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This scientific commentary discusses the importance of understanding carbon source-sink dynamics in masting tree species, particularly in the context of climate change and drought stress. The authors examine the responses of carbon allocation to environmental constraints in stone pine stands, focusing on reproductive effort, radial growth, and tree-ring isotope composition. They find evidence of a shift from sink-limited to source- and sink-co-limited reproduction, suggesting diminishing carbon resources due to drought stress. However, caution is needed in interpreting the data, and further research is necessary to fully understand carbon allocation in trees. [Extracted from the article]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Climate warming is expected to positively alter upward and poleward treelines which are controlled by low temperature and a short growing season. Despite the importance of treelines as a bioassay of climate change, a global field assessment and posterior forecasting of tree growth at annual scales is lacking. Using annually resolved tree‐ring data located across Eurasia and the Americas, we quantified and modeled the relationship between temperature and radial growth at treeline during the 20th century. We then tested whether this temperature–growth association will remain stable during the 21st century using a forward model under two climate scenarios (RCP 4.5 and 8.5). During the 20th century, growth enhancements were common in most sites, and temperature and growth showed positive trends. Interestingly, the relationship between temperature and growth trends was contingent on tree age suggesting biogeographic patterns in treeline growth are contingent on local factors besides climate warming. Simulations forecast temperature–growth decoupling during the 21st century. The growing season at treeline is projected to lengthen and growth rates would increase and become less dependent on temperature rise. These forecasts illustrate how growth may decouple from climate warming in cold regions and near the margins of tree existence. Such projected temperature–growth decoupling could impact ecosystem processes in mountain and polar biomes, with feedbacks on climate warming. [ABSTRACT FROM AUTHOR]

The negative impacts of drought on forest growth and productivity last for several years generating legacies, although the factors that determine why such legacies vary across sites and tree species remain unclear.We used an extensive network of tree‐ring width (RWI, ring‐width index) records of 16 tree species from 567 forests, and high‐resolution climate and normalized difference vegetation index (NDVI) datasets across Spain during the common period 1982‒2008 to test the hypothesis that climate conditions and growth features modulate legacy effects of drought on forests. Legacy effects of drought were calculated as the differences between detrended‐only RWI and NDVI series (i.e. after removing long‐term growth trends) and pre‐whitened RWI and NDVI series predicted by a model including drought intensity. Superposed Epoch Analysis (SEA) was used to estimate whether legacy effects differed from random. Finally, legacy effects were related to water balance, growth persistence and variability, and tree species identity.We found a widespread occurrence of drought legacy effects on both RWI and NDVI, but they were seldom significant. According to SEA, first‐year drought legacies were negative and different from random in 9% and 5% of the RWI and NDVI series respectively. The number of significant second‐ and third‐year legacies was substantially lower. Differences between RWI and NDVI legacies indicate that canopy greenness and radial growth responses to drought are decoupled. We found variations in legacies between tree species with gymnosperms presenting larger first‐year drought legacies than angiosperms, which were exposed to less severe droughts. Greater growth variability can explain the presence of first‐year RWI legacies in gymnosperms from dry sites despite that the relationship between growth variability and legacies was complex.Synthesis. Accounting for species and site responses to drought provides a better understanding of the magnitude and duration of drought legacies on forest growth and productivity. Despite the widespread occurrence of growth reductions in the years during and after drought occurrence, significant legacies were not very common, mostly lasted one year, and were more widespread in gymnosperms. These are relevant factors to be considered in the future when studying the consequences of drought on forest productivity and tree growth. [ABSTRACT FROM AUTHOR]

