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Forest protection and afforestation have been identified as a means to partially offset anthropogenic CO2 emissions. Yet, increasingly frequent observations of drought-induced tree mortality are reported. Here, we applied a risk analysis framework for global drought-induced forest mortality by examining extreme reductions in greenness and water content of forest canopies during past mortality events as well as growth recovery of surviving individual trees following stand-scale mortality events. We defined a drought-induced mortality risk index (DMR) that explains 80% of documented tree mortality. Rising CO2 alleviated the increase of DMR with short-term drought, however, the observed DMR increases with long-term drought no matter whether considering plant responses to CO2. DMR in sites where tree mortality has been observed significantly increased since the 1980s. More than that, drought exposure threatened 0.28 billion hectares of forested areas. Our framework highlights how climate change-induced drought, especially hotter-droughts, threatens the sustainability of global forests.

A warmer climate will increase aridity and threaten forest persistence in xeric areas. This is the case of some Atlas cedar (Cedrus atlantica) forests showing recent growth decline, dieback and high mortality rates in North Africa. A lower resistance to drought, manifested as stronger growth loss, could increase the drought-related mortality risk in these forests as has been found for gymnosperms worldwide. It could be also expected that changes in dryness are linked to large-scale atmospheric circulation patterns such as the North Atlantic Oscillation (NAO). We tested these hypotheses by analyzing growth resilience indices (resilience, recovery, resistance) derived from tree-ring data. We sampled nine Atlas cedar plots located in north-western Algeria, where climate shifted towards drier conditions in the 1980s. In these forests, drier winters, associated to positive NAO phases, constrained tree growth and resistance, two outputs of the drought impact. During dry years, the resilience index decreased as elevation increased. The association between cedar growth resistance and drought severity is strengthening: the drier the climate conditions, the lower the growth resistance. Resistance also showed a significant higher temporal variability in 8 out of 9 plots as drought intensified. These findings have allowed identifying the increase in the temporal variability of growth resistance as a new early-warning signal of drought stress.

Carbon accounting in the land sector requires a reference level from which to calculate past losses of carbon and potential for gains using a stock-based target. Carbon carrying capacity represented by the carbon stock in primary forests is an ecologically-based reference level that allows estimation of the mitigation potential derived from protecting and restoring forests to increase their carbon stocks. Here we measured and collated tree inventory data at primary forest sites including from research studies, literature and forest inventories (7982 sites, 288,262 trees, 27 countries) across boreal, temperate, and subtropical Global Ecological Zones within Europe. We calculated total biomass carbon stock per hectare (above- and below-ground, dead biomass) and found it was 1.6 times larger on average than modelled global maps for primary forests and 2.3 times for all forests. Large trees (diameter greater than 60 cm) accounted for 50% of biomass and are important carbon reservoirs. Carbon stock foregone by harvesting of 12–52% demonstrated the mitigation potential. Estimated carbon gain by protecting, restoring and ongoing growth of existing forests equated to 309 megatons carbon dioxide equivalents per year, additional to, and higher than, the current forest sink, and comparable to the Green Deal 2030 target for carbon dioxide removals.

Forests in tectonically active regions are disturbed by earthquakes. Besides direct injuries to trees, earthquakes also induce stand-wide changes in hydrological conditions, whose effects on long-term forest growth and resilience remain unknown. Here we establish spatio-temporal links between global tree-ring width series and earthquakes after 1900, disentangle seismic signals from climate-induced variations in ring width series, test growth changes using superposed epoch analysis and quantify post-earthquake resilience shifts along environmental gradients in seven regions around the world. We found sites with enhanced resilience locate in relatively dry areas of temperate regions, where the response of tree growth to growing-season precipitation also increased after earthquakes. Our results provide evidence that earthquake-induced soil cracks and fractures increased precipitation infiltration to deeper soil layers and enhanced the use of water and nutrients by trees. In contrast, reduced post-earthquake resilience in regions with abundant precipitation can be explained by increased soil erosion and nutrient leaching. We conclude that seismic disturbances cause decadal-scale shifts in forest resilience under specific environmental conditions, disentangling complex interactions between lithosphere, biosphere and atmosphere. These findings can contribute to a better understanding of how the Earth system functions.

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.

Severe droughts limit tree growth and forest productivity worldwide, a phenomenon which is expected to aggravate over the next decades. However, how drought intensity and climatic conditions before and after drought events modulate tree growth resilience remains unclear, especially when considering the range-wide phenotypic variability of a tree species. We gathered 4632 Aleppo pine (Pinus halepensis Mill.) tree-ring width series from 281 sites located in 11 countries across the Mediterranean basin, representing the entire geographic and bioclimatic range of the species. For each site and year of the period 1950–2020, we quantified tree-growth resilience and its two components, resistance and recovery, to account for the impact of drought and the capacity to recover from it. Relative drought intensity of each year was assessed using SPEI (Standardized Precipitation Evapotranspiration Index), a climatic water deficit index. Generalized additive mixed models were used to explore the non-linear relationships between resilience and its two components and drought intensity, preceding and following years climatic conditions. We found that P. halepensis radial growth was highly dependent on the SPEI from September of the previous year to June of the current year. Trees growing under more arid bioclimates showed higher inter-annual growth variability and were more sensitive to drought, resulting in an increased response magnitude to pre-, during and post-drought conditions. In contrast to our expectations, drought intensity only slightly affected resilience, which was rather negatively affected by favorable preceding conditions and improved by favorable following conditions. Resilience and its components are highly dependent on preceding and following years climatic conditions, which should always be taken into account when studying growth response to drought. With the observed and predicted increase in drought frequency, duration and intensity, favorable conditions followi

Understanding the chemical composition of our planet's crust was one of the biggest questions of the 20th century. More than 100 years later, we are still far from understanding the global patterns in the bioavailability and spatial coupling of elements in topsoils worldwide, despite their importance for the productivity and functioning of terrestrial ecosystems. Here, we measured the bioavailability and coupling of thirteen macro- and micronutrients and phytotoxic elements in topsoils (3–8 cm) from a range of terrestrial ecosystems across all continents (∼10,000 observations) and in response to global change manipulations (∼5,000 observations). For this, we incubated between 1 and 4 pairs of anionic and cationic exchange membranes per site for a mean period of 53 days. The most bioavailable elements (Ca, Mg, and K) were also amongst the most abundant in the crust. Patterns of bioavailability were biome-dependent and controlled by soil properties such as pH, organic matter content and texture, plant cover, and climate. However, global change simulations resulted in important alterations in the bioavailability of elements. Elements were highly coupled, and coupling was predictable by the atomic properties of elements, particularly mass, mass to charge ratio, and second ionization energy. Deviations from the predictable coupling-atomic mass relationship were attributed to global change and agriculture. Our work illustrates the tight links between the bioavailability and coupling of topsoil elements and environmental context, human activities, and atomic properties of elements, thus deeply enhancing our integrated understanding of the biogeochemical connections that underlie the productivity and functioning of terrestrial ecosystems in a changing world.

