Finding balance: Tree-ring isotopes differentiate between acclimation and stress-induced imbalance in a long-term irrigation experiment.

Scots pine (Pinus sylvestris L.) is a common European tree species, and understanding its acclimation to the rapidly changing climate through physiological, biochemical or structural adjustments is vital for predicting future growth. We investigated a long-term irrigation experiment at a naturally dry forest in Switzerland, comparing Scots pine trees that have been continuously irrigated for 17 years (irrigated) with those for which irrigation was interrupted after 10 years (stop) and non-irrigated trees (control), using tree growth, xylogenesis, wood anatomy, and carbon, oxygen and hydrogen stable isotope measurements in the water, sugars and cellulose of plant tissues. The dendrochronological analyses highlighted three distinct acclimation phases to the treatments: irrigated trees experienced (i) a significant growth increase in the first 4 years of treatment, (ii) high growth rates but with a declining trend in the following 8 years and finally (iii) a regression to pre-irrigation growth rates, suggesting the development of a new growth limitation (i.e. acclimation). The introduction of the stop treatment resulted in further growth reductions to below-control levels during the third phase. Irrigated trees showed longer growth periods and lower tree-ring δ13 C values, reflecting lower stomatal restrictions than control trees. Their strong tree-ring δ18 O and δ2 H (O-H) relationship reflected the hydrological signature similarly to the control. On the contrary, the stop trees had lower growth rates, conservative wood anatomical traits, and a weak O-H relationship, indicating a physiological imbalance. Tree vitality (identified by crown transparency) significantly modulated growth, wood anatomical traits and tree-ring δ13 C, with low-vitality trees of all treatments performing similarly regardless of water availability. We thus provide quantitative indicators for assessing physiological imbalance and tree acclimation after environmental stresses. We also show that tree vitality is crucial in shaping such responses. These findings are fundamental for the early assessment of ecosystem imbalances and decline under climate change.

Differential growth rate, water use efficiency and climate sensitivity between males and females of Ilex aquifolium in north-western Spain.

BACKGROUND AND AIMS: Dioecious plant species, i.e., those in which male and female functions are housed in different individuals, are particularly vulnerable to global environmental changes. For long-lived plant species, such as trees, long-term studies are imperative to understand how growth patterns and their sensitivity to climate variability differentially affect the sexes. METHODS: Here, we explore long-term intersexual differences in wood traits, namely radial growth rates, water use efficiency quantified as stable carbon isotope abundance of wood cellulose, and their climate sensitivity in Ilex aquifolium trees growing in a natural population in NW Spain. KEY RESULTS: We found that sex differences in secondary growth rates were variable over time, with males outperforming females in both radial growth rates and water use efficiency in recent decades. Summer water stress significantly reduced the growth of female trees in the following growing season, while the growth of male trees was primarily favoured by cloudy and rainy conditions the previous fall and winter combined with low cloud cover and warm conditions in summer. Sex-dependent lagged correlations between radial growth and water availability were found, with a strong association between tree growth and cumulative water availability in females at 30 months and in males at 10 months. CONCLUSIONS: Overall, our results point to greater vulnerability of female tress to increasing drought, which could lead to sex-ratio biases threatening population viability in the future.

Urbanization exacerbates climate sensitivity of eastern United States broadleaf trees.

Tree growth is a key mechanism driving carbon sequestration in forest ecosystems. Environmental conditions are important regulators of tree growth that can vary considerably between nearby urban and rural forests. For example, trees growing in cities often experience hotter and drier conditions than their rural counterparts while also being exposed to higher levels of light, pollution, and nutrient inputs. However, the extent to which these intrinsic differences in the growing conditions of trees in urban versus rural forests influence tree growth response to climate is not well known. In this study, we tested for differences in the climate sensitivity of tree growth between urban and rural forests along a latitudinal transect in the eastern United States that included Boston, Massachusetts, New York City, New York, and Baltimore, Maryland. Using dendrochronology analyses of tree cores from 55 white oak trees (Quercus alba), 55 red maple trees (Acer rubrum), and 41 red oak trees (Quercus rubra) we investigated the impacts of heat stress and water stress on the radial growth of individual trees. Across our three-city study, we found that tree growth was more closely correlated with climate stress in the cooler climate cities of Boston and New York than in Baltimore. Furthermore, heat stress was a significant hindrance to tree growth in higher latitudes while the impacts of water stress appeared to be more evenly distributed across latitudes. We also found that the growth of oak trees, but not red maple trees, in the urban sites of Boston and New York City was more adversely impacted by heat stress than their rural counterparts, but we did not see these urban-rural differences in Maryland. Trees provide a wide range of important ecosystem services and increasing tree canopy cover was typically an important component of urban sustainability strategies. In light of our findings that urbanization can influence how tree growth responds to a warming climate, we suggest that municipalities consider these interactions when developing their tree-planting palettes and when estimating the capacity of urban forests to contribute to broader sustainability goals in the future.