Dendrometers are being increasingly used to measure stem radius changes in trees and to unravel the mechanisms underlying stem daily rhythms of radial expansion and contraction. Nevertheless, automated dendrometers have not been often used to measure root radius dynamics, their relationship with environmental variables and the influence of endogenous processes, especially in drought-prone Mediterranean areas. Here, we measured root radius dynamics of two coexisting oak species (the evergreen Quercus ilex L. and the deciduous Quercus faginea Lam). Our goals were to describe annual, seasonal and diurnal scale root radius patterns and to disentangle the role of different environmental parameters as drivers. Long-term high-resolution measurements (every 15 min over 7 years) were collected with automated point dendrometers on the main tree roots of five individuals per species. Root radius annual change patterns were bimodal and similar for both oak species. Quercus faginea Lam showed three times larger root increment in the spring than Q. ilex , but the bimodal pattern was stronger in Q. ilex , which showed a larger root increment in autumn. Quercus faginea Lam showed an earlier root phenological activation in the spring and in late summer compared with Q. ilex. The effects of environmental drivers across species were similar at daily scales: root radius increased with air temperature and soil moisture, and it decreased with rising vapor pressure deficit. Furthermore, daily root radius variations for both oak species were maintained after extracting statistically the environmental effects, which points toward a significant role of endogenous drivers. These differences in root radius change patterns at seasonal to daily scales likely result from the differences in leaf phenology and growth strategy. Quercus faginea Lam is deciduous and has a faster growing rate in spring than the evergreen Q. ilex, which can grow more in summer. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Climate warming increases vulnerability to drought in Mediterranean water-limited forests. However, we still lack knowledge of the long-term physiological responses of coexisting pine species in these forests regarding their ability to cope with warming-induced drought stress., We investigated spatiotemporal patterns of tree performance for five isohydric pines with partially overlapping ecological niches in the eastern Iberian Peninsula along an altitudinal gradient: Pinus halepensis = Pinus pinaster ≤ Pinus nigra ≤ Pinus sylvestris ≤ Pinus uncinata., Using indexed tree-ring widths ( TRW i), we assessed the changes in the temporal coherence of radial growth (growth synchrony, â C) over the period 1902-2011 across three elevation belts: low ≈1100 m; mid = 1615 m; high = 2020 m. We also examined by mixed modelling whether TRW i showed an increased coupling with leaf-level gas exchange (inferred from indexed carbon isotope discrimination, Δ13C i) by enhanced stomatal regulation in response to an amplified regional drought stress., Increasingly negative annual water balances (decrease in annual precipitation minus evapotranspiration = −4·8 mm year−1; 1970-2011) prompted more synchronous growth of coexisting pines between low- and mid-elevation belts, with â C rising from 0·25 ± 0·04 (1902-1951) to 0·62 ± 0·05 (1962-2011). This effect was coupled with tighter stomatal regulation at mid-elevation as indicated by high correlations between TRW i and Δ13C i (>0·60 from the mid-1970s onwards) which resembled those found at low elevation. Simultaneously, TRW i vs. Δ13C i uncoupling occurred at the high-elevation belt across species., Weaker growth-climate relationships as elevation increased highlighted the major role of the altitude-dependent thermal gradient in growth responsiveness to drought; however, an intensified Δ13C i response to spring water availability across elevation belts observed from mid-1970s onwards suggested regional shifts in tree physiological activity linked to earlier seasonal drought impacts. Warming-induced drought stress is spreading to higher altitudes in Iberian pinewoods as multispecies growth is linked to progressively tighter stomatal control of water losses reflected in wood Δ13C. [ABSTRACT FROM AUTHOR]

Individual trees, not forests, respond to climate. Such an individual-scale approach has seldom been used to retrospectively track the radial growth responses of trees to climate in dendrochronology. The aim of this study was to adopt this individual view to retrospectively assess tree sensitivity to climate warming, and to evaluate and compare the potential drivers of tree growth responses to climate acting at species, site and individual scales., Following a dendroecological framework, we sampled a network of 29 Pinus uncinata forests in NE Spain and obtained tree-ring widths series from 642 trees. Individual features as northness, elevation, slope, basal area, sapwood area, tree height and tree age were used to evaluate the potential drivers of tree growth responses to climate. The analysed data set includes diverse ecological and biogeographical conditions. The tree growth responses to climate were assessed by relating growth indices to climatic variables using linear-mixed effects models., Maximum November temperatures during the year prior to tree-ring formation enhanced P. uncinata growth mainly in mid-elevation sites, whereas at higher elevations growth was more dependent on the positive effect of warmer minimum May temperatures during the year of tree-ring formation. Current June precipitation was the positive main climatic driver of growth in sites prone to water deficit such as the southernmost limit of the species distribution area or very steep sites. Elevation was the main factor controlling how much growth variability is explained by climate at the site and tree scales. Climate warming was more intense during the early 20th century, when the importance of elevation as an indirect modulator of growth declined as compared with the late 20th century., Synthesis. The individual-scale approach taken in this study allowed detecting that trees growing at southern and low-elevation sites were the most negatively affected by warm and dry summer conditions. Our results emphasize that both (i) an individual-scale approach to quantify tree growth responses to climate and (ii) a detailed evaluation of the potential biotic and abiotic drivers of those individual responses are necessary to understand climate sensitivity of trees. [ABSTRACT FROM AUTHOR]