Key message: Abies alba Mill.–Pinus uncinata Ramond. ecotone dynamics are examined along both altitudinal and protection level gradients by combining field inventories and Landsat data. An upward expansion of A. alba to the subalpine belt is observed in the last decades as a result of stand maturation after logging cessation. Context: High-mountain forests constitute sensitive locations to monitor the impacts of global change on tree-species composition and ecotone dynamics. In this study, we focus on the Spanish Pyrenees where silver fir (Abies alba Mill.) coexists with mountain pine (Pinus uncinata Ramond.) forming montane-subalpine ecotones. Aims: The main goal of this study is to assess the spatiotemporal dynamics of the silver fir–mountain pine ecotone and its underlying driving factors. Methods: We reconstructed the spatial distribution and dynamics of the species by combining remote sensing imagery and field plot data from 1989 to 2015, employing support vector machine techniques for image classification. Using variance analysis and mixed effects models, we then analyzed the evolution of basal area and replacement index, a measure of relative change in species composition, over time and altitude range. Additionally, we explored their relationship with site factors and protection level (National Park vs. protection buffer zone). Results: Silver fir has expanded its distribution in both the National Park and the protection buffer zone, whereas mountain pine has remained stable. Both species exhibit increased basal area associated with stand maturation and a higher level of protection. The replacement index indicates a rise in silver fir in the understory on North-facing slopes, attributed to stand densification. These findings are particularly noticeable in the area with the highest level of protection. Conclusion: The cessation of traditional land uses has led to ongoing stand densification, promoting succession and favoring the increased abundance of silver f

Recent drought-induced dieback alters forest dynamics, which are also shaped by past management. In western Pyrenean silver fir (Abies alba) stands, dieback concurs in space and time with the legacies of past management, but the impacts on forest growth, structure and composition are unknown. We aim to disentangle how dieback interacts with the legacies of past human use and modulates the recent dynamics of silver fir forests. To this end, we sampled eleven silver fir forests across wide climatic gradients and included declining and non-declining sites. We measured radial growth, structure, composition, understory cover and type and amount of deadwood. Silver fir growth declines in response to late-summer drought. In declining sites, most defoliated stands showed the lowest silver fir density and were those where growth depended more on water availability. Tree death enhanced the cover of dominant understory plants such as Buxus sempervirens. Past management activities leave an imprint in the growth of silver fir, such as releases due to past logging, but also affect the number of stumps and snags and the current tree density. A more extensive monitoring will be required to fully disentangle the multiple influences of past management legacies and current climate change on forest dynamics.

In seasonally dry forests, wildfires can reduce competition for soil water among trees and improve forest resilience to drought. We tested this idea by comparing tree-ring growth patterns of Pinus pinea stands subjected to two prescribed burning intensities (H, high; L, low) and compared them with unburned (U) control stands in southwestern Spain. Then, we assessed post-growth resilience to two droughts that occurred before (2005) and after (2012) the prescribed burning (2007). Resilience was quantified as changes in radial growth using resilience indices and as changes in cover and greenness using the NDVI. The NDVI sharply dropped after the fire, and minor drops were also observed after the 2005 and 2012 droughts. We found that post-drought growth and resilience were improved in the H stands, where growth also showed the lowest coherence among individual trees and the lowest correlation with water year precipitation. In contrast, trees from the L site showed the highest correlations with precipitation and the drought index. These findings suggest that tree growth recovered better after drought and responded less to water shortage in the H trees. Therefore, high-intensity fires are linked to reduced drought stress in Mediterranean pine forests.

Editorial on the Research Topic: Forests under pressure: The need for interdisciplinary approaches to address forest vulnerability to tree mortality in response to drought

This document provides supporting analysis and figures for the main article., Identifying Korean pine tree species from carbonized trees. In this study, the tree species of carbonized trees was identified by analyzing the microscopic anatomical features of wood on three planes (cross-, radial-, and tangential-section). For the identified Korean pine (Pinus koraiensis Siebold & Zucc.) in this study, the microscopic anatomical features were showed below, Cross-section features. The boundary of growth ring is slightly obvious, and the early wood gradually changes to late wood within the ring. The growth of early wood is uniform, and the early wood accounts for most of the ring width. The axial parenchyma tissue is absent. The resin channels are divided into axial and radial types. The axial channels are mostly single and usually distributed in the late wood or located at 2/3 of the ring width. The lipid cells are thin-walled and often contain quasi-invasion bodies. There are 3~6 lipid cells around the maximum axial resin channel. The transverse section of early wood tracheid is square, rectangular and polygon, and the axial parenchyma tissue is absent., Radial-section features. Radiative tracheids are located in 1~2 columns of the upper and lower edges, and are occasionally seen in the middle of the rays. The xylem low rays are sometimes made up entirely of radial tracheids. The inner wall is smooth and the outer edge is wavy. The cross field pattern between ray parenchyma cells and early wood tracheids was pane-like, or pine type was occasionally seen in late wood, with number of 1~4, 1 (rare 2) in horizontal line. Axial parenchyma cell horizontal wall perforation and end wall nodular thickening are usually absent or few, not obvious, no indentation., Tangential-section features. There are 4-8 wood rays per mm, including two types: single column and spindle-shaped. The single column of rays is 2~13 cells or more in height, most of them are 3~8 cells; the spindle rays have radial resin canals, 2~3 columns of ray cells above and below the approach; The upper and lower ends gradually sharpen into a single column, the height of which is 3-10 cells or more. The ray cells are usually oval in shape. The radial resin tract is much smaller, with 3-5 adipocytes around the tract., Conclusions. According to the microscopic anatomical characteristics of the above three planes, the carbonized logs, No. CBSXA-01, CBSXA-12, CBSXA-21, CBSXA-24, CBSXA-27, CBSXB-01, CBSXB-02, CBSXB-03, CBSXB-05, CBSXB-07, CBSXB-08, CBSXB-10, CBSXB-11, CBSXB-14, CBSXB-15, CBSXB-17, CBSXB-21, and CBSXB-26, were identified as Korean pine (Pinus koraiensis) of the family Pinaceae and the genus Pinus. The anatomical features of the tree planes of the 18 samples were separately showed below in the large Figure S1.