Thinning followed by slash burning enhances growth and reduces vulnerability to drought for Pinus nigra.

Increasingly frequent severe drought events are pushing Mediterranean forests to unprecedented responses. Lack of management leads to dense forests that are highly susceptible to drought stress, potentially resulting in extensive dieback and increased vulnerability to other disturbances. Forest treatments like thinning and slash burning reduce competition for resources and have the potential to enhance tree growth and vigor and minimize tree vulnerability to drought. Here, we used tree rings to study the growth and physiological response of black pine (Pinus nigra) to drought in northeastern Spain under different treatments, including two thinning intensities (light and heavy, with 10% and 40% basal area reduction, respectively) followed by two understory treatments (clearing alone and in combination with slash burning), resulting in a research design of four treatments plus an untreated control with three replicates. Specifically, we studied basal area increment (BAI), resilience indices, and intrinsic water use efficiency (iWUE) using carbon and oxygen isotope composition (δ13C and δ18O in tree-ring cellulose) before and after treatments. Our results showed that BAI and resistance to drought increased in the heavy-thin (burned and unburned) and light-thin burned units. Resilience increased in the burned units regardless of the thinning intensity, while recovery was not affected by treatment. Slash burning additionally increased BAI in the light-thin and resistance and resilience in the heavy-thin units compared with clearing alone. The stable isotope analysis revealed a minor effect of treatments on δ13C and δ18O. No change in iWUE among treatments was presumably linked to a proportional increase in both net CO2 assimilation and stomatal conductance, which particularly increased in the heavy-thin (burned and unburned) and light-thin burned units, indicating that these trees were the least affected by drought. This study shows that management approaches aimed at reducing wildfire hazard can also increase the vigor of dominant trees under drought stress. By reducing competition both from the overstory and the understory, thinning followed by clearing alone or in combination with slash burning promotes tree growth and vigor and increases its resistance and resilience to drought.

A comprehensive resilience assessment of Mexican tree species and their relationship with drought events over the last century.

The resilience of forests to drought events has become a major natural resource sustainability concern, especially in response to climate change. Yet, little is known about the legacy effects of repeated droughts, and tree species ability to respond across environmental gradients. In this study, we used a tree-ring database (121 sites) to evaluate the overall resilience of tree species to drought events in the last century. We investigated how climate and geography affected the response at the species level. We evaluated temporal trends of resilience using a predictive mixed linear modeling approach. We found that pointer years (e.g., tree growth reduction) occurred during 11.3% of the 20th century, with an average decrease in tree growth of 66% compared to the previous period. The occurrence of pointer years was associated with negative values of the Standardized Precipitation Index (SPI, 81.6%) and Palmer Drought Severity Index (PDSI, 77.3%). Tree species differed in their resilience capacity, however, species inhabiting xeric conditions were less resistant but with higher recovery rates (e.g., Abies concolor, Pinus lambertiana, and Pinus jeffreyi). On average, tree species needed 2.7 years to recover from drought events, with extreme cases requiring more than a decade to reach pre-drought tree growth rates. The main abiotic factor related to resilience was precipitation, confirming that some tree species are better adapted to resist the effects of droughts. We found a temporal variation for all tree resilience indices (scaled to 100), with a decreasing resistance (-0.56 by decade) and resilience (-0.22 by decade), but with a higher recovery (+1.72 by decade) and relative resilience rate (+0.33 by decade). Our results emphasize the importance of time series of forest resilience, particularly by distinguishing the species-level response in the context of legacy of droughts, which are likely to become more frequent and intense under a changing climate.