Aim The Mediterranean Basin is considered to be a climate-change hotspot, for which rising temperatures and associated aridification have been forecast. Such trends could affect the performance and growth of conifers in these drought-prone areas. We evaluated whether radial growth and wood density can act as proxy measures of precipitation and drought in a Mediterranean conifer. Location Iberian Juniperus thurifera forests in northern and eastern Spain. Methods We sampled 10 stands encompassing J. thurifera's distributional area. We related four annually resolved tree-ring variables (earlywood and latewood width, and maximum and minimum wood density) to climatic factors (temperature, precipitation and drought index) for the period 1951-2000. We then analysed the spatial and temporal patterns of those associations. Results Of all the four tree-ring variables analysed, minimum wood density presented the strongest response to climate. Higher values of minimum wood density were related to drier spring conditions, particularly in the most xeric sites, confirming that a drought-induced reduction in the radial expansion of tracheids increases earlywood density. Main conclusions Minimum wood density is a biogeographically meaningful proxy of spring water availability for the Mediterranean conifer J. thurifera. [ABSTRACT FROM AUTHOR]

Winter-drought induced forest diebacks in the low-latitude margins of species' distribution ranges can provide new insights into the mechanisms (carbon starvation, hydraulic failure) underlying contrasting tree reactions. We analysed a winter-drought induced dieback at the Scots pine's southern edge through a dual-isotope approach (Δ13 C and δ18 O in tree-ring cellulose). We hypothesized that a differential long-term performance, mediated by the interaction between CO2 and climate, determined the fates of individuals during dieback. Declining trees showed a stronger coupling between climate, growth and intrinsic water-use efficiency ( WUEi) than non-declining individuals that was noticeable for 25 years prior to dieback. The rising stomatal control of water losses with time in declining trees, indicated by negative Δ13 C- δ18 O relationships, was likely associated with their native aptitude to grow more and take up more water (suggested by larger tracheid lumen widths) than non-declining trees and, therefore, to exhibit a greater cavitation risk. Freeze-thaw episodes occurring in winter 2001 unveiled such physiological differences by triggering dieback in those trees more vulnerable to hydraulic failure. Thus, WUEi tightly modulated growth responses to long-term warming in declining trees, indicating that co-occurring individuals were differentially predisposed to winter-drought mortality. These different performances were unconnected to the depletion of stored carbohydrates. [ABSTRACT FROM AUTHOR]

Abstract: Acclimation to local conditions may produce adaptive responses in plants subjected to diverse climatic stresses. However, it has not been assessed how local adaptation and phenotypic plasticity contribute to photoprotection mechanisms in response to contrasting climatic conditions in Mediterranean tree species. We analyzed photoprotection mechanisms in mature trees of the Mediterranean evergreen oak Quercus ilex at three sites with contrasting climatic conditions, i.e. xeric, continental and mesic sites. We studied morphological and physiological parameters indicative of photoprotection in adult trees in the field. In order to establish whether these parameters were genetically determined we compared adults with seedlings germinated from acorns of the three sites and grown under common greenhouse conditions. In the field we found no significant differences in most of the physiological parameters in summer, but in winter the adult trees from the continental site were photoinhibited. In contrast, there were significant differences between seedlings in most photoprotective parameters evaluated. Morphological traits such as trichome density and leaf reflectance differed between populations, both in field-grown trees and in greenhouse-grown seedlings, being higher in all cases in plants from the xeric site than elsewhere. Our findings suggest the existence of constitutive differences in leaf photoprotection mechanisms among Q. ilex populations. These divergences may represent an inherent source for more stress tolerant ecotypes in the face of changing climatic conditions. [Copyright &y& Elsevier]

