Non-linear response of Norway spruce to climate variation along elevational and age gradients in the Carpathians.

Climate change, namely increased warming coupled with a rise in extreme events (e.g., droughts, storms, heatwaves), is negatively affecting forest ecosystems worldwide. In these ecosystems, growth dynamics and biomass accumulation are driven mainly by environmental constraints, inter-tree competition, and disturbance regimes. Usually, climate-growth relationships are assessed by linear correlation due to the simplicity and straightforwardness of modeling. However, applying this method may bias results, since the ecological and physiological responses of trees to environmental factors are non-linear, and usually bell-shaped. In the Eastern Carpathian, Norway spruce is at the southeasternmost edge of its natural occurrence; this region is thus potentially vulnerable to climate change. A non-linear assessment of climate-growth relationships using machine-learning techniques for Norway spruce in this area had not been conducted prior to this study. To address this knowledge gap, we analyzed a large tree-ring network from 158 stands, with over 3000 trees of varying age distributed along an elevational gradient. Our results showed that non-linearity in the growth-climate response of spruce was season-specific: temperatures from the previous autumn and current growing season, along with water availability during winter, induced a bell-shaped response. Moreover, we found that at low elevations, spruce growth was mainly limited by water availability in the growing season, while winter temperatures are likely to have had a slight influence along the entire elevational gradient. Furthermore, at elevations lower than 1400 m, spruce trees were also found to be sensitive to previous autumn water availability. Overall, our results shed new light on the response of Norway spruce to climate in the Carpathians, which may aid in management decisions.

Daytime warming triggers tree growth decline in the Northern Hemisphere.

Global warming has been linked to declines in tree growth. However, it is unclear how the asymmetry in daytime and nighttime warming influences this response. Here, we use 2947 residual tree-ring width chronologies covering 32 species at 2493 sites, between 1901 and 2018, across the Northern Hemisphere, to analyze the effects of daytime and nighttime temperatures, precipitation, and drought stress on the radial growth of trees. We show that drought stress was primarily triggered by daytime rather than nighttime warming. The radial growth of trees was more sensitive to drought stress in warm regions than in cold regions, especially for angiosperms. Our study provides robust evidence that daytime warming is the primary driver of the observed declines in forest productivity related to drought stress and that daytime and nighttime warming should be considered separately when modelling forest-climate interactions and feedbacks in a future, warmer world.

Applying space-for-time substitution to infer the growth response to climate may lead to overestimation of tree maladaptation: Evidence from the North American White Spruce Network.

Under climate change circumstances, increasing studies have reported the temporal instability of tree growth responses to climate, which poses a major challenge to linearly extrapolating past climate and future growth dynamics using tree-ring data. Space-for-time substitution (SFTS) is a potential solution to this problem that is widely used in the dendrochronology field to project past or future temporal growth response trajectories from contemporary spatial patterns. However, the projected accuracy of the SFTS in the climate effects on tree growth remains uncertain. Here, we empirically test the SFTS method by comparing the effect of spatial and temporal climate variations on climate responses of white spruce (Picea glauca), which has a transcontinental range in North America. We first applied a response surface regression model to capture the variations in growth responses along the spatial climate gradients. The results showed that the relationships between growth and June temperature varied along spatial climate gradients in a predictable way. And their relationships varied mainly along with local temperate condition. Then, the projected correlation coefficients between growth and climate using SFTS were compared against the observed. We found that the growth response changes caused by spatial versus temporal climate variations showed opposite trends. Moreover, the projected correlation coefficients using the SFTS were significantly lower than the observed. This finding suggests that applying the SFTS to project the growth response of white spruce might lead to an overestimation of the degree of tree maladaptation in future climate scenarios. And the overestimation is likely to get weaker from Alaska and Yukon Territory in the west to Quebec in the east. Although this is only a case study of the SFTS method for projecting tree growth response, our findings suggest that direct application of the SFTS method may not be applicable to all regions and all tree species.

