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Mistletoes are xylem-tapping hemiparasites that rely on their hosts for water and nutrient uptake. Thus, they impair tree performance in the face of environmental stress via altering the carbon and water relations and nutritional status of trees. To improve our understanding of physiological responses to mistletoe and ongoing climate change, we investigated radial growth, stable carbon and oxygen isotopic signals, and elemental composition of tree rings in silver fir (Abies alba Mill.) and Scots pine (Pinus sylvestris L.) forests infested with Viscum album L. We compared temporal series (1990–2020) of basal area increment (BAI), intrinsic water-use efficiency (iWUE), oxygen isotope composition (δ18O), nutrient concentrations and stoichiometric ratios between non-infested (NI) and severely infested (SI) fir and pine trees from populations located close to the xeric distribution limit of the species in north-eastern Spain. The SI trees showed historically higher growth, but the BAI trend was negative for more than three decades before 2020 and their growth rates became significantly lower than those of NI trees by the mid-2010s. Mistletoe infestation was related to an enhanced sensitivity of radial growth to vapour pressure deficit (atmospheric drought). The SI trees showed less pronounced iWUE increases (fir) and lower iWUE values (pine) than NI trees. The lower tree-ring δ18O values of SI trees may be the result of several superimposed effects operating simultaneously, including leaf-level evaporative enrichment, source water isotopic signals, and anatomical and phenological differences. We observed a deterioration of potassium (K) nutrition in tree-ring wood of both species in SI trees, along with accumulation of manganese (Mn). We suggest that such nutritional patterns are driven by the indirect effect of mistletoe-induced drought stress, particularly in pine. The combined analyses of different physiological indicators imprinted on tree rings provided evidence of the progressive onset of carbon, water and nutrient imbalances in mistletoe-infested conifers inhabiting seasonally dry regions. [ABSTRACT FROM AUTHOR]

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Summary: The climatic significance and ecosystem implications of latewood intra-annual density fluctuations (IADFs) are still not fully understood in Mediterranean ring-porous oak species. To solve this issue, we investigated: (i) the climate drivers of radial growth and latewood IADFs in the Mediterranean oak Quercus faginea , and (ii) whether they were correlated to increased canopy cover and greenness as recorded by the Normalized Difference Vegetation Index (NDVI). IADFs were formed in the mid to late latewood and they were characterized by rows of parenchyma cells. Such IADFs were uncommon and only present in 30–50% of sampled trees during 1999 and 2014, both years characterized by high precipitation in late July and early August. This relationship between IADFs formation and summer wet conditions differed from the conditions that enhance Q. faginea annual growth, wet winter-spring conditions, and low summer temperatures. Furthermore, IADFs formation showed a positive relationship with August NDVI values, indicating a correspondence of canopy greenness with secondary growth reactivation. We conclude that latewood IADFs in Q. faginea are a robust proxy of rare late-summer wet conditions and enhanced canopy activity as reflected by increased August NDVI values. Further monitoring of xylogenesis in other Mediterranean hardwoods along climatic gradients is suggested to mechanistically explain IADF production in seasonally dry biomes. [ABSTRACT FROM AUTHOR]

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]

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

Old-growth forests have a primary environmental and ecological value, as they closely recall primeval forests and the natural forest landscape that was spread throughout Italy and Europe before man intervention heavily altered the functioning of most natural ecosystems. These ecosystems are more and more recognized as keystones of environmental policies (i.e., the EU Biodiversity Strategy 2030) aimed to preserve biodiversity and mitigate the impacts of climate change. A recent report highlights that Europe's old-growth forests account for only 3 percent of the overall forest surface. The majority of italian old-growth forests are located along the Apennines, especially in the Pollino, Sila and Aspromonte National Parks, in remote montane areas characterized by irregular topographical conditions (i.e., high slopes and/or shallow soils) which hamper the access and exploitation of these less productive stands, thus allowing the survival of old-growth forests for many centuries. The ongoing climate change is a significant challenge to the conservation of old-growth forests, particularly in areas such as the Mediterranean basin, where an intensification of extreme climate events is expected. Heat waves can exacerbate drought's negative impacts on tree growth and survival, leading to reduced woody carbon stocks and biodiversity in old-growth forests. To date, there is poor understanding of such highly valuable and extremely fragile natural ecosystems in Italy. Insights are available based on data gathered as part of two research programs, funded by the Italian Ministry of the Environment and Land and Sea Protection, with the purpose of creating a Network of Old Growth Forests in Italy's National Parks. In this study, we aimed to highlight the role and importance of old-growth forests, focusing on those located within the Pollino National Park. A comprehensive overview of the key strengths and weaknesses, that mark these peculiar forest ecosystems, was followed by a discussion of the future outlook, grounded on strategies that will ensure not only the protection and conservation of these ecosystems, but also their development and promotion. [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]

