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Drought- and heat-driven tree mortality, along with associated insect outbreaks, have been observed globally in recent decades and are expected to increase in future climates. Despite its potential to profoundly alter ecosystem carbon and water cycles, how tree mortality scales up to ecosystem functions and fluxes is uncertain. We describe a framework for this scaling where the effects of mortality are a function of the mortality attributes, such as spatial clustering and functional role of the trees killed, and ecosystem properties, such as productivity and diversity. We draw upon remote-sensing data and ecosystem flux data to illustrate this framework and place climate-driven tree mortality in the context of other major disturbances. We find that emerging evidence suggests that climate-driven tree mortality impacts may be relatively small and recovery times are remarkably fast (~4 years for net ecosystem production). We review the key processes in ecosystem models necessary to simulate the effects of mortality on ecosystem fluxes and highlight key research gaps in modeling. Overall, our results highlight the key axes of variation needed for better monitoring and modeling of the impacts of tree mortality and provide a foundation for including climate-driven tree mortality in a disturbance framework.

The impacts of climate extremes on terrestrial ecosystems are poorly understood but important for predicting carbon cycle feedbacks to climate change. Coupled climate-carbon cycle models typically assume that vegetation recovery from extreme drought is immediate and complete, which conflicts with the understanding of basic plant physiology. We examined the recovery of stem growth in trees after severe drought at 1338 forest sites across the globe, comprising 49,339 site-years, and compared the results with simulated recovery in climate-vegetation models. We found pervasive and substantial "legacy effects" of reduced growth and incomplete recovery for 1 to 4 years after severe drought. Legacy effects were most prevalent in dry ecosystems, among Pinaceae, and among species with low hydraulic safety margins. In contrast, limited or no legacy effects after drought were simulated by current climate-vegetation models. Our results highlight hysteresis in ecosystem-level carbon cycling and delayed recovery from climate extremes.

The soil microbiome plays an important role in forest functioning. However, the impact of drought-induced dieback and tree death on soil microbial biomass, community structure, and functional composition is unknown. We also lack understanding on how soil microbiota varies seasonally in such declining stands. We used Phospholipid Fatty Acids (PLFA) analysis to quantify soil microbial biomass and study its seasonal changes in three Mediterranean forests showing dieback and dominated by three pine species (Pinus halepensis Mill, Pinus pinaster Ait. and Pinus sylvestris L.). We also measured microclimatic parameters and soil physical and chemical parameters under trees with different vigor, assessed as canopy defoliation degree, and related them to seasonal changes in the soil microbial community. We found marked differences in soil microbial community structure, total biomass, and the relative abundance of major functional groups among forests. First, soil microbial biomass peaked either in the dry summer (P. halepensis) or in autumn (P. pinaster and P. sylvestris). Accordingly, the relative abundance of most functional groups, excluding Arbuscular mycorrhizal fungi, displayed substantial variation between seasons. In addition, the relative abundance of fungi and Gram-positive bacteria exhibited an opposite pattern compared to actinomycetes and Gram-negative bacteria. Second, soil physical and chemical parameters had a significant impact on within-site PLFA variation, although their influence was less important than that of seasonal variation. Third, differences between defoliated and healthy trees were minor and restricted to averaged ratios between different PLFA markers. Overall, the structure, biomass and relative abundance of major functional groups of the soil microbiome vary considerably among stand types and seasons in forests showing ongoing dieback and high mortality. However, while the seasonal dynamics show predictable patterns, which should be accounted for in future studies, the within-site variation is highly variable and mainly depends on soil physical and chemical parameters., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Understanding how climate change influences succession is fundamental for predicting future forest composition. Warming is expected to accelerate species succession at their cold thermal ranges, such as alpine treelines. Here we examined how interactions and successional strategies of the early-successional birch (Betula utilis) and the late-successional fir (Abies spectabilis) affected treeline dynamics by combining plot data with an individual-based treeline model at treelines in the central Himalayas. Fir showed increasing recruitment and a higher upslope shift rate (0.11 ± 0.02 m yr -1 ) compared with birch (0.06 ± 0.03 m yr -1 ) over the past 200 years. Spatial analyses indicate strong interspecies competition when trees were young. Model outputs from various climatic scenarios indicate that fir will probably accelerate its upslope movement with warming, while birch recruitment will decline drastically, forming stable or even retreating treelines. Our findings point to accelerating successional dynamics with late-successional species rapidly outcompeting pioneer species, offering insight into future forest succession and its influences on ecosystem services., Competing Interests: Competing interests: The authors declare no competing interests., (© 2024. The Author(s), under exclusive licence to Springer Nature Limited.)

