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

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.

Spatial and temporal variation in functional traits allows trees to adjust to shifting environmental conditions such as water stress. However, the change of traits, both mean and variances, along water availability gradients and across growing seasons, as well as their covariation with tree performance, have been rarely assessed. We examined intraspecific trait variation in coexisting evergreen (Quercus ilex ssp. ilex and Q. ilex ssp. ballota) and deciduous (Quercus faginea and Quercus humilis) Mediterranean oaks along a wide water availability gradient in northeastern Spain during six years. We measured leaf area (LA), shoot twig mass (Sm), leaf mass per area (LMA) and the ratio of shoot twig to leaf biomass (Sm:Lm). We characterized tree performance through basal area increment (BAI) and drought resilience indices. Higher variation was found within individuals than between individuals across populations and years. Within species, we found trait adjustments toward more conservative water-use (low LA and Sm and high LMA) with increasing drier conditions. Intraspecific trait variation was constrained by water availability, particularly on the deciduous species. In Q. ilex, trait variance of LMA positively covaried with annual BAI, whereas variance of LA, Sm and Sm:Lm was positively related to resistance and resilience against the severe 2012 drought in deciduous oaks. Our results support a tradeoff between the ability to tolerate drought and the capacity to cope with unpredictable changes in the environment through increased intraspecific trait variation, which may have implications on tree performance in the face of increased extreme events.

[Context] Global change is leading to more frequent and intense tree damage and mortality events. Drought-induced tree mortality is occurring worldwide leading to broad-scale events, but the spatial patterns of tree damage and mortality, their underlying drivers and their variation over time is largely unknown., [Objectives] We investigated the spatial patterns of tree damage and mortality across Mediterranean forests of the Iberian Peninsula, the underlying effects of stand structure and climate, and how the spatial patterns and relationships with underlying drivers changed over time., [Methods] We used the Spanish Forest Inventory to analyse the autocorrelation in tree damage and mortality across forest types, hurdle-gamma models to quantify the effect of stand structure and climate on tree damage and mortality, and cross-correlograms to assess their spatial dependence and its change over time., [Results] We observed a greater magnitude and a stronger autocorrelation in tree damage than mortality, with positive aggregation up to 20 kms. There was a spatial dependence between tree damage and mortality with their drivers, with spatial aggregation increasing with water availability, drought intensity and stand structure. The spatial dependence of tree damage and mortality with the underlying drivers increased over time, particularly for drought intensity., [Conclusions] Our results suggest that the combined effect of intense competition and drought could favour more extensive die-off and tree mortality events, providing key information for identifying vulnerable areas and the planning of adaptation measures.

Recent drought-induced dieback alters forest dynamics, which are also shaped by past management. In western Pyrenean silver fir (Abies alba) stands, dieback concurs in space and time with the legacies of past management, but the impacts on forest growth, structure and composition are unknown. We aim to disentangle how dieback interacts with the legacies of past human use and modulates the recent dynamics of silver fir forests. To this end, we sampled eleven silver fir forests across wide climatic gradients and included declining and non-declining sites. We measured radial growth, structure, composition, understory cover and type and amount of deadwood. Silver fir growth declines in response to late-summer drought. In declining sites, most defoliated stands showed the lowest silver fir density and were those where growth depended more on water availability. Tree death enhanced the cover of dominant understory plants such as Buxus sempervirens. Past management activities leave an imprint in the growth of silver fir, such as releases due to past logging, but also affect the number of stumps and snags and the current tree density. A more extensive monitoring will be required to fully disentangle the multiple influences of past management legacies and current climate change on forest dynamics.

[Purpose of Review] The capacity of woody plants to cope with climate change depends on their adjustments to changing environmental conditions by phenotypic plasticity or by genotypic changes (i.e., local adaptation). To determine whether intraspecific trait variation (ITV) translates into resistance or tolerance to drought and eventually how it relates to performance when facing drought, we reviewed the recent literature on the intraspecific variation in woody plant functional traits along aridity gradients., [Recent Findings] At the intraspecific level, functional traits vary along aridity gradients, but this variation depends on the considered trait and species. While ITV is ubiquitous and of relevant magnitude, its relationship with tree performance in the case of growth or survival is unclear and very idiosyncratic., [Summary] ITV varies along aridity gradients and, for several traits, is of comparable magnitude as interspecific trait variation. However, the relationship between ITV and tree performance is not consistent among species. This makes the use of ITV challenging when quantifying its contribution to drought tolerance.

Increased winter and early spring temperatures due to climate change can enhance forest productivity due to earlier growth onset in temperate regions. However, drought-prone forests can be highly vulnerable to the combined effect of high summer temperatures and water stress. Understanding how water stress and rising temperatures along the growing season control tree growth in co-occurring species with contrasting drought tolerances is key to project climate change effects on forest dynamics. Here, we evaluated the interactive effect of seasonal temperature and water availability on annual radial growth and climate–growth instability over six decades (1951–2015) in three pine species with contrasting drought tolerances (Pinus pinaster, Pinus nigra, and Pinus sylvestris). For this, we retrospectively observed radial tree growth using dendroecological methods and evaluated seasonal temperature and water availability effects by using linear mixed models. Early growing season temperature and water availability had a positive effect on tree growth, but the positive effect of late season temperature was modulated by water availability. Moving time-window analyses revealed temporal instability in climate–growth relationships. Since the 1980s, pine species showed a higher growth sensitivity to both seasonal temperatures and annual water availability. Furthermore, growth reductions were more pronounced due to increased summer temperatures and reduced precipitation. Our results were similar for the three studied pine species despite their contrasting tolerance to drought. Overall, climate warming effects on pine growth are contingent upon water availability in Mediterranean continental forests. Synchronization among species, climate–growth instability, and negative growth trends suggests an increased vulnerability to drought of Mediterranean pine species in response to ongoing climate change.

Tree phenology is sensitive to climate warming and changes in seasonal precipitation. Long xylogenesis records are scarce, thus limiting our ability to analyse how radial growth responds to climate variability. Alternatively, process-based growth models can be used to simulate intra-annual growth dynamics and to better understand why growth bimodality varies along temperature and precipitation gradients. We used the Vaganov-Shashkin (VS) growth model to analyse the main climatic drivers of growth bimodality in eight trees and shrubs conifers (four pines and four junipers) across Spain. We selected eleven sites with different continentality degree and spring/autumn precipitation ratios since we expected to find pronounced bimodal growth in less continental sites with spring and autumn precipitation peaks. The VS model successfully simulated annual growth rates at all sites as a function of daily temperature and soil moisture data. Bimodal growth patterns clustered into less continental sites showing low spring/autumn precipitation ratios. This finding agrees with observed climate-growth associations showing that growth was enhanced by wet-cool winter-to-spring conditions, but also by wet autumn conditions in the most bimodal sites. We observed a stronger growth bimodality in pines compared to junipers. We discuss the spatial variability of climate drivers in bimodality growth pattern and how increasing continentality and shifts in seasonal precipitation could affect growth patterns. Bimodality could be an advantageous response to overcome summer drought in Mediterranean forests. The ability of some species to reactivate growth during autumn might determine their capacity to withstand increasing summer aridity.

