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Knowledge regarding the growth of trees is essential to understanding their response to predicted warmer and drier climate scenarios. We used the annual rings of three Mexican pines (Pinus montezumae Lamb., Pinus oocarpa Schiede ex Schltdl., and Pinus monophylla Torr. & Frém) to explore their drought responses. Correlation analyses showed that hydroclimatic factors differentially impact tree species in terms of the intensity and temporality. The negative influence of the maximum temperature and positive effect of the precipitation on the growth indices were notable, with P. montezumae being the most responsive species, followed by P. oocarpa and P. monophylla. The climate–growth relationships were specific and driven by the differential hydrothermal conditions across the study areas. SPEI analyses indicated that P. monophylla is better able to tolerate drought than P. montezumae or P. oocarpa, especially in recent years. The lower resilience of P. montezumae and P. oocarpa could predispose them to a higher mortality risk if warming and drying rates increase. Our findings strengthen the understanding of the responses of tree growth to seasonal drought, which is critical considering the biogeographic shifts that will potentially be experienced by these forests in the future. This knowledge improves the understanding of young Mexican stands and could contribute to the design of management strategies in the face of predicted climatic variations. [ABSTRACT FROM AUTHOR]

Evapotranspiration demand has increased rapidly as temperatures have risen, affecting forest productivity. Consequently, carbon (C) uptake by forests is being modified; therefore, a more refined knowledge of the relationships between C capture and hydroclimate variability is required, particularly in drought-prone regions. In this study, we analyzed the relationships between climate and C capture as stemwood through the dendroecological analyses of radial growth in 15 conifer species distributed along a broad bioclimatic gradient in Mexico. The C content was calculated using densitometry data; correlations and mixed models were then used to determine the influence of climatic variables (precipitation, mean maximum and minimum temperatures, SPEI drought index) on tree growth. Each species showed specific responses to the climate with Taxodium mucronatum being the most responsive species. Both precipitation and maximum temperature best explained changes in C capture, with minimum temperature and SPEI playing secondary roles. The winter before the growth period was the most important season for C capture, particularly through positive responses to wet-cool conditions. However, the climatic influences of the current fall and summer were also notable. Seasonal climatic influences have implications for C uptake and forest productivity in the face of the severe droughts that repeatedly affect the study region. [ABSTRACT FROM AUTHOR]

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]

Climate extremes such as cold spells are becoming more frequent as climate variability increases. However, few studies have evaluated the impacts of winter cold spells on forest cover, tree growth and leaf and sapwood non-structural carbohydrate (NSC) concentrations. We analyzed changes in tree cover using remote sensing data and compared the radial growth of coexisting and defoliated Pinus halepensis trees and non-defoliated P. halepensis and Pinus pinaster trees. We also compared NSC concentrations in leaves and sapwood of defoliated and non-defoliated P. halepensis and Quercus ilex trees. In January 2021, a rapid drop in temperatures led to minimum values (−21.3 °C) in eastern Spain and triggered canopy defoliation in several planted (P. halepensis) and native (Q. ilex) tree species. The cold spell led to a decrease in forest cover in the most defoliated stands and reduced radial growth of defoliated P. halepensis and sapwood NSC concentrations in P. halepensis and Q. ilex, particularly starch. Prior to the cold spell, defoliated P. halepensis trees significantly (p < 0.05) grew more (2.73 ± 1.70 mm) in response to wetter winter conditions than non-defoliated P. halepensis (2.29 ± 1.08 mm) and P. pinaster (1.39 mm) trees. Those P. halepensis individuals which grew faster at a young age were less resilient to the winter cold spell in later years. The study stands showed a high recovery capacity after the cold spell, but the Mediterranean drought-avoiding P. halepensis was the most affected species. [ABSTRACT FROM AUTHOR]

Intensive livestock management impacts forest and trees in different ways. Pig manure is a major source of nitrogen (N) pollution of surface and ground waters in some European regions such as north-eastern Spain, but it is understudied how manure application impacts agroforestry systems. How pig manure affects tree radial growth and the N cycle was assessed by measuring N concentrations in soil, leaves and wood and δ15N in tree-ring wood in two tree species widely planted for agricultural (Prunus dulcis) and reforestation (Pinus halepensis) purposes in the study area. Soil physicochemical characteristics and the biomass and structure of major soil microbial groups were also measured. Trees irrigated with pig effluent (manure application) and control trees not subjected to manure application were compared. Soil N, phosphorus (P) and potassium (K) concentrations of fertilized trees increased, but soil microbiota biomass decreased. Similar growth between fertilized and non-fertilized pine trees was found, but lower growth in fertilized almond trees was observed. Leaf N concentrations decreased but δ15N wood increased in trees subjected to pig manure application. Pig manure application alters the N cycling in the soil and within trees. [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]

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. [ABSTRACT FROM AUTHOR]