Traditional ditches ("acequias" in Spanish) derive meltwater and infiltrate groundwater providing ecological services downstream in the semi-arid Sierra Nevada range (SE Spain). Therefore, they may act as a nature-based solution by alleviating drought stress in trees growing near ditches by enhancing growth and reducing their intrinsic water-use efficiency (iWUE). Such a mitigation role of acequias is critical given that some oak (Quercus pyrenaica) and pine (Pinus sylvestris) stands reach their xeric distribution limits in Europe. We compared tree-ring width data and wood δ 13 C, a proxy of iWUE, in oak and pine stands located near or far (control) from ditches with different infiltration capacity in two watersheds. We assessed how trees responded to climate data, drought stress, and vegetation greenness through correlations and resilience indices. Oak trees located near ditches grew more and responded less to precipitation, soil moisture, a drought index, and greenness than control trees. In pines, we did not find this pattern, and ditch trees grew more than control trees only during an extremely dry year (1995). Climate-growth correlations suggested a longer growing season in ditch pines. Growth of ditch oaks from the "Acequia Nueva" (AN), with high infiltration capacity, responded more to autumn soil moisture and showed the lowest δ 13 C. Growth was enhanced by cool-wet spring conditions in pines and also by warm-wet conditions in the prior winter in the case of oaks. Control trees showed lower resistance to drought. Control trees presented higher wood δ 13 C values except for old oaks from the "Acequia Grande" (AG) site which may show long-term acclimation. Traditional ditches alleviate drought stress in oak and pine stands subjected to regional xeric climate conditions., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Stem growth responses to soil and atmospheric drought are critical to forecasting the tree carbon sink strength. Yet, responses of drought-prone forests remain uncertain despite global aridification trends. Stem diameter variations at an hourly resolution were monitored in five Mediterranean tree species from a mesic and a xeric site for 6 and 12 years. Stem growth and dehydration responses to soil (REW) and atmospheric (VPD) drought were explored at different timescales. Annually, growth was determined by the number of growing days and hours. Seasonally, growth was bimodal (autumn growth ≈ 8%-18% of annual growth), varying among species and sites across the hydrometeorological space, while dehydration consistently responded to REW. Sub-daily, substantial growth occurred during daytime, with nighttime-to-daytime ratios ranging between 1.2 and 3.5 (Arbutus unedo ≈ Quercus faginea < Quercus ilex < Pinus halepensis in the mesic site, and Juniperus thurifera < P. halepensis in the xeric site). Overall, time windows favourable for growth were limited by soil (rather than atmospheric) drought, modulating annual and seasonal growth in Mediterranean species, and stems maintained non-negligible growth during daytime. These patterns contrast with observations from wetter or cooler biomes, demonstrating the growth plasticity of drought-prone species to more arid climate conditions., (© 2024 The Author(s). Plant, Cell & Environment published by John Wiley & Sons Ltd.)

With ongoing global warming, increasing water deficits promote physiological stress on forest ecosystems with negative impacts on tree growth, vitality, and survival. How individual tree species will react to increased drought stress is therefore a key research question to address for carbon accounting and the development of climate change mitigation strategies. Recent tree-ring studies have shown that trees at higher latitudes will benefit from warmer temperatures, yet this is likely highly species-dependent and less well-known for more temperate tree species. Using a unique pan-European tree-ring network of 26,430 European beech (Fagus sylvatica L.) trees from 2118 sites, we applied a linear mixed-effects modeling framework to (i) explain variation in climate-dependent growth and (ii) project growth for the near future (2021-2050) across the entire distribution of beech. We modeled the spatial pattern of radial growth responses to annually varying climate as a function of mean climate conditions (mean annual temperature, mean annual climatic water balance, and continentality). Over the calibration period (1952-2011), the model yielded high regional explanatory power (R 2  = 0.38-0.72). Considering a moderate climate change scenario (CMIP6 SSP2-4.5), beech growth is projected to decrease in the future across most of its distribution range. In particular, projected growth decreases by 12%-18% (interquartile range) in northwestern Central Europe and by 11%-21% in the Mediterranean region. In contrast, climate-driven growth increases are limited to around 13% of the current occurrence, where the historical mean annual temperature was below ~6°C. More specifically, the model predicts a 3%-24% growth increase in the high-elevation clusters of the Alps and Carpathian Arc. Notably, we find little potential for future growth increases (-10 to +2%) at the poleward leading edge in southern Scandinavia. Because in this region beech growth is found to be primarily water-limited, a northward shift in its distributional range will be constrained by water availability., (© 2024 The Author(s). Global Change Biology published by John Wiley & Sons Ltd.)

Mistletoes are xylem-tapping hemiparasites that rely on their hosts for water and nutrient uptake. Thus, they impair tree performance in the face of environmental stress via altering the carbon and water relations and nutritional status of trees. To improve our understanding of physiological responses to mistletoe and ongoing climate change, we investigated radial growth, stable carbon and oxygen isotopic signals, and elemental composition of tree rings in silver fir (Abies alba Mill.) and Scots pine (Pinus sylvestris L.) forests infested with Viscum album L. We compared temporal series (1990-2020) of basal area increment (BAI), intrinsic water-use efficiency (iWUE), oxygen isotope composition (δ18O), nutrient concentrations and stoichiometric ratios between non-infested (NI) and severely infested (SI) fir and pine trees from populations located close to the xeric distribution limit of the species in north-eastern Spain. The SI trees showed historically higher growth, but the BAI trend was negative for more than three decades before 2020 and their growth rates became significantly lower than those of NI trees by the mid-2010s. Mistletoe infestation was related to an enhanced sensitivity of radial growth to vapour pressure deficit (atmospheric drought). The SI trees showed less pronounced iWUE increases (fir) and lower iWUE values (pine) than NI trees. The lower tree-ring δ18O values of SI trees may be the result of several superimposed effects operating simultaneously, including leaf-level evaporative enrichment, source water isotopic signals, and anatomical and phenological differences. We observed a deterioration of potassium (K) nutrition in tree-ring wood of both species in SI trees, along with accumulation of manganese (Mn). We suggest that such nutritional patterns are driven by the indirect effect of mistletoe-induced drought stress, particularly in pine. The combined analyses of different physiological indicators imprinted on tree rings provided evidence of the progressive onset of carbon, water and nutrient imbalances in mistletoe-infested conifers inhabiting seasonally dry regions., (© The Author(s) 2024. Published by Oxford University Press.)