[Purpose of Review] The capacity of woody plants to cope with climate change depends on their adjustments to changing environmental conditions by phenotypic plasticity or by genotypic changes (i.e., local adaptation). To determine whether intraspecific trait variation (ITV) translates into resistance or tolerance to drought and eventually how it relates to performance when facing drought, we reviewed the recent literature on the intraspecific variation in woody plant functional traits along aridity gradients., [Recent Findings] At the intraspecific level, functional traits vary along aridity gradients, but this variation depends on the considered trait and species. While ITV is ubiquitous and of relevant magnitude, its relationship with tree performance in the case of growth or survival is unclear and very idiosyncratic., [Summary] ITV varies along aridity gradients and, for several traits, is of comparable magnitude as interspecific trait variation. However, the relationship between ITV and tree performance is not consistent among species. This makes the use of ITV challenging when quantifying its contribution to drought tolerance.

Riparian alder forests are threatened by Phytophthora across Europe. Comparative studies of the pathogenicity of Phytophthora species are crucial for developing effective management strategies. Although only a limited number of species, particularly P. × alni, lead to tree decline in natural environments, many species demonstrate pathogenicity in inoculation trials. Phytophthora species vary in their ability to infect different tissues, such as phloem and xylem, and trigger defence responses in the host through the formation of tyloses and callose. By comparing the histological responses of alder to various Phytophthora species, we can gain insights into the success of P. × alni and the potential damage that could be caused by other species. To investigate the defence strategies of black alder (Alnus glutinosa) against attack by Phytophthora and Halophytophthora species present in Catalonia (NE Spain), we conducted inoculation trials on saplings using nine potentially pathogenic species and compared the histological responses. Phytophthora × alni and P. plurivora were the most aggressive species followed by other exotic species such as P. × cambivora and P. cactorum. Phytophthora × alni and P. plurivora were equally damaging despite the higher prevalence of P. × alni in declining alders in natural settings. Although P. × alni mainly invaded the phloem, P. plurivora also invaded the xylem and triggered the production of tyloses. Histological analyses revealed a diverse range of plant responses to infection by Phytophthora species, providing a better understanding of their adaptability in natural environments compared with solely observing lesions. The low level of callose production in saplings inoculated with P. × alni compared with callose production in saplings inoculated with less pathogenic species suggests that P. × alni can evade recognition by the host and, hence, could partially explain its success.

Tree phenology is sensitive to climate warming and changes in seasonal precipitation. Long xylogenesis records are scarce, thus limiting our ability to analyse how radial growth responds to climate variability. Alternatively, process-based growth models can be used to simulate intra-annual growth dynamics and to better understand why growth bimodality varies along temperature and precipitation gradients. We used the Vaganov-Shashkin (VS) growth model to analyse the main climatic drivers of growth bimodality in eight trees and shrubs conifers (four pines and four junipers) across Spain. We selected eleven sites with different continentality degree and spring/autumn precipitation ratios since we expected to find pronounced bimodal growth in less continental sites with spring and autumn precipitation peaks. The VS model successfully simulated annual growth rates at all sites as a function of daily temperature and soil moisture data. Bimodal growth patterns clustered into less continental sites showing low spring/autumn precipitation ratios. This finding agrees with observed climate-growth associations showing that growth was enhanced by wet-cool winter-to-spring conditions, but also by wet autumn conditions in the most bimodal sites. We observed a stronger growth bimodality in pines compared to junipers. We discuss the spatial variability of climate drivers in bimodality growth pattern and how increasing continentality and shifts in seasonal precipitation could affect growth patterns. Bimodality could be an advantageous response to overcome summer drought in Mediterranean forests. The ability of some species to reactivate growth during autumn might determine their capacity to withstand increasing summer aridity.

The growth responses to climate variability are still unknown in locally threatened conifers from dry regions, but this information is necessary for improving the conservation of relict populations under increasing aridification. We characterized the radial growth patterns and responses to climate of Tetraclinis articulata, a Cupressaceae tree endemic to the western Mediterranean Basin, in a relict population located in southwestern Spain (Doñana) and two populations from the northern Morocco where the species core habitat is found (Tétouan, Ifarten). We assessed climate-growth relationships by using tree-ring width, climate data, drought and North Atlantic Oscillation (NAO) indices. Climate-growth analyses were refined using the climwin R package to select the most informative statistical models. The main climatic constraints of growth were inferred by using the process-based Vaganov-Shashkin (VS) model explicitly considering non-linear climate-growth relationships. Tetraclinis articulata growth was favored by wet conditions from the prior autumn to the spring of the growth year. In Doñana, warmer May conditions led to growth decline but this negative effect could be reversed by wet-warm conditions in the prior late autumn. Growth in the two Moroccan sites was constrained by 6- to 18-month long droughts peaking in summer, which account for cumulative water deficit since the previous autumn. Winter and early spring precipitation were the main climate drivers of growth in the Moroccan T. articulata populations, and their year-to-year variability was linked to the NAO. The VS model simulations showed that T. articulata growth is enhanced by wet soil conditions in late winter and early spring, probably recharging shallow soil water pools. The VS model also confirmed that warmer spring-summer conditions could amplify drought stress and threaten the long-term persistence of the relict Doñana population.