Large old trees increase growth under shifting climatic constraints: Aligning tree longevity and individual growth dynamics in primary mountain spruce forests.

In a world of accelerating changes in environmental conditions driving tree growth, tradeoffs between tree growth rate and longevity could curtail the abundance of large old trees (LOTs), with potentially dire consequences for biodiversity and carbon storage. However, the influence of tree-level tradeoffs on forest structure at landscape scales will also depend on disturbances, which shape tree size and age distribution, and on whether LOTs can benefit from improved growing conditions due to climate warming. We analyzed temporal and spatial variation in radial growth patterns from ~5000 Norway spruce (Picea abies [L.] H. Karst) live and dead trees from the Western Carpathian primary spruce forest stands. We applied mixed-linear modeling to quantify the importance of LOT growth histories and stand dynamics (i.e., competition and disturbance factors) on lifespan. Finally, we assessed regional synchronization in radial growth variability over the 20th century, and modeled the effects of stand dynamics and climate on LOTs recent growth trends. Tree age varied considerably among forest stands, implying an important role of disturbance as an age constraint. Slow juvenile growth and longer period of suppressed growth prolonged tree lifespan, while increasing disturbance severity and shorter time since last disturbance decreased it. The highest age was not achieved only by trees with continuous slow growth, but those with slow juvenile growth followed by subsequent growth releases. Growth trend analysis demonstrated an increase in absolute growth rates in response to climate warming, with late summer temperatures driving the recent growth trend. Contrary to our expectation that LOTs would eventually exhibit declining growth rates, the oldest LOTs (>400 years) continuously increase growth throughout their lives, indicating a high phenotypic plasticity of LOTs for increasing biomass, and a strong carbon sink role of primary spruce forests under rising temperatures, intensifying droughts, and increasing bark beetle outbreaks.

Differences in leaf gas exchange strategies explain Quercus rubra and Liriodendron tulipifera intrinsic water use efficiency responses to air pollution and climate change.

Trees continuously regulate leaf physiology to acquire CO2 while simultaneously avoiding excessive water loss. The balance between these two processes, or water use efficiency (WUE), is fundamentally important to understanding changes in carbon uptake and transpiration from the leaf to the globe under environmental change. While increasing atmospheric CO2 (iCO2 ) is known to increase tree intrinsic water use efficiency (iWUE), less clear are the additional impacts of climate and acidic air pollution and how they vary by tree species. Here, we couple annually resolved long-term records of tree-ring carbon isotope signatures with leaf physiological measurements of Quercus rubra (Quru) and Liriodendron tulipifera (Litu) at four study locations spanning nearly 100 km in the eastern United States to reconstruct historical iWUE, net photosynthesis (Anet ), and stomatal conductance to water (gs ) since 1940. We first show 16%-25% increases in tree iWUE since the mid-20th century, primarily driven by iCO2 , but also document the individual and interactive effects of nitrogen (NOx ) and sulfur (SO2 ) air pollution overwhelming climate. We find evidence for Quru leaf gas exchange being less tightly regulated than Litu through an analysis of isotope-derived leaf internal CO2 (Ci ), particularly in wetter, recent years. Modeled estimates of seasonally integrated Anet and gs revealed a 43%-50% stimulation of Anet was responsible for increasing iWUE in both tree species throughout 79%-86% of the chronologies with reductions in gs attributable to the remaining 14%-21%, building upon a growing body of literature documenting stimulated Anet overwhelming reductions in gs as a primary mechanism of increasing iWUE of trees. Finally, our results underscore the importance of considering air pollution, which remains a major environmental issue in many areas of the world, alongside climate in the interpretation of leaf physiology derived from tree rings.

Radial growth of Korshinsk peashrub and its response to drought in different sub-arid climate regions of northwest China.