1. Long-term basal area increment (BAI) in Abies pinsapo was studied to investigate the way density-dependent factors modulate the responses of radial growth to climatic stresses in relict stands of a drought-sensitive Mediterranean fir. 2. First, we verified that spatially explicit competition predicts mean A. pinsapo BAI at our study site; i.e. it modulates the degree to which the average climate-driven potential for growth is expressed. Second, we verified that the long-term pattern of temperature predicts the long-term pattern of BAI, estimated as the main trend over a time period of c. 40 years. Finally, we assessed whether the intensity of tree-to-tree competition restrains the potential improvements achieved by our model of BAI when a short-term, high-frequency stressor such as drought (inter-annual precipitation variability) is introduced. 3. We applied Dynamic Factor Analysis (DFA) to characterize regional climatic trends and to test the hypothesis that trees subjected to contrasting competition intensity may differ in their growth pattern. Significant long-term climate trends obtained by DFA were used as predictors of long-term BAI. 4. The mean BAI was mainly determined by competition, whereas growth trends obtained by DFA did not differ among dominant, suppressed and dying trees. Common trends of growth decline were strongly related to long-term, late-winter to summer temperatures, while the residuals were related to total annual precipitation, although with decreasing significance as competition increased. Our results support the contention that the reported patterns of A. pinsapo growth decline and death occur as a result of the interacting effects of both competition and climate stressors acting at long- and short-term time scales. 5. Synthesis. Long-term climatic drought stress was the main driving factor of growth decline in A. pinsapo. Moreover, trees already suffering from competition (a long-term stress) were predisposed to decline given an additional short-term stress, such as a severe drought. [ABSTRACT FROM AUTHOR]

•Seasonal radial-increment and xylogenesis data can help to elucidate how climate modulates wood formation in conifers. Few xylogenesis studies have assessed how plastic xylogenesis is in sympatric conifer species from continental Mediterranean areas, where low winter temperatures and summer drought constrain growth. •Here, we analysed intra-annual patterns of secondary growth in sympatric conifer species ( Juniperus thurifera, Pinus halepensis and Pinus sylvestris). Two field sites (xeric and mesic) were evaluated using dendrometers, microcores and climatic data. •A bimodal pattern of xylogenesis characterized by spring and autumn precipitation and subsequent cambial reactivation was detected in J. thurifera at both study sites and in P. halepensis at the xeric site, but was absent in P. sylvestris where growth was largely controlled by day length. In the xeric site J. thurifera exhibited an increased response to water availability in autumn relative to P. halepensis and summer cambial suppression was more marked in J. thurifera than in P. halepensis. • Juniperus thurifera exhibited increased plasticity in its xylogenesis pattern compared with sympatric pines, enabling this species to occupy sites with more variable climatic conditions. The plastic xylogenesis patterns of junipers in drought-stressed areas may also provide them with a competitive advantage against co-occurring pines. [ABSTRACT FROM AUTHOR]

Abstract: Silver-fir (Abies alba Mill.) populations located at the south side of the main Pyrenean axis and Pre-Pyrenees constitute the south-western distribution limit of the species and, therefore, may be responding more noticeably to climate change than those in the centre of the range. The increasing aridity detected in the Pyrenees during the 20th century should affect more negatively the physiological performance of the southernmost silver-fir stand growing under more xeric conditions in comparison with stands growing within the main distribution area under mesic conditions. To evaluate the climatic influence on the performance of silver fir near its distribution limit, we studied several physiological and growth variables in shoots and needles from two silver fir populations located in nearby but climatically contrasting sites: Paco Ezpela (site E) and Gamueta (site G). Site E showed a stronger Mediterranean influence than site G, i.e. the former site was characterized by higher temperatures and leaf-to-air water vapour pressure difference and lower precipitation in summer than the latter site. Silver firs from site E showed lower values of primary and secondary growth, needle length, stomatal conductance, net photosynthesis and photosystem II (PSII) efficiency than individuals from site G. The reduction in net photosynthesis could be ascribed to a low CO2 availability and to a lower PSII efficiency. We conclude that the physiological differences found between both sites were caused by the more xeric conditions of site E as compared with the more mesic environment in site G. The predicted increase of severe droughts in the southern Pyrenees might cause a decrease in photosynthesis and growth in those silver-fir stands located near the ecological limit of the species. [Copyright &y& Elsevier]

To understand how tree growth has responded to recent climate warming, an understanding of the tree-climate-site complex is necessary. To achieve this, radial growth variability among 204 trees established before 1850 was studied in relation to both climatic and site factors. Seventeen forest stands were sampled in the Spanish Central Pyrenees. Three species were studied: Pinus uncinata, Abies alba, and Pinus sylvestris. For each tree, a ring-width residual chronology was built. All trees cross-dated well, indicating a common influence of the regional climate. For the 1952-1993 period, the radial growth of all species, especially P. uncinata, was positively correlated with warm Novembers during the year before ring formation and warm Mays of the year the annual ring formed. Differences in species-stand elevation modulated the growth-climate associations. Radial growth in P. uncinata at high elevation sites was reduced when May temperatures were colder and May precipitation more abundant. In the 20th century, two contrasting periods in radial growth were observed: one (1900-1949) with low frequency of narrow and wide rings, low mean annual sensitivity, and low common growth variation; and another (19501994) with the reverse characteristics. The increased variability in radial growth since the 1950s was observed for all species and sites, which suggests a climatic cause. The low shared variance among tree chronologies during the first half of the 20th century may result from a "relaxation" of the elevation gradient, allowing local site conditions to dominate macroclimatic influence. These temporal trends may be related to the recently reported increase of climatic variability and warmer conditions. This study emphasizes the need to carefully assess the relationships between radial growth and site conditions along ecological gradients to improve dendroclimatic reconstructions. [ABSTRACT FROM AUTHOR]