During the past years, growth and productivity of different oak species have been constrained by water shortage in seasonally dry regions such as the Mediterranean Basin. Thinning could improve oak radial growth in these drought-prone regions through the reduction of tree competition for soil water in summer. However, we still lack adequate, long-term assessments on how lasting are thinning treatments effects and to what extent they contribute to oak growth recovery after drought. Here we aim: (i) to study the radial growth sensitivity to drought of Quercus subpyrenaica in the Spanish Pre-Pyrenees, and (ii) to verify if thinning represents a suitable option to enhance growth resistance to drought and post-drought growth recovery. We analysed basal area increment (BAI) trends in the period 1960–2020 of formerly coppiced oak stands thinned in 1984 and compared them with unthinned plots and also with coexisting Scots pine (Pinus sylvestris) growing in thinned plots. We used the Standardized Precipitation Evapotranspiration Index (SPEI) to estimate the severity of droughts and we also assessed climate-growth relationships. Oaks in thinned plots showed higher BAI (369 mm2) than those in unthinned plots (221 mm2). Growth rates remained higher in thinned than in unthinned plots also under severe drought stress. A severe summer drought in 1986 caused abrupt BAI reductions in both oaks (- 40.5%) and pines (- 40.1%). The positive effect of thinning on growth lasted for over 20 years and slightly declined as canopies closed. In the thinned plots, trees with smaller diameter showed the greatest growth release. Oaks in unthinned plots and Scots pine were more sensitive to short-term droughts in terms of growth reduction than oaks in thinned plots, while long term droughts have similar effects on oaks from both thinned and unthinned plots. Oaks were resilient to drought, showing recovery periods lasting from 1 to 2 years in both thinned and unthinned plots. However, intense and p

Forest growth is driven by climate variability at continental to regional scales, but other factors play major roles at local scales (0.1–1 ha). Topography impacts on tree growth responses to climate stressors, including drought, by modifying radiation, evapotranspiration rates and the access to soil moisture. However, there is a lack of studies investigating how topographical factors (elevation, aspect, slope) affect climate-growth relationships considering both continuous (tree-ring width) and discrete wood-anatomical features (e.g., intra-annual density fluctuations–IADFs). Here, we investigated how topography modulated the influences of climate and drought on semi-arid Juniperus thurifera forests from north-eastern Spain. We compared two stands located in valley bottoms and two stands located in steep slopes. Radial growth was measured using dendrochronology which also allowed quantifying latewood IADFs. A proxy of topographic influence (incident radiation) was calculated. In addition, the Vaganov-Shashkin (VS) model was used to infer the main climatic constraints of growth at intra-annual scales. We found that junipers growing in valley bottoms were taller and produced more IADFs than junipers growing in steep slopes, but responded less to precipitation variability. This was confirmed by the VS model which showed that low soil moisture in June limited growth, particularly in steep-slope sites and during dry periods. Wet-cool late-summer conditions induced the formation of IADFs. The topoclimatic modulation of growth was explained by the interaction between incident radiation and tree age. Furthermore, climate-growth associations are changing as temperatures rise. Drought impacts on growth are strengthening, but less responsive junipers growing in valley bottoms may be buffered against intensified aridification.

Floodplain forests are sensitive to climate warming and increased drought, as showed by recent oak (Quercus robur) dieback and mortality episodes. However, a comprehensive comparison of coexisting tree species under different climate settings or biomes are lacking. Herein, we compared growth rates, growth responses to climate and drought severity, and modeled climate mediated growth of oak and three coexisting tree species (ash, Fraxinus angustifolia; alder, Alnus glutinosa; elm, Ulmus minor). Two floodplain forests subjected to cooler (temperate climate, Ticino) and warmer (Mediterranean climate, Bosco Pantano) conditions in northern and southern Italy, respectively, were analyzed. Ash seemed to be the most sensitive to drought, particularly at the Mediterranean site where oak and elm growth were also negatively affected by water shortages. Alder appeared to be the least sensitive species in terms of growth variability to drought under both temperate and Mediterranean climate conditions. Furthermore, the growth model revealed the influence of soil moisture in spring and summer on the constrained growth of ash and oak and illustrated how oak growth could be severely reduced during drastic hotter droughts. Alder seemed to be the most drought-resistant species under both environmental conditions. These results could represent the first attempts in documenting the ecological consequences of drought in terms of projected climate trends in less investigated Mediterranean floodplain forests. Furthermore, these results highlight how climate and tree-ring data combined with growth models could be useful tools to detect early warning signals of growth decline and impending dieback in floodplain forests in response to dry spells.

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.

Drought reduces canopy cover, productivity and tree growth in forests. However, there is still little knowledge on how drought affects coupling between canopy greenness assessed by remote sensing and hydraulic conductivity detected by wood anatomy. This combination could improve the understanding of forest response to climate change. Thus, we investigated the impacts of a hot drought, which occurred in summer 2017, on radial growth, earlywood hydraulic diameter (Dh), a proxy of conductivity, and several remote-sensing indices in mixed Mediterranean hardwood forests (Quercus pubescens Willd. – Fraxinus ornus L.). In general, growth showed a higher coherence among trees and a higher responsiveness to climate. Growth decreased during the drought year, particularly for Q. pubescens, which showed high defoliation and dieback intensity. Both species showed a decline of Dh in 2018 after the drought and subsequent warm winter conditions. We found positive relationships between Dh and remote-sensing data for Q. pubescens in some of these vulnerable sites, where (i) growth was constrained by dry spring-summer conditions and (ii) Dh and growth covaried. These findings indicate a high variability among sites and tree species in their responses to drought considering earlywood anatomy, growth canopy cover and water content. However, some common patterns emerge such as links between potential hydraulic conductivity (Dh), tree cover and Dh-growth covariation in the most impacted sites. Further, F. ornus seem to perform better in terms of growth under drought conditions, showing less mortality and dieback than Q. pubescens. Future studies could explore how water transport and changes in canopy cover respond to dry and warm conditions and if that covariation indicates vulnerability to drought.

Tree-ring studies are still lacking in tropical African forests. This is the case in the seasonally dry miombo forests located in Southern Africa. In the Angolan miombo, subject to intense charcoal exploitation, tree-ring data is urgently needed to estimate the age at which the minimum permitted cutting diameter is reached. Further, climate-growth relationships must be also investigated to understand how major miombo tree species respond to climate constraints and teleconnections such as the El Niño Southern Oscillation (ENSO). To achieve both aims, we studied radial growth data of two miombo tree legume species (Brachystegia spiciformis, Julbernardia paniculata) in wet (Bailundo) and dry (Caála) Angolan sites using dendrochronological methods. Both species have diffuse porous wood and conspicuous ring boundaries delimited by terminal parenchyma in latewood. Sampled individuals had ages (at 1.3 m) between 28 and 34 years with ring widths ranging 3.8–4.3 and 5.5–6.0 mm in dry and wet sites, respectively. In the wet (dry) site, Brachystegia and Julbernardia reached maximum diameter increment rates of 1.05–1.32 (0.74–0.91) cm yr−1 at an age of 12 (14−20) years. Both species took 12–15 years to reach a minimum cutting diameter of 15 cm. The growth variability among conspecific individuals was lower in the dry (mean standard error, 1.4 cm) than in the wet site (mean standard error, 2.7 cm). We also found that wet conditions from November to February, often linked to El Niño events, enhanced growth for both species, with greater growth consistency among individuals and higher sensitivity to climate found in the dry site. This information may help to estimate the optimal age for minimum cutting diameter that guarantees the sustainable use of charcoal and fuelwood.

