Drought legacies in mixed Mediterranean forests: Analysing the effects of structural overshoot, functional traits and site factors.

Previous favorable climate conditions stimulate tree growth making some forests more vulnerable to hotter droughts. This so-called structural overshoot may contribute to forest dieback, but there is little evidence on its relative importance depending on site conditions and tree species because of limited field data. Here, we analyzed remote sensing (NDVI) and tree-ring width data to evaluate the impacts of the 2017 drought on canopy cover and growth in mixed Mediterranean forests (Fraxinus ornus, Quercus pubescens, Acer monspessulanum, Pinus pinaster) located in southern Italy. Legacy effects were assessed by calculating differences between observed and predicted basal area increment (BAI). Overall, the growth response of the study stands to the 2017 drought was contingent on site conditions and species characteristics. Most sites presented BAI and canopy cover reductions during the drought. Growth decline was followed by a quick recovery and positive legacy effects, particularly in the case of F. ornus. However, we found negative drought legacies in some species (e.g., Q. pubescens, A. monspessulanum) and sites. In those sites showing negative legacies, high growth rates prior to drought in response to previous wet winter-spring conditions may have predisposed trees to drought damage. Vice versa, the positive drought legacy found in some F. ornus site was linked to post-drought growth release due to Q. pubescens dieback and mortality. Therefore, we found evidences of structural drought overshoot, but it was restricted to specific sites and species. Our findings highlight the importance of considering site settings such as stand composition, pre-drought conditions and different tree species when studying structural overshoot. Droughts contribute to modify the composition and dynamics in mixed forests.

Revealing legacy effects of extreme droughts on tree growth of oaks across the Northern Hemisphere.

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

Growth of tree (Pinus sylvestris) and shrub (Amelanchier ovalis) species is constrained by drought with higher shrub sensitivity in dry sites.

We lack understanding of how variable is radial growth of coexisting tree and shrub species, and how growth is constrained by drought depending on site aridity. Here, we compared the radial growth of two widespread and coexisting species, a winter deciduous shrub (Amelanchier ovalis Medik.) and an evergreen conifer tree (Pinus sylvestris L.). We sampled four sites in Northeastern Spain subjected to different aridity levels and used dendrochronological methods to quantify growth patterns and responses to climate variables. The growth of the two species varied between regions, being lower in the driest sites. The first-order autocorrelation (growth persistence) was higher in more mesic sites but without clear differences between species. Tree and shrub growth negatively responded to elevated summer temperatures and positively to spring-summer precipitation and wet conditions. However, negative growth responses of the shrub to drought were only observed in the two driest sites in contrast to widespread responses of the tree. Abrupt growth reductions were common in the drier sites, but resilience indices show that the two species rapidly recovered pre-drought growth levels. The lower growth synchrony of the shrub as compared to the tree can be due to the multistemmed architecture, fast growth and low stature of the shrub. Besides, the high dependency of the shrub growth on summer rainfall can explain why drought limitations were only apparent in the two driest sites. In any case, results point out to the dendrochronological potential of shrubs, which is particularly relevant giving its ability to inhabit woodlands and treeless regions under harsh climatic conditions. Nevertheless, further research is required to elucidate the capacity of shrub species to tolerate drought, as well as to understand how shrubs thrive in water- and cold-limited environments.

Climatic drivers of cork growth depend on site aridity.

Cork is one of the main non-timber forest products in the world. Most of its production is concentrated in the Iberian Peninsula, a climate change hotspot. Climate warming may lead to increased aridification and reduce cork production in that region. However, we still lack assessments of climate-cork relationships across ample geographical and climatic gradients explicitly considering site aridity. We quantified cork growth by measuring cork ring width and related it to climate variables and a drought index using dendrochronology. Four cork oak (Quercus suber) forests located from north eastern Spain to south western Morocco (31.5-41.5° N) and subjected to different aridity levels were sampled. Warm conditions in spring to early summer, when cork is formed, reduced cork width, whereas high precipitation in winter and spring enhanced it. The response of cork to increased water availability in summer peaked (r = 0.89, p = 0.00002) in the most arid and continental site considering 14-month long droughts. A severe drought caused a disproportionate loss of cork production in this site, where for every five-fold decrease in the drought index, the cork-width index declined by a factor of thirteen. Therefore, site aridity determines the responses of cork growth to the soil water availability resulting from accumulated precipitation during winter and spring previous to cork growth and until summer. In general, this cumulative water balance, which is very dependent on temperature and evapotranspiration rate, is critical for cork production, especially in continental, dry sites. The precipitation during the hydrological year can be used as a proxy of cork production in similar sites. Assessments of climate-cork relationships in the western Mediterranean basin could be used as analogues to forecast the impacts of aridification on future cork production.