Non-linear modelling reveals a predominant moisture limit on juniper growth across the southern Tibetan Plateau.

BACKGROUND AND AIMS: Tree growth in plateau forests is critically limited by harsh climatic conditions. Many mathematical statistical methods have been used to identify the relationships between tree growth and climatic factors, but there is still uncertainty regarding the relative importance of these factors across different regions. We tested major climatic limits at 30 sites to provide insights into the main climatic limits for juniper trees (Juniperus tibetica Kom.) across the southern Tibetan Plateau. METHODS: We analysed the linear and non-linear relationships between tree growth and climatic factors using Pearson correlation statistics and a process-based forward Vaganov-Shashkin-Lite (VS-Lite) model, respectively. These relationships were used to identify the strength of the influence of different climatic factors throughout the species' growing season and to identify the main climatic factors limiting tree growth. KEY RESULTS: Growth of juniper trees began in April and ended in October in the study area. The radial growth of juniper trees was limited by soil moisture throughout the summer (June-August) of the current year at 24 sampling sites and was limited by temperature at the other six sites on the southern Tibetan Plateau. CONCLUSIONS: Soil moisture limited juniper growth at the majority of sites. Temperature in the current summer limited the growth of juniper trees at a few sampling sites in the western part of the study area. Local climate conditions may contribute to different limiting factors in the growth response of trees on the southern Tibetan Plateau. These findings may contribute to our understanding of divergent forest dynamics and to sustainable forest management under future climate scenarios.

Tree-ring-based seasonal temperature reconstructions and ecological implications of recent warming on oak forest health in the Zagros Mountains, Iran.

Abrupt changes in temperature have especially strong impacts on fragile ecosystems located in semi-arid regions. In this study, we analyzed tree-ring widths (TRW) of Mediterranean cypress (Cupressus sempervirens var. horizontalis) in the Zagros Mountains, Iran. Furthermore, we separately measured earlywood width (EWW) and latewood width (LWW) of Persian oak (Quercus brantii Lindl.) to examine if intra-annual resolution of tree-ring parameters of Q. brantii tree rings can be used as high-resolution paleoclimate proxies. Climate-growth relationships revealed that mean monthly maximum temperatures (Tmax) are a dominant factor determining radial tree growth and negatively affect both oak and cypress in the Zagros Mountains. Accordingly, we reconstructed two different seasonal windows of past Tmax variability, namely, January-March and June-August over the periods 1860-2015 and 1560-2015, respectively. Regime shift detection identified twelve warm and nine cold significant regime shifts in our summer Tmax reconstructions. The longest hot summer period occurred in the twentieth century, and two warm regime shifts occurred in 1999 and 2008. The highest values of the warm summer regime shift index occurred in 2008, which coincided with fungal pathogen attacks and insect outbreak of the oak leaf roller moth (Tortrix viridana L.) in the Zagros oak woodlands. Interestingly, we found common warm and cold periods in historic climate variability between the summer and winter Tmax reconstructions. Warm and cold regime shifts occurred simultaneously from 1955 to 2015, and significant regional warm summer and winter regime shifts have occurred between 2008 and 2015. The winter and summer Tmax reconstructions show high spatial correlations with large areas in West Asia, North Africa, and the eastern Mediterranean region. Our results strengthen initial studies on past climate variability in Iran and contribute to an enhanced understanding of past temperature variability in West Asia.

A 168-year temperature recording based on tree rings and latitude differences in temperature changes in northeast China.