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.

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

[Aims] Shifts in xylem phenology directly determine the forest capacity for carbon sequestration. However, a systematic understanding of the spatial patterns and the underpinning drivers in determining the cessation of wood formation (Ccw) is lacking at a pan-continental scale. Here, we addressed this knowledge gap by compiling a new dataset of multiple xylem phenology timings for northern conifers., [Locations] Sixty-two study sites, Northern Hemisphere (25–55° N)., [Time period] 2003–2018 (16 years)., [Taxa] Thirty-three conifer species., [Methods] A generalized additive model was fitted to characterize the latitudinal pattern in Ccw. Structural equation modelling and a linear mixed-effects model were applied to determine the main drivers underlying the latitudinal pattern in Ccw., [Results] The Ccw followed a flat S-shaped pattern with increasing latitude. Photoperiod was the dominant determinant of the latitudinal pattern of Ccw, and a longer photoperiod was associated with an earlier Ccw. Both mean growing-season temperature and total growing-season precipitation exhibited significantly positive relationships to the cessation of cell elongation and thus the Ccw across all study sites. In arid regions, the pre-growing-season temperature had a significantly negative effect on Ccw. In humid regions, Ccw was positively affected by the mean growing-season temperature. The onset of wood formation showed significantly positive coupling with Ccw at arid sites but not at humid sites. Early successional species were sensitive to hydrothermal variations during the pre-growing season., [Main conclusions] We reveal the dominant role of photoperiod in determining the cessation of wood formation for northern conifers and highlight differentiated interactive effects between photoperiod and seasonal climatic factors and the preceding xylem phenophases in determining Ccw among ecoregions and tree species. These insights provide evidence to reduce uncertainty in prediction of the forest carbon uptake potential and the consequent biophysical feedbacks of northern forests.

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.

Some widely distributed timber species, such as oaks, are vulnerable to low soil water availability and drought. Therefore, selecting broadleaved minor species with lower sensitivity to drought could be an alternative in seasonally dry areas. However, the growth performance of these minor hardwood species is unknown, particularly under warm and dry conditions at the southern limit of their distribution range. We assessed the radial growth response to drought by correlating tree-ring and climate data in coexisting service trees (Sorbus domestica) and two oak species (Quercus faginea, Quercus pyrenaica). Trees were sampled in three Mediterranean sites located in Spain with different precipitation patterns. We used the Vaganov–Shashkin (VS) growth model to infer the main climate constraints of growth. To ascertain whether climate is changing tree phenology, we also simulated changes in xylem onset and cessation timings and compared them with leaf falling dates in service trees. Service trees showed a trend towards advancing leaf fall, but this was not related to xylem growth. Oaks responded more to a drought index than service trees. The strongest responses corresponded to droughts peaking from spring to summer, but oaks tended to respond to longer droughts (9–13 months) than service trees (2–7 months). These different responses are due to the positive responses of all species to high precipitation in the growing season, but the sensitivity of oak growth to warm summer conditions and increased atmospheric water demand. The VS model indicated a more bimodal growth pattern in services trees than in oaks, which could allow the former to recover better after a dry summer. Service trees could be more widely used as a source of valuable timber wood under Mediterranean continental conditions.

Assessing post-fire recovery is essential to forecast how ecosystems will respond to future warmer conditions and higher fire severity. Such assessments must consider site conditions and the post-fire recovery of trees and shrubs. We used tree-ring data and intrinsic water-use efficiency (WUEi) to quantify the post-fire responses of a tree (Pinus pinea) and a shrub (Juniperus phoenicea) in Mediterranean coastal dunes located in the Doñana protected area, SW Spain. We compared pines and junipers growing in an inter-dune slack with those growing in a nearby dune ridge. We quantified crown damage and bark char in pines impacted by a wildfire. Pines with lower crown damage after the fire showed a higher growth rate one year later. Growth decreased afterwards in the case of fast-growing pines from the slack site, whereas slow-growing dune pines showed increasing growth rates one to three years after the fire. The fire did not change the pines’ WUEi, whereas the junipers located in dunes presented the highest WUEi values. Severe crown damage (damage > 60%) impairs long-term growth recovery in P. pinea. Open and heterogeneous landscapes can reduce the wildfire risk in the study Mediterranean area, where slack vs. dune and tree vs. shrub differences should be considered in post-fire management plans.

Lack of tree fecundity data across climatic gradients precludes the analysis of how seed supply contributes to global variation in forest regeneration and biotic interactions responsible for biodiversity. A global synthesis of raw seedproduction data shows a 250-fold increase in seed abundance from cold-dry to warm-wet climates, driven primarily by a 100-fold increase in seed production for a given tree size. The modest (threefold) increase in forest productivity across the same climate gradient cannot explain the magnitudes of these trends. The increase in seeds per tree can arise from adaptive evolution driven by intense species interactions or from the direct effects of a warm, moist climate on tree fecundity. Either way, the massive differences in seed supply ramify through food webs potentially explaining a disproportionate role for species interactions in the wet tropics.

The loss of resilience and prolonged recovery times after extreme climate events can be used as early warning signals of impending tipping points or abrupt irreversible changes. However, evidence of such critical slowing down in growth series is still lacking in terrestrial ecosystems under harsh environmental conditions such as alpine shrublands. Alpine shrublines, at which shrublands shift into grasslands under climate thresholds, constitute unique settings to look for critical slowing down phenomena. On the northeastern Tibetan Plateau, we sampled Salix oritrepha Schneid. (willow shrubs) in three elevational sites along each of two transects (QL, Qilian; HSX, Huashixia) running from alpine treeline position to shrubline. We then quantified the radial growth resilience and recovery indices of alpine willow shrubs to cold spells along elevational gradients by using dendrochronology. Willow growth was primarily constrained by low winter temperatures in QL and low summer temperatures in HSX. Cold spell events were detected in 2008 and 2011 in HSX and QL sites, respectively. Because temperature decreases with elevation, both growth resilience and recovery indices differed within each transect, showing a decreasing trend with elevation. Growth in 78% and 56% willow shrubs recovered back to the level before cold spell within two years in QL and HSX sites, respectively. Moreover, the average growth recovery period was longer at the shrubline (ca. 2–4 years) than near the treeline position (ca. 1–2 years). Critical slowing down theory could thus be applied to assess growth resilience towards the alpine shrubline, a thermal threshold for shrub growth. Our results and framework highlight the priority to directly measure the resilience metrics and also offer a promising perspective that the critical slowing down phenomenon may be universal in various ecosystems.