Old-growth forests have a primary environmental and ecological value, as they closely recall primeval forests and the natural forest landscape that was spread throughout Italy and Europe before man intervention heavily altered the functioning of most natural ecosystems. These ecosystems are more and more recognized as keystones of environmental policies (i.e., the EU Biodiversity Strategy 2030) aimed to preserve biodiversity and mitigate the impacts of climate change. A recent report highlights that Europe's old-growth forests account for only 3 percent of the overall forest surface. The majority of italian old-growth forests are located along the Apennines, especially in the Pollino, Sila and Aspromonte National Parks, in remote montane areas characterized by irregular topographical conditions (i.e., high slopes and/or shallow soils) which hamper the access and exploitation of these less productive stands, thus allowing the survival of old-growth forests for many centuries. The ongoing climate change is a significant challenge to the conservation of old-growth forests, particularly in areas such as the Mediterranean basin, where an intensification of extreme climate events is expected. Heat waves can exacerbate drought's negative impacts on tree growth and survival, leading to reduced woody carbon stocks and biodiversity in old-growth forests. To date, there is poor understanding of such highly valuable and extremely fragile natural ecosystems in Italy. Insights are available based on data gathered as part of two research programs, funded by the Italian Ministry of the Environment and Land and Sea Protection, with the purpose of creating a Network of Old Growth Forests in Italy's National Parks. In this study, we aimed to highlight the role and importance of old-growth forests, focusing on those located within the Pollino National Park. A comprehensive overview of the key strengths and weaknesses, that mark these peculiar forest ecosystems, was followed by a discussion of the future outlook, grounded on strategies that will ensure not only the protection and conservation of these ecosystems, but also their development and promotion. [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]

Old-growth forests have a primary environmental and ecological value, as they closely recall primeval forests and the natural forest landscape that was spread throughout Italy and Europe before man intervention heavily altered the functioning of most natural ecosystems. These ecosystems are more and more recognized as keystones of environmental policies (i.e., the EU Biodiversity Strategy 2030) aimed to preserve biodiversity and mitigate the impacts of climate change. A recent report highlights that Europe's old-growth forests account for only 3 percent of the overall forest surface. The majority of italian old-growth forests are located along the Apennines, especially in the Pollino, Sila and Aspromonte National Parks, in remote montane areas characterized by irregular topographical conditions (i.e., high slopes and/or shallow soils) which hamper the access and exploitation of these less productive stands, thus allowing the survival of old-growth forests for many centuries. The ongoing climate change is a significant challenge to the conservation of old-growth forests, particularly in areas such as the Mediterranean basin, where an intensification of extreme climate events is expected. Heat waves can exacerbate drought's negative impacts on tree growth and survival, leading to reduced woody carbon stocks and biodiversity in old-growth forests. To date, there is poor understanding of such highly valuable and extremely fragile natural ecosystems in Italy. Insights are available based on data gathered as part of two research programs, funded by the Italian Ministry of the Environment and Land and Sea Protection, with the purpose of creating a Network of Old Growth Forests in Italy's National Parks. In this study, we aimed to highlight the role and importance of old-growth forests, focusing on those located within the Pollino National Park. A comprehensive overview of the key strengths and weaknesses, that mark these peculiar forest ecosystems, was followed by a discussion of the future outlook, grounded on strategies that will ensure not only the protection and conservation of these ecosystems, but also their development and promotion. [ABSTRACT FROM AUTHOR]