During the past years, growth and productivity of different oak species have been constrained by water shortage in seasonally dry regions such as the Mediterranean Basin. Thinning could improve oak radial growth in these drought-prone regions through the reduction of tree competition for soil water in summer. However, we still lack adequate, long-term assessments on how lasting are thinning treatments effects and to what extent they contribute to oak growth recovery after drought. Here we aim: (i) to study the radial growth sensitivity to drought of Quercus subpyrenaica in the Spanish Pre-Pyrenees, and (ii) to verify if thinning represents a suitable option to enhance growth resistance to drought and post-drought growth recovery. We analysed basal area increment (BAI) trends in the period 1960–2020 of formerly coppiced oak stands thinned in 1984 and compared them with unthinned plots and also with coexisting Scots pine (Pinus sylvestris) growing in thinned plots. We used the Standardized Precipitation Evapotranspiration Index (SPEI) to estimate the severity of droughts and we also assessed climate-growth relationships. Oaks in thinned plots showed higher BAI (369 mm2) than those in unthinned plots (221 mm2). Growth rates remained higher in thinned than in unthinned plots also under severe drought stress. A severe summer drought in 1986 caused abrupt BAI reductions in both oaks (- 40.5%) and pines (- 40.1%). The positive effect of thinning on growth lasted for over 20 years and slightly declined as canopies closed. In the thinned plots, trees with smaller diameter showed the greatest growth release. Oaks in unthinned plots and Scots pine were more sensitive to short-term droughts in terms of growth reduction than oaks in thinned plots, while long term droughts have similar effects on oaks from both thinned and unthinned plots. Oaks were resilient to drought, showing recovery periods lasting from 1 to 2 years in both thinned and unthinned plots. However, intense and p

Impacts of climate warming on forests vigour are forecasted to increase in magnitude. Yet it remains unclear how stand characteristics and competition modulate the relationship between tree growth and gross primary production with drought. Here, we studied how the spatial variation in stand density, basal area and height modulates tree growth (Basal Area Increment, BAI and stand growth), summer NDVI, as well as their responses to drought (Standardized Precipitation-Evapotranspiration Index, SPEI) in 56 Aleppo pine (Pinus halepensis Mill.) planted forests located in Northeast Spain. Long-term BAI responses to SPEI were strongly determined by stand density, suggesting that competition modulates Aleppo pine growth responses to drought. Along this, summer NDVI also displayed strong associations with SPEI. NDVI was mostly related with stand growth, suggesting canopy densification drives NDVI pattern and trends. Short-term BAI and NDVI responses to severe droughts were mainly independent of stand characteristics. In the studied region, drought is a universal factor limiting Aleppo pine secondary growth and canopy greening. However, the results suggest that stand density modulates Aleppo pine growth responses to drought on the long-term, reducing the growth in densest stands. Denser stands with larger trees are the ones that present higher NDVI values, suggesting that canopy activity depends more on stand canopy coverage than on secondary growth rate and its response to drought. In these Mediterranean pines, canopy activity and secondary growth are temporally coupled but spatially decoupled.

Forest growth is driven by climate variability at continental to regional scales, but other factors play major roles at local scales (0.1–1 ha). Topography impacts on tree growth responses to climate stressors, including drought, by modifying radiation, evapotranspiration rates and the access to soil moisture. However, there is a lack of studies investigating how topographical factors (elevation, aspect, slope) affect climate-growth relationships considering both continuous (tree-ring width) and discrete wood-anatomical features (e.g., intra-annual density fluctuations–IADFs). Here, we investigated how topography modulated the influences of climate and drought on semi-arid Juniperus thurifera forests from north-eastern Spain. We compared two stands located in valley bottoms and two stands located in steep slopes. Radial growth was measured using dendrochronology which also allowed quantifying latewood IADFs. A proxy of topographic influence (incident radiation) was calculated. In addition, the Vaganov-Shashkin (VS) model was used to infer the main climatic constraints of growth at intra-annual scales. We found that junipers growing in valley bottoms were taller and produced more IADFs than junipers growing in steep slopes, but responded less to precipitation variability. This was confirmed by the VS model which showed that low soil moisture in June limited growth, particularly in steep-slope sites and during dry periods. Wet-cool late-summer conditions induced the formation of IADFs. The topoclimatic modulation of growth was explained by the interaction between incident radiation and tree age. Furthermore, climate-growth associations are changing as temperatures rise. Drought impacts on growth are strengthening, but less responsive junipers growing in valley bottoms may be buffered against intensified aridification.

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

The time required for an ecosystem to recover from severe drought is a key component of ecological resilience. The phenology effects on drought recovery are, however, poorly understood. These effects centre on how phenology variations impact biophysical feedbacks, vegetation growth and, ultimately, recovery itself. Using multiple remotely sensed datasets, we found that more than half of ecosystems in mid- and high-latitudinal Northern Hemisphere failed to recover from extreme droughts within a single growing season. Earlier spring phenology in the drought year slowed drought recovery when extreme droughts occurred in mid-growing season. Delayed spring phenology in the subsequent year slowed drought recovery for all vegetation types (with importance of spring phenology ranging from 46% to 58%). The phenology effects on drought recovery were comparable to or larger than other well-known postdrought climatic factors. These results strongly suggest that the interactions between vegetation phenology and drought must be incorporated into Earth system models to accurately quantify ecosystem resilience.

[Background] Climate and land-use changes, which include the application of various types of organic and inorganic fertilisers, have been reducing the species diversity of Mediterranean grasslands and threatening its conservation. Annual plants are one of the most diverse functional groups of species in these grasslands, despite suffering competitive pressure from perennial herbaceous and woody species, and they are essential for ecosystem functioning and stability., [Aims] To quantify how fertilisation modulates the impact of plant-to-plant interactions and climate fluctuations on the dynamics of annuals in Mediterranean grasslands. We hypothesised that the application of sewage sludge would increase competition between functional groups, reducing the abundance of annuals in the long-term, but would buffer the negative impacts of drought on the year-to-year fluctuation of the diversity of annuals., [Methods] In a semi-natural species-rich Mediterranean grassland in northern Spain, we analysed the changes in the taxonomical and functional composition and diversity of annuals over 14 years in response to variations in the abundance of perennial herbaceous and woody species, climate fluctuations and fertilisation with sewage sludge. We quantified separately the patterns of year-to-year fluctuations and long-term trends., [Results] The frequency and diversity of annuals decreased with higher abundance of perennial herbaceous species, drought in June and cold winters. The addition of sewage sludge decreased the abundance of annuals in the long-term, seemed to promote competition between annuals and other functional groups at an interannual scale, and mitigated the negative effects of drought and cold., [Conclusions] Fertilisation influences differently the temporal response of annuals to climate fluctuations and plant-to-plant interactions.