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. Plain Language Summary: The occurrence patterns of seasonal extreme drought and wetness events are dramatically shifting with climate warming. However, how will different seasonal extreme climate regimes affect the bioclimatic sensitivity of tree growth remains poorly understood. In this study, we investigated the sensitivity of tree growth to different seasonal climate factors and preceding tree growth conditions during 1951–2013 under different seasonal extreme drought/wetness regimes, using 1,032 tree ring chronologies covering 121 gymnosperm and angiosperm species. We found the magnitude in tree growth reduction caused by seasonal extreme drought events is much larger than that in tree growth stimulation by seasonal extreme wetness events in arid and temperate regions. Tree growth in arid and temperate dry regions is more negatively impacted by extreme drought events in pre‐growing‐seasons (PGSs) than in growing‐seasons. We further found that angiosperms are more sensitive to PGS water availability, while gymnosperms are more sensitive to legacy effects of the preceding tree growth conditions in temperate dry and humid regions. These findings highlight divergent bioclimatic legacy effects on tree growth under different seasonal extreme climate regimes, and provide valuable insights into the future trajectories of forest growth across diverse ecoregions and functional groups in a more extreme climate. Key Points: Negative asymmetry in tree growth was observed under regimes with seasonal extreme droughtsTree growth in arid and temperate dry regions is more vulnerable to extreme droughts in pre‐growing‐seasons than in growing‐seasonsAngiosperms are more sensitive to pre‐growing‐season water availability, and gymnosperms to legacy effects of the preceding tree growth [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]

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. [ABSTRACT FROM AUTHOR]

The Tibetan Plateau hosts a continuous distribution of alpine treelines from the Qilian Mountains to the Hengduan Mountains and the Himalaya Mountains. However, not much is known about the broadscale alpine treeline dynamics and their responses to climate warming across the Tibetan Plateau. Herein, we collected a total of 59 treeline sites across different forest regions of the Tibetan Plateau and the related field data (i.e., upward advance magnitude, tree recruitment and height growth), expansion potential (i.e., elevational difference between the current treeline and the tree species line (EP)) and vegetation TI (an index of species interactions) from the published references. Site characteristics (e.g., elevation, slope and aspect) and the related environmental factors were used to analyze the relationships between treeline shifts and environmental variables. Despite increases in the recruitment and growth of trees at most treeline sites, alpine treeline positions showed heterogeneous responses to climate warming. Most treelines advanced over the last century, while some treelines showed long-term stability. EP was significantly and positively linked to the summer warming rate and treeline shifts, suggesting that the position of current tree species line is of crucial importance in evaluating treeline dynamics under climate change. In addition, warming-induced treeline advances were modulated by plant–plant interactions. Overall, this study highlighted the heterogeneous responses of regional-scale alpine treelines to climate warming on the Tibetan Plateau. [ABSTRACT FROM AUTHOR]

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. [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]

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]

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]

Abstract: The causal factors and effects of forest declines are not well understood in temperate conifer forests. Most studies have focused on climatic and environmental stressors and have obviated the potential role of historical forest management as a predisposing factor of decline. Here, we assess if the recent silver fir (Abies alba) decline observed in the Spanish Pyrenees was predisposed by historical logging and incited by warming-induced drought stress. We analysed a dataset of environmental, structural, and historical variables at the tree and stand level including 32 sites with contrasting degrees of defoliation distributed over 5600km2. We followed a dendroecological approach to reconstruct historical logging and to infer the effects of warming-induced drought stress on growth. The silver fir decline was more severe and widespread in western low-elevation mixed forests dominated by trees of small size and slow growth. These sites were subject to higher water deficits than eastern sites, where late-summer rainfall as the key climatic variable controlling silver fir growth was higher. Declining sites showed more frequent growth releases induced by historical logging than non-declining sites. Historical logging and warming-induced drought acted as long-term predisposing and short-term inciting factors of silver fir decline in the Pyrenees, respectively. We suggest that biomass increases caused by past intense logging affected the vulnerability of silver fir against late-summer water deficit. Future research in declining temperate conifer forests should consider the interacting role of predisposing historical management and inducing climatic stressors such as droughts. [Copyright &y& Elsevier]

An understanding of the interactions between climate change and forest structure on tree growth are needed for decision making in forest conservation and management. In this paper, we investigated the relative contribution of tree features and stand structure on Atlas cedar (Cedrus atlantica) radial growth in forests that have experienced heavy grazing and logging in the past. Dendrochronological methods were applied to quantify patterns in basal-area increment and drought sensitivity of Atlas cedar in the Middle Atlas, northern Morocco. We estimated the tree-to-tree competition intensity and quantified the structure in Atlas cedar stands with contrasting tree density, age, and decline symptoms. The relative contribution of tree age and size and stand structure to Atlas cedar growth decline was estimated by variance partitioning using partial-redundancy analyses. Recurrent drought events and temperature increases have been identified from local climate records since the 1970s. We detected consistent growth declines and increased drought sensitivity in Atlas cedar across all sites since the early 1980s. Specifically, we determined that previous growth rates and tree age were the strongest tree features, while Quercus rotundifolia basal area was the strongest stand structure measure related to Atlas cedar decline. As a result, we suggest that Atlas cedar forests that have experienced severe drought in combination with grazing and logging may be in the process of shifting dominance toward more drought-tolerant species such as Q. rotundifolia. [ABSTRACT FROM AUTHOR]