Trees grow at night, when the vapor pressure deficit (VPD) is low enough. Therefore, intra-annual density fluctuations (IADFs) should be formed when the VPD drops below a certain threshold. This idea is tested by assessing climate-latewood IADF relationships in six conifer species under Mediterranean climate conditions. Hourly climate and dendrometer data were analyzed for years with elevated IADF production in two species (Pinus halepensis, Juniperus thurifera). Lastly, climate–growth relationships were evaluated in two drought-prone sites to assess the relative role of minimum vs. maximum temperatures as growth drivers. Latewood IADF production was positively related to growth rate. IADFs were more abundant when monthly or 10-day long precipitation was high in the late growing season (August and September). According to dendrometer data, growth mainly occurred in early night (20–2 h) and early morning (6–8 h). This growth window corresponded to rainy periods with VPD below a minimum threshold associated with summer storms. Latewood IADFs are produced in response to these wet late-summer conditions, which could be related to bimodal growth. These associations are in line with correlations showing that high minimum (night) rather than maximum (day) temperatures reduce growth. This last idea should be further checked in drought-prone forests using global tree-ring databases.

Wood encodes environmental information that can be recovered through the study of tree-ring width and wood anatomical variables such as lumen area or cell-wall thickness. Anatomical variables often provide a stronger hydroclimate signal than tree-ring width, but they show a low tree-to-tree coherence. We investigate the sources of variation in tree-ring width, lumen area, and cell-wall thickness in three pine species inhabiting sites with contrasting climate conditions: Pinus lumholtzii in wet-summer northern Mexico, and Pinus halepensis and Pinus sylvestris in dry-summer north-eastern Spain. We quantified the amount of variance of these three variables explained by spring and summer water balance and how it varied among trees. Wood anatomical variables accounted for a larger inter-individual variability than tree-ring width data. Anatomical traits responded to hydroclimate more individualistically than tree-ring width. This individualistic response represents an important issue in long-term studies on wood anatomical characteristics. We emphasized the degree of variation among individuals of the same population, which has far-reaching implications for understanding tree species’ responses to climate change. Dendroclimatic and wood anatomical studies should focus on trees rather than on the mean population series.

Drought-induced dieback is a matter of global concern. It has been widely reported and could compromise the climate warming mitigation potential of forests. That is why we need reliable early-warning proxies to achieve better forecasts of forest dieback and tree death. Tree-ring data can provide some of these needed proxies. Here I propose considering minimum (MND) and maximum (MXD) wood density values as proxies of tree vulnerability against drought. This hypothesis is evaluated in silver fir (Abies alba) forests from the western Spanish Pyrenees showing ongoing dieback processes after the severe 1985 drought. MXD increased in response to warm summer conditions in 1985 whereas MND increased in response to growing-season water deficit in 1986. The average between prior MXD (MXDt-1) and subsequent MND (MNDt) could be used to detect severe drought impacts and forecast dieback.

Shrub recruitment, a key component of vegetation dynamics beyond forests, is a highly sensitive indicator of climate and environmental change. Warming-induced tipping points in Arctic and alpine treeless ecosystems are, however, little understood. Here, we compare two long-term recruitment datasets of 2,770 shrubs from coastal East Greenland and from the Tibetan Plateau against atmospheric circulation patterns between 1871 and 2010 Common Era. Increasing rates of shrub recruitment since 1871 reached critical tipping points in the 1930s and 1960s on the Tibetan Plateau and in East Greenland, respectively. A recent decline in shrub recruitment in both datasets was likely related to warmer and drier climates, with a stronger May to July El Niño Southern Oscillation over the Tibetan Plateau and a stronger June to July Atlantic Multidecadal Oscillation over Greenland. Exceeding the thermal optimum of shrub recruitment, the recent warming trend may cause soil moisture deficit. Our findings suggest that changes in atmospheric circulation explain regional climate dynamics and associated response patterns in Arctic and alpine shrub communities, knowledge that should be considered to protect vulnerable high-elevation and high-latitude ecosystems from the cascading effects of anthropogenic warming.

There is large uncertainty on the future of European silver fir (Abies alba) forests and how they will respond to climate warming. Many studies, such as Walder et al. (2021), suggest they will show positive growth trends and a larger distribution area in response to warmer conditions whenever summer drought is not severe and long. This contradicts localized dieback events in continental, drought-prone Mediterranean sites mainly located in the southwestern margin of the current species distribution range. We discuss these disagreements and propose compound climate extremes and elevated atmospheric drought as drivers of dieback in similar marginal silver fir populations.

Forest biomass is an important component of terrestrial carbon pools. However, how climate, biodiversity, and structural attributes co-determine spatiotemporal variation in forest biomass remains not well known. We aimed to shed light on these drivers of forest biomass by measuring diversity and structural attributes of tree species in 400-m2 plots located every 100 m along a 4200-m elevational gradient in the eastern Himalayas. We applied structural equation models to test how climate, species richness, structural attributes, and their interactions influence forest biomass. Importantly, species richness was a stronger driver of biomass than environmental and structural attributes such as annual air temperature or stem density. Integrating the availability of energy and the demand for water, potential evapotranspiration was more strongly correlated with biomass than water availability, likely due to the strong influence of the Indian summer monsoon. Thus, interactions between climate and tree community composition ultimately control how much carbon is stored in woody biomass across bioclimatic gradients. This fundamental understanding will support predictive efforts of the forest carbon sink in this hydroclimatically important region and help preserving regional forests as a potent natural solution for climate change mitigation.