A ring-width series was developed from Dahurian larch (Larix gmelinii) in the northeastern forest area of Inner Mongolia, China. By analyzing the relationships between tree-ring data and climate records, an August-September mean maximum temperature (T89) series during 1845 and 2012 was reconstructed based on a simple linear regression equation. This reconstructed series explained 40.9% variance of the observed temperature from 1959 to 2012. The reconstructed T89 series was consistent with the historical disaster events caused by extreme climate (e.g., flood, frost disaster, and cold damage). Besides, the temperature comparisons showed that the year in which the warm months (April-September) in northeast China began to warm up has latitude differences. It started with a gradual delay from north to south, starting 1980 in the south region, after 1950 AD in the central region and after 1940 in the north region. Our study can enrich high-resolution temperature series in Northeast China and help clarify the characteristic of recent warming in northeast China.

Moisture-driven shift in the climate sensitivity of white spruce xylem anatomical traits is coupled to large-scale oscillation patterns across northern treeline in northwest North America.

Tree growth at northern treelines is generally temperature-limited due to cold and short growing seasons. However, temperature-induced drought stress was repeatedly reported for certain regions of the boreal forest in northwestern North America, provoked by a significant increase in temperature and possibly reinforced by a regime shift of the pacific decadal oscillation (PDO). The aim of this study is to better understand physiological growth reactions of white spruce, a dominant species of the North American boreal forest, to PDO regime shifts using quantitative wood anatomy and traditional tree-ring width (TRW) analysis. We investigated white spruce growth at latitudinal treeline across a >1,000 km gradient in northwestern North America. Functionally important xylem anatomical traits (lumen area, cell-wall thickness, cell number) and TRW were correlated with the drought-sensitive standardized precipitation-evapotranspiration index of the growing season. Correlations were computed separately for complete phases of the PDO in the 20th century, representing alternating warm/dry (1925-1946), cool/wet (1947-1976) and again warm/dry (1977-1998) climate regimes. Xylem anatomical traits revealed water-limiting conditions in both warm/dry PDO regimes, while no or spatially contrasting associations were found for the cool/wet regime, indicating a moisture-driven shift in growth-limiting factors between PDO periods. TRW reflected only the last shift of 1976/1977, suggesting different climate thresholds and a higher sensitivity to moisture availability of xylem anatomical traits compared to TRW. This high sensitivity of xylem anatomical traits permits to identify first signs of moisture-driven growth in treeline white spruce at an early stage, suggesting quantitative wood anatomy being a powerful tool to study climate change effects in the northwestern North American treeline ecotone. Projected temperature increase might challenge growth performance of white spruce as a key component of the North American boreal forest biome in the future, when drier conditions are likely to occur with higher frequency and intensity.

What prevails in climatic response of Pinus sylvestris in-between its range limits in mountains: slope aspect or elevation?

The roles of slope orientation and elevational temperature gradient were investigated for Scots pine (Pinus sylvestris L.) growth in the middle of its growth range, where these factors can significantly modulate microclimate and thus plant growth. We assumed that slope orientation causes more complex and severe effects than elevation because it influences all three main factors of plant growth: light, heat, and moisture. In addition to the total ring width, the earlywood and latewood width and latewood ratio were considered variables that contain information about tree ring growth during the season and wood structure over all tree life span on three sampling sites at different elevations and opposite slopes. Despite the observed dependence of pine growth rate on temperature and solar radiation, the mean latewood ratio is stable and similar between all sampling sites, being presumably defined by the genotype of individual trees. The seasonality of the climatic response of tree growth is bound to spatiotemporal variation of the vegetative season timing due to the elevational temperature lapse and local warming. However, its direction is primarily defined by slope orientation, where southern slope is moisture-limited, even at adjacent sites, and divergent climatic reactions of earlywood (weak moisture-limited in the last decades) and latewood growth (temperature-limited) were revealed on the northern slope.

Contrasting resistance and resilience to extreme drought and late spring frost in five major European tree species.