The fundamental trade-off between current and future reproduction has long been considered to result in a tendency for species that can grow large to begin reproduction at a larger size. Due to the prolonged time required to reach maturity, estimates of tree maturation size remain very rare and we lack a global view on the generality and the shape of this trade-off. Using seed production from five continents, we estimate tree maturation sizes for 486 tree species spanning tropical to boreal climates. Results show that a species' maturation size increases with maximum size, but in a non-proportional way: the largest species begin reproduction at smaller sizes than would be expected if maturation were simply proportional to maximum size. Furthermore, the decrease in relative maturation size is steepest in cold climates. These findings on maturation size drivers are key to accurately represent forests' responses to disturbance and climate change., (© 2024 John Wiley & Sons Ltd.)

Pinus pinea is an important Mediterranean species due to its adaptability and tolerance to aridity and its high-quality pine nuts. Different forest types located in Mediterranean native and non-native environments provide the opportunity to perform comparative studies on the species' response to climate change. The aims of this study were to elucidate growth patterns of the species growing in native and exotic habitats and to analyze its response to climatic fluctuations, particularly drought, in both geographical contexts. Understanding stone pine ( Pinus pinea ) growth responses to climate variability in native and exotic habitats by comparing natural stands and plantations may provide useful information to plan adequate management under climate change. By doing so, we enhance the understanding of P. pinea 's adaptability and provide practical approaches to its sustainable management. In this study, we reconstructed and compared the stem radial growth of seven stone pine stands, two in southern Spain and five in central-southern Chile, growing under different climatic conditions. We quantified the relationships between growth variability and climate variables (total rainfall, mean temperature, and SPEI drought index). Growth was positively correlated with autumn rainfall in plantations and with autumn-winter rainfall in natural stands. Growth was also enhanced by high autumn-to-spring rainfall in the driest Chilean plantation, whereas in the wettest and coolest plantation, such correlation was found in winter and summer. A negative impact of summer temperature was found only in one of the five Chilean plantations and in a Spanish site. The correlation between SPEI and tree-ring width indices showed different patterns between and within countries. Overall, exotic plantations showed lower sensitivity to climate variability than native stands. Therefore, stone pine plantations may be useful to assist in mitigating climate change.

The future performance of the widely abundant European beech (Fagus sylvatica L.) across its ecological amplitude is uncertain. Although beech is considered drought-sensitive and thus negatively affected by drought events, scientific evidence indicating increasing drought vulnerability under climate change on a cross-regional scale remains elusive. While evaluating changes in climate sensitivity of secondary growth offers a promising avenue, studies from productive, closed-canopy forests suffer from knowledge gaps, especially regarding the natural variability of climate sensitivity and how it relates to radial growth as an indicator of tree vitality. Since beech is sensitive to drought, we in this study use a drought index as a climate variable to account for the combined effects of temperature and water availability and explore how the drought sensitivity of secondary growth varies temporally in dependence on growth variability, growth trends, and climatic water availability across the species' ecological amplitude. Our results show that drought sensitivity is highly variable and non-stationary, though consistently higher at dry sites compared to moist sites. Increasing drought sensitivity can largely be explained by increasing climatic aridity, especially as it is exacerbated by climate change and trees' rank progression within forest communities, as (co-)dominant trees are more sensitive to extra-canopy climatic conditions than trees embedded in understories. However, during the driest periods of the 20th century, growth showed clear signs of being decoupled from climate. This may indicate fundamental changes in system behavior and be early-warning signals of decreasing drought tolerance. The multiple significant interaction terms in our model elucidate the complexity of European beech's drought sensitivity, which needs to be taken into consideration when assessing this species' response to climate change., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Author(s). Published by Elsevier B.V. All rights reserved.)

As major terrestrial carbon sinks, forests play an important role in mitigating climate change. The relationship between the seasonal uptake of carbon and its allocation to woody biomass remains poorly understood, leaving a significant gap in our capacity to predict carbon sequestration by forests. Here, we compare the intra-annual dynamics of carbon fluxes and wood formation across the Northern hemisphere, from carbon assimilation and the formation of non-structural carbon compounds to their incorporation in woody tissues. We show temporally coupled seasonal peaks of carbon assimilation (GPP) and wood cell differentiation, while the two processes are substantially decoupled during off-peak periods. Peaks of cambial activity occur substantially earlier compared to GPP, suggesting the buffer role of non-structural carbohydrates between the processes of carbon assimilation and allocation to wood. Our findings suggest that high-resolution seasonal data of ecosystem carbon fluxes, wood formation and the associated physiological processes may reduce uncertainties in carbon source-sink relationships at different spatial scales, from stand to ecosystem levels., (© 2024. The Author(s).)

Given the context of significant global warming and the intensification of extreme climate events in the last century, large-scale reforestation and afforestation have been recognized as effective strategies to mitigate the climate crisis. Since the 1970s, China has launched several afforestation programs aimed at regional ecological protection, playing an important role in reaching carbon neutrality by 2060. This study provided a detailed analysis of the growth suitability of the main planted conifers (Pinus sylvestris var. mongolica and Pinus tabulaeformis) and broadleaves (Populus spp., Robinia pseudoacacia) in the semi-arid northern China. We compared the radial growth trends of plantations and their responses to extreme droughts from 1980 to 2018. Growth of most plantations has significantly increased over time, but broadleaves showed recent growth reductions in the past decade, which may be related to tree age and reduced soil moisture. Nevertheless, under warmer climate scenarios, the growth of plantations is forecasted to continue increasing. Broadleaves showed a better post-drought recovery, probably linked to their anisohydric behavior, than conifers, which presented a better resistance to drought. Growth of conifers depended more on warmer temperature and better precipitation conditions during the growing season, whereas broadleaves mainly reacted to warm temperature. Additionally, pre-drought growth levels weakened resilience components, while post-drought precipitation compensated for drought-induced growth deficit. Growth and resilience were negatively related to tree age, while higher stand density reduced growth. This assessment and projections of growth and drought resilience indicate the sustainability of most plantations in semi-arid regions, but future warmer and drier conditions may lead to an uncertain future regarding forest health and reduce their carbon sink potential., (© 2024 John Wiley & Sons Ltd.)