Historical disconnection from floodplain alters riparian forest composition, tree growth and deadwood amount.

Riparian forests are among the most dynamic but threatened terrestrial ecosystems. Their dynamism and conservation depend on historical changes in river geomorphology, which can be evaluated through changes in channel sinuosity. However, we lack long-term assessments on sinuosity and how they impact riparian forest composition, tree growth and deadwood amount. To fill this research gap, we reconstructed river sinuosity in 14 sites across the middle Ebro basin, north-eastern Spain, using historical aerial photographs taken in 1927, 1956, 1998-2003 and 2014-2015. Relationships between sinuosity, stand composition and deadwood amount and decay degree were calculated. We also reconstructed radial growth of the major tree species (Populus alba, Populus nigra, Fraxinus angustifolia, Salix alba and Ulmus minor) in two sites to evaluate how coupled it was with changes in river flow after dam building. From 1927 to 2015, sinuosity decreased passing from 1.39 to 1.20. The river dynamics were altered in the 1950s and 1960s after dam and dyke building. Sites with high sinuosity values in 1956 corresponded to mature stands with large P. nigra individuals. Sinuosity was negatively related to F. angustifolia (rs = -0.83, p < 0.001) and P. alba (rs = -0.64, p = 0.02) abundance, whereas sites dominated by P. alba and U. minor presented abundant decayed deadwood. A loss of sinuosity and a contraction of the riverbank gradient increased disconnection of active channel from floodplain, with a mixing of more (e.g., P. nigra) and less phreatophytic species (e.g., U. minor). River flow diversion reduced growth and increased the tree-to-tree P. alba growth coherence. Hydrological droughts contributed to growth decline and dieback of U. minor, which is sensitive to spring river flow. Conservation and restoration of riparian forests must consider historical changes in river geomorphology related to human activities.

Tree-ring and remote sensing analyses uncover the role played by elevation on European beech sensitivity to late spring frost.

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

Weak genetic differentiation but strong climate-induced selective pressure toward the rear edge of mountain pine in north-eastern Spain.

Local differentiation at distribution limits may influence species' adaptive capacity to environmental changes. However, drivers, such gene flow and local selection, are still poorly understood. We focus on the role played by range limits in mountain forests to test the hypothesis that relict tree populations are subjected to genetic differentiation and local adaptation. Two alpine treelines of mountain pine (Pinus uncinata Ram. ex DC) were investigated in the Spanish Pyrenees. Further, an isolated relict population forming the species' southernmost distribution limit in north-eastern Spain was also investigated. Using genotyping by sequencing, a genetic matrix conformed by single nucleotide polymorphisms (SNPs) was obtained. This matrix was used to perform genotype-environment and genotype-phenotype associations, as well as to model risk of non-adaptedness. Increasing climate seasonality appears as an essential element in the interpretation of SNPs subjected to selective pressures. Genetic differentiations were overall weak. The differences in leaf mass area and radial growth rate, as well as the identification of several SNPs subjected to selective pressures, exceeded neutral predictions of differentiation among populations. Despite genetic drift might prevail in the isolated population, the Fst values (0.060 and 0.066) showed a moderate genetic drift and Nm values (3.939 and 3.555) indicate the presence of gene flow between the relict population and both treelines. Nonetheless, the SNPs subjected to selection pressures provide evidences of possible selection in treeline ecotones. Persistence in range boundaries seems to involve several selective pressures in species' traits, which were significantly related to enhanced drought seasonality at the limit of P. uncinata distribution range. We conclude that gene flow is unlikely to constrain adaptation in the P. uncinata rear edge, although this species shows vulnerability to future climate change scenarios involving warmer and drier conditions.

The role of nutritional impairment in carbon-water balance of silver fir drought-induced dieback.