Diplodia shoot blight is an emergent forest disease in Europe caused by Diplodia sapinea. The short-term impacts of the pathogen on tree physiology are well known, but its capacity to cause mortality has been poorly documented. We compared the survival of four pine species affected by Diplodia shoot blight following a hailstorm: Pinus sylvestris, P. nigra, P. pinea and P. halepensis. In the case of P. sylvestris, survival in the hail-affected sites was compared with survival in other sites affected by Diplodia shoot blight with no hailstorm records. Mortality and crown condition were recorded over two years. Dendrochronological analyses were conducted to assess growth responses to drought and to test the influence of radial growth before the outbreak of Diplodia shoot blight on survival. The endophytic community, as well as the abundance of D. sapinea, was quantified by metabarcoding and qPCR respectively, and were correlated with crown damage. Crown damage was the best predictor of mortality across species. Pinus sylvestris experienced a much higher rate of mortality than P. nigra, P. pinea and P. halepensis. Two years after the outbreak, P. halepensis was the only species that could recover crown condition. Mortality was found to be unrelated to radial growth prior the outbreak. Drought responses did not correlate with mortality differences across pine species or sites. In the case of P. sylvestris, mortality was initially higher amongst diseased trees in areas affected by hailstorms than in sites not affected by hailstorms, however it tended to equalise after two years. Amongst P. sylvestris trees, crown damage correlated with a higher abundance of the pathogen. Signs of competition amongst endophytes were observed between non-defoliated and defoliated Scots pine trees following hailstorms. Our study shows that D. sapinea can cause a significant mortality to P. sylvestris. The legacy effects of crown damage can last for at least two years after an outbreak. Durin

Drought-induced forest dieback can lead to a tipping point in community dominance, but the coupled response at the tree and stand-level response has not been properly addressed. New spatially and temporally integrated monitoring approaches that target different biological organization levels are needed. Here, we compared the temporal responses of dendrochronological and spectral indices from 1984 to 2020 at both tree and stand levels, respectively, of a drought-prone Mediterranean Pinus pinea forest currently suffering strong dieback. We test the influence of climate on temporal patterns of tree radial growth, greenness and wetness spectral indices; and we address the influence of major drought episodes on resilience metrics. Tree-ring data and spectral indices followed different spatio-temporal patterns over the study period (1984–2020). Combined information from tree growth and spectral trajectories suggests that a reduction in tree density during the mid-1990s could have promoted tree growth and reduced dieback risk. Additionally, over the last decade, extreme and recurrent droughts have resulted in crown defoliation greater than 40% in most plots since 2019. We found that tree growth and the greenness spectral index were positively related to annual precipitation, while the wetness index was positively related to mean annual temperature. The response to drought, however, was stronger for tree growth than for spectral indices. Our study demonstrates the value of long-term retrospective multiscale analyses including tree and stand-level scales to disentangle mechanisms triggering and driving forest dieback.

Tree-ring data has been widely used to inform about tree growth responses to drought at the individual scale, but less is known about how tree growth sensitivity to drought scales up driving changes in forest dynamics. Here, we related tree-ring growth chronologies and stand-level forest changes in basal area from two independent data sets to test if tree-ring responses to drought match stand forest dynamics (stand basal area growth, ingrowth, and mortality). We assessed if tree growth and changes in forest basal area covary as a function of spatial scale and tree taxa (gymnosperm or angiosperm). To this end, we compared a tree-ring network with stand data from the Spanish National Forest Inventory. We focused on the cumulative impact of drought on tree growth and demography in the period 1981–2005. Drought years were identified by the Standardized Precipitation Evapotranspiration Index, and their impacts on tree growth by quantifying tree-ring width reductions. We hypothesized that forests with greater drought impacts on tree growth will also show reduced stand basal area growth and ingrowth and enhanced mortality. This is expected to occur in forests dominated by gymnosperms on drought-prone regions. Cumulative growth reductions during dry years were higher in forests dominated by gymnosperms and presented a greater magnitude and spatial autocorrelation than for angiosperms. Cumulative drought-induced tree growth reductions and changes in forest basal area were related, but initial stand density and basal area were the main factors driving changes in basal area. In drought-prone gymnosperm forests, we observed that sites with greater growth reductions had lower stand basal area growth and greater mortality. Consequently, stand basal area, forest growth, and ingrowth in regions with large drought impacts was significantly lower than in regions less impacted by drought. Tree growth sensitivity to drought can be used as a predictor of gymnosperm demographic rates in

The consecutive occurrence of drought and insect outbreaks could lead to cumulative, negative impacts on boreal forest productivity. To disentangle how both stressors affected productivity, we compared changes in tree canopy cover and radial growth after a severe outbreak in Siberian spruce (Picea obovata) southern taiga forests. Specifically, we studied the impacts of the 2012 severe drought followed by a Siberian silk moth (Dendrolimus sibiricus, hereafter SSM) outbreak, which started in 2016, on spruce forests by comparing one non-defoliated site and two, nearby fully defoliated sites, using remote sensing and tree-ring data. The SSM outbreak caused total defoliation and death of trees in the infested stands. We found a sharp drop (−32%) in the normalized difference infrared index and reduced radial growth in the defoliated sites in 2018. The growth reduction due to the 2012 drought was −37%, whereas it dropped to 4% of pre-outbreak growth in 2018. Tree growth was constrained by warm and dry conditions from June to July, but such a negative effect of summer water shortage was more pronounced in the defoliated sites than in the non-defoliated site. This suggests a predisposition of sites where trees show a higher growth responsivity to drought to SSM-outbreak defoliation. Insect defoliation and drought differently impacted taiga forest productivity since tree cover dropped due to the SSM outbreak, whereas tree growth was reduced either by summer drought or by the SSM outbreak. The impacts of abiotic and biotic stressors on boreal forests could be disentangled by combining measures or proxies of canopy cover and radial growth which also allow the investigation of drought sensitivity predisposes to insect damage.

