/year. Spring phenological phases (onset of cambial cell growth and tracheid enlargement) show significant shifts toward earlier occurrence in the year (for 0.28-0.34 days/year). In addition, there is a significant increase in simulated growth rates during entire growing season associated with the intra-annual redistribution of the dominant climatic controls over cambial activity. Results suggest that higher growth rates at treeline are driven by (i) temperature-stimulated intensification of spring cambial kinetics, and (ii) decoupling of summer growth rates from the limiting effect of low summer temperature due to higher frequency of climatically optimal days. Our results highlight that the cambial kinetics stimulation by increasing spring and summer temperatures and shifting spring phenology determine the recent growth trends of treeline ecosystems. Redistribution of individual climatic factors controlling cambial activity during the growing season questions the temporal stability of climatic signal of cold forest chronologies under ongoing climate change.

Growth-limiting factors and climate response variability in Norway spruce (Picea abies L.) along an elevation and precipitation gradients in Slovenia.

Norway spruce (Picea abies L.) is among the most sensitive coniferous species to ongoing climate change. However, previous studies on its growth response to increasing temperatures have yielded contrasting results (from stimulation to suppression), suggesting highly site-specific responses. Here, we present the first study that applies two independent approaches, i.e. the nonlinear, process-based Vaganov-Shashkin (VS) model and linear daily response functions. Data were collected at twelve sites in Slovenia differing in climate regimes and ranging elevation between 170 and 1300 m a.s.l. VS model results revealed that drier Norway spruce sites at lower elevations are mostly moisture limited, while moist high-elevation sites are generally more temperature limited. Daily response functions match well the pattern of growth-limiting factors from the VS model and further explain the effect of climate on radial growth: prevailing growth-limiting factors correspond to the climate variable with higher correlations. Radial growth correlates negatively with rising summer temperature and positively with higher spring precipitation. The opposite response was observed for the wettest site at the highest elevation, which positively reacts to increased summer temperature and will most likely benefit from a warming climate. For all other sites, the future radial growth of Norway spruce largely depends on the balance between spring precipitation and summer temperature.

Climate warming induced synchronous growth decline in Norway spruce populations across biogeographical gradients since 2000.

Covering large parts of Europe, Norway spruce (Picea abies L Karst.) plays an important role in the adaptation strategy of forest services to future climate change. Although dendroecology can provide valuable information on the past relationships between tree growth and climate, most previous studies were biased towards species-specific distribution limits, where old individuals grow slowly under extreme conditions. In the present study, we investigated the growth variability and climate sensitivity of 2851 Norway spruce trees along longitudinal (E 12-26°), latitudinal (N 45-51°), and elevation (118-1591 m a.s.l.) gradients in central-eastern Europe. We reveal that summer weather significantly affects the radial growth of spruce trees, but the effects strongly vary along biogeographical gradients. Extreme summer heatwaves in 2000 and 2003 reduced the growth rates by 10-35%, most pronounced in the southern Carpathians. In contrast to the population in the Czech Republic, climate warming induced a synchronous decline in the growth rates across biogeographical gradients in the Carpathian arc. By demonstrating the increased vulnerability of Norway spruce under warmer climate conditions, we recommended that the forest services and conservation managers replace or admix monocultures of this species with more drought-resilient mixtures including fir, beech and other broadleaved species.

Disentangling the effects of disturbance, climate and tree age on xylem hydraulic conductivity of Betula pendula.

BACKGROUND AND AIMS: The increasing frequency of disturbances in temperate forests is responsible for the greater numbers of trees with mechanically damaged cambial zones. Adjustment of wood anatomical structure to balance between safe and efficient water conductivity is one mechanism trees employ to cope with mechanical damage. The relative role of disturbances, tree age and climate in shaping xylem conduits and affecting xylem hydraulic conductivity remains unknown. METHODS: We performed an experiment with five different mechanical treatments simulating natural disturbances of juvenile Betula pendula trees (stem scarring, tilting, decapitation, root exposure and stem-base burial). After 3 years, trees were cut down, conduit size and density were measured, and specific hydraulic conductivity of each tree ring was calculated. Between-tree and between-year variability in xylem conductivity was decomposed into effects of tree age, climate and disturbances using linear mixed-effects models. KEY RESULTS: Xylem-specific hydraulic conductivity decreased significantly after treatment in decapitated, tilted and scarred trees. In the last treatment, wood anatomical adjustment was restricted to the area next to the callus tissue zone; in contrast, specific hydraulic conductivity declined over the entire stem circumference after tilting or decapitation. The response of trees with buried stems and exposed roots was generally weak. The overall effect of disturbances on inter-annual variability of wood anatomical structure was greater than the contribution of tree age and climate. CONCLUSIONS: The results indicate that disturbances are important drivers of xylem hydraulic conductivity. Expected increases in the frequency and intensity of disturbances may alter the theoretical capacity of forest stands for water conductance with a feedback to climate.

