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

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

Climate and Soil Microsite Conditions Determine Local Adaptation in Declining Silver Fir Forests.

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.

Climate change-related growth improvements in a wide niche-breadth tree species across contrasting environments.

BACKGROUND AND AIMS: The vulnerability and responsiveness of forests to drought are immensely variable across biomes. Intraspecific tree responses to drought in species with wide niche breadths that grow across contrasting climatically environments might provide key information regarding forest resistance and changes in species distribution under climate change. Using a species with an exceptionally wide niche breath, we tested the hypothesis that tree populations thriving in dry environments are more resistant to drought than those growing in moist locations. METHODS: We determined temporal trends in tree radial growth of 12 tree populations of Nothofagus antarctica (Nothofagaceae) located across a sharp precipitation gradient (annual precipitation of 500-2000 mm) in Chile and Argentina. Using dendrochronological methods, we fitted generalized additive mixed-effect models to predict the annual basal area increment as a function of year and dryness (De Martonne aridity index). We also measured carbon and oxygen isotope signals (and estimated intrinsic water-use efficiency) to provide potential physiological causes for tree growth responses to drought. KEY RESULTS: We found unexpected improvements in growth during 1980-1998 in moist sites, while growth responses in dry sites were mixed. All populations, independent of site moisture, showed an increase in their intrinsic water-use efficiency in recent decades, a tendency that seemed to be explained by an increase in the photosynthetic rate instead of drought-induced stomatal closure, given that δ18O did not change with time. CONCLUSIONS: The absence of drought-induced negative effects on tree growth in a tree species with a wide niche breadth is promising because it might relate to the causal mechanisms tree species possess to face ongoing drought events. We suggest that the drought resistance of N. antarctica might be attributable to its low stature and relatively low growth rate.

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

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

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

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

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

Tree growth response to drought partially explains regional-scale growth and mortality patterns in Iberian forests.

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 terms of stand basal area growth and ingrowth at regional scales, but further studies may try to disentangle how initial stand density modulates such relationships. Drought-induced growth reductions and their cumulative impacts have strong potential to be used as early-warning indicators of regional forest vulnerability.

Wood density and hydraulic traits influence species' growth response to drought across biomes.

Tree species display a wide variety of water-use strategies, growth rates and capacity to tolerate drought. However, if we want to forecast species capacity to cope with increasing aridity and drought, we need to identify which measurable traits confer resilience to drought across species. Here, we use a global tree ring network (65 species; 1931 site series of ring-width indices-RWI) to evaluate the relationship of long-term growth-drought sensitivity (RWI-SPEI drought index relationship) and short-term growth response to extreme drought episodes (resistance, recovery and resilience indices) with functional traits related to leaf, wood and hydraulic properties. Furthermore, we assess the influence of climate (temperature, precipitation and climatic water deficit) on these trait-growth relationships. We found a close correspondence between the long-term relationship between RWI and SPEI and resistance and recovery of tree growth to severe drought episodes. Species displaying a stronger RWI-SPEI relationship to drought and low resistance and high recovery to extreme drought episodes tended to have a higher wood density (WD) and more negative leaf minimum water potential (Ψmin). Such associations were largely maintained when accounting for direct climate effects. Our results indicate that, at a cross-species level and global scale, wood and hydraulic functional traits explain species' growth responses to drought at short- and long-term scales. These trait-growth response relationships can improve our understanding of the cross-species capacity to withstand climate change and inform models to better predict drought effects on forest ecosystem dynamics.

Pine processionary moth outbreaks cause longer growth legacies than drought and are linked to the North Atlantic Oscillation.

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.

Compound climate events increase tree drought mortality across European forests.

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.

Mediterranean old-growth forests exhibit resistance to climate warming.

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

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

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

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

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

Climate sensitivity and drought seasonality determine post-drought growth recovery of Quercus petraea and Quercus robur in Europe.

Recent studies have identified strong relationships between delayed recovery of tree growth after drought and tree mortality caused by subsequent droughts. These observations raise concerns about forest ecosystem services and post-drought growth recovery given the projected increase in drought frequency and extremes. For quantifying the impact of extreme droughts on tree radial growth, we used a network of tree-ring width data of 1689 trees from 100 sites representing most of the distribution of two drought tolerant, deciduous oak species (Quercus petraea and Quercus robur). We first examined which climatic factors and seasons control growth of the two species and if there is any latitudinal, longitudinal or elevational trend. We then quantified the relative departure from pre-drought growth during droughts, and how fast trees were able to recover the pre-drought growth level. Our results showed that growth was more related to precipitation and climatic water balance (precipitation minus potential evapotranspiration) than to temperature. However, we did not detect any clear latitudinal, longitudinal or elevational trends except a decreasing influence of summer water balance on growth of Q. petraea with latitude. Neither species was able to maintain the pre-drought growth level during droughts. However, both species showed rapid recovery or even growth compensation after summer droughts but displayed slow recovery in response to spring droughts where none of the two species was able to fully recover the pre-drought growth-level over the three post-drought years. Collectively, our results indicate that oaks which are considered resilient to extreme droughts have also shown vulnerability when droughts occurred in spring especially at sites where long-term growth is not significantly correlated with climatic factors. This improved understanding of the role of drought seasonality and climate sensitivity of sites is key to better predict trajectories of post-drought growth recovery in response to the drier climate projected for Europe.