Rear-edge populations at the xeric distribution limit of tree species are particularly vulnerable to forest dieback triggered by drought. This is the case of silver fir (Abies alba) forests located in Southwestern Europe. While silver fir drought-induced dieback patterns have been previously explored, information on the role played by nutritional impairment is lacking despite its potential interactions with tree carbon-water balances. We performed a comparative analysis of radial growth, intrinsic water-use efficiency (iWUE), oxygen isotopes (δ18 O) and nutrient concentrations in leaves of declining (DD) and non-declining (ND) trees in silver fir in four forests in the Spanish Pyrenees. We also evaluated the relationships among dieback predisposition, intraspecific trait variation (wood density and leaf traits) and rhizosphere soil physical-chemical properties. The onset of growth decline in DD trees occurred more than two decades ago, and they subsequently showed low growth resilience against droughts. The DD trees presented consistently lower foliar concentrations of nutrients such as P, K, Cu and Ni than ND trees. The strong effects of foliar nutrient status on growth resilience indices support the key role played by mineral nutrition in tree functioning and growth before, during and after drought. In contrast, variability in wood density and leaf morphological traits, as well as soil properties, showed weak relationships with tree nutritional status and drought performance. At the low elevation, warmer sites, DD trees showed stronger climate-growth relationships and lower δ18 O than ND trees. The uncoupling between iWUE and δ18 O, together with the positive correlations between P and K leaf concentrations and δ18 O, point to deeper soil/bedrock water sources and vertical decoupling between nutrient and water uptake in DD trees. This study provides novel insights into the mechanisms driving silver fir dieback and highlights the need to incorporate tree nutrition into forest dieback studies.

Las poblaciones del límite xérico de distribución de las especies de árboles son particularmente vulnerables al decaimiento forestal inducido por sequía. Este es el caso de los bosques de abeto (Abies alba) situados en el suroeste de Europa. Si bien los patrones de decaimiento provocado por sequía del abeto se han explorado previamente, falta información sobre el papel que desempeña el deterioro nutricional a pesar de sus interacciones potenciales con los balances de agua y carbono de los árboles. En este estudio, hemos realizado un análisis comparativo del crecimiento radial, la eficiencia intrínseca del uso del agua (iWUE), los isótopos de oxígeno (δ18O) y las concentraciones de nutrientes en hojas de árboles decaídos (DD) y no decaídos (ND) en cuatro abetares de los Pirineos españoles. También evaluamos las relaciones entre la predisposición al decaimiento, la variación de rasgos intraespecíficos (densidad de la madera y rasgos de las hojas) y las propiedades físico-químicas de la rizosfera. El inicio de la disminución del crecimiento en los árboles DD ocurrió hace más de dos décadas y posteriormente mostraron una baja resiliencia de crecimiento frente a las sequías. Los árboles DD presentaron concentraciones foliares consistentemente más bajas de nutrientes como P, K, Cu y Ni que los árboles ND. Los fuertes efectos del estado de los nutrientes foliares en los índices de resiliencia del crecimiento respaldan el papel clave que desempeña la nutrición mineral en el funcionamiento y el crecimiento de los árboles antes, durante y después de la sequía. En contraste, la variabilidad en la densidad de la madera y los rasgos morfológicos de las hojas, así como las propiedades del suelo, mostraron una relación débil con el estado nutricional de los árboles y la respuesta del crecimiento a la sequía. En los sitios más cálidos y de baja elevación, los árboles DD mostraron relaciones clima-crecimiento más fuertes y un δ18 O más bajo que los árboles ND. El desacoplamiento entre iWUE y δ18 O, junto con las correlaciones positivas entre las concentraciones foliares de P y K y δ18 O, apuntan a fuentes de agua más profundas del suelo/lecho rocoso y un desacoplamiento vertical entre la absorción de nutrientes y agua en los árboles DD. Este estudio proporciona información novedosa sobre los mecanismos que impulsan el decaimiento del abeto y destaca la necesidad de incorporar la nutrición de los árboles en los estudios de muerte regresiva del bosque.

Declines in canopy greenness and tree growth are caused by combined climate extremes during drought-induced dieback.