Increased spruce tree growth in Central Europe since 1960s.

Tree growth response to recent environmental changes is of key interest for forest ecology. This study addressed the following questions with respect to Norway spruce (Picea abies, L. Karst.) in Central Europe: Has tree growth accelerated during the last five decades? What are the main environmental drivers of the observed tree radial stem growth and how much variability can be explained by them? Using a nationwide dendrochronological sampling of Norway spruce in the Czech Republic (1246 trees, 266 plots), novel regional tree-ring width chronologies for 40(±10)- and 60(±10)-year old trees were assembled, averaged across three elevation zones (break points at 500 and 700m). Correspondingly averaged drivers, including temperature, precipitation, nitrogen (N) deposition and ambient CO2 concentration, were used in a general linear model (GLM) to analyze the contribution of these in explaining tree ring width variability for the period from 1961 to 2013. Spruce tree radial stem growth responded strongly to the changing environment in Central Europe during the period, with a mean tree ring width increase of 24 and 32% for the 40- and 60-year old trees, respectively. The indicative General Linear Model analysis identified CO2, precipitation during the vegetation season, spring air temperature (March-May) and N-deposition as the significant covariates of growth, with the latter including interactions with elevation zones. The regression models explained 57% and 55% of the variability in the two tree ring width chronologies, respectively. Growth response to N-deposition showed the highest variability along the elevation gradient with growth stimulation/limitation at sites below/above 700m. A strong sensitivity of stem growth to CO2 was also indicated, suggesting that the effect of rising ambient CO2 concentration (direct or indirect by increased water use efficiency) should be considered in analyses of long-term growth together with climatic factors and N-deposition.

Environmental factors exert strong control over the climate-growth relationships of Picea abies in Central Europe.

The growth response of trees to changing climate is frequently discussed as increasing temperatures and more severe droughts become major risks for forest ecosystems. However, the ability of trees to cope with the changing climate and the effects of other environmental factors on climate-growth relationships are still poorly understood. There is thus an increasing need to understand the ability of individual trees to cope with changing climate in various environments. To improve the current understanding, a large tree-ring network covering the whole area of the Czech Republic (in 7×7km grids) was utilized to investigate how the climate-growth relationships of Norway spruce are affected by 1) various geographical variables, 2) changing levels of acidic deposition, 3) soil characteristics and 4) age, tree diameter and neighbourhood competition. The period from 1930 to 2013 was divided into four, 21-year long intervals of differing levels of acidic deposition, which peaked in the 1972-1993 period. Our individual-based, spatiotemporal, multivariate analyses revealed that spruce growth was mostly affected by drought and warm summers. Drought plays the most important negative role at lower altitudes, while the positive effect of higher temperature was identified for trees at higher altitudes. Increased levels of acidic deposition, together with geographical variables, were identified as the most important factors affecting climate-growth association. Tree age, tree size and soil characteristics also significantly modulate climate-growth relationships. The importance of all environmental variables on climate-growth relationships was suppressed by acidic deposition during periods when this was at a high level; growth was significantly more enhanced by spring and summer temperatures during these periods. Our results suggest that spruce will undergo significant growth reduction under the predicted climate changes, especially at the lower altitudes which lie outside of its natural range.

Discerning environmental factors affecting current tree growth in Central Europe.

We examined the effect of individual environmental factors on the current spruce tree growth assessed from a repeated country-level statistical landscape (incl. forest) survey in the Czech Republic. An extensive set of variables related to tree size, competition, site characteristics including soil texture, chemistry, N deposition and climate was tested within a random-effect model to explain growth in the conditions of dominantly managed forest ecosystems. The current spruce basal area increment was assessed from two consecutive landscape surveys conducted in 2008/2009 and six years later in 2014/2015. Tree size, age and competition within forest stands were found to be the dominant explanatory variables, whereas the expression of site characteristics, environmental and climatic drives was weaker. The significant site variables affecting growth included soil C/N ratio and soil exchangeable acidity (pH KCl; positive response) reflecting soil chemistry, long-term N-deposition (averaged since 1975) in combination with soil texture (clay content) and Standardized Precipitation Index (SPI), a drought index expressing moisture conditions. Sensitivity of growth to N-deposition was positive, although weak. SPI was positively related to and significant in explaining tree growth when expressed for the growth season. Except SPI, no significant relation of growth was determined to altitude-related variables (temperature, growth season length). We identified the current spruce growth optimum at elevations about 800ma.s.l. or higher in the conditions of the country. This suggests that at lower elevations, limitation by a more pronounced water deficit dominates, whereas direct temperature limitation may concern the less frequent higher elevations. The mixed linear model of spruce tree growth explained 55 and 65% of the variability with fixed and random effects included, respectively, and provided new insights on the current spruce tree growth and factors affecting it within the environmental gradients of the country.