Old-growth forests have a primary environmental and ecological value, as they closely recall primeval forests and the natural forest landscape that was spread throughout Italy and Europe before man intervention heavily altered the functioning of most natural ecosystems. These ecosystems are more and more recognized as keystones of environmental policies (i.e., the EU Biodiversity Strategy 2030) aimed to preserve biodiversity and mitigate the impacts of climate change. A recent report highlights that Europe's old-growth forests account for only 3 percent of the overall forest surface. The majority of italian old-growth forests are located along the Apennines, especially in the Pollino, Sila and Aspromonte National Parks, in remote montane areas characterized by irregular topographical conditions (i.e., high slopes and/or shallow soils) which hamper the access and exploitation of these less productive stands, thus allowing the survival of old-growth forests for many centuries. The ongoing climate change is a significant challenge to the conservation of old-growth forests, particularly in areas such as the Mediterranean basin, where an intensification of extreme climate events is expected. Heat waves can exacerbate drought's negative impacts on tree growth and survival, leading to reduced woody carbon stocks and biodiversity in old-growth forests. To date, there is poor understanding of such highly valuable and extremely fragile natural ecosystems in Italy. Insights are available based on data gathered as part of two research programs, funded by the Italian Ministry of the Environment and Land and Sea Protection, with the purpose of creating a Network of Old Growth Forests in Italy's National Parks. In this study, we aimed to highlight the role and importance of old-growth forests, focusing on those located within the Pollino National Park. A comprehensive overview of the key strengths and weaknesses, that mark these peculiar forest ecosystems, was followed by a discussion of the future outlook, grounded on strategies that will ensure not only the protection and conservation of these ecosystems, but also their development and promotion. [ABSTRACT FROM AUTHOR]

Seasonal patterns of wood formation (xylogenesis) remain understudied in mixed pine–oak forests despite their contribution to tree coexistence through temporal niche complementarity. Xylogenesis was assessed in three pine species (Pinus cembroides, Pinus leiophylla, Pinus engelmannii) and one oak (Quercus grisea) coexisting in a semi-arid Mexican forest. The main xylogenesis phases (production of cambium cells, radial enlargement, cell-wall thickening and maturation) were related to climate data considering 5–15-day temporal windows. In pines, cambium activity maximized from mid-March to April as temperature and evaporation increased, whereas cell radial enlargement peaked from April to May and was constrained by high evaporation and low precipitation. Cell-wall thickening peaked from June to July and in August–September as maximum temperature and vapour pressure deficit (VPD) increased. Maturation of earlywood and latewood tracheids occurred in May–June and June–July, enhanced by high minimum temperatures and VPD in P. engelmannii and P. leiophylla. In oak, cambial onset started in March, constrained by high minimum temperatures, and vessel radial enlargement and radial increment maximized in April as temperatures and evaporation increased, whereas earlywood vessels matured from May to June as VPD increased. Overall, 15-day wet conditions enhanced cell radial enlargement in P. leiophylla and P. engelmannii, whereas early-summer high 15-day temperature and VPD drove cell-wall thickening in P. cembroides. Warm night conditions and high evaporation rates during spring and summer enhanced growth. An earlier growth peak in oak and a higher responsiveness to spring–summer water demand in pines contributed to their coexistence. [ABSTRACT FROM AUTHOR]

Different leaf (evergreen vs. deciduous habit) and xylem (diffuse-vs. ring-porous wood) traits represent contrasting strategies to face seasonal changes in water availability and temperature. However, how contrasting leaf and xylem habits of coexisting tree species affect stem wood formation and tree-ring development remains poorly understood. Here, we investigated the spatio-temporal patterns of wood formation in two deciduous oaks (Quercus faginea and Quercus petraea) and two evergreen oaks (Quercus ilex and Quercus suber) coexisting in seasonally dry Mediterranean forests along an aridity gradient in Spain. We hypothesized that growth responses to drought and intra- and inter-annual growth patterns would differ between functional groups. We simulated intra- and interannual growth using a modified version of the Vaganov-Shashkin (VS) process-based, growth model. The VS model simulations were used to estimate growth changes under a high emission scenario (RCP 8.5) for the current distribution of the study oak species and to forecast their future performance under warm (4.8 °C) conditions in the Iberian Peninsula. Our simulations indicate that climate warming would induce a shortening of the ringgrowth season and a reduction of radial growth in evergreen and deciduous Mediterranean oaks, particularly in dry sites from southern and eastern Iberia currently occupied by Q. ilex and Q. faginea. Evergreen oaks may better recover after dry periods than deciduous oaks by resuming growth after the summer drought. Low soil water availability in spring would be more detrimental to growth of deciduous oaks. Process-based growth models should be refined and validated to better forecast changes in tree growth as a function of climate. [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]