The impact of air pollution on forests, especially in urban areas, has been increasingly discussed recently. Many pollutants, including heavy metals, are released into the atmosphere from various sources, such as mining, non-ferrous metal processing plants, and fossil fuel combustion. These pollutants can adversely affect not only tree growth but also other species, including humans. This study compared the concentrations of several elements in tree-ring wood from two conifer species (Silver fir, Abies alba; Norway spruce, Picea abies) growing in polluted and unpolluted areas. Two regions in northern Romania (Bicaz and Tarnița) that were subjected to historical pollution changes were selected. Two chemical analyses were used: inductively coupled plasma mass spectrometry (ICP-MS) and X-ray fluorescence spectrometry (XRF). The silver fir trees from the intensively polluted area in the Tarnița region were negatively impacted by industrial pollution: the Mn concentrations were, on average, three times higher in polluted areas than in unpolluted areas (ca. 30 vs. 10 mg kg -1 ). This finding was consistent for both ICP-MS and XRF analyses. However, in Norway spruce, this difference was found only in the XRF data, which detected Mn concentrations seven times higher in trees from polluted areas than those from unpolluted areas (ca. 700 vs. 100 mg kg -1 ). In the Tarnița region, Norway spruce accumulated more heavy metals than silver fir, but the most pronounced differences between polluted and unpolluted areas were found in silver fir. The two analytical methods are commonly used to determine metal concentrations in wood, and they complement each other, with ICP-MS having a low detection limit for some elements and XRF having higher detection limits and better accuracy. Each method has its advantages and disadvantages, and the optimal method depends on many factors, such as the type of heavy metal analyzed, its concentration in wood, sample type, cost, analysis time, and sample preparation., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Inc. All rights reserved.)

Previous favorable climate conditions stimulate tree growth making some forests more vulnerable to hotter droughts. This so-called structural overshoot may contribute to forest dieback, but there is little evidence on its relative importance depending on site conditions and tree species because of limited field data. Here, we analyzed remote sensing (NDVI) and tree-ring width data to evaluate the impacts of the 2017 drought on canopy cover and growth in mixed Mediterranean forests (Fraxinus ornus, Quercus pubescens, Acer monspessulanum, Pinus pinaster) located in southern Italy. Legacy effects were assessed by calculating differences between observed and predicted basal area increment (BAI). Overall, the growth response of the study stands to the 2017 drought was contingent on site conditions and species characteristics. Most sites presented BAI and canopy cover reductions during the drought. Growth decline was followed by a quick recovery and positive legacy effects, particularly in the case of F. ornus. However, we found negative drought legacies in some species (e.g., Q. pubescens, A. monspessulanum) and sites. In those sites showing negative legacies, high growth rates prior to drought in response to previous wet winter-spring conditions may have predisposed trees to drought damage. Vice versa, the positive drought legacy found in some F. ornus site was linked to post-drought growth release due to Q. pubescens dieback and mortality. Therefore, we found evidences of structural drought overshoot, but it was restricted to specific sites and species. Our findings highlight the importance of considering site settings such as stand composition, pre-drought conditions and different tree species when studying structural overshoot. Droughts contribute to modify the composition and dynamics in mixed forests., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Introduction: Plantations located outside the species distribution area represent natural experiments to assess tree tolerance to climate variability. Climate change amplifies warming-related drought stress but also leads to more climate extremes., Methods: We studied plantations of the European larch (Larix decidua), a conifer native to central and eastern Europe, in northern Spain. We used climate, drought and tree-ring data from four larch plantations including wet (Valgañón, site V; Santurde, site S), intermediate (Ribavellosa, site R) and dry (Santa Marina, site M) sites. We aimed to benchmark the larch tolerance to climate and drought stress by analysing the relationships between radial growth increment (hereafter growth), climate data (temperature, precipitation, radiation) and a drought index., Results: Basal area increment (BAI) was the lowest in the driest site M (5.2 cm2 yr-1; period 1988-2022), followed by site R (7.5 cm2 yr-1), with the youngest and oldest and trees being planted in M (35 years) and R (150 years) sites. BAI peaked in the wettest sites (V; 10.4 cm2 yr-1; S, 10.8 cm2 yr-1). We detected a sharp BAI reduction (30% of the regional mean) in 2001 when springto-summer conditions were very dry. In the wettest V and S sites, larch growth positively responded to current March and June-July radiation, but negatively to March precipitation. In the R site, high April precipitation enhanced growth. In the driest M site, warm conditions in the late prior winter and current spring improved growth, but warm-sunny conditions in July and dry-sunny conditions in August reduced it. Larch growth positively responded to spring-summer wet conditions considering short (1-6 months) and long (9-24 months) time scales in dry (site M) and wet-intermediate (sites S and R) sites, respectively., Discussion: Larch growth is vulnerable to drought stress in dry slow-growing plantations, but also to extreme spring wet-cloudy events followed by dry-hot conditions in wet fast-growing plantations., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2024 Camarero, Gazol, Valeriano, Vergarechea and Cattaneo.)