Tree growth and treeline responses to temperature: Different questions and concepts.

Climate warming is expected to enhance tree growth at alpine treelines. A higher growth rate is forecasted as temperatures rise and growth becomes less dependent on the temperature rise. Since radial growth is just one component of treeline dynamics those forecasts do not necessarily apply to treeline elevation or latitude; treelines can shift upward or poleward or remain stable.

Run to the hills: Forest growth responsiveness to drought increased at higher elevation during the late 20th century.

Climate warming is expected to enhance forest growth in cold-limited biomes while triggering reductions in drought-limited biomes. However, as temperature raises, it is unclear how temperature- and drought-growth couplings shift across elevation gradients in different biomes. We still lack comprehensive analyses on how altitude modulates the influence of temperature and drought on tree growth during the second half of the 20th century when climate warming accelerated. We compared the worldwide responses of tree growth (RWI, ring-width indices) to two of its major climatic constraints, growing-season minimum temperatures and drought (SPEI index), across biomes and elevation gradients during two periods with different warming rates (1960-1980 vs. 1980-2000). We found a decrease in the correlations of minimum temperatures with growth, but a strengthening of drought-growth relationships. However, these patterns varied across biomes because correlations between growth and temperature decreased in temperate forests and woodland shrubland, while correlations between growth and SPEI increased in boreal forests and decreased in temperate forests. Differences in growth responsiveness to climate between the two periods were more marked for mid-latitude forests situated between 1200 and 1600 m. The slopes of the relationships between growth-temperature correlations and elevation decreased in late spring and midsummer. The slopes of the relationships between growth-drought correlations and elevation increased in temperate forests and woodland shrubland suggesting that drought impacts are "climbing" in these biomes. Temperature controls on forest growth are relaxing as the climate warms, while drought is becoming a more significant constraint for tree growth, particularly for mid-elevation forests and in drought-prone woodland and shrubland. The strengthening of drought-growth coupling should be considered in vegetation models to reduce the uncertainty on forest climate mitigation.

Global fading of the temperature-growth coupling at alpine and polar treelines.

Climate warming is expected to positively alter upward and poleward treelines which are controlled by low temperature and a short growing season. Despite the importance of treelines as a bioassay of climate change, a global field assessment and posterior forecasting of tree growth at annual scales is lacking. Using annually resolved tree-ring data located across Eurasia and the Americas, we quantified and modeled the relationship between temperature and radial growth at treeline during the 20th century. We then tested whether this temperature-growth association will remain stable during the 21st century using a forward model under two climate scenarios (RCP 4.5 and 8.5). During the 20th century, growth enhancements were common in most sites, and temperature and growth showed positive trends. Interestingly, the relationship between temperature and growth trends was contingent on tree age suggesting biogeographic patterns in treeline growth are contingent on local factors besides climate warming. Simulations forecast temperature-growth decoupling during the 21st century. The growing season at treeline is projected to lengthen and growth rates would increase and become less dependent on temperature rise. These forecasts illustrate how growth may decouple from climate warming in cold regions and near the margins of tree existence. Such projected temperature-growth decoupling could impact ecosystem processes in mountain and polar biomes, with feedbacks on climate warming.

Evidence of non-stationary relationships between climate and forest responses: Increased sensitivity to climate change in Iberian forests.

Climate and forest structure are considered major drivers of forest demography and productivity. However, recent evidence suggests that the relationships between climate and tree growth are generally non-stationary (i.e. non-time stable), and it remains uncertain whether the relationships between climate, forest structure, demography and productivity are stationary or are being altered by recent climatic and structural changes. Here we analysed three surveys from the Spanish Forest Inventory covering c. 30 years of information and we applied mixed and structural equation models to assess temporal trends in forest structure (stand density, basal area, tree size and tree size inequality), forest demography (ingrowth, growth and mortality) and above-ground forest productivity. We also quantified whether the interactive effects of climate and forest structure on forest demography and above-ground forest productivity were stationary over two consecutive time periods. Since the 1980s, density, basal area and tree size increased in Iberian forests, and tree size inequality decreased. In addition, we observed reductions in ingrowth and growth, and increases in mortality. Initial forest structure and water availability mainly modulated the temporal trends in forest structure and demography. The magnitude and direction of the interactive effects of climate and forest structure on forest demography changed over the two time periods analysed indicating non-stationary relationships between climate, forest structure and demography. Above-ground forest productivity increased due to a positive balance between ingrowth, growth and mortality. Despite increasing productivity over time, we observed an aggravation of the negative effects of climate change and increased competition on forest demography, reducing ingrowth and growth, and increasing mortality. Interestingly, our results suggest that the negative effects of climate change on forest demography could be ameliorated through forest management, which has profound implications for forest adaptation to climate change.