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 (d18O) 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 d18O than ND trees. The uncoupling between iWUE and d18O, together with the positive correlations between P and K leaf concentrations and d18O, 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. [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 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. [ABSTRACT FROM AUTHOR]

Several forest dieback and mortality phenomena reported across the Italian Peninsula over the last years underline an enhanced forest vulnerability in response to climate extremes, mainly heat waves and dry spells. Hundreds or even thousands hectares of forests completely die out over few years, causing considerable negative impacts on ecosystem services such as carbon sequestration and biodiversity. Thus, there is a compelling need to monitor these phenomena for understanding the future dynamics of forest stands: these crucial assumptions have led to SilvaCuore's development, the first App designed in Italy to report the forests health status. The support of an active users' community can not only allow researchers to survey declining forest sites, but also to better plan research activities and management measures. Silvacuore is an App designed with the willingness to strenghten the link between the citizens and the scientific community, contributing to the development of a valuable scientific database, available on a website and linked to the App. Through a user-friendly interface, the App leads the users throughout a simplified reporting procedure. SilvaCuore may therefore be perceived by a dual aspect: a resource for the scientific community and a unique opportunity for citizens to play an active role in research activities. [ABSTRACT FROM AUTHOR]

We propose some ideas for an adaptive silviculture in the Mediterranean biogeographic region that could assist in shaping forests which are less vulnerable to extreme climatic events, such as those drought and heat waves that are expected to be more intense and frequent in the future. We discuss about silvicultural approaches that would aim to: regulate competition and densitydependent effects; enhance functional diversity; increase the complexity of forest communities. These approaches are especially advisable in the Mediterranean region, which is a hotspot for climate change and where forests have been exploited for many centuries and are only recently taking advantage of a recovery phase. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Key message: Juglans, Fraxinus, Quercus and Pinus species seem to better maximize the carbon–water ratio providing useful indications on species selection for forestry plantations in areas with increasing drought risk. Maximizing carbon sequestration for a given water budget is extremely important in the contest of climate change in the Mediterranean region, which is characterized by increasing temperatures and rising water stress. This issue is fundamental for plantation stands, where limited water availability during the growing season reduces CO2 assimilation and, consequently, tree growth. In this study, the main objective was to investigate the performances in terms of carbon–water balance of conifer (Pinus halepensis and Cupressus sempervirens) and hardwood (Quercus robur, Juglans regia, Fraxinus excelsior and Populus spp.) mixed plantations. To this aim, we used carbon isotope signatures to evaluate the intrinsic water-use efficiency (iWUE) and the species-specific relationship between basal area increments (BAI) and iWUE. At the species level, the highest iWUE values corresponded to the lowest carbon accumulation in terms of BAI, for water-saving species such as Cupressus. Conversely, Populus had the lowest iWUE and the highest BAI accumulation. Juglans, Fraxinus, and Pinus showed the most balanced ratio between BAI and iWUE. Overall, no clear correlation of iWUE and BAI was evident within all species, except for Populus and Cupressus. Considering projected aridification and increased temperatures that will negatively impact the growth, our data suggest that Pinus, for conifers, and Quercus, Juglans, Fraxinus for hardwood species should be preferred when choosing species for forestry plantation, as they performed better in terms of BAI and iWUE ratio. [ABSTRACT FROM AUTHOR]