Forests are undergoing increasing risks of drought-induced tree mortality. Species replacement patterns following mortality may have a significant impact on the global carbon cycle. Among major hardwoods, deciduous oaks (Quercus spp.) are increasingly reported as replacing dying conifers across the Northern Hemisphere. Yet, our knowledge on the growth responses of these oaks to drought is incomplete, especially regarding post-drought legacy effects. The objectives of this study were to determine the occurrence, duration, and magnitude of legacy effects of extreme droughts and how that vary across species, sites, and drought characteristics. The legacy effects were quantified by the deviation of observed from expected radial growth indices in the period 1940-2016. We used stand-level chronologies from 458 sites and 21 oak species primarily from Europe, north-eastern America, and eastern Asia. We found that legacy effects of droughts could last from 1 to 5 years after the drought and were more prolonged in dry sites. Negative legacy effects (i.e., lower growth than expected) were more prevalent after repetitive droughts in dry sites. The effect of repetitive drought was stronger in Mediterranean oaks especially in Quercus faginea. Species-specific analyses revealed that Q. petraea and Q. macrocarpa from dry sites were more negatively affected by the droughts while growth of several oak species from mesic sites increased during post-drought years. Sites showing positive correlations to winter temperature showed little to no growth depression after drought, whereas sites with a positive correlation to previous summer water balance showed decreased growth. This may indicate that although winter warming favors tree growth during droughts, previous-year summer precipitation may predispose oak trees to current-year extreme droughts. Our results revealed a massive role of repetitive droughts in determining legacy effects and highlighted how growth sensitivity to climate, drought seasonality and species-specific traits drive the legacy effects in deciduous oak species., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have influenced the research reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Species interactions such as facilitation and competition play a crucial role in driving species range shifts. However, density dependence as a key feature of these processes has received little attention in both empirical and modelling studies. Herein, we used a novel, individual-based treeline model informed by rich in situ observations to quantify the contribution of density-dependent species interactions to alpine treeline dynamics, an iconic biome boundary recognized as an indicator of global warming. We found that competition and facilitation dominate in dense versus sparse vegetation scenarios respectively. The optimal balance between these two effects was identified at an intermediate vegetation thickness where the treeline elevation was the highest. Furthermore, treeline shift rates decreased sharply with vegetation thickness and the associated transition from positive to negative species interactions. We thus postulate that vegetation density must be considered when modelling species range dynamics to avoid inadequate predictions of its responses to climate warming., (© 2024 John Wiley & Sons Ltd.)

We lack understanding of how variable is radial growth of coexisting tree and shrub species, and how growth is constrained by drought depending on site aridity. Here, we compared the radial growth of two widespread and coexisting species, a winter deciduous shrub (Amelanchier ovalis Medik.) and an evergreen conifer tree (Pinus sylvestris L.). We sampled four sites in Northeastern Spain subjected to different aridity levels and used dendrochronological methods to quantify growth patterns and responses to climate variables. The growth of the two species varied between regions, being lower in the driest sites. The first-order autocorrelation (growth persistence) was higher in more mesic sites but without clear differences between species. Tree and shrub growth negatively responded to elevated summer temperatures and positively to spring-summer precipitation and wet conditions. However, negative growth responses of the shrub to drought were only observed in the two driest sites in contrast to widespread responses of the tree. Abrupt growth reductions were common in the drier sites, but resilience indices show that the two species rapidly recovered pre-drought growth levels. The lower growth synchrony of the shrub as compared to the tree can be due to the multistemmed architecture, fast growth and low stature of the shrub. Besides, the high dependency of the shrub growth on summer rainfall can explain why drought limitations were only apparent in the two driest sites. In any case, results point out to the dendrochronological potential of shrubs, which is particularly relevant giving its ability to inhabit woodlands and treeless regions under harsh climatic conditions. Nevertheless, further research is required to elucidate the capacity of shrub species to tolerate drought, as well as to understand how shrubs thrive in water- and cold-limited environments., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Wood growth is key to understanding the feedback of forest ecosystems to the ongoing climate warming. An increase in spatial synchrony (i.e., coincident changes in distant populations) of spring phenology is one of the most prominent climate responses of forest trees. However, whether temperature variability contributes to an increase in the spatial synchrony of spring phenology and its underlying mechanisms remains largely unknown. Here, we analyzed an extensive dataset of xylem phenology observations of 20 conifer species from 75 sites over the Northern Hemisphere. Along the gradient of increase in temperature variability in the 75 sites, we observed a convergence in the onset of cell enlargement roughly toward the 5 th of June, with a convergence in the onset of cell wall thickening toward the summer solstice. The increase in rainfall since the 5 th of June is favorable for cell division and expansion, and as the most hours of sunlight are received around the summer solstice, it allows the optimization of carbon assimilation for cell wall thickening. Hence, the convergences can be considered as the result of matching xylem phenological activities to favorable conditions in regions with high temperature variability. Yet, forest trees relying on such consistent seasonal cues for xylem growth could constrain their ability to respond to climate warming, with consequences for the potential growing season length and, ultimately, forest productivity and survival in the future., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2024 Elsevier Inc. All rights reserved.)

Cork is one of the main non-timber forest products in the world. Most of its production is concentrated in the Iberian Peninsula, a climate change hotspot. Climate warming may lead to increased aridification and reduce cork production in that region. However, we still lack assessments of climate-cork relationships across ample geographical and climatic gradients explicitly considering site aridity. We quantified cork growth by measuring cork ring width and related it to climate variables and a drought index using dendrochronology. Four cork oak (Quercus suber) forests located from north eastern Spain to south western Morocco (31.5-41.5° N) and subjected to different aridity levels were sampled. Warm conditions in spring to early summer, when cork is formed, reduced cork width, whereas high precipitation in winter and spring enhanced it. The response of cork to increased water availability in summer peaked (r = 0.89, p = 0.00002) in the most arid and continental site considering 14-month long droughts. A severe drought caused a disproportionate loss of cork production in this site, where for every five-fold decrease in the drought index, the cork-width index declined by a factor of thirteen. Therefore, site aridity determines the responses of cork growth to the soil water availability resulting from accumulated precipitation during winter and spring previous to cork growth and until summer. In general, this cumulative water balance, which is very dependent on temperature and evapotranspiration rate, is critical for cork production, especially in continental, dry sites. The precipitation during the hydrological year can be used as a proxy of cork production in similar sites. Assessments of climate-cork relationships in the western Mediterranean basin could be used as analogues to forecast the impacts of aridification on future cork production., Competing Interests: Declaration of competing interest Authors declare no conflict of interest., (Copyright © 2023 The Author(s). Published by Elsevier B.V. All rights reserved.)