[Background] Climate and land-use changes, which include the application of various types of organic and inorganic fertilisers, have been reducing the species diversity of Mediterranean grasslands and threatening their conservation. Annual plants are one of the most diverse functional groups of species in these grasslands, despite suffering competitive pressure from perennial herbaceous and woody species, and they are essential for ecosystem functioning and stability., [Aims] To quantify how fertilisation modulates the impact of plant-to-plant interactions and climate fluctuations on the dynamics of annuals in Mediterranean grasslands. We hypothesised that the application of sewage sludge would increase competition between functional groups, reducing the abundance of annuals in the long-term, but would buffer the negative impacts of drought on the year-to-year fluctuation of the diversity of annuals., [Methods] In a semi-natural species-rich Mediterranean grassland in northern Spain, we analysed the changes in the taxonomical and functional composition and diversity of annuals over 14 years in response to variations in the abundance of perennial herbaceous and woody species, climate fluctuations, and fertilisation with sewage sludge. We quantified separately the patterns of year-to-year fluctuations and long-term trends., [Results] The frequency and diversity of annuals decreased with a higher abundance of perennial herbaceous species, drought in June, and cold winters. The addition of sewage sludge decreased the abundance of annuals in the long-term, seemed to promote competition between annuals and other functional groups at an interannual scale, and mitigated the negative effects of drought and cold., [Conclusions] Fertilisation influences differently the temporal response of annuals to climate fluctuations and plant-to-plant interactions.

Understanding the mechanisms underlying community stability has become an urgent need in order to protect ecosystems from global change and resulting biodiversity loss. While community stability can be influenced by richness, synchrony in annual fluctuations of species, species stability and functional traits, the relative contributions of these drivers to stability are still unclear. In semi-natural grasslands, land-use changes such as fertilization might affect stability by decreasing richness and influencing year-to-year fluctuations. In addition, they can promote long-term directional trends, shifting community composition and influencing grassland maintenance. Thus, it is important to consider how species and community stability vary year-to-year but also in the long term. Using a 14-year vegetation time series of a species-rich semi-natural Mediterranean grassland, we studied the relative importance of richness, synchrony, species stability and functional traits on community stability. To assess land-use change effects on stability, we applied a fertilization treatment. To distinguish stability patterns produced by year-to-year fluctuations from those caused by long-term trends, we compared the results obtained using a detrending approach from those without detrending. Stability is influenced by richness, synchrony and functional traits. Fertilization decreases species and community stability by promoting long-term trends in species composition, favouring competitive species and decreasing richness. Studying stability at the community and species level, and accounting for the effect of trends is essential to understand stability and its drivers more comprehensively.

There is large uncertainty on the future of European silver fir (Abies alba) forests and how they will respond to climate warming. Many studies, such as Walder et al. (2021), suggest they will show positive growth trends and a larger distribution area in response to warmer conditions whenever summer drought is not severe and long. This contradicts localized dieback events in continental, drought-prone Mediterranean sites mainly located in the southwestern margin of the current species distribution range. We discuss these disagreements and propose compound climate extremes and elevated atmospheric drought as drivers of dieback in similar marginal silver fir populations.

Tree-ring data has been widely used to inform about tree growth responses to drought at the individual scale, but less is known about how tree growth sensitivity to drought scales up driving changes in forest dynamics. Here, we related tree-ring growth chronologies and stand-level forest changes in basal area from two independent data sets to test if tree-ring responses to drought match stand forest dynamics (stand basal area growth, ingrowth, and mortality). We assessed if tree growth and changes in forest basal area covary as a function of spatial scale and tree taxa (gymnosperm or angiosperm). To this end, we compared a tree-ring network with stand data from the Spanish National Forest Inventory. We focused on the cumulative impact of drought on tree growth and demography in the period 1981–2005. Drought years were identified by the Standardized Precipitation Evapotranspiration Index, and their impacts on tree growth by quantifying tree-ring width reductions. We hypothesized that forests with greater drought impacts on tree growth will also show reduced stand basal area growth and ingrowth and enhanced mortality. This is expected to occur in forests dominated by gymnosperms on drought-prone regions. Cumulative growth reductions during dry years were higher in forests dominated by gymnosperms and presented a greater magnitude and spatial autocorrelation than for angiosperms. Cumulative drought-induced tree growth reductions and changes in forest basal area were related, but initial stand density and basal area were the main factors driving changes in basal area. In drought-prone gymnosperm forests, we observed that sites with greater growth reductions had lower stand basal area growth and greater mortality. Consequently, stand basal area, forest growth, and ingrowth in regions with large drought impacts was significantly lower than in regions less impacted by drought. Tree growth sensitivity to drought can be used as a predictor of gymnosperm demographic rates in

Understanding the mechanisms underlying community stability has become an urgent need in order to protect ecosystems from global change and resulting biodiversity loss. While community stability can be influenced by richness, synchrony in annual fluctuations of species, species stability and functional traits, the relative contributions of these drivers to stability are still unclear. In semi-natural grasslands, land-use changes such as fertilization might affect stability by decreasing richness and influencing year-to-year fluctuations. In addition, they can promote long-term directional trends, shifting community composition and influencing grassland maintenance. Thus, it is important to consider how species and community stability vary year-to-year but also in the long term. Using a 14-year vegetation time series of a species-rich semi-natural Mediterranean grassland, we studied the relative importance of richness, synchrony, species stability and functional traits on community stability. To assess land-use change effects on stability, we applied a fertilization treatment. To distinguish stability patterns produced by year-to-year fluctuations from those caused by long-term trends, we compared the results obtained using a detrending approach from those without detrending. Stability is influenced by richness, synchrony and functional traits. Fertilization decreases species and community stability by promoting long-term trends in species composition, favouring competitive species and decreasing richness. Studying stability at the community and species level, and accounting for the effect of trends is essential to understand stability and its drivers more comprehensively., [Methods] Study site and experimental design: In 2003, 12 plots of 15x5 m (hereafter called macro-plots) were established inside an area of 5500 m2. Half of the macro-plots (six) were used as control plots and half were fertilized with sewage sludge in a single event in 2003, applying manually to the soil surface 5 kg m-2 . The sludge came from a municipal urban wastewater treatment plant located in Tudela (Navarra, Spain), and it was sludge previously dried to 28% dry matter by centrifugation. To accurately assess vegetation changes, a 1x1 m permanent plot was placed in the centre of each macro-plot. Every year for 14 consecutive years (from 2004 to 2017), at the end of June, vegetation was sampled by R. Ibáñez, who identified and recorded every vascular plant species present in each of the permanent plots. The 1x1 m permanent plots were divided into 100 10x10 cm subplots to measure species abundance (frequency) by counting the number of 10x10 cm subplots in which the species was present (presence was recorded if shoots overlapped with the sampling unit/subplot, not according to rooted plants). Richness, synchrony and stability measures: Species richness in each permanent plot was measured both as cumulative species richness, counting the number of species found at least once in a permanent plot during the 14 years of the study, and as mean species richness, averaging the number of species found in a permanent plot over the 14 years (Lepš et al., 2018). Community-level synchrony for each permanent plot was calculated using the log variance ratio index (“Logvar”), which is the log-transformation of the ratio of observed to expected variance (i.e. the ratio of variance of the total community abundance to the sum of variances of the abundance of each species; Lepš et al., 2018; Roscher et al., 2011). Stability at both the community and the species level was calculated as the inverse of the coefficient of variation (CV-1) across years of cumulative or individual specie

