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Global warming modulates the spatial and temporal occurrence of insect outbreaks, leading to as-yet-unknown effects on forests ecosystems. Warmer trends may favour the upward and northward expansion of pests, albeit increased exposure to heat events and droughts may also hamper insect growth and development. Thus far, further research is needed about the balance of climate warming on pine processionary moth (PPM) (Thaumetopoea pityocampa) development stages, especially at broader scales, despite being the main conifer-defoliating insect in the Mediterranean area. This research analyses the relationships of PPM with the climatic conditions during its most vulnerable development phases in Spain to provide base-information for forest managers. We hypothesize that warmer winter temperatures will increase PPM growth at their larval feeding stage, whereas summer-heat events will reduce PPM rates from egg and early larval stages, leading to counterbalanced effects of climate warming. Furthermore, we expect dry springs will allow an earlier development of PPM and higher incidence of outbreaks. To analyse climate effects on PPM populations in Andalusia, Castilla-La Mancha, and Navarre (37 to 42°N), we compiled information from the Regional Forest Services comprising up to 26 years of monitoring data from 2465 forest stands dominated by Pinus nigra and Pinus sylvestris. We fitted Cumulative Link Mixed Models (CLMM) to test the climate effects on four PPM defoliation severity levels. Our results support higher PPM larvae survival and incidence due to warmer winter conditions, but also the reduction of PPM population size under summer heat conditions. Particularly, the increasing incidence associated to warmer winters during the larval defoliating stage is counterbalanced by the decreasing population at earlier life stages during summer heat events. Spring drought also plays an important role on PPM incidence rates, but its effect is idiosyncratic to each region, suggesting an

Defoliating insects’ outbreaks play a critical role in trees’ carbon cycle. The pine processionary moth (PPM; Thaumetopoea pityocampa) is the major defoliating insect of Mediterranean coniferous forests. The frequency and intensity of PPM outbreaks is projected to increase as winter temperatures become milder due to climate warming. An accurate evaluation of this projection requires a wide spatial baseline of the historical PPM incidence. PPM outbreaks affect tree secondary growth leading to narrow rings, providing a tree level signal. However, PPM defoliation rings can be confounded with drought rings, the most frequent cause of narrow rings in Mediterranean environments. Thus, an accurate identification of PPM rings demands the consideration of additional tree ring traits. Here, we introduce a multiproxy approach to identify and distinguish PPM and drought events. We sampled four Pinus nigra (3) and P. sylvestris (1) stands in Spain. We identified and verified years of PPM defoliation using remote sensing analysis and field observations of Regional Forest Service. We identified drought events through the combination of climatic data with radial growth reductions. We considered climate growth residuals, among-trees growth variability, latewood percentage (%LW), intrinsic water-use efficiency (iWUE) and minimum blue intensity (BI) to discern between droughts and PPM outbreaks. In comparison with drought rings, PPM rings showed 1) more negative residuals in climate growth models, 2) higher secondary growth variance, 3) higher percentage of latewood and 4) lower iWUE. Minimum BI did not differ between drought and PPM rings, but was lower than in the rest of the rings. The combination of these traits provides a signature to identify PPM rings, opening the opportunity to reconstruct PPM incidence on a broad spatio-temporal scale.

Tree pollarding was a dominant management strategy of European forests for centuries creating open agroforestry landscapes with important cultural and environmental values. This traditional practice has been widely abandoned in last decades with a subsequent impact in terms of biodiversity and cultural loss. Central Spain hosts the largest and best-preserved area of pollarded narrow-leaved ash (Fraxinus angustifolia Vahl.) woodlands in Europe. The main aim of this research is to obtain rigorous historical records of pollarding frequency to get adequate information for traditional ash management. We used dendrochronological techniques to evaluate temporal changes of pollarding frequency and rotation length. We analysed the stand level synchrony and the effect of land property on pollarding activity from 322 trees growing in eight pollard stands in Central Spain. Pollarding events were unequivocally identified at tree level by a characteristic change in growth pattern. We identified 2426 tree-level pruning events with the first event dated in 1777. Historical pruning recurrence ranged between 5 and 10 years with higher pollarding frequency on private lands. Pruning events within each site were synchronous, suggesting the existence of a rotational schema within each stand. Pruning frequency decreased drastically in the 1970s matching with the depopulation of rural areas and the general abandonment of traditional practices. Pollarding practices have recovered in recent decades although with lower intensity and lacking the synchronic historical patterns. Providing technical and economic support to make this traditional activity profitably would have strong environmental revenue due the multiple ecosystem services provided by pollarded ashes.