Thinning focused on achieving growth and diameter management objectives has typically led to stands with reduced climate sensitivity compared to unthinned stands. We integrated dendrochronological with Airborne Laser Scanner (LiDAR) data and growth models to assess the long-term impact of thinning intensity on Canary pine (Pinus canariensis) radial growth. In 1988, 18 permanent treatment units were established in 73-year-old Canary pine plantations and three thinning treatments were applied (C–control-unthinned; 0% basal area removal; MT–moderate thinning: 10% and 15% basal area removal, and HT–heavy thinning: 46% and 45% basal area removal on the windward and leeward slopes, respectively). Dendrochronological data were measured in 2022 and expressed as basal area increment (BAI). The impact of climate on growth was examined by fitting linear regression models considering two different Representative Concentration Pathway (RCP) climate scenarios, RCP 2.6 and RCP 4.5. Finally, LiDAR data were used for standing segmentation to evaluate changes in overall growth under different climatic scenarios. The LiDAR–stand attributes differed between aspects. The BAI of the most recent 20 years (BAI20) after thinning was significantly higher for the moderate and heavy treatments on the leeward plots (F = 47.31, p < 0.001). On the windward plots, BAI decreased after moderate thinning. Considerable thinning treatments resulted in stronger changes in growth when compared to RCP climatic scenarios. From a silviculture perspective, the mapping of canopy structure and growth response to thinning under different climatic scenarios provides managers with opportunities to conduct thinning strategies for forest adaptation. Combining dendrochronological and LiDAR data at a landscape scale substantially improves the value of the separate datasets as forecasted growth response maps allow improving thinning management plans. [ABSTRACT FROM AUTHOR]

A higher frequency and intensity of droughts will impair forest productivity. Therefore, improving our understanding of which factors enhance tree growth resilience against drought has become a crucial issue, but we lack information at the intraspecific level.In this study, we investigate the role played by climatic conditions and tree characteristics in the growth response to severe droughts in two deciduous Fagaceae species near their southern distribution limit: the temperate European beech Fagus sylvatica and the Mediterranean pubescent oak Quercus pubescens. The study area is located in Catalonia, NE Spain, where 149 trees were cored covering the water availability gradient of the species in the region. In addition, tree size (diameter and height) and an ensemble of stem, leaf and hydraulic traits were collected for each tree. Growth responses to extreme droughts during the period 1980–2015 were assessed using resilience indicators based on residual basal area increment series predicted by a model including growing season water balance.Although resistance was unrelated to drought intensity, higher water availability improved recovery and reduced drought legacy effects. Taller trees showed higher resilience against drought, which could be explained by the relationship between the variation in tree height and functional traits, such as leaf nitrogen concentration, leaf turgor loss point and resistance to xylem embolism. For both species, trees with lower cavitation resistance, higher leaf nitrogen concentration and tighter stomatal regulation (suggested by less negative turgor loss point and carbon isotope composition) displayed better performance during and after severe droughts. Both species showed a progressive decrease in resilience and increase in legacy effects.Synthesis. This study highlights the relevance of analysing intraspecific variation of functional traits involved in tree water and carbon balance to improve our understanding of the resilience and variability of growth responses to drought within species. [ABSTRACT FROM AUTHOR]

Keywords: alpine regions; altitudinal distribution; climate change; Juniperuspingii var. wilsonii; moisture; shrub ring; shrubline; temperature EN alpine regions altitudinal distribution climate change Juniperuspingii var. wilsonii moisture shrub ring shrubline temperature 1 4 4 06/09/21 20210601 NES 210601 The genus I Juniperus i , composed of about 76 species, has a broad altitudinal distribution over the northern hemisphere (Adams 2014). (e) Wilson juniper shrub population at the study site. gl To characterize the natural history and responses to climate of this uniquely high shrubline, we estimated the age and quantified the growth patterns of a random sample of its shrubs. We collected basal discs (diameter range: 20-40 mm) from the largest stems of 14 young and isolated juniper shrub patches growing above 5,250 m a. s. l. A plot survey (size: 30 × 150 m) was also conducted near this shrubline site to estimate the age structure of juniper shrub population (see Appendix S1 for details). Soil temperatures during the growing season were more than 3.1°C higher than air temperatures, at least partly highlighting the importance of soil temperature to the juniper shrub growth. [Extracted from the article]