Climatic warming is assumed to expand the geographic range of insect pests whose distribution is mainly constrained by low temperatures. This is the case of the pine processionary moth (Thaumetopoea pityocampa), which is one of the main conifer defoliators in the Mediterranean Basin. Warmer winters may lead to a northward/upward expansion of this insect, as short-term studies have shown. However, no long-term data, i.e. spanning at least one century, has been used to examine these projections. We test the hypotheses that climatic warming (i) has caused an upward shift of the pine processionary moth, and (ii) has increased the frequency of severe defoliations. We used dendrochronological methods to reconstruct defoliations over the period 1900–2006 in 14 sites spanning a wide altitudinal range (1070–1675 m) in Teruel, eastern Spain. We built local ring-width chronologies for four co-occurring pine species with different degree of susceptibility against the moth defoliations, from highly suitable or palatable species (Pinus nigra) to moderately (Pinus sylvestris, Pinus halepensis) or rarely defoliated species (Pinus pinaster). We validated the tree-ring reconstructions of outbreaks using a field record of stand defoliations spanning the period 1971–2006. Outbreaks in the most affected P. nigra stands corresponded to abrupt one- to two-year growth reductions (70–90% growth loss). Reconstructed outbreaks occurred on average every 9–14 years. The growth memory of outbreaks was weaker but lasted longer (1–6 years) than that due to droughts (1–3 years). Neither an upward expansion nor an increase in outbreak frequency was observed. Severe PPM defoliations did not increase as climate warmed, rather they were positively related to the winter North Atlantic Oscillation.

1. Understanding the mechanisms underlying community stability has become an urgent need to protect ecosystems from global change and resulting biodiversity loss. While community stability can be influenced by species richness, synchrony in annual fluctuations of species, species stability and functional traits, the relative contributions of these drivers to stability are still unclear. In semi-natural grasslands, land-use changes such as fertilization might affect stability by decreasing richness and influencing year-to-year fluctuations. In addition, they can promote long-term directional trends, shifting community composition and influencing grassland maintenance. Thus, it is important to consider how species and community stability vary year-to-year but also in the long term., 2. Using a 14-year vegetation time series of a species-rich semi-natural Mediterranean grassland, we studied the relative importance of richness, synchrony, species stability and functional traits on community stability. To assess land-use change effects on stability, we applied a fertilization treatment. To distinguish stability patterns produced by year-to-year fluctuations from those caused by long-term trends, we compared the results obtained using a detrending approach from those without detrending., 3. Independently of the treatment and approach applied, the most stable communities were those composed of asynchronous species with low specific leaf area. Fertilization decreased year-to-year and long-term community stability by increasing community-weighted mean of specific leaf area, decreasing species stability or also reducing richness in the case of year-to-year stability. Additionally, traits such as seed mass had an indirect effect on stability through synchrony. Long-term trends appeared in control and fertilized plots (due to fertilization), decreasing community and species stability and leading to differences in the relationships found between community stability and some of its drivers. This reflects the importance of accounting for the effect of temporal trends on community and species stability using both a long-term and a year-to-year approach., 4. Synthesis. Stability is influenced by richness, synchrony and functional traits. Fertilization decreases species and community stability by promoting long-term trends in species composition, favouring competitive species and decreasing richness. Studying stability at the community level and species level, and accounting for the effect of trends is essential to understand stability and its drivers more comprehensively.

Drought is an important driver of forest dynamics in the Mediterranean region. The forecasted increase in drought frequency and severity can notably influence tree growth, forest structure, composition and productivity. Understanding how coexisting tree species respond to drought is thus crucial to understand which are less vulnerable and will perform better in a warmer and drier world. To assess drought vulnerability, we used dendrochronology to study the radial growth trends and responses to a drought index of four pine species (Pinus halepensis, Pinus pinea, Pinus nigra, and Pinus sylvestris) coexisting in North-eastern Spain. We reconstructed the growth of each species and evaluated their short- and long-term growth response to drought for the common period 1980–2017. The growth of the four pine species depended on water availability and high early spring temperatures impacted the growth of P. nigra and P. sylvestris negatively. The occurrence of a severe drought between 2005 and 2007 lead to marked growth reductions in the four species, but it was greater in magnitude in P. pinea and P. halepensis in 2005, and in P. nigra in 2007. The results of basal area increment models at the individual tree level suggested that P. halepensis trees grow more than the rest of species. After accounting for age and drought effects, P. nigra and P. sylvestris displayed negative growth trends in the 2008–2017 period while P. pinea and P. halepensis displayed positive growth trends. P. sylvestris was the most resistant species and P. pinea the less resistant. Conversely, P. halepensis and P. pinea were slightly more resilient than P. sylvestris. Moreover, P. sylvestris was the species displaying the highest autocorrelation and the lowest coefficient of variation in ring-width indices. A marked drop in the autocorrelation of P. pinea ring-width index was observed in response to the 2005 drought. These results indicate that all study species are vulnerable to drought but in differe

Drought is an important driver of forest dynamics in the Mediterranean region. The forecasted increase in drought frequency and severity can notably influence tree growth, forest structure, composition and productivity. Understanding how coexisting tree species respond to drought is thus crucial to understand which are less vulnerable and will perform better in a warmer and drier world. To assess drought vulnerability, we used dendrochronology to study the radial growth trends and responses to a drought index of four pine species (Pinus halepensis, Pinus pinea, Pinus nigra, and Pinus sylvestris) coexisting in North-eastern Spain. We reconstructed the growth of each species and evaluated their short- and long-term growth response to drought for the common period 1980–2017. The growth of the four pine species depended on water availability and high early spring temperatures impacted the growth of P. nigra and P. sylvestris negatively. The occurrence of a severe drought between 2005 and 2007 lead to marked growth reductions in the four species, but it was greater in magnitude in P. pinea and P. halepensis in 2005, and in P. nigra in 2007. The results of basal area increment models at the individual tree level suggested that P. halepensis trees grow more than the rest of species. After accounting for age and drought effects, P. nigra and P. sylvestris displayed negative growth trends in the 2008–2017 period while P. pinea and P. halepensis displayed positive growth trends. P. sylvestris was the most resistant species and P. pinea the less resistant. Conversely, P. halepensis and P. pinea were slightly more resilient than P. sylvestris. Moreover, P. sylvestris was the species displaying the highest autocorrelation and the lowest coefficient of variation in ring-width indices. A marked drop in the autocorrelation of P. pinea ring-width index was observed in response to the 2005 drought. These results indicate that all study species are vulnerable to drought but in differe