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

Legacies of past climate conditions and historical management govern forest productivity and tree growth. Understanding how these processes interact and the timescales over which they influence tree growth is critical to assess forest vulnerability to climate change. Yet, few studies address this issue, likely because integrated long-term records of both growth and forest management are uncommon. We applied the stochastic antecedent modelling (SAM) framework to annual tree-ring widths from mixed forests to recover the ecological memory of tree growth. We quantified the effects of antecedent temperature and precipitation up to 4 years preceding the year of ring formation and integrated management effects with records of harvesting intensity from historical forest management archives. The SAM approach uncovered important time periods most influential to growth, typically the warmer and drier months or seasons, but variation among species and sites emerged. Silver fir responded primarily to past climate conditions (25–50 months prior to the year of ring formation), while European beech and Scots pine responded mostly to climate conditions during the year of ring formation and the previous year, although these responses varied among sites. Past management and climate interacted in such a way that harvesting promoted growth in young silver fir under wet and warm conditions and in old European beech under drier and cooler conditions. Our study shows that the ecological memory associated with climate legacies and historical forest management is species-specific and context-dependent, suggesting that both aspects are needed to properly evaluate forest functioning under climate change.

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

Rear-edge tree populations are experiencing a combination of higher temperatures and more intense droughts that might push individuals beyond their tolerance limits. This trend towards rising atmospheric [CO] is concurrent with an increase in intrinsic water use efficiency (iWUE), which theoretically enhances photosynthesis and decrease evapotranspiration rates, consequently improving tree resistance to drought. However, it remains unclear whether iWUE is favouring tree growth under current climate conditions, particularly when climate and iWUE legacy effects are simultaneously considered. We evaluated this question with an extensive sampling along Iberian rear-edge (dry) populations comprising four mountain ranges and two distinct altitudes. We simultaneously examined the effects of climate and iWUE on secondary growth using annually resolved basal area increments (BAIs) for the period 1901–2017. We used linear mixed models including second-order autocorrelation and 1-year legacy effects of iWUE and summer drought. BAI and iWUE increased across the studied period. iWUE increase was driven by changes in atmospheric CO concentration and water availability during the growing season. Climate and iWUE exerted direct and lagged effects on beech growth. Water availability during growing season was the main driver of tree growth, combining direct and indirect effects through its impact on iWUE. Legacy effects of water availability and iWUE were more important than growing season conditions. The net effect of iWUE shifted when lagged effects were considered, resulting in a net negative impact on tree growth. Synthesis: Our results reveal that climate and iWUE legacy effects must be considered to assess the net iWUE effect on secondary growth. Considering lagged effects, the current increase in iWUE is constraining tree growth. Modelling efforts of tree growth response to climate warming should include climate and iWUE legacy effects to adequately assess terrestrial ecosyste

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

Severe droughts have the potential to reduce forest productivity and trigger tree mortality. Most trees face several drought events during their life and therefore resilience to dry conditions may be crucial to long-term survival. We assessed how growth resilience to severe droughts, including its components resistance and recovery, is related to the ability to survive future droughts by using a tree-ring database of surviving and now-dead trees from 118 sites (22 species, >3,500 trees). We found that, across the variety of regions and species sampled, trees that died during water shortages were less resilient to previous non-lethal droughts, relative to coexisting surviving trees of the same species. In angiosperms, drought-related mortality risk is associated with lower resistance (low capacity to reduce impact of the initial drought), while it is related to reduced recovery (low capacity to attain pre-drought growth rates) in gymnosperms. The different resilience strategies in these two taxonomic groups open new avenues to improve our understanding and prediction of drought-induced mortality.