Mountain forests affect spatial and temporal variability of snow processes through snow interception and by modifying the energy balance of snowpack. The high sensitivity of snow cover to seasonal temperatures in mid–latitude mountains is well known and is of particular interest with regard to a future warmer climate. The snowpack in the Pyrenees is expected to be the most impacted by climate change in the Mediterranean mountains, where future climate trends project rising temperatures and decreasing precipitation. This study analyzes how changes in temperature and precipitation can affect current forest–snow interactions in four forests, located near each other but under contrasting topographic settings, in the Spanish Pyrenees. This understanding will allow us to anticipate the future hydrological responses of Pyrenean forested mountain basins. The research was accomplished by performing a sensitivity analysis using simulations from the Cold Regions Hydrological Model (CRHM) and by comparing forest canopy sites (F) vs. openings (O). The CRHM platform focuses on the incorporation of physically based descriptions of snow–dominated regions hydrological processes. It was found that forest cover induced different snowpack sensibility to climatic change conditions in the studied forests. Delayed onset of snow accumulation (F: 13 days·°C−1; O: 5 days·°C−1) and reduced snowpack duration (F: 28 %·°C−1; O: 23 %·°C−1) under warmer temperatures were more intense in areas beneath the forest canopy compared to openings. A lower annual peak of snow water equivalent (SWE) (F: 81 mm·°C−1; O: 129 mm·°C−1), earlier melt-out date (F: 8 days·°C−1; O: 10 days·°C−1) and slower melting rates (F: 0.4 mm·day−1·°C−1; O: 0.5 mm·day−1·°C−1) with increasing temperatures were more intense in forest openings. The forest–driven reduction in snowpack duration (40%) was significantly enhanced with warming (10% per °C). Lower precipitation (20% precipitation reduction) could increase the response of

Tree resistance to extreme droughts and post-drought recovery are sensitive to the drought timing. However, how the bioclimatic sensitivity of tree growth may vary with the timing and order of extreme droughts and wetness is still poorly understood. Here, we quantified the bioclimatic sensitivity of tree growth in the period of 1951–2013 under different seasonal extreme drought/wetness regimes over the extra-tropical Northern Hemisphere, using 1,032 tree ring chronologies from 121 gymnosperm and angiosperm species. We found a negative asymmetry in tree growth under regimes with seasonal extreme droughts. With extreme drought, tree growth in arid and temperate dry regions is more negatively impacted by pre-growing-season (PGS) extreme droughts. Clade-wise, angiosperms are more sensitive to PGS water availability, and gymnosperms to legacy effects of the preceding tree growth conditions in temperate dry and humid regions. Our finding of divergent bioclimatic legacy effects underscores contrasting trends in forest responses to drought across different ecoregions and functional groups in a more extreme climate.

Climatic warming is assumed to expand the geographic range of insect pests whose distribution is mainly constrained by low temperatures. This is the case of the pine processionary moth (Thaumetopoea pityocampa), which is one of the main conifer defoliators in the Mediterranean Basin. Warmer winters may lead to a northward/upward expansion of this insect, as short-term studies have shown. However, no long-term data, i.e. spanning at least one century, has been used to examine these projections. We test the hypotheses that climatic warming (i) has caused an upward shift of the pine processionary moth, and (ii) has increased the frequency of severe defoliations. We used dendrochronological methods to reconstruct defoliations over the period 1900–2006 in 14 sites spanning a wide altitudinal range (1070–1675 m) in Teruel, eastern Spain. We built local ring-width chronologies for four co-occurring pine species with different degree of susceptibility against the moth defoliations, from highly suitable or palatable species (Pinus nigra) to moderately (Pinus sylvestris, Pinus halepensis) or rarely defoliated species (Pinus pinaster). We validated the tree-ring reconstructions of outbreaks using a field record of stand defoliations spanning the period 1971–2006. Outbreaks in the most affected P. nigra stands corresponded to abrupt one- to two-year growth reductions (70–90% growth loss). Reconstructed outbreaks occurred on average every 9–14 years. The growth memory of outbreaks was weaker but lasted longer (1–6 years) than that due to droughts (1–3 years). Neither an upward expansion nor an increase in outbreak frequency was observed. Severe PPM defoliations did not increase as climate warmed, rather they were positively related to the winter North Atlantic Oscillation.

In mountains, shrubs forming their uppermost distributional limits (shrubline) are highly sensitive to increasing cold limitations upwards. However, the constraints imposed by a shorter growing season due to lower temperatures upslope could interact with drought stress in mountainous regions. Here, we aim to investigate that question by comparing radial growth patterns of shrubs in wet (forest region of Nangqian County, BZ hereafter) vs. dry (forest region of Leiwuqi County, LWQ hereafter) regions located on the eastern Tibetan plateau. Considering the climate trend in this area towards warmer and drier conditions, we hypothesize that growth of high-elevation shrubs will be mainly limited by low summer temperatures under sufficient soil moisture. To test this hypothesis, we collected ring-width data from 123 alpine shrubs (Salix oritrepha Schneid.) along an altitudinal gradient from 4200 to 4600 m a.s.l. Dendroecological analyses showed that shrub growth was positively related to mean June or July temperature during the past decades in both wet (BZ) and dry (LWQ) regions regardless of site elevation. By contrast, correlations between shrub growth and seasonal precipitation were not significant. Shrub growth trends were positive before 2010 but turned negative afterwards. Overall, radial growth of shrubs along the altitudinal gradients up to the shrubline could shift from being temperature to moisture limited if warming-induced drought stress intensifies.

Drought is an important driver of forest dynamics in the Mediterranean region. The forecasted increase in drought frequency and severity can notably influence tree growth, forest structure, composition and productivity. Understanding how coexisting tree species respond to drought is thus crucial to understand which are less vulnerable and will perform better in a warmer and drier world. To assess drought vulnerability, we used dendrochronology to study the radial growth trends and responses to a drought index of four pine species (Pinus halepensis, Pinus pinea, Pinus nigra, and Pinus sylvestris) coexisting in North-eastern Spain. We reconstructed the growth of each species and evaluated their short- and long-term growth response to drought for the common period 1980–2017. The growth of the four pine species depended on water availability and high early spring temperatures impacted the growth of P. nigra and P. sylvestris negatively. The occurrence of a severe drought between 2005 and 2007 lead to marked growth reductions in the four species, but it was greater in magnitude in P. pinea and P. halepensis in 2005, and in P. nigra in 2007. The results of basal area increment models at the individual tree level suggested that P. halepensis trees grow more than the rest of species. After accounting for age and drought effects, P. nigra and P. sylvestris displayed negative growth trends in the 2008–2017 period while P. pinea and P. halepensis displayed positive growth trends. P. sylvestris was the most resistant species and P. pinea the less resistant. Conversely, P. halepensis and P. pinea were slightly more resilient than P. sylvestris. Moreover, P. sylvestris was the species displaying the highest autocorrelation and the lowest coefficient of variation in ring-width indices. A marked drop in the autocorrelation of P. pinea ring-width index was observed in response to the 2005 drought. These results indicate that all study species are vulnerable to drought but in differe