Several dieback episodes triggered by droughts are revealing the high vulnerability of Mediterranean forests, manifested as declines in growth, increased defoliation, and rising mortality rates. Understanding forest responses to such climate extreme events is of high priority for predicting their future vegetation dynamics. We examined how remotely sensed measures of vegetation activity (NDVI, Normalized Difference Vegetation Index) and radial growth (BAI, basal area increment) responded to climate extreme events. We considered tree (Pinus sylvestris, Quercus pubescens, Quercus frainetto) and shrub (Juniperus phoenicea) populations from Italy and Spain showing recent dieback phenomena. Two components of drought, namely elevated atmospheric demand (VPD, vapor pressure deficit) and low soil moisture were analyzed in nearby stands showing or not showing dieback symptoms. Dieback stands exhibited lower NDVI values than non-dieback stands. NDVI and BAI were positively related in all sites except for the dieback stand of Q. frainetto that was negatively related. Such NDVI-BAI linkages were related to specific time windows, which could be useful for identifying when climatic conditions have the greatest influence on vegetation. Growth decline occurred in response to increasing VPD, but responses differed among species. J. phoenicea was the most negatively impacted by higher VPD, whereas oaks responded to soil moisture. A high VPD was related to stronger growth reduction in dieback P. sylvestris trees regardless of soil moisture changes. We highlighted that coupling between proxies of forest productivity (NDVI, BAI) allows better understanding and forecasting of drought-induced dieback phenomena in forests and shrublands. Scaling up from tree to stand levels might be feasible when using the maximum growing season NDVI, which can be applied for retrospective modeling of the impact of drought stress on forest productivity and tree growth.

Mediterranean old-growth forests exhibit resistance to climate warming.

Old-growth mountain forests represent an ideal setting for studying long-term impacts of climate change. We studied the few remnants of old-growth forests located within the Pollino massif (southern Italy) to evaluate how the growth of conspecific young and old trees responded to climate change. We investigated two conifer species (Abies alba and Pinus leucodermis) and two hardwood species (Fagus sylvatica and Quercus cerris). We sampled one stand per species along an altitudinal gradient, ranging from a drought-limited low-elevation hardwood forest to a cold-limited subalpine pine forest. We used a dendrochronological approach to characterize the long-term growth dynamics of old (age > 120 years) versus young (age < 120 years) trees. Younger trees grew faster than their older conspecifics during their juvenile stage, regardless of species. Linear mixed effect models were used to quantify recent growth trends (1950-2015) and responses to climate for old and young trees. Climate sensitivity, expressed as radial growth responses to climate during the last three decades, partially differed between species because high spring temperatures enhanced conifer growth, whereas F. sylvatica growth was negatively affected by warmer spring conditions. Furthermore, tree growth was negatively impacted by summer drought in all species. Climate sensitivity differed between young and old trees, with younger trees tending to be more sensitive in P. leucodermis and A. alba, whereas older F. sylvatica trees were more sensitive. In low-elevation Q. cerris stands, limitation of growth due to drought was not related to tree age, suggesting symmetric water competition. We found evidence for a fast-growth trend in young individuals compared with that in their older conspecifics. Notably, old trees tended to have relatively stable growth rates, showing remarkable resistance to climate warming. These responses to climate change should be recognized when forecasting the future dynamics of old-growth forests for their sustainable management.

Modeling Climate Impacts on Tree Growth to Assess Tree Vulnerability to Drought During Forest Dieback.

Forest dieback because of drought is a global phenomenon threatening particular tree populations. Particularly vulnerable stands are usually located in climatically stressing locations such as xeric sites subjected to seasonal drought. These tree populations show a pronounced loss of vitality, growth decline, and high mortality in response to extreme climate events such as heat waves and droughts. However, dieback events do not uniformly affect stands, with some trees showing higher symptoms of drought vulnerability than other neighboring conspecifics. In this study, we investigated if trees showing different vulnerabilities to dieback showed lower growth rates (Grs) and higher sensitivities to the climate in the past using dendroecology and the Vaganov-Shashkin (VS) process-based growth model. We studied two Pinus pinaster stands with contrasting Grs showing recent dieback in the Iberian System, north-eastern Spain. We compared coexisting declining (D) and non-declining (ND) trees with crown defoliation values above and below the 50% threshold, respectively. The mean growth rate was lower in D than in ND trees in the two stands. The two vigor classes showed a growth divergence prior to the dieback onset and different responsiveness to climate. The ND trees were more responsive to changes in spring water balance and soil moisture than D trees, indicating a loss of growth responsiveness to the climate in stressed trees. Such an interaction between water availability and vigor was reflected by the VS-model simulations, which provided evidence for the observation that growth was mainly limited by low soil moisture in both sites. Such an interaction between water availability and vigor was reflected by the VS-model simulations, which provided evidence for the observation that growth was mainly limited by low soil moisture in both sites. The presented comparisons indicated different stand vulnerabilities to drought contingent on-site conditions. Further research should investigate the role played by environmental conditions and individual features such as access to soil water or hydraulic traits and implement them in process-based growth models to better forecast dieback.