Riparian forests are among the most dynamic but threatened terrestrial ecosystems. Their dynamism and conservation depend on historical changes in river geomorphology, which can be evaluated through changes in channel sinuosity. However, we lack long-term assessments on sinuosity and how they impact riparian forest composition, tree growth and deadwood amount. To fill this research gap, we reconstructed river sinuosity in 14 sites across the middle Ebro basin, north-eastern Spain, using historical aerial photographs taken in 1927, 1956, 1998-2003 and 2014-2015. Relationships between sinuosity, stand composition and deadwood amount and decay degree were calculated. We also reconstructed radial growth of the major tree species (Populus alba, Populus nigra, Fraxinus angustifolia, Salix alba and Ulmus minor) in two sites to evaluate how coupled it was with changes in river flow after dam building. From 1927 to 2015, sinuosity decreased passing from 1.39 to 1.20. The river dynamics were altered in the 1950s and 1960s after dam and dyke building. Sites with high sinuosity values in 1956 corresponded to mature stands with large P. nigra individuals. Sinuosity was negatively related to F. angustifolia (r s  = -0.83, p < 0.001) and P. alba (r s  = -0.64, p = 0.02) abundance, whereas sites dominated by P. alba and U. minor presented abundant decayed deadwood. A loss of sinuosity and a contraction of the riverbank gradient increased disconnection of active channel from floodplain, with a mixing of more (e.g., P. nigra) and less phreatophytic species (e.g., U. minor). River flow diversion reduced growth and increased the tree-to-tree P. alba growth coherence. Hydrological droughts contributed to growth decline and dieback of U. minor, which is sensitive to spring river flow. Conservation and restoration of riparian forests must consider historical changes in river geomorphology related to human activities., Competing Interests: Declaration of competing interest Authors declare no conflict of interest., (Copyright © 2023 The Author(s). Published by Elsevier B.V. All rights reserved.)

Although the global climate is warming, external forcing driven by explosive volcanic eruptions may still cause abrupt cooling. The 1809 and 1815 Tambora eruptions caused lasting cold extremes worldwide, providing a unique lens that allows us to investigate the magnitude of global forest resilience to and recovery from volcanic cooling. Here, we show that growth resilience inferred from tree-ring data was severely impacted by cooling in high latitudes and elevations: the average tree growth decreased substantially (up to 31.8%), especially in larch forests, and regional-scale probabilities of severe growth reduction (below -2σ) increased up to 1390%. The influence of the eruptions extended longer (beyond the year 1824) in mid- than in high-latitudes, presumably due to the combined impacts of cold and drought stress. As Tambora-size eruptions statistically occur every 200-400 years, assessing their influences on ecosystems can help humankind mitigate adverse impacts on natural resources through improved management, especially in high latitude and elevation regions., (© 2023. Springer Nature Limited.)

Humans have shaped open oak forests for centuries through pollarding and grazing. Nowadays, these cultural landscapes face the abandonment of their traditional uses and new threats, including rising temperatures and increasing drought stress, especially in southern Europe. We need precise data on the long-term radial growth changes of these oak woodlands to better characterize and preserve them. To fill this research gap, we compared the growth patterns and responses to climate variables and a drought index of three traditionally pollarded deciduous oaks (Quercus subpyrenaica, Quercus faginea, Quercus pyrenaica) and one previously pruned, evergreen oak (Quercus ilex) in central and northeastern Spain. In the three deciduous oaks, we reconstructed radial growth suppressions which were mainly attributed to past pollarding events. Recent post-pollarding growth improvement was transitory but long-term growth enhancement could be maintained by periodic pollarding. Formerly pollarded oaks were old reaching maximum ages of 313 years in the case of Q. faginea. Formerly pruned Q. ilex trees were also old reaching ages of at least 384 years. Peaks in major growth suppressions of Q. faginea sites corresponded to periods of intense timber demand following abrupt socioeconomic changes (land tenure and land use changes, local population growth, wars) such as the 1820s, 1840s, 1910s and 1940s. However, other growth suppressions corresponded to dry periods such as the 1870s and 1950s. Oak growth was constrained by warm-dry conditions in spring and by short- to long-term summer droughts (4-18 months). Pollarding abandonment and increased aridification threaten the survival of such old pollarded oak stands that preserve unique cultural, ecological and biological values., Competing Interests: Declaration of competing interest Authors declare no conflict of interest., (Copyright © 2023 The Author(s). Published by Elsevier B.V. All rights reserved.)