The increased frequency and intensity of droughts have seriously affected the stability of plantation ecosystems in the Chinese Loess Plateau. Caragana korshinskii Kom. was the dominant afforested shrub species in this region. Evaluating the radial growth of C. korshinskii and its response to drought can provide valuable information for sustainable management of plantations in the context of climate change. In this study, based on 237 shrub C. korshinskii annual ring samples from nine sites in different climate regions, we investigated the response of C. korshinskii radial growth to climate (temperature, precipitation, and monthly resolved standardized precipitation evapotranspiration index (SPEI_01)), and evaluated the differences between them using calculated indices of drought resistance, recovery, and resilience. The results demonstrate that the radial growth of C. korshinskii was mainly limited by drought stress in the previous September in arid regions and in March and June in semi-arid regions, whereas C. korshinskii in semi-humid regions was less influenced by drought stress. Recovery after drought decreased with increasing resistance, and resilience increased significantly with increasing resistance and recovery. Differences in precipitation were found to be the main factor generating variations in shrub resilience; with an increase in precipitation, the recovery and resilience after drought gradually increased. For plantation management, this study suggests that efficient utilization of precipitation resources and site-specific afforestation in different climate and site conditions may help to enhance resilience and improve the ecological service function of plantation forests in the Loess Plateau.

Tree-ring evidence of ecological stress memory.

Plants experiencing stress could develop the ability to reshape their response toward present stress based on past stress experience, called 'ecological stress memory' (ESM), which is important for plant acclimation to repeated stresses. Although ESM has been largely reported, it remains unclear whether ESM could improve tree resistance to recurrent stress in subsequent decades. Here, we explore it from a tree-ring network of 1491 trees from 50 long-living juniper forests on the Tibetan Plateau. Through comparing performances of tree radial growth in past sequential growth stresses, we found that trees could obtain ESM under antecedent stresses and elevate resistance to subsequent stress after several years or even decades. Such positive effects of ESM are associated with post-stress recovery. Trees with slow recovery trajectories after antecedent stress show significantly improved resistance to subsequent stress, while trees with extremely fast post-stress recovery showed decreased resistance to subsequent stress. These results imply that temporary depressive tree radial growth after antecedent stress might be a trigger of long storage of ESM. Incorporating positive effects of ESM and relationship between ESM activation and post-stress recovery into future Earth system models could advance our capacity to predict forest dynamics and forest ecosystem stabilization under future stress conditions.

Common mechanisms explain nitrogen-dependent growth of Arctic shrubs over three decades despite heterogeneous trends and declines in soil nitrogen availability.

Heterogeneity has been observed in the responses of Arctic shrubs to climate variability over recent decades, which may reflect landscape-scale variability in belowground resources. At a northern fringe of tall shrub expansion (Yuribei, Yamal Peninsula, Russia), we sought to determine the mechanisms relating nitrogen (N) limitation to shrub growth over decadal time. We analysed the ratio of 15 N to 14 N isotopes in wood rings of 10 Salix lanata individuals (399 measurements) to reconstruct annual point-based bioavailable N between 1980 and 2013. We applied a model-fitting/model-selection approach with a suite of competing ecological models to assess the most-likely mechanisms that explain each shrub's individual time-series. Shrub δ15 N time-series indicated declining (seven shrubs), increasing (two shrubs) and no trend (one shrub) in N availability. The most appropriate model for all shrubs included N-dependent growth of linear rather than saturating form. Inclusion of plant-soil feedbacks better explained ring width and δ15 N for eight of 10 individuals. Although N trajectories were individualistic, common mechanisms of varying strength confirmed the N-dependency of shrub growth. The linear mechanism may reflect intense scavenging of scarce N; the importance of plant-soil feedbacks suggests that shrubs subvert the microbial bottleneck by actively controlling their environment.

Local groundwater decline exacerbates response of dryland riparian woodlands to climatic drought.