Differences in temperature sensitivity and drought recovery between natural stands and plantations of conifers are species-specific.

Forests are being impacted by climate and land-use changes which have altered their productivity and growth. Understanding how tree growth responds to climate in natural and planted stands may provide valuable information to prepare management in sight of climate change. Plantations are expected to show higher sensitivity to climate and lower post-drought resilience than natural stands, due to their lower compositional and structural diversity. We reconstructed and compared the radial growth of six conifers with contrasting ecological and climatic niches (Abies pinsapo, Cedrus atlantica, Pinus sylvestris, Pinus nigra, Pinus pinea, Pinus pinaster) in natural and planted stands subjected to seasonal drought in 40 sites. We quantified the relationships between individual growth variability and climate variables (temperature, precipitation and the SPEI drought index), as well as post-drought resilience. Elevated precipitation during the previous autumn-winter and current spring to early summer enhanced growth in both natural and planted stands of all species. Temperature effects on growth were less consistent: only plantations of A. pinsapo, C. atlantica, P. nigra, P. pinea, P. sylvetris and a natural stand of P. nigra showed negative impacts of summer temperature on growth. Drought reduced growth of all species in both plantations and natural stands, with variations in the temporal scale of the response. Drought constrained growth more severely in natural stands than in plantations of C. atlantica, P. pinaster and P. nigra, whereas the inverse pattern was found for A. pinsapo. Resilience to drought varied between species: natural stands of A. pinsapo, C. atlantica and P. pinaster recovered faster than plantations, while P. pinea plantations recovered faster than natural stands. Overall, plantations did not consistently show a higher sensitivity to climate and a lower capacity to recover after drought. Therefore, plantations are potential tools for mitigating climate warming.

Inter and intra-annual links between climate, tree growth and NDVI: improving the resolution of drought proxies in conifer forests.

The inter- and intra-annual variability in radial growth reflects responses to climatic variability and water shortage, especially in areas subjected to seasonal drought. However, it is unknown how this variability is related to forest productivity, which can be assessed by measuring changes in canopy greenness and cover through remote sensing products as the Normalized Difference Vegetation Index (NDVI). We combine xylogenesis with measurements of inter-annual changes in seasonal wood production (earlywood width, adjusted latewood width) and NDVI to improve the understanding of climate and drought impacts on growth and forest productivity in a Pinus teocote stand located in northern Mexico. Cambial dynamics accelerated in March and a high production of radially enlarging and thickening tracheids were observed from April to October and from June to October, respectively. Tracheid maturation was very active in October when latewood production peaked. Wet conditions in winter-spring and summer-autumn enhanced earlywood and latewood production, respectively. Earlywood and latewood were constrained by long (4-10 months) and short (2-3 months) droughts, respectively. The earlywood production depended on April soil moisture, which agrees with the peak of radially enlarging tracheid production found during that month. Aligning drought proxies at inter- and intra-annual scales by using growth and productivity measures improves our understanding of conifer forest responses to water shortage.

Forest and woodland replacement patterns following drought-related mortality.

Forest vulnerability to drought is expected to increase under anthropogenic climate change, and drought-induced mortality and community dynamics following drought have major ecological and societal impacts. Here, we show that tree mortality concomitant with drought has led to short-term (mean 5 y, range 1 to 23 y after mortality) vegetation-type conversion in multiple biomes across the world (131 sites). Self-replacement of the dominant tree species was only prevalent in 21% of the examined cases and forests and woodlands shifted to nonwoody vegetation in 10% of them. The ultimate temporal persistence of such changes remains unknown but, given the key role of biological legacies in long-term ecological succession, this emerging picture of postdrought ecological trajectories highlights the potential for major ecosystem reorganization in the coming decades. Community changes were less pronounced under wetter postmortality conditions. Replacement was also influenced by management intensity, and postdrought shrub dominance was higher when pathogens acted as codrivers of tree mortality. Early change in community composition indicates that forests dominated by mesic species generally shifted toward more xeric communities, with replacing tree and shrub species exhibiting drier bioclimatic optima and distribution ranges. However, shifts toward more mesic communities also occurred and multiple pathways of forest replacement were observed for some species. Drought characteristics, species-specific environmental preferences, plant traits, and ecosystem legacies govern postdrought species turnover and subsequent ecological trajectories, with potential far-reaching implications for forest biodiversity and ecosystem services.