Ongoing climatic change is threatening the survival of drought-sensitive tree species, such as silver fir ( Abies alba ). Drought-induced dieback had been previously explored in this conifer, although the role played by tree-level genetic diversity and its relationship with growth patterns and soil microsite conditions remained elusive. We used double digest restriction-site-associated DNA sequencing (ddRADseq) to describe different genetic characteristics of five silver fir forests in the Spanish Pyrenees, including declining and non-declining trees. Single nucleotide polymorphisms (SNPs) were used to investigate the relationships between genetics, dieback, intraspecific trait variation (functional dendrophenotypic traits and leaf traits), local bioclimatic conditions, and rhizosphere soil properties. While there were no noticeable genetic differences between declining and non-declining trees, genome-environment associations with selection signatures were abundant, suggesting a strong influence of climate, soil physicochemical properties, and soil microbial diversity on local adaptation. These results provide novel insights into how genetics and diverse environmental factors are interrelated and highlight the need to incorporate genetic data into silver fir forest dieback studies to gain a better understanding of local adaptation.

Background and Aims: The vulnerability and responsiveness of forests to drought are immensely variable across biomes. Intraspecific tree responses to drought in species with wide niche breadths that grow across contrasting climatically environments might provide key information regarding forest resistance and changes in species distribution under climate change. Using a species with an exceptionally wide niche breath, we tested the hypothesis that tree populations thriving in dry environments are more resistant to drought than those growing in moist locations., Methods: We determined temporal trends in tree radial growth of 12 tree populations of Nothofagus antarctica (Nothofagaceae) located across a sharp precipitation gradient (annual precipitation of 500-2000 mm) in Chile and Argentina. Using dendrochronological methods, we fitted generalized additive mixed-effect models to predict the annual basal area increment as a function of year and dryness (De Martonne aridity index). We also measured carbon and oxygen isotope signals (and estimated intrinsic water-use efficiency) to provide potential physiological causes for tree growth responses to drought., Key Results: We found unexpected improvements in growth during 1980-1998 in moist sites, while growth responses in dry sites were mixed. All populations, independent of site moisture, showed an increase in their intrinsic water-use efficiency in recent decades, a tendency that seemed to be explained by an increase in the photosynthetic rate instead of drought-induced stomatal closure, given that δ18O did not change with time., Conclusions: The absence of drought-induced negative effects on tree growth in a tree species with a wide niche breadth is promising because it might relate to the causal mechanisms tree species possess to face ongoing drought events. We suggest that the drought resistance of N. antarctica might be attributable to its low stature and relatively low growth rate., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

The benefits of masting (volatile, quasi-synchronous seed production at lagged intervals) include satiation of seed predators, but these benefits come with a cost to mutualist pollen and seed dispersers. If the evolution of masting represents a balance between these benefits and costs, we expect mast avoidance in species that are heavily reliant on mutualist dispersers. These effects play out in the context of variable climate and site fertility among species that vary widely in nutrient demand. Meta-analyses of published data have focused on variation at the population scale, thus omitting periodicity within trees and synchronicity between trees. From raw data on 12 million tree-years worldwide, we quantified three components of masting that have not previously been analysed together: (i) volatility, defined as the frequency-weighted year-to-year variation; (ii) periodicity, representing the lag between high-seed years; and (iii) synchronicity, indicating the tree-to-tree correlation. Results show that mast avoidance (low volatility and low synchronicity) by species dependent on mutualist dispersers explains more variation than any other effect. Nutrient-demanding species have low volatility, and species that are most common on nutrient-rich and warm/wet sites exhibit short periods. The prevalence of masting in cold/dry sites coincides with climatic conditions where dependence on vertebrate dispersers is less common than in the wet tropics. Mutualist dispersers neutralize the benefits of masting for predator satiation, further balancing the effects of climate, site fertility and nutrient demands., (© 2023. The Author(s), under exclusive licence to Springer Nature Limited.)

Despite the importance of species interaction in modulating the range shifts of plants, little is known about the responses of coexisting life forms to a warmer climate. Here, we combine long-term monitoring of cambial phenology in sympatric trees and shrubs at two treelines of the Tibetan Plateau, with a meta-analysis of ring-width series from 344 shrubs and 575 trees paired across 11 alpine treelines in the Northern Hemisphere. Under a spring warming of +1°C, xylem resumption advances by 2-4 days in trees, but delays by 3-8 days in shrubs. The divergent phenological response to warming was due to shrubs being 3.2 times more sensitive than trees to chilling accumulation. Warmer winters increased the thermal requirement for cambial reactivation in shrubs, leading to a delayed response to warmer springs. Our meta-analysis confirmed such a mechanism across continental scales. The warming-induced phenological mismatch may give a competitive advantage to trees over shrubs, which would provide a new explanation for increasing alpine treeline shifts under the context of climate change., (© The Author(s) 2023. Published by Oxford University Press on behalf of China Science Publishing & Media Ltd.)

Climate warming may induce growth decline in warm-temperate areas subjected to seasonal soil moisture deficit, whereas increasing atmospheric CO 2 concentration (C a ) is expected to enhance tree growth. An accurate understanding of tree growth and physiological processes responding to climate warming and increasing C a is critical. Here, we analyzed tree-ring stable carbon isotope and wood anatomical traits of Pinus tabuliformis from Qinling Mountains in China to understand how lumen diameter (LD) determining potential hydraulic conductivity and cell-wall thickness (CWT) determining carbon storage responded to climate and C a . The effects of climate and C a on intrinsic water-use efficiency (iWUE) were isolated, and iWUE values due to only-climate (iWUE Clim ) and only-CO 2 effects (iWUE CO2 ) were obtained. During a low-iWUE period, the influences of climate on earlywood (EW) LD and latewood (LW) CWT prevailed. During a high-iWUE period, CO 2 fertilization promoted cell enlargement and carbon storage but this was counteracted by a negative influence of climate warming. The limiting direct effects of iWUE Clim and indirect effects of climate on EW LD were greater than on LW CWT. P. tabuliformis in temperate forests will face a decline of growth and carbon fixation, but will produce embolism-resistant tracheids with narrow lumen responding to future hotter droughts., (© 2023. The Author(s) under exclusive licence to International Society of Biometeorology.)