There is debate on which tree species can sustain forest ecosystem services in a drier and warmer future. In Europe, the use of non-native timber species, such as Douglas fir (Pseudotsuga menziesii [Mirb.] Franco), is suggested as a solution to mitigate climate change impacts because of their high growth resilience to drought. However, the biogeographical, climatic and ecological limits for widely planted timber species still need to be defined. Here, we study the growth response to climate variables and drought of four Douglas fir plantations in northern Spain subjected to contrasting climate conditions. Further, we measure wood density in one of the sites to obtain a better understanding of growth responses to climate. Correlative analyses and simulations based on the Vaganov–Shaskin process-based model confirm that growth of Douglas fir is constrained by warm and dry conditions during summer and early autumn, particularly in the driest study site. Minimum wood density increased in response to dry spring conditions. Therefore, planting Douglas fir in sites with a marked summer drought will result in reduced growth but a dense earlywood. Stands inhabiting dry sites are vulnerable to late-summer drought stress and can act as “sentinel plantations”, delineating the tolerance climate limits of timber species.

Climate change can lead to the simultaneous occurrence of extreme droughts and heat waves increasing the frequency of compound events with unknown impacts on forests. Here we use two independent datasets, a compiled database of tree drought mortality events and the ICP-Forest level I plots, to study the impacts of the simultaneous occurrence of hot summers, with elevated vapour pressure deficit (VPD), and dry years on forest defoliation and mortality across Europe. We focused on tree drought mortality and background mortality rates, and we studied their co-occurrence with compound events of hot summers and dry years. In total, 143 out of 310 mortality events across Europe, i.e. 46% of cases, corresponded with rare compound events characterized by hot summers and dry years. Over the past decades, summer temperature increased in most sites and severe droughts resulted in compound events not observed before the 1980s. From the ICP-Forest plots we identified 291 (1718 trees) and 61 plots (128 trees) where severe defoliation and mortality, respectively, were caused by drought. The analyses of these events showed that 34% and 27% of the defoliation and mortality cases corresponded with rare compound climate events, respectively. Background mortality rates across Europe in the period 1993–2013 presented higher values in regions where summer temperature and VPD more steeply rose, where drought frequency increased. The steady increase in summer temperatures and VPD in Southern and Eastern Europe may favor the occurrence of compound events of hot summers and dry conditions. Giving that both, local and intense tree drought mortality events and background forest mortality rates, are linked to such compound events we can expect an increase in forest drought mortality in these European regions over the next decades.

Wood formation during the growing season is shaped by the intra-annual variability of climatic conditions. In the Mediterranean, the cambial activity is seasonally constrained by winter low temperature and summer drought, resulting in bimodal growth patterns. Although bimodal growth is an ecologically important adaptation of woody species to seasonally dry environments, its variability across space and among species remains unclear. We combined direct monitoring of intra-annual radial growth using dendrometers and xylogenesis micro-sampling with indirect wood formation modeling to assess growth bimodality for two coexisting oak species across six sites in North-Eastern Spain. We focused on species with two contrasting life strategies, evergreen with diffuse-porous wood (Quercus ilex) and winter deciduous with ring-porous wood (Q. faginea/Q. pubescens). Study sites spanned from oceanic areas with high autumn precipitation to dry continental inland regions. We hypothesized that growth bimodality reflects the interaction between local precipitation patterns and species-specific traits. Our results revealed cambial activity in autumn after summer quiescence across the entire region and for both oak species. However, growth bimodality and the relative contribution of the autumn growth peak to the total annual growth was geographically structured and species-specific. Growth was unimodal under continental and Atlantic conditions. By contrast, growth bimodality was significant along the Mediterranean coast, where precipitation showed a bimodal pattern. Moreover, evergreen Q. ilex showed more growth bimodality compared with deciduous Q. faginea/Q. pubescens at Mediterranean sites. Different intra-annual distribution of radial growth could facilitate coexistence of both oak species through temporal niche partitioning. The growth bimodality helps to compensate for reduced growth in summer by autumn cambial resumption and, therefore, the differences in its intensity between si

Ongoing climatic change is threatening the survival of drought-sensitive tree species, such as silver fir (Abies alba). Drought-induced dieback had been previously explored in this conifer, although the role played by tree-level genetic diversity and its relationship with growth patterns and soil microsite conditions remained elusive. We used double digest restriction-site-associated DNA sequencing (ddRADseq) to describe different genetic characteristics of five silver fir forests in the Spanish Pyrenees, including declining and non-declining trees. Single nucleotide polymorphisms (SNPs) were used to investigate the relationships between genetics, dieback, intraspecific trait variation (functional dendrophenotypic traits and leaf traits), local bioclimatic conditions, and rhizosphere soil properties. While there were no noticeable genetic differences between declining and non-declining trees, genome–environment associations with selection signatures were abundant, suggesting a strong influence of climate, soil physicochemical properties, and soil microbial diversity on local adaptation. These results provide novel insights into how genetics and diverse environmental factors are interrelated and highlight the need to incorporate genetic data into silver fir forest dieback studies to gain a better understanding of local adaptation.

Land-use and climate change are leading to more frequent and intense tree damage and mortality events. Drought-induced tree mortality is occurring worldwide leading to broad-scale events, but the spatial patterns of tree damage and mortality, their underlying drivers and their variation over time is largely unknown. We investigated the spatial patterns of tree damage and mortality across Mediterranean forests of the Iberian Peninsula, the underlying effects of competition and climate, and how the spatial patterns and relationships with underlying drivers changed over time. We used the Spanish Forest Inventory to analyse the autocorrelation in tree damage and mortality across forest types, hurdle-gamma models to quantify the effect of competition and climate on tree damage and mortality, and cross-correlograms to assess their spatial dependence and its change over time. We observed a greater magnitude and a stronger autocorrelation in tree damage than mortality, with positive aggregation up to 20 kilometres. There was a spatial dependence between tree damage and mortality with their drivers, with spatial aggregation increasing with water availability, drought intensity and competition. The spatial dependence of tree damage and mortality with the underlying drivers increased over time, particularly for drought intensity. Our results suggest that the combined effect of intense competition and drought could favour more extensive die-off and tree mortality events, providing key information for identifying vulnerable areas and the planning of adaptation measures.