Long-term nutrient imbalances linked to drought-triggered forest dieback.

Drought-induced forest dieback is causing reductions in productivity, increasing tree mortality and impairing terrestrial carbon uptake worldwide. However, the role played by long-term nutrient imbalances during drought-induced dieback is still unknown. To improve our knowledge on the relationships between dieback and nutrient imbalances, we analysed wood anatomical traits (tree-ring width and wood density), soil properties and long-term chemical information in tree-ring wood (1900-2010) by non-destructive Micro X-ray fluorescence (µXRF) and destructive (ICP-OES) techniques. We studied two major European conifers with ongoing drought-induced dieback in mesic (Abies alba, silver fir) and xeric (Pinus sylvestris, Scots pine) sites. In each site we compared coexisting declining (D) and non-declining (ND) trees. We used dendrochronology and generalized additive and linear mixed models to analyse trends in tree-ring nutrients and their relationships with wood traits. The D trees presented lower growth and higher minimum wood density than ND trees, corresponding to a smaller lumen area of earlywood tracheids and thus a lower theoretical hydraulic conductivity. These differences in growth and wood-anatomy were more marked in silver fir than in Scots pine. Moreover, most of the chemical elements showed higher concentrations in D than in ND trees during the last two-five decades (e.g., Mn, K and Mg), while Ca and Na increased in the sapwood of ND trees. The Mn concentrations, and related ratios (Ca:Mn, Mn:Al and P:Mn) showed the highest differences between D and ND trees for both tree species. These findings suggest that a reduced hydraulic conductivity, consistent with hydraulic impairment, is affecting the use of P in D trees, making them more prone to drought-induced damage. The retrospective quantifications of Mn ratios may be used as early-warning signals of impending dieback.

Post-drought Resilience After Forest Die-Off: Shifts in Regeneration, Composition, Growth and Productivity.

A better understanding on the consequences of drought on forests can be reached by paying special attention to their resilience capacity, i.e., the ability to return to a state similar to pre-drought conditions. Nevertheless, extreme droughts may surpass the threshold for the resilience capacity triggering die-off causing multiple changes at varying spatial and temporal scales and affecting diverse processes (tree growth and regeneration, ecosystem productivity). Combining several methodological tools allows reaching a comprehensive characterization of post-drought forest resilience. We evaluated the changes in the abundance, regeneration capacity (seedling abundance), and radial growth (annual tree rings) of the main tree species. We also assessed if drought-induced reductions in growth and regeneration of the dominant tree species scale-up to drops in vegetation productivity by using the Normalized Difference Vegetation Index (NDVI). We studied two conifer forests located in north-eastern Spain which displayed drought-induced die-off during the last decades: a Scots pine (Pinus sylvestris) forest under continental Mediterranean conditions and a Silver fir (Abies alba) forest under more temperate conditions. We found a strong negative impact of a recent severe drought (2012) on Scots pine growth, whereas the coexisting Juniperus thurifera showed positive trends in basal area increment (0.02 ± 0.003 cm2 yr-1). No Scots pine recruitment was observed in sites with intense die-off, but J. thurifera and Quercus ilex recruited. The 2012 drought event translated into a strong NDVI reduction (32% lower than the 1982-2014 average). In Silver fir we found a negative impact of the 2012 drought on short-term radial growth, whilst long-term growth of Silver fir and the coexisting Fagus sylvatica showed positive trends. Growth rates were higher in F. sylvatica (0.04 ± 0.003 cm2 yr-1) than in A. alba (0.02 ± 0.004 cm2 yr-1). These two species recruited beneath declining and non-declining Silver fir trees. The 2012 drought translated into a strong NDVI reduction which lasted until 2013. The results presented here suggest two different post-drought vegetation pathways. In the Scots pine forest, the higher growth and recruitment rates of J. thurifera correspond to a vegetation shift where Scots pine is being replaced by the drought-tolerant juniper. Conversely, in the Silver fir forest there is an increase of F. sylvatica growth and abundance but no local extinction of the Silver fir. Further research is required to monitor the evolution of these forests in the forthcoming years to illustrate the cumulative impacts of drought on successional dynamics.