Extreme climate events (ECEs) such as severe droughts, heat waves, and late spring frosts are rare but exert a paramount role in shaping tree species distributions. The frequency of such ECEs is expected to increase with climate warming, threatening the sustainability of temperate forests. Here, we analyzed 2,844 tree-ring width series of five dominant European tree species from 104 Swiss sites ranging from 400 to 2,200 m a.s.l. for the period 1930-2016. We found that (a) the broadleaved oak and beech are sensitive to late frosts that strongly reduce current year growth; however, tree growth is highly resilient and fully recovers within 2 years; (b) radial growth of the conifers larch and spruce is strongly and enduringly reduced by spring droughts-these species are the least resistant and resilient to droughts; (c) oak, silver fir, and to a lower extent beech, show higher resistance and resilience to spring droughts and seem therefore better adapted to the future climate. Our results allow a robust comparison of the tree growth responses to drought and spring frost across large climatic gradients and provide striking evidence that the growth of some of the most abundant and economically important European tree species will be increasingly limited by climate warming. These results could serve for supporting species selection to maintain the sustainability of forest ecosystem services under the expected increase in ECEs.

Water stress limits transpiration and growth of European larch up to the lower subalpine belt in an inner-alpine dry valley.

Climate change will further constrain water availability in dry inner-alpine environments and affect water relations and growth conditions in mountain forests, including the widespread larch forests. To estimate the effects of climate conditions on water balance and growth, variation in sap flow and stem radius of European larch was measured for 3 yr along an elevation transect from 1070 to 2250 m above sea level (asl) in an inner-alpine dry valley in South Tyrol/Italy. Additionally, long-term climate-growth relations were derived from tree cores. Sap flow and radial growth were reduced in dry periods up to an elevation of 1715 m, leading to maximum annual growth at 2000 m. In a wet year no growth difference between elevations was observed. Long-term tree ring data showed a positive growth response to precipitation up to 1715 m and to temperature only above 2000 m. Our results demonstrate that reduced water availability and higher atmospheric water demand limit larch at low elevation within dry Alpine regions. This indicates a general upward shift of this species' elevational amplitude upon climate change, and respective negative effects on future silvicultural use and ecosystem services at lower elevations in the European Alps.

Drought timing influences the legacy of tree growth recovery.

Whether and how the timing of extreme events affects the direction and magnitude of legacy effects on tree growth is poorly understood. In this study, we use a global database of Ring-Width Index (RWI) from 2,500 sites to examine the impact and legacy effects (the departure of observed RWI from expected RWI) of extreme drought events during 1948-2008, with a particular focus on the influence of drought timing. We assessed the recovery of stem radial growth in the years following severe drought events with separate groupings designed to characterize the timing of the drought. We found that legacies from extreme droughts during the dry season (DS droughts) lasted longer and had larger impacts in each of the 3 years post drought than those from extreme droughts during the wet season (WS droughts). At the global scale, the average integrated legacy from DS droughts (0.18) was about nine times that from WS droughts (0.02). Site-level comparisons also suggest stronger negative impacts or weaker positive impacts of DS droughts on tree growth than WS droughts. Our results, therefore, highlight that the timing of drought is a crucial factor determining drought impacts on tree recovery. Further increases in baseline aridity could therefore exacerbate the impact of punctuated droughts on terrestrial ecosystems.

Past crops yield dynamics reconstruction from tree-ring chronologies in the forest-steppe zone based on low- and high-frequency components.

Interrelations of the yield variability of the main crops (wheat, barley, and oats) with hydrothermal regime and growth of conifer trees (Pinus sylvestris and Larix sibirica) in forest-steppes were investigated in Khakassia, South Siberia. An attempt has been made to understand the role and mechanisms of climatic impact on plants productivity. It was found that amongst variables describing moisture supply, wetness index had maximum impact. Strength of climatic response and correlations with tree growth are different for rain-fed and irrigated crops yield. Separated high-frequency variability components of yield and tree-ring width have more pronounced relationships between each other and with climatic variables than their chronologies per se. Corresponding low-frequency variability components are strongly correlated with maxima observed after 1- to 5-year time shift of tree-ring width. Results of analysis allowed us to develop original approach of crops yield dynamics reconstruction on the base of high-frequency variability component of the growth of pine and low-frequency one of larch.

