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We still lack information on the long-term growth responses to climate of relict tree populations, which often persist in topoclimatic refugia. To fill that research gap, we studied three relict cork oak (Quercus suber) populations located in northern Spain using dendrochronology. The sites were subjected to humid (Zarautz), continental (Bozoó) and xeric (Sestrica) climate conditions. Cool–wet conditions during the current spring enhanced growth in Bozoó and Sestrica, whereas wet conditions in the previous October enhanced growth in Zarautz. In this site, growth also increased in response to dry conditions in the prior winter linked to high North Atlantic Oscillation indices. Correlations between the precipitation summed from the previous September to the current May peaked at the driest site (Sestrica). The strongest growth responses to drought severity were also found at this site, where growth negatively responded to 9-month early-summer droughts, followed by the continental Bozoó site, where growth was constrained by 1-month July droughts. Growth declined in response to 6-month January droughts in the wettest site (Zarautz), where cork oak was vulnerable to previous late-summer to autumn drought stress. Despite warmer and drier spring conditions that would negatively impact cork oak at the Bozoó and Sestrica sites, trees from these populations could tolerate further aridity. [ABSTRACT FROM AUTHOR]

South rear-edge populations of widely distributed temperate and boreal tree species such as birches (Betula pubescens and Betula pendula) are considered particularly vulnerable to climate warming, and at the same time, they constitute genetic reservoirs of drought-adapted ecotypes. Here, we compared radial growth patterns and responses to climate, river, or reservoir flows and a drought index of rear-edge (southernmost) populations (Toledo Mountains, central-southern Spain) with populations located in northern Spain of B. pubescens and B. pendula. Then, we performed a comparative analysis across Europe of B. pendula populations. The main climatic constraint of birch growth was a high summer water deficit, although the effect of local hydrological conditions was particularly important in rear-edge populations. We found declining growth trends in rear-edge stands dating from the early 21st century, related to decreasing water availability and increasing aridity. Our results also suggested distinct growth patterns and climate-growth associations of B. pendula across Europe that show how populations further south and in warmer locations were more sensitive to drought stress. Drought-induced growth decline can be exacerbated by local human land uses, leading to reduced river inflow, thus endangering birch populations at their southern distribution limit. Protection of threatened rear-edge birch populations requires adequate management of local water resources. [ABSTRACT FROM AUTHOR]

Forests around the world are facing climate change. Increased drought stress and severe heat waves in recent decades have negatively impacted on forest health, making them more vulnerable and prone to dieback and mortality phenomena. Although the term vulnerability is used to indicate an increased susceptibility of forests to climate change with a worsening of their vigour status that can compromise their ability to respond to further climate extreme events, there are still uncertainties on how to evaluate it. Indeed, evaluation of forest vulnerability is complex both because of some critical issues in the estimation methods used and because of the multiple factors influencing the response of forests to ongoing climate change. A way to assess the vulnerability to environmental stresses is by combining remote sensing and dendroecological data. However, these two approaches entail multiple uncertainties, including growth/photosynthetic relationships, carbon allocation dynamics, biases of tree-ring data and noisy remote sensing data, which require further clarification for proper monitoring of pre- and post-drought forest trajectories. Our review aims to create an overview of the current literature and knowledge to understand the critical issues, needs and possible solutions that forest vulnerability research is addressing. We focus on Mediterranean forests located in a climate warming hotspot and showing a high vulnerability to increased aridification. [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]

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]

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]

Tree plantations have been proposed as suitable carbon sinks to mitigate climate change. Drought may reduce their carbon uptake, increasing their vulnerability to stress and affecting their growth recovery and resilience. We investigated the recent growth rates and responses to the climate and drought in eight Atlas cedar (Cedrus atlantica) plantations located along a wide climate gradient from wetter sites in south-eastern France and north Spain to dry sites in south-eastern Spain. The cedar growth increased in response to the elevated precipitation from the prior winter to the current summer, but the influence of winter precipitation on growth gained importance in the driest sites. The growth responsiveness to climate and drought peaked in those dry sites, but the growth resilience did not show a similar gradient. The Atlas cedar growth was driven by the total precipitation during the hydrological year and this association strengthened from the 1980s onwards, a pattern related to the winter North Atlantic Oscillation (NAO). High winter NAO indices and drier conditions were associated with lower growth. At the individual level, growth resilience was related to tree age, while growth recovery and year-to-year growth variability covaried. Plantations' resilience to drought depends on both climate and tree-level features. [ABSTRACT FROM AUTHOR]