Abstract: Questions: What are the differences in tree growth and resilience in response to extreme droughts between three co‐existing pine tree species? Are growth–resilience components more influenced by intrinsic (age, tree height) or extrinsic factors (mass effects or functional diversity of the neighbourhood)? Is the tree's vulnerability to drought buffered by the functional diversity of the surrounding plant community? Location: Mediterranean forest, northeast Spain. Methods: Trees from three co‐existing pine species with different drought tolerances (Aleppo pine – Pinus halepensis Mill.; Black pine – Pinus nigra subsp. salzmannii (Dunal) Franco; Scots pine – Pinus sylvestris L.) were sampled. The average age of the studed pines was ~30 years. Radial growth from 1980 to 2015 was measured and the pines’ responses to the most severe recent droughts (2005 and 2012) were studied by relating different growth resilience components (Rt, resistance; Rc, recovery; RRs, relative resilience) with intrinsic (age and height) and extrinsic factors of the neighbouring community around each tree. The considered extrinsic factors were CWM and niche complementarity of tree (height), leaf (specific leaf area, leaf dry matter content) and wood (wood specific gravity) traits. The way in which these factors affected growth resilience to drought was assessed. Results: For the whole study period we found lower growth of Scots pine than Aleppo pine, but similar resilience components (except for higher Rt in 2005 for the former species). Despite each drought year resulting in distinct responses, the growth resilience to drought was mostly dependent on tree age for Aleppo pine, while functional diversity modulated a high percentage of the resilience variance of Scots pine. Black pine had an intermediate strategy. CWM explained, in general, less variance of the resilience components than functional diversity. Conclusions: The positive effect of surrounding functional diversity on Scots pine could be related to a greater resistance to drought, likely due to improved resource acquisition favouring its performance at the dry limit of its distribution area. Intrinsic tree features, mass effects and functional diversity differently shape growth resilience to different droughts for co‐existing tree species. [ABSTRACT FROM AUTHOR]

Tree species display a wide variety of water-use strategies, growth rates and capacity to tolerate drought. However, if we want to forecast species capacity to cope with increasing aridity and drought, we need to identify which measurable traits confer resilience to drought across species. Here, we use a global tree ring network (65 species; 1931 site series of ring-width indices--RWI) to evaluate the relationship of long-term growth-drought sensitivity (RWI-SPEI drought index relationship) and short-term growth response to extreme drought episodes (resistance, recovery and resilience indices) with functional traits related to leaf, wood and hydraulic properties. Furthermore, we assess the influence of climate (temperature, precipitation and climatic water deficit) on these trait-growth relationships. We found a close correspondence between the long-term relationship between RWI and SPEI and resistance and recovery of tree growth to severe drought episodes. Species displaying a stronger RWI-SPEI relationship to drought and low resistance and high recovery to extreme drought episodes tended to have a higher wood density (WD) and more negative leaf minimum water potential (Ψmin). Such associations were largely maintained when accounting for direct climate effects. Our results indicate that, at a cross-species level and global scale, wood and hydraulic functional traits explain species' growth responses to drought at short-and long-term scales. These trait-growth response relationships can improve our understanding of the cross-species capacity to withstand climate change and inform models to better predict drought effects on forest ecosystem dynamics. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

We evaluated the response of the Earth land biomes to drought by correlating a drought index with three global indicators of vegetation activity and growth: vegetation indices from satellite imagery, tree-ring growth series, and Aboveground Net Primary Production (ANPP) records. Arid and humid biomes are both affected by drought, and we suggest that the persistence of the water deficit (i.e., the drought time-scale) could be playing a key role in determining the sensitivity of land biomes to drought. We found that arid biomes respond to drought at short time-scales; that is, there is a rapid vegetation reaction as soon as water deficits below normal conditions occur. This may be due to the fact that plant species of arid regions have mechanisms allowing them to rapidly adapt to changing water availability. Humid biomes also respond to drought at short time-scales, but in this case the physiological mechanisms likely differ from those operating in arid biomes, as plants usually have a poor adaptability to water shortage. On the contrary, semiarid and subhumid biomes respond to drought at long time-scales, probably because plants are able to withstand water deficits, but they lack the rapid response of arid biomes to drought. These results are consistent among three vegetation parameters analyzed and across different land biomes, showing that the response of vegetation to drought depends on characteristic drought time-scales for each biome. Understanding the dominant time-scales at which drought most influences vegetation might help assessing the resistance and resilience of vegetation and improving our knowledge of vegetation vulnerability to climate change. [ABSTRACT FROM AUTHOR]