Recent warming evidence inferred from a tree-ring-based winter-half year minimum temperature reconstruction in northwestern Yichang, South Central China, and its relation to the large-scale circulation anomalies.

High-resolution winter temperature reconstructions in China are rare, yet vital for the comprehensive understanding of past climate change. In the present work, the first winter-half year minimum mean temperature from previous November to current April in northwestern Yichang, South Central China, was reconstructed back to 1875 based on tree-ring material. The reconstruction can explain 55 % of the variance over the calibration period during 1955-2011. The temperature maintained at comparatively low level before 1958, and an abnormal warming was seen since 1959. However, the warming trend stagnated after 2000 AD. 2001-2010 was the warmest decade not only during the instrumental period but also during the whole reconstructed period. The reconstruction indicates good spatial resemblance to other temperatures series in adjacent areas and Northern Hemisphere, yet the recent warming in this study is earlier and more prominent than that of Southeast China. This work also manifests that the winter-half year minimum temperature in study area has good agreement with summer (June-September) maximum temperature variation in Southeast China at decadal scale, except that the winter-half year warming in recent decades is more evident than summer. This reconstruction is not only useful in improving our knowledge of long-term temperature variation but also useful in predicting the tree growth dynamics in the future in the study area.

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Populus euphratica is an important tree species in the arid regions of Northwest China, which is sensitive to climate changes. Climate of the Northwest China is changing to be "warm and humid", but how it would affect the regional forest growth is not clear. In this study, the radial growth response of P. euphratica to major climatic factors and their temporal changes during 1984-2021 were analyzed by using dendrochronology method in the desert oasis ecotone of Cele in the southern Tarim basin. The results showed that tree-ring width index of P. euphratica had a significant negative correlation with temperature in September of the previous year, and in February and May of current year, had significant positive correlation with precipitation in September of previous year and March and May of current year, and had significant positive correlations with SPEI in February and May of current year. The relationships between tree-ring width index and combined month climatic factors were more obvious. The results of moving correlation analysis showed that the correlation between tree-ring width index and temperature in the growing season tended to be strengthened in recent years, while the correlation between tree-ring width index and precipitation, SPEI tended to be declined or remain stable. The variations of the relationships between tree-ring width index and combined month climatic factors were more obvious compared that with single month. Current regional climate is conducive to the growth and development, as well as the improvement of ecological shelter function of P. euphratica forest in the desert oasis ecotone of Cele.

Recent warming and increasing CO2 stimulate growth of dominant trees under no water limitation in South Korea.

Increases in temperatures and atmospheric CO2 concentration influence the growth performance of trees worldwide. The direction and intensity of tree growth and physiological responses to changing climate do, however, vary according to environmental conditions. Here we present complex, long-term, tree-physiological responses to unprecedented temperature increase in East Asia. For this purpose, we studied radial growth and isotopic (δ13C and δ18O) variations using tree-ring data for the past 100 yr of dominant Quercus mongolica trees from the cool-temperate forests from Hallasan, South Korea. Overall, we found that tree stem basal area increment, intercellular CO2 concentration and intrinsic water-use efficiency significantly increased over the last century. We observed, however, short-term variability in the trends of these variables among four periods identified by change point analysis. In comparison, δ18O did not show significant changes over time, suggesting no major hydrological changes in this precipitation-rich area. The strength and direction of growth-climate relationships also varied during the past 100 yr. Basal area increment (BAI) did not show significant relationships with the climate over the 1924-1949 and 1975-1999 periods. However, over 1950-1974, BAI was negatively affected by both temperature and precipitation, while after 2000, a temperature stimulus was observed. Finally, over the past two decades, the increase in Q. mongolica tree growth accelerated and was associated with high spring-summer temperatures and atmospheric CO2 concentrations and decreasing intrinsic water-use efficiency, δ18O and vapour pressure deficit, suggesting that the photosynthetic rate continued increasing under no water limitations. Our results indicate that the performance of dominant trees of one of the most widely distributed species in East Asia has benefited from recent global changes, mainly over the past two decades. Such findings are essential for projections of forest dynamics and carbon sequestration under climate change.