[Background] Plasticity in response to environmental drivers can help trees cope with droughts. However, our understanding of the importance of plasticity and physiological adjustments in trees under global change is limited., [Methods] We used the International Tree-Ring Data Bank (ITRDB) to examine 20th century growth responses in conifer trees during (resistance) and following (resilience) years of severe soil and atmospheric droughts occurring in isolation or as compound events. Growth resilience indices were calculated using observed growth divided by expected growth to avoid spurious correlations, in which the expected values were obtained by the autoregressive moving average (ARIMA) model. We used high atmospheric vapour pressure deficit (VPD) to select years of atmospheric drought and low annual values of the Standardized Precipitation-Evapotranspiration Index (SPEI) to select years with soil drought. We acquired the sensitivities (i.e., the slopes of the relationships) by fitting the resilience indices as a function of environmental drivers, and assessed how these sensitivities changed over time for different types of drought events using linear mixed models. We also checked whether plasticity in growth responses was sufficient to prevent long-term trends of growth reductions during or after severe droughts. We acknowledge that by focusing on the response of surviving trees from the ITRDB we are potentially biasing our results towards higher resilience, as stand level responses (e.g., mortality) may result in lowered competition after the disturbance event., [Results] Sensitivities of resilience to VPD and SPEI changed throughout the 20th century, with the directions of these changes often reversing in the second half of the century. For the 1961–2010 period, changing sensitivities had positive effects on resilience, especially following years of high-VPD and compound events, avoiding growth losses that would have occurred if sensitivities had remained constant. Despite sensitivity changes, resilience was still lower at the end of the 20th century compared to the beginning of the century., [Conclusions] Future adjustments to low-SPEI and high-VPD events are likely to continue to compensate for the trends in climate only partially, leading to further generalized reductions in tree growth of conifers. An improved understanding of these plastic adjustments and their limits, as well as potential compensatory processes at the stand level, is needed to project forest responses to climate change.

Assessing tree growth patterns and deviations from expected climate baselines across wide environmental gradients is fundamental to determine forest vulnerability to drought. This need is particularly compelling for the southernmost limit of the tree species distribution where hot droughts often trigger forest dieback processes. This is the case of some silver fir (Abies alba) populations located in southwestern Europe (Spanish Pyrenees) which present ongoing dieback processes since the 1980s. We sampled 21 silver fir stands showing different dieback intensity, assessed using defoliation levels, quantified their growth patterns and characterized their responses to climate. Then, we assessed growth deviations from climatic predictions using the process-based Vaganov-Shashkin (VS) growth model. The forests showing most intense dieback, i.e. highest defoliation levels, were mainly located in low-elevation sites of the western Pyrenees. Trees in these stands displayed the lowest growth rates and the highest year-to-year variability in growth and their growth was limited by late-summer evaporative demand. In eastern and central Pyrenees, we detected a mild growth limitation by low soil moisture during the late growing season and positive growth recovery in recent years with respect to a climate baseline. Decreasing growth trajectories were the most common pattern, while rising trends were common in stands with low dieback in eastern and central Pyrenees. Our results portend systematic spatial variability of growth trends across the Pyrenean silver fir populations forming the south-western distribution limit of the species in Europe. Decoupling of growth between eastern and western populations observed in the recent decades suggests contrasting vulnerability to climate change, and more importantly, the decoupling of growth patterns in western clusters could be used as an early-warning signal of impending dieback. Consequently, we foresee future dieback events to have more

The efficient conservation of vulnerable ecosystems in the face of global change requires a complete understanding of how plant communities respond to various environmental factors. We aim to demonstrate that a combined use of different approaches, traits, and indices representing each of the taxonomic and functional characteristics of plant communities will give complementary information on the factors driving vegetation assembly patterns. We analyzed variation across an environmental gradient in taxonomic and functional composition, richness, and diversity of the herb-layer of a temperate beech-oak forest that was located in northern Spain. We measured species cover and four functional traits: leaf dry matter content (LDMC), specific leaf area (SLA), leaf size, and plant height. We found that light is the most limiting resource influencing herb-layer vegetation. Taxonomic changes in richness are followed by equivalent functional changes in the diversity of leaf size but by opposite responses in the richness of SLA. Each functional index is related to different environmental factors even within a single trait (particularly for LDMC and leaf size). To conclude, each characteristic of a plant community is influenced by different and even contrasting factors or processes. Combining different approaches, traits, and indices simultaneously will help us understand how plant communities work.

The efficient conservation of vulnerable ecosystems in the face of global change requires a complete understanding of how plant communities respond to various environmental factors. We aim to demonstrate that a combined use of different approaches, traits, and indices representing each of the taxonomic and functional characteristics of plant communities will give complementary information on the factors driving vegetation assembly patterns. We analyzed variation across an environmental gradient in taxonomic and functional composition, richness, and diversity of the herb-layer of a temperate beech-oak forest that was located in northern Spain. We measured species cover and four functional traits: leaf dry matter content (LDMC), specific leaf area (SLA), leaf size, and plant height. We found that light is the most limiting resource influencing herb-layer vegetation. Taxonomic changes in richness are followed by equivalent functional changes in the diversity of leaf size but by opposite responses in the richness of SLA. Each functional index is related to different environmental factors even within a single trait (particularly for LDMC and leaf size). To conclude, each characteristic of a plant community is influenced by different and even contrasting factors or processes. Combining different approaches, traits, and indices simultaneously will help us understand how plant communities work.

Alpine treelines are expected to shift upward due to climate warming, whereas warmer conditions can have negative impacts on forests located near the xeric, equatorward limit of the distribution of tree species (rear edge). We compare tree populations forming the distribution limits of mountain pine (Pinus uncinata) in north-eastern Spain: two cold-limited alpine treeline populations, and a rear-edge drought-prone stand. In the treelines, trees were sampled in three altitudinal belts to evaluate within-site variations considering the forest, the transitional ecotone and the treeline. Tree growth was markedly higher in one of the treeline sites (Tesso), particularly in the forest belt, as compared to the other treeline (Las Cutas), and to the warmest rear-edge stand. Tree growth in the rear-edge population depended on June precipitation. Trees presented also comparatively smaller leaves with higher Specific Leaf Area (SLA) there than in the two treelines. Regarding the differences between altitudinal belts within the treelines, treeline soils showed lower N concentrations and a lower content of sand than in forests. Soil microbiota was dominated by bacteria in the treeline and by fungi in the forest, although with differences between sites. Soil characteristics and microbial composition were strongly related, whereas its relationship with tree growth and functional traits was less clear. The marked differences in leaf traits and growth response to climate found when comparing treeline and rear-edge sites highlights the biogeographical uniqueness of sites forming the equatorward distribution limit. The greater tree growth in the Tesso treeline as compared to Las Cutas indicate that interactions between climate and physical and chemical properties of the soil influence tree growth and its interactions with soil microbial communities in the treeline. A better understanding of plant-soil feedbacks may help to understand the future dynamics of tree populations forming the