Long-term thinning effects on tree growth, drought response and water use efficiency at two Aleppo pine plantations in Spain.

In Mediterranean areas where drought-induced forest dieback and tree mortality have been widely reported, it is still under debate how the likely risks of climate change will affect tree growth and consequently forest productivity. Increasing tree mortality has been associated not only to increased drought, but also to a lack of management in many dense pine forests and plantations, where warming may intensify tree-to-tree competition for soil water. This emphasizes the need of using silviculture to adapt dense stands of Mediterranean pine reforestations to warmer and drier conditions. Here we combined dendrochronology and C and O isotope analyses of wood in two Aleppo pine (Pinus halepensis) plantations, growing under semiarid conditions and experimentally thinned at high and moderate intensities along with control. The main aim was to understand the responses of radial growth and water use efficiency (WUEi) to different thinning intensities, and to analyze the effectiveness of thinning to enhance post-drought growth resilience. Thinning had a positive effect on growth, produced an increase of δ18O, reduced growth sensitivity to drought and decreased WUEi, suggesting a reduction of drought stress. These results were consistent across sites, and were significant even 20 years after the intervention took place. Considering the climate effects on growth through the SPEI drought index to calculate resistance and recovery indices, an increase of resistance after thinning was observed. We conclude that high thinning intensity (50% of basal area removed) is a useful silviculture intervention on Mediterranean Aleppo pine plantations that enhances their growth, and makes them less dependent on harsh climatic conditions, improving their resilience against drought and consequently making them better adapted to more unfavourable conditions.

Variation in the access to deep soil water pools explains tree-to-tree differences in drought-triggered dieback of Mediterranean oaks.

Individual differences in the access to deep soil water pools may explain the differential damage among coexisting, conspecific trees as a consequence of drought-induced dieback. We addressed this issue by comparing the responses to a severe drought of three Mediterranean oak species with different drought tolerance, Quercus pubescens L. and Quercus frainetto Ten., mainly thriving at xeric and mesic sites, respectively, and Quercus cerris L., which dominates at intermediate sites. For each species, we compared coexisting declining (D) and non-declining (ND) trees. The stable isotope composition (δ2H, δ18O) of xylem and soil water was used to infer a differential use of soil water sources. We also measured tree size and radial growth to quantify the long-term divergence of wood production between D and ND trees and non-structural carbohydrates (NSCs) in sapwood to evaluate if D trees presented lower NSC values. The ND trees had access to deeper soil water than D trees except in Q. frainetto, as indicated by significantly more depleted xylem water values. However, a strong δ2H offset between soil and xylem water isotopes observed in peak summer could suggest that both tree types were not physiologically active under extreme drought conditions. Alternative processes causing deuterium fractionation, however, could not be ruled out. Tree height and recent (last 15-25 years) growth rates in all species studied were lower in D than in ND trees by 22 and 44%, respectively. Lastly, there was not a consistent pattern of NSC sapwood concentration; in Q. pubescens, it was higher in ND trees while in Q. frainetto, the D trees were the ones exhibiting the higher NSC concentration. We conclude that the vulnerability to drought among conspecific Mediterranean oaks depends on the differential access to deep soil water pools, which may be related to differences in rooting depth, tree size and growth rate.

Drought and Phytophthora Are Associated With the Decline of Oak Species in Southern Italy.