Dryland riparian woodlands are considered to be locally buffered from droughts by shallow and stable groundwater levels. However, climate change is causing more frequent and severe drought events, accompanied by warmer temperatures, collectively threatening the persistence of these groundwater dependent ecosystems through a combination of increasing evaporative demand and decreasing groundwater supply. We conducted a dendro-isotopic analysis of radial growth and seasonal (semi-annual) carbon isotope discrimination (Δ13 C) to investigate the response of riparian cottonwood stands to the unprecedented California-wide drought from 2012 to 2019, along the largest remaining free-flowing river in Southern California. Our goals were to identify principal drivers and indicators of drought stress for dryland riparian woodlands, determine their thresholds of tolerance to hydroclimatic stressors, and ultimately assess their vulnerability to climate change. Riparian trees were highly responsive to drought conditions along the river, exhibiting suppressed growth and strong stomatal closure (inferred from reduced Δ13 C) during peak drought years. However, patterns of radial growth and Δ13 C were quite variable among sites that differed in climatic conditions and rate of groundwater decline. We show that the rate of groundwater decline, as opposed to climate factors, was the primary driver of site differences in drought stress, and trees showed greater sensitivity to temperature at sites subjected to faster groundwater decline. Across sites, higher correlation between radial growth and Δ13 C for individual trees, and higher inter-correlation of Δ13 C among trees were indicative of greater drought stress. Trees showed a threshold of tolerance to groundwater decline at 0.5 m year-1 beyond which drought stress became increasingly evident and severe. For sites that exceeded this threshold, peak physiological stress occurred when total groundwater recession exceeded ~3 m. These findings indicate that drought-induced groundwater decline associated with more extreme droughts is a primary threat to dryland riparian woodlands and increases their susceptibility to projected warmer temperatures.

Cumulative growth and stress responses to the 2018-2019 drought in a European floodplain forest.

Droughts increasingly threaten the world's forests and their potential to mitigate climate change. In 2018-2019, Central European forests were hit by two consecutive hotter drought years, an unprecedented phenomenon that is likely to occur more frequently with climate change. Here, we examine tree growth and physiological stress responses (increase in carbon isotope composition; Δδ13 C) to this consecutive drought based on tree rings of dominant tree species in a Central European floodplain forest. Tree growth was not reduced for most species in 2018, indicating that water supply in floodplain forests can partly buffer meteorological water deficits. Drought stress responses in 2018 were comparable to former single drought years but the hotter drought in 2018 induced drought legacies in tree growth while former droughts did not. We observed strong decreases in tree growth and increases in Δδ13 C across all tree species in 2019, which are likely driven by the cumulative stress both consecutive hotter droughts exerted. Our results show that consecutive hotter droughts pose a novel threat to forests under climate change, even in forest ecosystems with comparably high levels of water supply.

Harnessing tree-ring phenotypes to disentangle gene by environment interactions and their climate dependencies in a circum-Mediterranean pine.

BACKGROUND AND AIMS: Understanding the genetic basis of adaptation and plasticity in trees constitutes a knowledge gap. We linked dendrochronology and genomics [single nucleotide polymorphisms (SNPs)] for a widespread conifer (Pinus halepensis Mill.) to characterize intraspecific growth differences elicited by climate. METHODS: The analysis comprised 20-year tree-ring series of 130 trees structured in 23 populations evaluated in a common garden. We tested for genotype by environment interactions (G × E) of indexed ring width (RWI) and early- to latewood ratios (ELI) using factorial regression, which describes G × E as differential gene sensitivity to climate. KEY RESULTS: The species' annual growth was positively influenced by winter temperature and spring moisture and negatively influenced by previous autumn precipitation and warm springs. Four and five climate factors explained 10 % (RWI) and 16 % (ELI) of population-specific interannual variability, respectively, with populations from drought-prone areas and with uneven precipitation experiencing larger growth reductions during dry vegetative periods. Furthermore, four and two SNPs explained 14 % (RWI) and 10 % (ELI) of interannual variability among trees, respectively. Two SNPs played a putative role in adaptation to climate: one identified from transcriptome sequencing of P. halepensis and another involved in response regulation to environmental stressors. CONCLUSIONS: We highlight how tree-ring phenotypes, obtained from a common garden experiment, combined with a candidate-gene approach allow the quantification of genetic and environmental effects determining adaptation for a conifer with a large and complex genome.

Connecting tree-ring phenotypes, genetic associations and transcriptomics to decipher the genomic architecture of drought adaptation in a widespread conifer.