Basal area increment and climate data over the period 1951-2015 for three pine species (Pinus pinaster, P. nigra and P. sylvestris) in the Iberian Northern Plateau

Forest ecosystems are increasingly exposed to the combined pressure of climate change and attacks by pests and pathogens. These stress factors can threaten already vulnerable species triggering dieback and rising defoliation and mortality rates. To characterize abiotic (drought, climate warmings) and biotic (pathogens) risks and their spatiotemporal patterns we quantified the recent loss of vitality for the endangered and relict Abies pinsapo forests from Andalusia, south-eastern Spain. Abies pinsapo is an iconic Mediterranean fir showing a high vulnerability to drought stress and also to several pests (Cryphalus numidicus) and root rot fungi (Armillaria mellea). We analyzed a monitoring network dataset of radial growth, defoliation and mortality from 2001 to 2017 including 1025 trees situated in three major mountain ranges (Sierra de Grazalema, Sierra de las Nieves, and Sierra Bermeja). We fitted several statistical models to determine the main drivers of changes in defoliation, a proxy of tree vigor, and mortality. Defoliation and mortality rates were much higher towards the East of the study area, mirroring the gradient from Atlantic to Mediterranean climatic conditions. In the most affected stands tree defoliation increased in response to a combination of long and severe droughts, with attacks by the beetle C. numidicus. Mortality rates increased in response to a higher defoliation rate, a lower relative radial-growth rate, long and severe droughts and a higher incidence of A. mellea. Our findings illustrate the value of monitoring networks recording changes in forest health to quantify and forecast future vulnerability of threatened tree species.

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

We analyzed the impacts of drought severity on a variety of sectors in a topographically complex basin (the upper Aragón basin 2181 km2) in the Central Spanish Pyrenees. Using diverse data sources including meteorological and hydrological observations, remote sensing and tree rings, we analyze the possible hydrological implications of drought occurrence and severity on water availability in various sectors, including downstream impacts on irrigation water supply for crop production. Results suggest varying responses in forest activity, secondary growth, plant phenology, and crop yield to drought impacts. Specifically, meteorological droughts have distinct impacts downstream, mainly due to water partitioning between streamflow and irrigation channels that transport water to crop producing areas. This implies that drought severity can extend beyond the physical boundaries of the basin, with impacts on crop productivity. This complex response to drought impacts makes it difficult to develop objective basin-scale operational definitions for monitoring drought severity. Moreover, given the high spatial variability in responses to drought across sectors, it is difficult to establish reliable drought thresholds from indices that are relevant across all socio-economic sectors. The anthropogenic impacts (e.g. water regulation projects, ecosystem services, land cover and land use changes) pose further challenges to assessing the response of different systems to drought severity. This study stresses the need to consider the seasonality of drought impacts and appropriate drought time scales to adequately assess and understand their complexity.

Snow dynamics are key to understanding tree growth in mountain forests and future response to climate change. However, precise monitoring of microclimate conditions and variables related to tree growth and functioning are lacking. To advance on those issues, snow cover and microclimate conditions, tree phenology, xylogenesis, intra-annual radial growth and the concentration of sapwood and needle non-structural carbohydrates were intensively monitored in four Pinus uncinata forests along an altitudinal gradient over three years in a Pyrenean valley (NE Spain). Snow dynamics exerted strong influence on soil temperature and moisture, particularly before and during the early growing season. Soil temperature was the most relevant microclimate variable during the overall xylogenesis, mainly influencing the production of mature tracheids. Large snow accumulation resulted in later snow depletion and a consequent delay in soil warming onset. Low soil temperatures in the spring, related to prolonged snow persistence, retarded cambial reactivation and led to lower growth rate. Despite strong spatial variability among plots, wood production was determined by snow dynamics in three out of the four studied plots. This study highlights the major role played by early and late growing season soil temperatures on radial growth of mountain conifers. The results of this study suggest that a future shallower and more transitory snowpack in the studied forests, together with warmer soil and air temperatures, may increase radial growth and productivity of similar mid-latitude, young mountain forests.

Mixed forests are expected to be more productive and resilient against disturbances than pure forests. However, tree to tree interactions are complex and vary depending on tree characteristics and multiple site features including soil properties. Such complexity is not normally accounted for in studies of mixing effects on tree response to drought. Here, we evaluate if neighbourhood tree identity influence soil biotic and abiotic characteristics in three mature, mixed Silver fir (Abies alba) forests. Further, we investigate the relationships between radial growth response to drought, tree characteristics (growth rate, size and competition pressure), tree neighbourhood, and soil physico-chemical and biological properties. Patterns of covariation between tree size, competition pressure, soil microbial composition, and growth recovery after drought were observed among different neighbourhood types. Slow-growing, small Silver fir trees experiencing high competitive pressure were associated with higher proportions of Scots pine (Pinus sylvestris) in the neighbourhood, high soil C:N ratios, and showed a faster growth recovery after drought. A neighbourhood dominated by broadleaf species (mainly European beech, Fagus sylvatica) was associated with high biomass of arbuscular mycorrhizal fungi, and low biomass of Gram-positive bacteria and Eukaryote, together with lower tree growth recovery. Therefore, coexistence with other Silver fir, Scots pine or European beech trees is associated to different covariation patterns of tree, soil and drought performance variables of the target Silver fir trees. Finally, our study remarks that combining tree, soil and neighbour variables at individual level helps to understand patterns of tree growth and growth response to drought under different stand mixtures

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.

Dieback in temperate forests is understudied, despite this biome is predicted to be increasingly affected by more extreme climate events in a warmer world. To evaluate the potential drivers of dieback we reconstructed changes in radial growth and intrinsic water-use efficiency (iWUE) from stable isotopes in tree rings. Particularly, we compared tree size, radial-growth trends, growth responses to climate (temperature, precipitation, cloudiness, number of foggy days) and drought, and changes in iWUE of declining and non-declining trees showing contrasting canopy dieback and defoliation. This comparison was done in six temperate forests located in northern Spain and based on three broadleaved tree species (Quercus robur, Quercus humilis, Fagus sylvatica). Declining trees presented lower radial-growth rates than their non-declining counterparts and tended to show lower growth variability, but not in all sites. The growth divergence between declining and non-declining trees was significant and lasted more in Q. robur (15–30 years) than in F. sylvatica (5–10 years) sites. Dieback was linked to summer drought and associated atmospheric patterns, but in the wettest Q. robur sites cold spells contributed to the growth decline. In contrast, F. sylvatica was the species most responsive to summer drought in terms of growth reduction followed by Q. humilis which showed coupled changes in growth and iWUE as a function of tree vigour. Low growth rates and higher iWUE characterized declining Q. robur and F. sylvatica trees. However, declining F. sylvatica trees became less water-use efficient close to the dieback onset, which could indicate impending tree death. In temperate forests, dieback and growth decline can be triggered by climate extremes such as dry and cold spells, and amplified by climate warming and rising drought stress.