Key Message: Pinus leiophylla production of radially enlarging tracheids is bimodal with peaks in June and October. The production of thickening tracheids and NDVI were coupled. In drought-prone areas, xylem production and forest productivity are assumed to depend on the amount and timing of precipitation. However, few studies have addressed if xylogenesis and proxies of forest productivity as the Normalized Difference Vegetation Index (NDVI) are coupled and respond to moisture availability. In these areas, drought seasonality and timing could lead to bimodal growth patterns. We tested if xylogenesis was bimodal and coupled with climate and NDVI at the intra-annual scale in Pinus leiophylla from NW Mexico. We found a bimodal growth pattern with a major peak of production of radially enlarging tracheids in June and a secondary peak in October. The June peak preceded a coupled rise of the amount of cell-wall-thickening tracheids and the NDVI which peaked from September to October. The timing in the production of thickening and mature tracheids matched the NDVI pattern. This coupling suggests active carbon uptake after wet summer conditions and carbohydrate usage in tracheid maturation and cell-wall thickening. The production of radially enlarging tracheids, which determines the final ring width, was enhanced by warmer minimum temperatures whenever precipitation was abundant. In contrast, more mature latewood tracheids were produced in response to cool-wet summer conditions. The bimodal behavior of P. leiophylla radially enlarging tracheids illustrates xylem plasticity in response to seasonal drought. Xylogenesis and NDVI data were uncoupled. Therefore, the assessment of forest responses to climate warming in similar drought-prone areas must explicitly consider shifts in productivity and the timing of wood production. [ABSTRACT FROM AUTHOR]

Several forest dieback and mortality phenomena reported across the Italian Peninsula over the last years underline an enhanced forest vulnerability in response to climate extremes, mainly heat waves and dry spells. Hundreds or even thousands hectares of forests completely die out over few years, causing considerable negative impacts on ecosystem services such as carbon sequestration and biodiversity. Thus, there is a compelling need to monitor these phenomena for understanding the future dynamics of forest stands: these crucial assumptions have led to SilvaCuore's development, the first App designed in Italy to report the forests health status. The support of an active users' community can not only allow researchers to survey declining forest sites, but also to better plan research activities and management measures. Silvacuore is an App designed with the willingness to strenghten the link between the citizens and the scientific community, contributing to the development of a valuable scientific database, available on a website and linked to the App. Through a user-friendly interface, the App leads the users throughout a simplified reporting procedure. SilvaCuore may therefore be perceived by a dual aspect: a resource for the scientific community and a unique opportunity for citizens to play an active role in research activities. [ABSTRACT FROM AUTHOR]

We propose some ideas for an adaptive silviculture in the Mediterranean biogeographic region that could assist in shaping forests which are less vulnerable to extreme climatic events, such as those drought and heat waves that are expected to be more intense and frequent in the future. We discuss about silvicultural approaches that would aim to: regulate competition and densitydependent effects; enhance functional diversity; increase the complexity of forest communities. These approaches are especially advisable in the Mediterranean region, which is a hotspot for climate change and where forests have been exploited for many centuries and are only recently taking advantage of a recovery phase. [ABSTRACT FROM AUTHOR]

We propose some ideas for an adaptive silviculture in the Mediterranean biogeographic region that could assist in shaping forests which are less vulnerable to extreme climatic events, such as those drought and heat waves that are expected to be more intense and frequent in the future. We discuss about silvicultural approaches that would aim to: regulate competition and densitydependent effects; enhance functional diversity; increase the complexity of forest communities. These approaches are especially advisable in the Mediterranean region, which is a hotspot for climate change and where forests have been exploited for many centuries and are only recently taking advantage of a recovery phase. [ABSTRACT FROM AUTHOR]

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 showthat 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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Warming-related growth decrease on southern Fagus sylvatica forests has been observed in different regions; however, whether it is a generalized fact or not remains unclear. Here we investigate the geographical pattern on growth response of the southwestern European beech forests to the warming climate shift which started in the 1980s. We sampled 15 beech forests (215 trees) across four climatically contrasting regions (Mediterranean, Pyrenean, low- and high-elevation Atlantic areas) near the southern distribution limit of the species in the Iberian Peninsula. Dendrochronological analyses were carried out to evaluate the growth of European beech since the 1950s. Growth responses quantified as pointer years, abrupt growth changes and long-term growth trends were compared between periods (before and after the 1980s climate shift), geographical regions and tree sizes. Analyses of the studied variables indicated a growth decrease in basal area increment after the climate shift in three of the four studied regions. Pyrenean stands were not negatively influenced by the climate shift, although an increase in the frequency of negative abrupt growth changes was also found there. Growth after the climate shift presented divergent patterns depending on the geographical region. Although Mediterranean and Atlantic stands presented different indicators of constrained growth, Pyrenean stands showed rising long-term growth trends. Such results suggest that regional characteristics differentially determine the growth response of the southern European beech forests to recent warming periods. Iberian beech forests located at the Pyrenees would benefit from forecasted warming conditions, whereas Atlantic and Mediterranean forests would be more prone to suffer warming-related growth decline. [ABSTRACT FROM AUTHOR]