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

The effects of climate change on forest growth are not homogeneous across tree species distribution ranges because of inter-population variability and spatial heterogeneity. Although latitudinal and thermal gradients in growth patterns have been widely investigated, changes in these patterns along longitudinal gradients due to the different timing and severity of regional droughts are less studied. Here, we investigated these responses in Mediterranean Black pine (Pinus nigra Arn.). We built a tree-ring width dataset comprising 77 forests (1202 trees) across the Mediterranean Basin. The biogeographical patterns in growth patterns and the relationships between growth and mean temperature, precipitation, drought and atmospheric circulations patterns (NAO -North Atlantic Oscillation-, SOI -Southern Oscillation Index- and MOI -Mediterranean Oscillation index-) were analyzed. Then, we evaluated the spatial and temporal growth synchrony between and within east and west populations. We found different growth and climate patterns in west vs. east Black pine populations, although in both regions growth was driven by similar temperature and precipitation variables. MOI significantly influenced tree growth, whilst NAO and SOI showed weaker effects. Growth of east and west Black pine populations desynchronized after the 1970s when several and uncoupled regional droughts occurred across the Mediterranean Basin. We detected a climate shift from the 1970s to the 1980s affecting growth patterns, changing growth-climate relationships, and reducing forest growth from west to east Black pine forests. Afterwards, climate and growth of east and west populations became increasingly more divergent. Our findings imply that integral bioclimatic and biogeographical analyses across the species distribution area must be considered to adequately assess the impact of climate change on tree growth under warming and more arid conditions.

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 wa

Global warming and land use changes, contributing to landscape level fuel increments, could threatenMediterranean pine forest resilience to wildfire disturbances. Reconstructions of historicalfire regimes allow forthe disentanglement of these two drivers by comparing the influence of climatic and anthropogenic variables onfire. Here we combine three sources of historical data: charcoal accumulation rates from a peat bog, detailedhistorical records offire incidence and tree-ring width data fromfive relict black pine (Pinus nigra) forests withfire-scarred trees located in Sierra de Gredos (central Spain). We found growth suppression in 1893 and 1894 inall the sites which coincided with a peak offire incidence in historical records and an increase in charcoalaccumulation rates. The occurrence of these three synchronous events suggests increased wildfire incidence inthe area which shaped the current stand structure of relict black pine forests. These late 19th century devel-opments, we argue, can be mainly attributed to anthropogenic factors and contributing climatic drivers. Weargue that the dissolution of the“Mesta”, the biggest transhumance livestock organization in Europe lasting fromthe 13th to the 19th centuries, led to more extensive grazing and uncontrolled use of forests and grasslandswhich likely contributed to increased wildfire incidence. Additionally, 1893 was characterized by anomalouslywarm spring temperatures which may have facilitated vegetationflammability. Our approach couples humanand climate systems as drivers of historicalfire incidence in Mediterranean pine forests.

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

Severe droughts have the potential to reduce forest productivity and trigger tree mortality. Most trees face several drought events during their life and therefore resilience to dry conditions may be crucial to long-term survival. We assessed how growth resilience to severe droughts, including its components resistance and recovery, is related to the ability to survive future droughts by using a tree-ring database of surviving and now-dead trees from 118 sites (22 species, '3,500 trees). We found that, across the variety of regions and species sampled, trees that died during water shortages were less resilient to previous non-lethal droughts, relative to coexisting surviving trees of the same species. In angiosperms, drought-related mortality risk is associated with lower resistance (low capacity to reduce impact of the initial drought), while it is related to reduced recovery (low capacity to attain pre-drought growth rates) in gymnosperms. The different resilience strategies in these two taxonomic groups open new avenues to improve our understanding and prediction of drought-induced mortality. © 2020, The Author(s).

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

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.