As boreal forests face significant threats from climate change, understanding evolutionary trajectories of coniferous species has become fundamental to adapting management and conservation to a drying climate. We examined the genomic architecture underlying adaptive variation related to drought tolerance in 43 populations of a widespread boreal conifer, white spruce (Picea glauca [Moench] Voss), by combining genotype-environment associations, genotype-phenotype associations, and transcriptomics. Adaptive genetic variation was identified by correlating allele frequencies for 6,153 single nucleotide polymorphisms from 2,606 candidate genes with temperature, precipitation and aridity gradients, and testing for significant associations between genotypes and 11 dendrometric and drought-related traits (i.e., anatomical, growth response and climate-sensitivity traits) using a polygenic model. We identified a set of 285 genes significantly associated with a climatic factor or a phenotypic trait, including 110 that were differentially expressed in response to drought under greenhouse-controlled conditions. The interlinked phenotype-genotype-environment network revealed eight high-confidence genes involved in white spruce adaptation to drought, of which four were drought-responsive in the expression analysis. Our findings represent a significant step toward the characterization of the genomic basis of drought tolerance and adaptation to climate in conifers, which is essential to enable the establishment of resilient forests in view of new climate conditions.

Long-term nitrogen addition does not sustain host tree stem radial growth but doubles the abundance of high-biomass ectomycorrhizal fungi.

Global change has altered nitrogen availability in boreal forest soils. As ectomycorrhizal fungi play critical ecological functions, shifts in their abundance and community composition must be considered in the response of forests to changes in nitrogen availability. Furthermore, ectomycorrhizas are symbiotic, so the response of ectomycorrhizal fungi to nitrogen cannot be understood in isolation of their plant partners. Most previous studies, however, neglect to measure the response of host trees to nitrogen addition simultaneously with that of fungal communities. In addition to being one-sided, most of these studies have also been conducted in coniferous forests. Deciduous and "dual-mycorrhizal" tree species, namely those that form ecto- and arbuscular mycorrhizas, have received little attention despite being widespread in the boreal forest. We applied nitrogen (30 kg ha-1  year-1 ) for 13 years to stands dominated by aspen (Populus tremuloides Michx.) and hypothesized that tree stem radial growth would increase, ectomycorrhizal fungal biomass would decrease, ectomycorrhizal fungal community composition would shift, and the abundance of arbuscular mycorrhizal (AM) fungi would increase. Nitrogen addition initially increased stem radial growth of aspen, but it was not sustained at the time we characterized their mycorrhizas. After 13 years, the abundance of fungi possessing extramatrical hyphae, or "high-biomass" ectomycorrhizas, doubled. No changes occurred in ectomycorrhizal and AM fungal community composition, or in ecto- and AM abundance measured as root colonization. This dual-mycorrhizal tree species did not shift away from ectomycorrhizal fungal dominance with long-term nitrogen input. The unexpected increase in high-biomass ectomycorrhizal fungi with nitrogen addition may be due to increased carbon allocation to their fungal partners by growth-limited trees. Given the focus on conifers in past studies, reconciling results of plant-mycorrhizal fungal relationships in stands of deciduous trees may demand a broader view on the impacts of nitrogen addition on the structure and function of boreal forests.

Inter and intra-annual links between climate, tree growth and NDVI: improving the resolution of drought proxies in conifer forests.

The inter- and intra-annual variability in radial growth reflects responses to climatic variability and water shortage, especially in areas subjected to seasonal drought. However, it is unknown how this variability is related to forest productivity, which can be assessed by measuring changes in canopy greenness and cover through remote sensing products as the Normalized Difference Vegetation Index (NDVI). We combine xylogenesis with measurements of inter-annual changes in seasonal wood production (earlywood width, adjusted latewood width) and NDVI to improve the understanding of climate and drought impacts on growth and forest productivity in a Pinus teocote stand located in northern Mexico. Cambial dynamics accelerated in March and a high production of radially enlarging and thickening tracheids were observed from April to October and from June to October, respectively. Tracheid maturation was very active in October when latewood production peaked. Wet conditions in winter-spring and summer-autumn enhanced earlywood and latewood production, respectively. Earlywood and latewood were constrained by long (4-10 months) and short (2-3 months) droughts, respectively. The earlywood production depended on April soil moisture, which agrees with the peak of radially enlarging tracheid production found during that month. Aligning drought proxies at inter- and intra-annual scales by using growth and productivity measures improves our understanding of conifer forest responses to water shortage.