Shrubs growing at higher latitudes and altitudes are considered to be highly sensitive to climate warming. In particular, alpine shrubs are potential climate proxies for understanding responses of high-elevation treeless ecosystems to warmer conditions. However, little is known about how alpine shrub radial growth responds to climate variables, specifically temperature and precipitation. This lack of knowledge is particularly notable in the Himalayas where shrubs reach some of their worldwide uppermost limits. Herein, we investigated the climatic response of alpine juniper shrub (Juniperus indica) to climate in two areas (dry Manang valley; wet Everest valley) situated in the central Himalayas. In spite of different sites and elevations, the radial growth of juniper shrub is positively correlated with spring precipitation in both areas, and also with summer precipitation in the dry area. Juniper shrub shares common climatic responses with Himalayan treeline tree populations, whose growth dynamics are also controlled by moisture availability. Thus, radial growth of juniper in the central Himalayas may experience drought stress if climate warming leads to drier conditions. We conclude that alpine shrubs are crucial indicators of the responses of alpine ecosystem to global climate warming. [ABSTRACT FROM AUTHOR]

Aim: The spatial patterns of tree populations reflect multiple ecological processes. However, little is known on whether these patterns mediate responses to climate in marginal tree populations such as those forming alpine tree lines. We examined the influence of tree‐to‐tree interactions on the responsiveness of tree lines to climate warming. Location: Central Himalayas. Taxon: Betula utilis; Abies spectabilis. Methods: We analysed a network of 17 tree line sites located across the central Himalayas, encompassing a wide longitudinal gradient characterized by increasing precipitation eastwards. We quantified the changes in density and the spatial patterns of three 50‐year age classes of the two main tree species found at the tree line (Betula utilis and Abies spectabilis), and related them to reconstructed shifts in tree line elevation. Results: Younger trees showed clustering near the tree line, while older trees tended to show random spatial distribution. Clustering decreased as climate conditions ameliorated, that is, in the wetter eastern sites. Lower rate of tree line elevation change was observed at the sites with higher clustering intensity. Main Conclusions: Our study indicates that tree aggregation weakens tree line responsiveness to climate warming, and thus warming‐induced drought stress tends to lower tree line shift rates by enhancing clustering. It also highlights the complexity and contingency of site‐dependent tree line responses to climate. Hence, to advance our understanding of tree line processes, we should consider both direct and indirect influences of relevant biotic (tree‐to‐tree interactions) and abiotic (climate) drivers of tree line dynamics. [ABSTRACT FROM AUTHOR]

The alpine treeline is commonly regarded as being sensitive to climatic warming because regeneration and growth of trees at treeline generally are limited by low temperature. The alpine treelines of the Tibetan Plateau (TP) occur at the highest elevations (4,900 m above sea level) in the Northern Hemisphere. Ongoing climatic warming is expected to shift treelines upward. Studies of treeline dynamics at regional and local scales, however, have yielded conflicting results, indicating either unchanging treeline elevations or upward shifts. To reconcile this conflict, we reconstructed in detail a century of treeline structure and tree recruitment at sites along a climatic gradient of 4 °C and mean annual rainfall of 650 mm on the eastern TP. Species interactions interacted with effects of warming on treeline and could outweigh them. Densification of shrubs just above treeline inhibited tree establishment, and slowed upward movement of treelines on a time scale of decades. Interspecific interactions are major processes controlling treeline dynamics that may account for the absence of an upward shift at some TP treelines despite continued climatic warming. [ABSTRACT FROM AUTHOR]