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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. [ABSTRACT FROM AUTHOR]

Abstract: Questions: Do vascular plant species richness and beta‐diversity differ between managed and structurally complex unmanaged stands? To what extent do species richness and beta‐diversity relate to forest structural attributes and heterogeneity? Location: Five national parks in central and southern Italy. Methods: We sampled vascular plant species composition and forest structural attributes in eight unmanaged temperate mesic forest stands dominated or co‐dominated by beech, and in eight comparison stands managed as high forests with similar environmental features. We compared plant species richness, composition and beta‐diversity across pairs of stands (unmanaged vs managed) using GLMMs. Beta‐diversity was quantified both at the scale of each pair of stands using plot‐to‐plot dissimilarity matrices (species turnover), and across the whole data set, considering the distance in the multivariate species space of individual plots from their centroid within the same stand (compositional heterogeneity). We modelled the relationship between species diversity (richness and beta‐diversity) and forest structural heterogeneity and individual structural variables using GLMMs and multiple regression on distance matrices. Results: Species composition differed significantly between managed and unmanaged stands, but not richness and beta‐diversity. We found weak evidence that plant species richness increased with increasing levels of structural heterogeneity and canopy diversification. At the scale of individual stands, species turnover was explained by different variables in distinct stands, with variables related to deadwood quantity and quality being selected most often. We did not find support for the hypothesis that compositional heterogeneity varies as a function of forest structural characteristics at the scale of the whole data set. Conclusions: Structurally complex unmanaged stands have a distinct herb layer species composition from that of mature stands in similar environmental conditions. Nevertheless, we did not find significantly higher levels of vascular plant species richness and beta‐diversity in unmanaged stands. Beta‐diversity was related to patterns of deadwood accumulation, while for species richness the evidence that it increases with increasing levels of canopy diversification was weak. These results suggest that emulating natural disturbance, and favouring deadwood accumulation and canopy diversification may benefit some, but not all, facets of plant species diversity in Apennine beech forests. [ABSTRACT FROM AUTHOR]

Increased forest vulnerability is being reflected as more widespread and severe drought-induced decline episodes. In particular, the Mediterranean area is revealing a high susceptibility to phenomena of loss in tree vitality across species. Within tree species, oaks (Quercus spp.) are experiencing extensive decline in many countries. However, in the wake of the so-called "oak decline phenomenon", the attention on these species has generally been limited. In this paper, we review the current available literature on oakdecline cases reported within the Mediterranean Basin, with particular remark for those occurred in Italy and Spain. More specifically our main aims were to: (i) provide an update on the patterns and mechanisms of decline by focusing on tree-ring and wood-anatomical variables; (ii) provide some hints for improving the resistance and resilience of oak stands experiencing decline. Our review reveals that drought is reported as the main driver triggering oak decline within the Mediterranean Basin, although other causes (i.e., increasing temperature, pathogens attack or excessive stand density) could exacerbate decline. In most reported cases, drought induced a substantial reduction of growth and changes in some wood anatomical properties. Indeed, growth decline prior death is also indicated as an early-warning signal of impending death. In ring-porous oak species, declining trees were often characterized by a very low production of latewood and a decrease in lumen area of the widest earlywood vessels, suggesting a potential reduction of hydraulic conductivity. Moreover, hydraulic dysfunction is reported as the main cause of decline. Finally, regarding management actions that should be considered for improving the resilience of declining stands and preserve the species-specific stand composition, it could be useful to shorten the rotation period of coppice stands or promoting their gradual conversion towards high forests, and favoring more drought-resistant species should also be considered. In addition, regeneration prior to regeneration cuts should be improved by anticipating seed dispersal or by planting oak seedlings obtained from local germoplasm. [ABSTRACT FROM AUTHOR]

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 nondeclining 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. [ABSTRACT FROM AUTHOR]

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. [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]