Climate warming and the occurrence of more severe dry spells are causing widespread drought-induced forest die-off events. Despite research on drought-triggered die-off processes is rapidly increasing, little is known on how soil conditions and rhizosphere features are affected by canopy dieback and tree death. We studied the soils in the rhizosphere of three coniferous forests where die-off was induced by a severe drought in 2012. We selected three forest types subjected to contrasting climatic and edaphic conditions dominated by three different tree species: silver fir (Abies alba; temperate conditions), Scots pine (Pinus sylvestris; continental and Mediterranean conditions) and Aleppo pine (Pinus halepensis; semi-arid and Mediterranean conditions). In each forest, we analyzed soil physical characteristics such as water retention capacity and soil texture, nutrient availability and microbial community structure (Phospholipid fatty acids, PLFA) below non-declining and declining or dying trees. We did not observe differences in nutrient availability between the two vigor classes. Conversely, we found strong differences in soil microbial community structure below non-declining and declining trees in the Silver fir and Aleppo pine stands. Soils in the Scots pine stand presented extremely low values of soil saturated sorptivity indicating a reduction of soil water infiltration after prolonged dry periods which could exacerbate drought stress. We conclude that forest dieback impacts the soil microbial community structure in the short term. Further research is required to understand the linkages between a reduced capacity of soil water infiltration after prolonged droughts, short-term changes in the soil microbiota, long-term nutrient imbalances and tree death. Soil conditions shall be considered as an important part of forest management strategies after drought-induced die off.

The negative impacts of drought on forest growth and productivity last for several years generating legacies, although the factors that determine why such legacies vary across sites and tree species remain unclear. We used an extensive network of tree‐ring width (RWI, ring‐width index) records of 16 tree species from 567 forests, and high‐resolution climate and normalized difference vegetation index (NDVI) datasets across Spain during the common period 1982‒2008 to test the hypothesis that climate conditions and growth features modulate legacy effects of drought on forests. Legacy effects of drought were calculated as the differences between detrended‐only RWI and NDVI series (i.e. after removing long‐term growth trends) and pre‐whitened RWI and NDVI series predicted by a model including drought intensity. Superposed Epoch Analysis (SEA) was used to estimate whether legacy effects differed from random. Finally, legacy effects were related to water balance, growth persistence and variability, and tree species identity. We found a widespread occurrence of drought legacy effects on both RWI and NDVI, but they were seldom significant. According to SEA, first‐year drought legacies were negative and different from random in 9% and 5% of the RWI and NDVI series respectively. The number of significant second‐ and third‐year legacies was substantially lower. Differences between RWI and NDVI legacies indicate that canopy greenness and radial growth responses to drought are decoupled. We found variations in legacies between tree species with gymnosperms presenting larger first‐year drought legacies than angiosperms, which were exposed to less severe droughts. Greater growth variability can explain the presence of first‐year RWI legacies in gymnosperms from dry sites despite that the relationship between growth variability and legacies was complex. Synthesis. Accounting for species and site responses to drought provides a better understanding of the magnitude and durat

Systematic forest networks of health monitoring have been established to follow changes in tree vigor and mortality. These networks often lack long-term growth data, but they could be complemented with tree ring data, since both defoliation and radial growth are proxies of changes in tree vigor. For instance, a severe water shortage should reduce growth and increase tree defoliation in drought-prone areas. However, the effects of climatic stress and drought on growth and defoliation could also depend on tree age. To address these issues, we compared growth and defoliation data with recent climate variability and drought severity in Abies pinsapo old and young trees sampled in Southern Spain, where a systematic health network (Andalucía Permanent Plot Network) was established. Our aims were: (i) to assess the growth sensitivity of old and young A. pinsapo trees and (ii) to test if relative changes in radial growth were related with recent defoliation, for instance, after severe droughts. We also computed the resilience indices to quantify how old and young trees recovered growth after recent droughts. Wet-cool conditions during the prior autumn and the current early summer improved the growth of old trees, whereas late-spring wet conditions enhanced the growth of young trees. Old trees were more sensitive to wet and sunny conditions in the early summer than young trees. Old and young trees were more responsive to the Standardized Precipitation-Evapotranspiration Index drought index of June–July and July–August calculated at short (one–three months) and mid (three–six months) time scales, respectively. Old trees presented a higher resistance to a severe drought in 1995 than young trees. A positive association was found between stand defoliation and relative growth. Combining monitoring and tree ring networks is useful for the detection of early warning signals of dieback in similar drought-prone forests.

This study links tree-ring growth and gross primary production for a variety of forest types under different environmental conditions across Spain. NOAA-AVHRR satellite imagery data were combined with dendrochronological records and climate data at a fine spatial resolution (1.21 km2) to analyze the interannual variability of tree-ring growth and vegetation activity for different forest biomes from 1981 to 2015. Specifically, we assessed the links between tree-ring width indices (TRWi), the Normalized Difference Vegetation Index (NDVI) and a variety of environmental conditions, represented by climatic variables (air temperature, precipitation, evapotranspiration and water balance) and elevation. The impact of these variables on tree growth was assessed by means of the Predictive Discriminant Analysis (PDA). Results reveal a general positive and significant relationship between inter-annual variability of the NDVI at a high spatial resolution (1.21 km2) and tree-ring growth. Maximum correlations between NDVI and tree-ring growth were recorded when cumulative NDVI values were considered, in some cases covering long time periods (6–10 months), suggesting that tree growth is mainly related to Gross Primary Production (GPP) at annual scale. The relationship between tree-ring growth and inter-annual variability of the NDVI, however, strongly varies between forest types and environmental conditions.

Climate warming is responsible for observed reduction in snowpack depth and an earlier and faster melt-out in many mountains of the Northern Hemisphere. Such changes in mountain hydroclimate could negatively affect productivity and tree growth in high-elevation forests, but few studies have investigated how and where recent warming trends and changes in snow cover influence forest growth. A network comprising 36 high-elevation Pinus uncinata forests was sampled in the NE Iberian Peninsula, mainly across the Spanish Pyrenees, using dendrochronology to relate tree radial growth to a detailed air temperature and snow depth data. Radial growth was negatively influenced by a longer winter snow season and a higher late-spring snowpack depth. Notably, the effect of snow on tree growth was found regardless the widely reported positive effect of growing-season air temperatures on P. uncinata growth. No positive influence of moisture from spring snowmelt on annual growth of P. uncinata was detected in sampled forests. Tall trees showed a lower growth responsiveness to snow than small trees. Decreasing trends in winter and spring snow depths were detected at most Pyrenean forests, suggesting that the growth of high-elevation P. uncinata forests can beneficiate for a shallower and of shorter duration snowpack associated with warmer conditions. However, water-limited sites located on steep slopes or on rocky substrates, with poor soil-water holding capacity, could experience drought stress because of early depleted snow-related soil moisture.