Drought constrains tree growth in regions with seasonal water deficit where growth decline can lead to tree death. This has been observed in regions such as the western Mediterranean Basin, which is a climate-warming hotspot. However, we lack information on intra- and inter-specific comparisons of growth rates and responses to water shortage in these hotspots, considering tree species with different drought tolerance. We sampled several sites located in north-eastern Spain showing dieback and high mortality rates of three pine species (Pinus sylvestris, Pinus pinaster, Pinus halepensis). We dated death years and reconstructed the basal area increment of coexisting living and recently dead trees using tree ring data. Then, we calculated bootstrapped Pearson correlations between a drought index and growth. Finally, we used linear mixed-effects models to determine differences in growth trends and the response to drought of living and dead trees. Mortality in P. sylvestris and P. pinaster peaked in response to the 2012 and 2017 droughts, respectively, and in sites located near the species' xeric distribution limits. In P. halepensis, tree deaths occurred most years. Dead trees showed lower growth rates than living trees in five out of six sites. There was a strong growth drop after the 1980s when climate shifted towards warmer and drier conditions. Tree growth responded positively to wet climate conditions, particularly in the case of living trees. Accordingly, growth divergence between living and dead trees during dry periods reflected cumulative drought impacts on trees. If aridification continues, tree drought mortality would increase, particularly in xeric distribution limits of tree species. [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]

Drought stress has induced dieback episodes affecting many forest types and tree species worldwide. However, there is scarce information regarding drought-triggered growth decline and canopy dieback in Mediterranean deciduous oaks. These species face summer drought but have to form new foliage every spring which can make them vulnerable to hotter and drier conditions during that season. Here, we investigated two stands dominated by Quercus frainetto Ten. and Quercus canariensis Willd. and situated in southern Italy and Spain, respectively, showing drought-induced dieback since the 2000s. We analyzed how radial growth and its responses to climate differed between non-declining (ND) and declining (D) trees, showing different crown defoliation and coexisting in each stand by: (i) characterizing growth variability and its responsiveness to climate and drought through time, and (ii) simulating growth responses to soil moisture and temperature thresholds using the Vaganov–Shashkin VS-lite model. Our results show how growth responsiveness to climate and drought was higher in D trees for both oak species. Growth has become increasingly limited by warmer-drier climate and decreasing soil moisture availability since the 1990s. These conditions preceded growth drops in D trees indicating they were more vulnerable to warming and aridification trends. Extremely warm and dry conditions during the early growing season trigger dieback. Changes in the seasonal timing of water limitations caused contrasting effects on long-term growth trends of D trees after the 1980s in Q. frainetto and during the 1990s in Q. canariensis. Using growth models allows identifying early-warning signals of vulnerability, which can be compared with shifts in the growth responses to warmer and drier conditions. Our approach facilitates establishing drought-vulnerability thresholds by combining growth models with field records of dieback. [ABSTRACT FROM AUTHOR]

Forest dieback and mortality events induced by drought stress are widely reported. However, few studies have jointly examined the role played by drought on growth and mortality in tree species inhabiting floodplain forests. Here, we focused on mortality events occurring since the early 2000s on large areas in a floodplain forest located within the Ticino regional park in Northwest Italy, where affected native (pedunculate oak, Quercus robur L.) and introduced tree species (black locust, Robinia pseudoacacia L.) coexist. We related growth with climate data and drought severity to discern if these species were similarly affected by drought. Then, we: (i) evaluated the presence of pathogens of the genus Phytophthora in recently dead oak trees since this was the most affected species and pathogens are often associated with oak decline cases; and (ii) compared xylem vessel diameter and tree-ring C isotope discrimination (δ13C) to highlight differences in water-use strategies between living and dead trees in both species. The radial growth of living and dead trees started diverging in the 1970s, although only after warm-drought periods occurred during 1990s did this divergence become significant. Growth of trees that died responded more negatively to drought than in the case of living trees. Moreover, trees that died formed smaller xylem vessels in the past than living trees and also showed more negative δ13C values in both tree species, indicating a higher intrinsic water-use efficiency in living than in dead trees. The pathogen Phytophthora cinnamomi Rands was only detected in one recently dead tree, suggesting that it is unlikely that dead oaks were predisposed to drought damage by the pathogen. We conclude that a climate shift from wet to warm-dry summer conditions in the early 1990s triggered forest dieback and induced mortality in both tree species. Temperate floodplain forests are susceptible to drought-induced dieback. The drought-sensitivity of both species could lead to successional shifts driven by a reduction of N inputs through N-fixing by black locust and the replacement of oak by drought-tolerant species. [ABSTRACT FROM AUTHOR]