Despite growing interest in predicting plant phenological shifts, advanced spring phenology by global climate change remains debated. Evidence documenting either small or large advancement of spring phenology to rising temperature over the spatio-temporal scales implies a potential existence of a thermal threshold in the responses of forests to global warming. We collected a unique data set of xylem cell-wall-thickening onset dates in 20 coniferous species covering a broad mean annual temperature (MAT) gradient (-3.05 to 22.9°C) across the Northern Hemisphere (latitudes 23°-66° N). Along the MAT gradient, we identified a threshold temperature (using segmented regression) of 4.9 ± 1.1°C, above which the response of xylem phenology to rising temperatures significantly decline. This threshold separates the Northern Hemisphere conifers into cold and warm thermal niches, with MAT and spring forcing being the primary drivers for the onset dates (estimated by linear and Bayesian mixed-effect models), respectively. The identified thermal threshold should be integrated into the Earth-System-Models for a better understanding of spring phenology in response to global warming and an improved prediction of global climate-carbon feedbacks., (© 2022 John Wiley & Sons Ltd.)

Local differentiation at distribution limits may influence species' adaptive capacity to environmental changes. However, drivers, such gene flow and local selection, are still poorly understood. We focus on the role played by range limits in mountain forests to test the hypothesis that relict tree populations are subjected to genetic differentiation and local adaptation. Two alpine treelines of mountain pine (Pinus uncinata Ram. ex DC) were investigated in the Spanish Pyrenees. Further, an isolated relict population forming the species' southernmost distribution limit in north-eastern Spain was also investigated. Using genotyping by sequencing, a genetic matrix conformed by single nucleotide polymorphisms (SNPs) was obtained. This matrix was used to perform genotype-environment and genotype-phenotype associations, as well as to model risk of non-adaptedness. Increasing climate seasonality appears as an essential element in the interpretation of SNPs subjected to selective pressures. Genetic differentiations were overall weak. The differences in leaf mass area and radial growth rate, as well as the identification of several SNPs subjected to selective pressures, exceeded neutral predictions of differentiation among populations. Despite genetic drift might prevail in the isolated population, the Fst values (0.060 and 0.066) showed a moderate genetic drift and Nm values (3.939 and 3.555) indicate the presence of gene flow between the relict population and both treelines. Nonetheless, the SNPs subjected to selection pressures provide evidences of possible selection in treeline ecotones. Persistence in range boundaries seems to involve several selective pressures in species' traits, which were significantly related to enhanced drought seasonality at the limit of P. uncinata distribution range. We conclude that gene flow is unlikely to constrain adaptation in the P. uncinata rear edge, although this species shows vulnerability to future climate change scenarios involving warmer and drier conditions., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 The Authors. Published by Elsevier B.V. All rights reserved.)

Extreme climate events such as late spring frosts (LSFs) negatively affect productivity and tree growth in temperate beech forests. However, detailed information on how these forests recover after such events are still missing. We investigated how LSFs affected forest cover and radial growth in European beech (Fagus sylvatica L.) populations located at different elevations at four sites in the Italian Apennines, where LSFs have been recorded. We combined tree-ring and remote-sensing data to analyse the sensitivity and recovery capacity of beech populations to LSFs. Using daily temperature records, we reconstructed LSF events and assessed legacy effects on growth. We also evaluated the role played by elevation and stand structure as modulators of LSFs impacts. Finally, using satellite images we computed Normalized Difference Vegetation Index (NDVI), Enhanced Vegetation Index (EVI) and LAI (Leaf Area Index) to evaluate the post-LSF canopy recovery. The growth reduction in LSF-affected trees ranged from 36 % to 84 %. We detected a negative impact of LSF on growth only during the LSF year, with growth recovery occurring within 1-2 years after the event. LSF-affected stands featured low vegetation indices until late June, i.e. on average 75 days after the frost events. We did not find a clear relationship between beech forest elevation and occurrence of LSFs defoliations. Our results indicate a high recovery capacity of common beech and no legacy effects of LSFs., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Alteration of forest by climate change and human activities modify the growth response of trees to temperature and moisture. Growth trends of young forests with even-aged stands recruited recently when the climate became warmer and drier are not well known. We analyze the radial growth response of young conifer trees (37-63 years old) to climatic parameters and drought stress employing Pearson correlations and the Vaganov-Shashkin Lite (VS-Lite) model. This study uses tree rings of six species of conifer trees (Pinus teocote, Pinus pseudostrobus, Pinus pinceana, Pinus montezumae, Pinus ayacahuite, and Taxodium mucronatum) collected from young forests with diverse growth conditions in northern and central Mexico. Seasonal ring growth and earlywood width (EW) were modeled as a function of temperature and soil moisture using the VS-Lite model. Wet and cool conditions in the previous winter and current spring enhance ring growth and EW production, mainly in sensitive species from dry sites (P. teocote, P. pseudostrobus, P. pinceana, and P. montezumae), whereas the growth of species from mesic sites (P. ayacahuite and T. mucronatum) shows little responsiveness to soil moisture. In P. ayacahuite and T. mucronatum, latewood growth is enhanced by warm summer conditions. The VS-Lite model shows that low soil moisture during April and May constrains growth in the four sensitive species, particularly in P. pinceana, the species dominant in the most xeric sites. Assessing seasonal ring growth and combining its response to climate with process-based growth models could complement xylogenesis data. Such framework should be widely applied, given the predicted warming and its impact on young forests., (© 2022. The Author(s).)