Growth trends clustering: A novel method for detecting forest disturbances and extracting climate signals in tree rings.

Tree-ring widths contain valuable historical information related to both forest disturbances and climate variability and changes within forests. However, current methods are still unable to accurately distinguish between disturbances and climate signals in tree rings, especially in the case of climate anomalies. To address this issue, we developed a novel method, called Growth Trends Clustering (GTC) that uses the distribution characteristics of tree-ring widths within a stand to distinguish the effects of climate and other forest disturbances. GTC employed a Gaussian mixture model to fit the probability density distribution of annual ring-width index (RWI) in a stand. Discriminative criteria were established to cluster diverse sub-distributions from the Gaussian mixture model into categories of growth release, suppression, or normal trends. This approach allowed us to identify the occurrence, duration, and severity of forest disturbances based on percentage changes in the growth release or suppression categories of trees. And the effect of climate on tree growth was assessed according to the mean statistics of the growth normal categories. Using common forest disturbances such as defoliating insects and thinning as examples, we validated our method using tree-ring collections from six sites in British Columbia and Quebec, Canada. We found that the GTC method was superior to traditional time-series analysis methods (e.g., Radial Growth Averaging, Boundary Line, Absolute Increase, and Curve Intervention Detection) for detecting past forest disturbances and was able to significantly enhance climate signals. In summary, the GTC method presented in this study introduces a novel statistical approach for accurately distinguishing between forest disturbances and climate signals in tree rings. This is particularly important for understanding forest disturbance regimes under climate change and for developing future disturbance mitigation strategies.

Hydrothermal environments lead to differences in the radial growth response of Pinus tabulaeformis to climate in the monsoon marginal zone, Northwestern China.

Regional climatic differences increase the complexity of tree radial growth responses to climate change in the monsoon marginal zones and may alter the carbon sequestration capacity of forests. In this study, we collected cores of Pinus tabulaeformis trees at nine sampling sites across different regions. We analysed the relationship between tree-ring width chronology and climatic factors at different sites using dendroecological methods. We used the tree-ring index to calculate resistance, recovery, and resilience as well as to explore the capacity of radial growth to cope with drought events. The results indicate that (1) Drought was the primary factor limiting tree growth, and tree-ring climate response patterns varied across three regions. Tree growth was sensitive to both temperature and precipitation in the eastern Qilian Mountains, while it was more sensitive to temperature in the Hassan Mountains and more sensitive to precipitation in the Helan Mountains. (2) The tree-ring climate response pattern remained unstable over time, and the relative influence of current climate on tree growth increased. (3) The ecological resilience of trees to extreme events varies across three regions, which could be attributed to regional moisture conditions and the duration of drought. In the context of the management and protection of trees in the study area in the future, more attention should be paid to the elasticity of tree growth after drought events.

Factors Limiting Radial Growth of Conifers on Their Semiarid Borders across Kazakhstan.

The forests of Central Asia are biodiversity hotspots at risk from rapid climate change, but they are understudied in terms of the climate-growth relationships of trees. This classical dendroclimatic case study was performed for six conifer forest stands near their semiarid boundaries across Kazakhstan: (1-3) Pinus sylvestris L., temperate forest steppes; (4-5) Picea schrenkiana Fisch. & C.A. Mey, foothills, the Western Tien Shan, southeast; (6) Juniperus seravschanica Kom., montane zone, the Western Tien Shan, southern subtropics. Due to large distances, correlations between local tree-ring width (TRW) chronologies are significant only within species (pine, 0.19-0.50; spruce, 0.55). The most stable climatic response is negative correlations of TRW with maximum temperatures of the previous (from -0.37 to -0.50) and current (from -0.17 to -0.44) growing season. The strength of the positive response to annual precipitation (0.10-0.48) and Standardized Precipitation Evapotranspiration Index (0.15-0.49) depends on local aridity. The timeframe of climatic responses shifts to earlier months north-to-south. For years with maximum and minimum TRW, differences in seasonal maximal temperatures (by ~1-3 °C) and precipitation (by ~12-83%) were also found. Heat stress being the primary factor limiting conifer growth across Kazakhstan, we suggest experiments there on heat protection measures in plantations and for urban trees, alongside broadening the coverage of the dendroclimatic net with accents on the impact of habitat conditions and climate-induced long-term growth dynamics.