Tree mortality is a key driver of forest dynamics and its occurrence is projected to increase in the future due to climate change. Despite recent advances in our understanding of the physiological mechanisms leading to death, we still lack robust indicators of mortality risk that could be applied at the individual tree scale. Here, we build on a previous contribution exploring the differences in growth level between trees that died and survived a given mortality event to assess whether changes in temporal autocorrelation, variance, and synchrony in time-series of annual radial growth data can be used as early warning signals of mortality risk. Taking advantage of a unique global ring-width database of 3065 dead trees and 4389 living trees growing together at 198 sites (belonging to 36 gymnosperm and angiosperm species), we analyzed temporal changes in autocorrelation, variance, and synchrony before tree death (diachronic analysis), and also compared these metrics between trees that died and trees that survived a given mortality event (synchronic analysis). Changes in autocorrelation were a poor indicator of mortality risk. However, we found a gradual increase in interannual growth variability and a decrease in growth synchrony in the last ~20 years before mortality of gymnosperms, irrespective of the cause of mortality. These changes could be associated with drought-induced alterations in carbon economy and allocation patterns. In angiosperms, we did not find any consistent changes in any metric. Such lack of any signal might be explained by the relatively high capacity of angiosperms to recover after a stress-induced growth decline. Our analysis provides a robust method for estimating early-warning signals of tree mortality based on annual growth data. In addition to the frequently reported decrease in growth rates, an increase in inter-annual growth variability and a decrease in growth synchrony may be powerful predictors of gymnosperm mortality risk, but not nece

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.

Post-fire forest resilience must be quantified in a long-term perspective considering changes in land-use related to fire dynamics. Historical land-use changes leading to increased wildfire severity may produce no analogue regime shifts including a loss in post-fire growth recovery. Here we reconstruct the historical fire dynamics by combining paleoecological proxies, historical fire records and tree-ring width data of relict Pinus nigra subsp. salzmannii forests in the Sierra de Gredos (central Spain). A high incidence of historical fires was recorded in the 1890s, coinciding with a peak in charcoal accumulation rates and a sharp decrease in pollen of P. nigra/Pinus sylvestris with a rapid increase of pollen of more flammable Pinus pinaster and shrubs. The shift observed in pollen assemblages, coupled with a peak in charcoal influx, support the occurrence of high-severity fires during the 1890s, when abrupt growth suppressions were observed. Trees took 2 years to recover to their pre-fire growth rates. Lasting growth-recovery periods or no growth suppression were observed in the 1920s and 1980s, when fire frequency was also high but the study sites were fragmented or protected. We documented an abrupt regime shift in the fire record during the 1890s affecting pine forests, which rapidly recovered pre-fire growth rates.

Drought is one of the key natural hazards impacting net primary production and tree growth in forest ecosystems. Nonetheless, tree species show different responses to drought events, which make it difficult to adopt fixed tools for monitoring drought impacts under contrasting environmental and climatic conditions. In this study, we assess the response of forest growth and a satellite proxy of the net primary production (NPP) to drought in peninsular Spain and the Balearic Islands, a region characterized by complex climatological, topographical, and environmental characteristics. Herein, we employed three different indicators based on in situ measurements and satellite image-derived vegetation information (i.e., tree-ring width, maximum annual greenness, and an indicator of NPP). We used seven different climate drought indices to assess drought impacts on the tree variables analyzed. The selected drought indices include four versions of the Palmer Drought Severity Index (PDSI, Palmer Hydrological Drought Index (PHDI), Z-index, and Palmer Modified Drought Index (PMDI)) and three multi-scalar indices (Standardized Precipitation Evapotranspiration Index (SPEI), Standardized Precipitation Index (SPI), and Standardized Precipitation Drought Index (SPDI)). Our results suggest that—irrespective of drought index and tree species—tree-ring width shows a stronger response to interannual variability of drought, compared to the greenness and the NPP. In comparison to other drought indices (e.g., PDSI), and our results demonstrate that multi-scalar drought indices (e.g., SPI, SPEI) are more advantageous in monitoring drought impacts on tree-ring growth, maximum greenness, and NPP. This finding suggests that multi-scalar indices are more appropriate for monitoring and modelling forest drought in peninsular Spain and the Balearic Islands.