Windstorms are forest disturbances which generate canopy gaps. However, their effects on Mediterranean forests are understudied. To fill that research gap, changes in tree, cover, growth and soil features in Pinus halepensis and Pinus sylvestris plantations affected by windthrows were quantified. In each plantation, trees and soils in closed-canopy stands and gaps created by the windthrow were sampled. Changes in tree cover and radial growth were assessed by using the Normalized Difference Vegetation Index (NDVI) and dendrochronology, respectively. Soil features including texture, nutrients concentration and soil microbial community structure were also analyzed. Windthrows reduced tree cover and enhanced growth, particularly in the P. halepensis site, which was probably more severely impacted. Soil characteristics were also more altered by the windthrow in this site: the clay percentage increased in gaps, whereas K and Mg concentrations decreased. The biomass of Gram positive bacteria and actinomycetes increased in gaps, but the biomass of Gram negative bacteria and fungi decreased. Soil gaps became less fertile and dominated by bacteria after the windthrow in the P. halepensis site. We emphasize the relevance of considering post-disturbance time recovery and disturbance intensity to assess forest resilience within a multi-scale approach. [ABSTRACT FROM AUTHOR]

Background and Objectives—Coexisting tree and shrub species will have to withstand more arid conditions as temperatures keep rising in the Mediterranean Basin. However, we still lack reliable assessments on how climate and drought affect the radial growth of tree and shrub species at intra- and interannual time scales under semi-arid Mediterranean conditions. Materials and Methods—We investigated the growth responses to climate of four co-occurring gymnosperms inhabiting semi-arid Mediterranean sites in northeastern Spain: two tree species (Aleppo pine, Pinus halepensis Mill.; Spanish juniper, Juniperus thurifera L.) and two shrubs (Phoenicean juniper, Juniperus phoenicea L.; Ephedra nebrodensis Tineo ex Guss.). First, we quantified the intra-annual radial-growth rates of the four species by periodically sampling wood samples during one growing season. Second, we quantified the climate–growth relationships at an interannual scale at two sites with different soil water availability by using dendrochronology. Third, we simulated growth responses to temperature and soil moisture using the forward, process-based Vaganov‒Shashkin (VS-Lite) growth model to disentangle the main climatic drivers of growth. Results—The growth of all species peaked in spring to early summer (May–June). The pine and junipers grew after the dry summer, i.e., they showed a bimodal growth pattern. Prior wet winter conditions leading to high soil moisture before cambium reactivation in spring enhanced the growth of P. halepensis at dry sites, whereas the growth of both junipers and Ephedra depended more on high spring–summer soil moisture. The VS-Lite model identified these different influences of soil moisture on growth in tree and shrub species. Conclusions—Our approach (i) revealed contrasting growth dynamics of co-existing tree and shrub species under semi-arid Mediterranean conditions and (ii) provided novel insights on different responses as a function of growth habits in similar drought-prone regions. [ABSTRACT FROM AUTHOR]

The increased frequency and intensity of warming-induced droughts have triggered dieback episodes affecting many forest types and tree species worldwide. Tree plantations are not exempt as they can be more vulnerable to drought than natural forests because of their lower structural and genetic diversity. Therefore, disentangling the physiological mechanisms leading to growth decline and tree mortality can provide tools to adapt forest management to climate change. In this study, we investigated a Pinus nigra Arn. plantation situated in northern Spain, in which some trees showed canopy dieback and radial-growth decline. We analyzed how radial growth and its responses to drought events differed between non-declining (ND) and declining (D) trees showing low and high canopy defoliation, respectively, in combination with carbon (δ13C) and oxygen (δ18O) isotope ratios in tree rings. The radial growth of P. nigra was constrained by water availability during the growing season and the previous autumn. The radial growth of D trees showed higher sensitivity to drought than ND trees. This fact is in accordance with the lower drought resilience and negative growth trends observed in D trees. Both tree classes differed in their growth from 2012 onwards, with D trees showing a reduced growth compared to ND trees. The positive δ13C-δ18O relationship together with the uncoupling between growth and intrinsic water-use efficiency suggest that D trees have less tight stomatal regulation than ND trees, which could involve a high risk of xylem embolism in the former class. Our results suggest that different water use strategies between coexisting ND and D trees were behind the differences in growth patterns and point to hydraulic failure as a possible mechanism triggering dieback and growth decline. [ABSTRACT FROM AUTHOR]