Comparison of the responses of radial growth to climate change for two dominant coniferous tree species in the eastern Qilian Mountains, northwestern China.

The temperature in northwestern China has increased significantly since the 1990s. However, the responses of mountainous forests to warming have not been extensively examined. We collected tree rings of two dominant coniferous species of Qinghai spruce (Picea crassifolia) and Chinese pine (Pinus tabulaeformis) in the eastern part of the Qilian Mountains, and analyzed the differences in the response dynamic of the radial growth of two species to climate change. The results showed that (1) the annual radial growth of Qinghai spruce was mainly restricted by the minimum temperature in July and October, and the growth of Chinese pine was mainly restricted by the mean temperature in September of the previous year, January, and July and the maximum temperature in March, May, and July. In particular, Qinghai spruce increased its sensitivity to total precipitation in the growing seasons in March, May, and July after the temperature abruptly increased. (2) In comparison to Qinghai spruce, Chinese pine showed a consistent response to the main climatic factors and was more severely affected by drought stress. Qinghai spruce had divergent responses to mean temperatures in March and May and minimum temperatures in April and June. (3) The growth of Qinghai spruce increased with a significant fluctuation at the end of the twentieth century, while the growth of Chinese pine first showed an increase and then a significant decreasing trend. At present, the increase in temperature has adversely affected the growth of Chinese pine in the eastern Qilian Mountains and promoted the growth of Qinghai spruce. However, a continuous temperature increase could negatively affect the growth of Qinghai spruce because of the increasing probability of drought stress. Therefore, we should pay more attention to the growth dynamics of Qinghai spruce, especially with the different water supply and demand, and to the effects of drought on Chinese pine in forest ecosystems in arid and semiarid areas.

Adaptive genetic variation to drought in a widely distributed conifer suggests a potential for increasing forest resilience in a drying climate.

Drought intensity and frequency are increasing under global warming, with soil water availability now being a major factor limiting tree growth in circumboreal forests. Still, the adaptive capacity of trees in the face of future climatic regimes remains poorly documented. Using 1481 annually resolved tree-ring series from 29-yr-old trees, we evaluated the drought sensitivity of 43 white spruce (Picea glauca (Moench) Voss) populations established in a common garden experiment. We show that genetic variation among populations in response to drought plays a significant role in growth resilience. Local genetic adaptation allowed populations from drier geographical origins to grow better, as indicated by higher resilience to extreme drought events, compared with populations from more humid geographical origins. The substantial genetic variation found for growth resilience highlights the possibility of selecting for drought resilience in boreal conifers. As a major research outcome, we showed that adaptive genetic variation in response to changing local conditions can shape drought vulnerability at the intraspecific level. Our findings have wide implications for forest ecosystem management and conservation.

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.

Climate-human interactions contributed to historical forest recruitment dynamics in Mediterranean subalpine ecosystems.

Long-term tree recruitment dynamics of subalpine forests mainly depend on temperature changes, but little is known about the feedbacks between historical land use and climate. Here, we analyze a southern European, millennium-long dataset of tree recruitment from three high-elevation pine forests located in Mediterranean mountains (Pyrenees, northeastern Spain; Pollino, southern Italy; and Mt. Smolikas, northern Greece). We identify synchronized recruitment peaks in the late 15th and early 16th centuries, following prolonged periods of societal and climate instability. Major European population crises in the 14th and 15th centuries associated with recurrent famines, the Black Death pandemic, and political turmoil are likely to have reduced the deforestation of subalpine environments and caused widespread rewilding. We suggest that a distinct cold phase in the Little Ice Age around 1450 ce could also have accelerated the cessation of grazing pressure, particularly in the Pyrenees, where the demographic crisis was less severe. Most pronounced in the Pyrenees, the enhanced pine recruitment from around 1500-1550 ce coincides with temporarily warmer temperatures associated with a positive phase of the North Atlantic Oscillation. We diagnose that a mixture of human and climate factors has influenced past forest recruitment dynamics in Mediterranean subalpine ecosystems. Our results highlight how complex human-climate interactions shaped forest dynamics during pre-industrial times and provide historical analogies to recent rewilding.