Forest decline induced by climate change is a global phenomenon that affects many tree species, mainly in drought-prone areas as the Mediterranean region. In southern Italy, several oak species have shown decline symptoms and elevated mortality since the 2000s due to drought stress. However, it remains to be answered whether decline occurred alone or whether a pathogen was also involved. To this aim, we compared two coexisting oak species in a forest located in southern Italy which are assumed to be less (Quercus cerris) and more tolerant to drought (Quercus pubescens). We sampled fifteen couples of neighboring declining (D) and non-declining (ND) trees of both species. Wood cores were taken from all trees to perform dendrochronological analyses to detect the decline onset and link it to potential climatic drivers. Carbon isotope ratios (d13C) were analyzed in wood of the two vigor classes to compare their water-use efficiency. Phytophthora presence was also assessed in soil samples from ten D-ND couples of trees per species. The oak species most affected by drought-induced decline in terms of leaf shedding and mortality was Q. cerris, i.e., the least tolerant to drought. In both species, the D trees showed a reduced growth rate compared with ND trees from 2000 onward when drought and warming intensified. Q. pubescens showed higher growth sensitivity to precipitation, temperature and drought than Q. cerris. This sensitivity to climate was magnified in D trees whose growth decreased in response to warm and dry conditions during the prior winter and the late summer. The Q. pubescens D trees were more efficient in their water use than ND trees before the growth divergence between D and ND trees amplified. In the studied area, Phytophthora quercina was isolated from 40% of the sampled trees, and tended to be more frequent amongst ND than amongst D trees. Our data suggests that droughts and warm summer conditions triggered oak decline. The high prevalence of P. quercina in the studied area warrants further study as a potential predisposing factor.

Size Matters a Lot: Drought-Affected Italian Oaks Are Smaller and Show Lower Growth Prior to Tree Death.

Hydraulic theory suggests that tall trees are at greater risk of drought-triggered death caused by hydraulic failure than small trees. In addition the drop in growth, observed in several tree species prior to death, is often interpreted as an early-warning signal of impending death. We test these hypotheses by comparing size, growth, and wood-anatomy patterns of living and now-dead trees in two Italian oak forests showing recent mortality episodes. The mortality probability of trees is modeled as a function of recent growth and tree size. Drift-diffusion-jump (DDJ) metrics are used to detect early-warning signals. We found that the tallest trees of the anisohydric Italian oak better survived drought contrary to what was predicted by the theory. Dead trees were characterized by a lower height and radial-growth trend than living trees in both study sites. The growth reduction of now-dead trees started about 10 years prior to their death and after two severe spring droughts during the early 2000s. This critical transition in growth was detected by DDJ metrics in the most affected site. Dead trees were also more sensitive to drought stress in this site indicating different susceptibility to water shortage between trees. Dead trees did not form earlywood vessels with smaller lumen diameter than surviving trees but tended to form wider latewood vessels with a higher percentage of vessel area. Since living and dead trees showed similar competition we did not expect that moderate thinning and a reduction in tree density would increase the short-term survival probability of trees.

A multi-proxy assessment of dieback causes in a Mediterranean oak species.

Drought stress causes forest dieback that is often explained by two interrelated mechanisms, namely hydraulic failure and carbon starvation. However, it is still unclear which functional and structural alterations, related to these mechanisms, predispose to dieback. Here we apply a multi-proxy approach for the characterization of tree structure (radial growth, wood anatomy) and functioning (δ13C, δ18O and non-structural carbohydrates (NSCs)) in tree rings before and after drought-induced dieback. We aim to discriminate which is the main mechanism and to assess which variables can act as early-warning proxies of drought-triggered damage. The study was tailored in southern Italy in two forests (i.e., San Paolo (SP) and Oriolo (OR)) where declining and non-declining trees of a ring-porous tree species (Quercus frainetto Ten.) showing anisohydric behavior coexist. Both stands showed growth decline in response to warm and dry spring conditions, although the onset of dieback was shifted between them (2002 in SP and 2009 in OR). Declining trees displayed a sharp growth drop after this onset with reductions of 49% and 44% at SP and OR sites, respectively. Further, contrary to what we expected, declining trees showed a lower intrinsic water-use efficiency compared with non-declining trees after the dieback onset (with reductions of 9.7% and 5.6% at sites SP and OR, respectively), due to enhanced water loss through transpiration, as indicated by the lower δ18O values. This was more noticeable at the most drought-affected SP stand. Sapwood NSCs did not differ between declining and non-declining trees, indicating no carbon starvation in affected trees. Thus, the characterized structural and functional alterations partially support the hydraulic failure mechanism of dieback. Finally, we show that growth data are reliable early-warning proxies of drought-triggered dieback.