Reversal in the drought stress response of the Scots pine forest ecosystem: Local soil water regime as a key to improving climate change resilience.

In a changing climate, forest ecosystems have become increasingly vulnerable to continuously exacerbating heat and associated drought conditions. Climate stress resilience is governed by a complex interplay of global, regional, and local factors, with hydrological conditions being among the key players. We studied a Scots pine (Pinus sylvestris L.) forest ecosystem located near the southern edge of the boreal ecotone, which is particularly subjected to frequent and prolonged droughts. By comparing the dendrochronological series of pines growing in apparently contrasting hydrological conditions ranging from the waterlogged peat bog area to the dry soil at the surrounding elevations, we investigated how the soil water regime affects the climate response and drought stress resilience of the forest ecosystem. We found that in the dry land area, a significant fraction of the trees were replaced after two major climate extremes: prolonged drought and extremely low winter temperatures. The latter has also been followed by a three- to ten-fold growth reduction of the trees that survived in the next year, whereas no similar effect has been observed in the peat bog area. Multi-scale detrended partial cross-correlation analysis (DPCCA) indicated that tree-ring width (TRW) was negatively correlated with spring and summer temperatures and positively correlated with the Palmer drought severity index (PDSI) for the same year. For the elevated dry land area, the above effect extends to interannual scales, indicating that prolonged heatwaves and associated droughts are among the factors that limit tree growth. In marked contrast, in the waterlogged peat bog area, a reversed tendency was observed, with prolonged dry periods as well as warmer springs and summers over several consecutive years, leading to increasing tree growth with a one- to three-year time lag. Altogether, our results indicate that the pessimal conditions of a warming climate could become favorable through the preservation of the soil water regime.

Sap flow and growth response of Norway spruce under long-term partial rainfall exclusion at low altitude.

Introduction: Under ongoing climate change, more frequent and severe drought periods accompanied by heat waves are expected in the future. Under these conditions, the tree's survival is conditioned by fast recovery of functions after drought release. Therefore, in the presented study, we evaluated the effect of long-term water reduction in soil on tree water use and growth dynamics of Norway spruce. Methods: The experiment was conducted in two young Norway spruce plots located on suboptimal sites at a low altitude of 440 m a.s.l. In the first plot (PE), 25% of precipitation throughfall was excluded since 2007, and the second one represented the control treatment with ambient conditions (PC). Tree sap flow, stem radial increment, and tree water deficit were monitored in two consecutive growing seasons: 2015-2016, with contrasting hydro-climatic conditions. Results: Trees in both treatments showed relatively isohydric behavior reflected in a strong reduction of sap flow under the exceptional drought of 2015. Nevertheless, trees from PE treatment reduced sap flow faster than PC under decreasing soil water potential, exhibiting faster stomatal response. This led to a significantly lower sap flow of PE, compared to PC in 2015. The maximal sap flow rates were also lower for PE treatment, compared to PC. Both treatments experienced minimal radial growth during the 2015 drought and subsequent recovery of radial growth under the more the humid year of 2016. However, treatments did not differ significantly in stem radial increments within respective years. Discussion: Precipitation exclusion treatment, therefore, led to water loss adjustment, but did not affect growth response to intense drought and growth recovery in the year after drought.