Forests play a key role in the carbon balance of terrestrial ecosystems. One of the main uncertainties in global change predictions lies in how the spatiotemporal dynamics of forest productivity will be affected by climate warming. Here we show an increasing influence of climate on the spatial variability of tree growth during the last 120 y, ultimately leading to unprecedented temporal coherence in ring-width records over wide geographical scales (spatial synchrony). Synchrony in growth patterns across cold-constrained (central Siberia) and droughtconstrained (Spain) Eurasian conifer forests have peaked in the early 21st century at subcontinental scales (~1,000 km). Such enhanced synchrony is similar to that observed in trees co-occurring within a stand. In boreal forests, the combined effects of recent warming and increasing intensity of climate extremes are enhancing synchrony through an earlier start of wood formation and a stronger impact of year-to-year fluctuations of growing-season temperatures on growth. In Mediterranean forests, the impact of warming on synchrony is related mainly to an advanced onset of growth and the strengthening of droughtinduced growth limitations. Spatial patterns of enhanced synchrony represent early warning signals of climate change impacts on forest ecosystems at subcontinental scales. [ABSTRACT FROM AUTHOR]

Tocochromanols are the most abundant lipid-soluble antioxidants in plants. Among them, α-tocopherol (α-Toc) shows a particularly high sensitivity to environmental stressors and its content is used as a stress biomarker even in non-photosynthetic tissues. Nevertheless, the presence of tocochromanols has not been described yet in the xylem of woody plants, even when their functions regarding cell membrane protection and the transport of photoassimilates may be crucial in this tissue and despite its potential utility in dendrometabolomics. Considering all these, we aimed to determine the presence and distribution of tocochromanols in the xylem of woody plants, to examine their responsiveness to high temperature and to evaluate their potential as environmental bioindicators. The analysis of 29 phyllogenetically diverse species showed that α-Toc is the most abundant and frequent tocochromanol in the xylem and is ubiquitously present in all the studied species, with a concentration ranging from 0.5 to 39.3 μg g−1 of dry weight. α-Tocopherol appeared to be mainly located in the parenchyma rays and was found in both the sapwood and the heartwood, suggesting that it is present even in dead parenchyma cells. The levels of α-Toc in the xylem did not change in response to locally induced xylem heating, but responded positively to the 3-year moving average of annual precipitation. The present findings suggest that α-Toc may be linked to changes in climatic stress. This should enhance further research on the environmental controls of α-Toc variation in the xylem as a first step towards a deeper understanding of dendrometabolomics. [ABSTRACT FROM PUBLISHER]

In this study, the authors provide a global assessment of the performance of different drought indices for monitoring drought impacts on several hydrological, agricultural, and ecological response variables. For this purpose, they compare the performance of several drought indices [the standardized precipitation index (SPI); four versions of the Palmer drought severity index (PDSI); and the standardized precipitation evapotranspiration index (SPEI)] to predict changes in streamflow, soil moisture, forest growth, and crop yield. The authors found a superior capability of the SPEI and the SPI drought indices, which are calculated on different time scales than the Palmer indices to capture the drought impacts on the aforementioned hydrological, agricultural, and ecological variables. They detected small differences in the comparative performance of the SPI and the SPEI indices, but the SPEI was the drought index that best captured the responses of the assessed variables to drought in summer, the season in which more drought-related impacts are recorded and in which drought monitoring is critical. Hence, the SPEI shows improved capability to identify drought impacts as compared with the SPI. In conclusion, it seems reasonable to recommend the use of the SPEI if the responses of the variables of interest to drought are not known a priori. [ABSTRACT FROM AUTHOR]

• This study assesses how different phases of shoot growth underlie seasonal change in leaf and stem dry matter content (LDMC and SDMC, respectively) of 12 woody Mediterranean species. The relationship between LDMC and nonstructural carbohydrate (NSC) concentrations is also explored and the seasonal vs interspecies variability of LDMC compared. • LDMC, SDMC and shoot elongation rate (SER) were measured on a monthly basis for a minimum of 12 months. Bud growth rate (BGR) and NSC concentrations were also assessed in several of the study species. • LDMC and SDMC decreased during shoot elongation in spring and increased in summer, showing a significant negative correlation with SER, but were unrelated to BGR. Half of the species analysed showed a positive relationship between LDMC and NSC. Seasonal fluctuations of LDMC within species were higher than interspecies differences, and species ranking was significantly affected by the month of sampling, except during winter months. • Seasonal changes in LDMC and SDMC are mainly related to shoot elongation phenology, and NSC sink–source relationships between old and growing organs can explain this relationship in some species. Owing to the high seasonal variability in LDMC, it is recommended that samples for comparative purposes should be collected as close to the winter as possible. [ABSTRACT FROM AUTHOR]