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.

Forecasted increase drought frequency and severity may drive worldwide declines in forest productivity. Species‐level responses to a drier world are likely to be influenced by their functional traits. Here, we analyse forest resilience to drought using an extensive network of tree‐ring width data and satellite imagery. We compiled proxies of forest growth and productivity (TRWi, absolutely dated ring‐width indices; NDVI, Normalized Difference Vegetation Index) for 11 tree species and 502 forests in Spain corresponding to Mediterranean, temperate, and continental biomes. Four different components of forest resilience to drought were calculated based on TRWi and NDVI data before, during, and after four major droughts (1986, 1994–1995, 1999, and 2005), and pointed out that TRWi data were more sensitive metrics of forest resilience to drought than NDVI data. Resilience was related to both drought severity and forest composition. Evergreen gymnosperms dominating semi‐arid Mediterranean forests showed the lowest resistance to drought, but higher recovery than deciduous angiosperms dominating humid temperate forests. Moreover, semi‐arid gymnosperm forests presented a negative temporal trend in the resistance to drought, but this pattern was absent in continental and temperate forests. Although gymnosperms in dry Mediterranean forests showed a faster recovery after drought, their recovery potential could be constrained if droughts become more frequent. Conversely, angiosperms and gymnosperms inhabiting temperate and continental sites might have problems to recover after more intense droughts since they resist drought but are less able to recover afterwards.

Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1-100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark

Humans have historically played a critical role in the management of Mediterranean-type ecosystems (MTEs) through traditional fire use. Although chestnut forests are widespread across the Mediterranean Basin, little is known about their historical fire regimes. Our goal here is to generate testable hypotheses about the drivers of fire regime dynamics in chestnut dominated ecosystems. To examine anthropogenic fire management we selected two sites in Spain that have similar biophysical characteristics but divergent levels of economic development and fire management policies. Fire regime-landscape feedbacks were characterized through a pilot dendroecological study, official fire statistics, aerial photography and forest inventory data. Our results suggest that fire incidence in both sites has increased since the pre-industrial era but fire season, fire size, and forest structure have changed to a greater extent in the more developed site. These changes are probably driven by the decline in annual anthropogenic burning of litterfall by local communities at the more developed site during the non-vegetative season.

Climate warming is predicted to amplify drought stress. Thus, it is important to understand how coexisting plant species respond to severe droughts. Here we study how seven Mediterranean woody plant species with different evolutionary history and functional characteristics (Pinus halepensis Mill., Juniperus phoenicea L., Pistacia lentiscus L., Rhamnus lycioides L., Rosmarinus officinalis L., Genista scorpius (L.) DC., and Globularia alypum L.) responded to a severe winter drought during 2011-2012 in Spain. The study site is located in the Valcuerna valley, Monegros desert, northeastern Spain. We evaluated how the drought affected the annual growth-ring formation of the species by using dendrochronology and quantified the intensity of drought-induced defoliation and mortality and compared it between species and groups of species with different evolutionary history. Radial growth of all species was strongly reduced by the 2012 drought. The pre-Mediterranean species (P. halepensis, J. phoenicea, P. lentiscus and R. lycioides) reduced growth more than the Mediterranean species (R. officinalis, G. scorpius and G. alypum). Defoliation was significantly higher in pre-Mediterranean than in Mediterranean species. When species were analyzed separately we found that P. halepensis was the species with the highest growth reduction but J. phoenicea was defoliated more severely and showed higher mortality rates as a consequence of drought. In the case of the Mediterranean shrubs, drought-induced mortality was only noticeable in R. officinalis. Drought impacted growth of all species but this did not induce mortality in all of them. Growth reduction was dependent on evolutionary history. However, functional characteristics of the species such as leaf stomatal regulation and root architecture may be more important than evolutionary history on explaining drought-induced mortality. Indeed, species with shallow root systems such as J. phoenicea and R. officinalis were the most adversely

Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1–100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark

Forest dieback is one of the most widespread responses to global-change drivers, such as climate warming-related drought stress and the spread of pathogens. Although both climatic and biotic stressors have been studied separately, much less is known on how drought and pathogens interact and induce dieback, particularly in formerly used forests. We determine the roles played by each of those drivers as factors causing recent dieback in three Pyrenean silver fir stands a managed site subjected to past logging and two unmanaged sites not logged for the past 50 years. The age, size, recent competition, and basal-area increment (BAI) trends of non-declining and declining trees, and the presence of fungal pathogens were investigated. Growth patterns at yearly to decadal time scales were compared to distinguish the roles and interactions played by the different stressors. In the managed site, declining trees displayed low growth already before logging (1950-1970s). In both unmanaged sites, declining and non-declining trees displayed divergent growth patterns after extreme droughts, indicating that dieback was triggered by severe water deficit. We did not find indications that fungal pathogens are the primary drivers of dieback, since a low proportion of declining trees were infested by primary pathogens (10%). However, trees with the primary fungal pathogen Heterobasidion showed lower BAI than non-declining trees. On the other hand, the secondary fungal pathogen Amylostereum was isolated from a higher number of trees than expected by chance. These findings highlight the importance of legacies, such as the past use in driving recent forest dieback. Past forest use could predispose to dieback by selecting slow-growing trees and thus, making some them more vulnerable to drought and fungal pathogens. © 2015 Elsevier B.V.

Droughts negatively impact forests by reducing growth and increasing defoliation leading to forest dieback as the climate becomes warmer and drier. However, the timing and severity of droughts determine how differently or intensively water shortage affects primary (shoot and leaf formation) and secondary growth (stem radial growth based on tree-ring widths). We compare the impact of two severe droughts (2005, 2012), showing different climatic characteristics on the growth responses of three Mediterranean holm oak stands in northeastern Spain. We also quantify climate trends and drought severity. Then, we use remote sensing data to infer how those droughts impacted forest productivity. Both droughts were characterized by warm and dry spring conditions leading to reduced budburst, low shoot production, asynchrony in primary growth and decreased productivity and scarce radial growth, particularly in 2005. However, defoliation peaked in 2012 when radial growth showed minimum values and early spring and late summer temperatures reached maximum values. We discuss how uncoupled and resilient are the responses of primary and secondary growth to drought. Finally, these findings are used to gain insight into the drought-related drivers of defoliation in Spanish holm oak forests.

[ES] El aumento de la mortalidad y la pérdida de vitalidad de los bosques son procesos que afectan a diversas especies de árboles y biomas. Estos fenómenos de decaimiento forestal se han asociado a la interacción de múltiples factores relacionados con el cambio global, como el aumento de las sequías o la mayor incidencia de factores bióticos que generan estrés. Sin embargo, el papel que juegan estos organismos en el decaimiento del bosque en relación a la sequía no está del todo comprendido. En este trabajo caracterizamos retrospectivamente los efectos que ejercen dichos organismos sobre el crecimiento radial de árboles estresados por sequía utilizando dendrocronología. El efecto combinado del muérdago y las sequías reducen el crecimiento del pino silvestre y aumentan la defoliación, siendo estos efectos más evidentes en el ápice del árbol. La procesionaria induce pérdidas de crecimiento en pino laricio, pero en general se observa una recuperación dos años después. No encontramos un papel determinante de los hongos patógenos de raíz en el decaimiento del abeto. Finalmente, pinos carrasco muertos que presentaban ataques de insectos perforadores mostraron patrones de crecimiento similares a pinos vivos. Este estudio nos permite caracterizar algunos de los complejos y múltiples efectos de diferentes factores bióticos de estrés sobre el crecimiento y la vulnerabilidad de los bosques susceptibles a los efectos negativos de las sequías., [EN] Increasing mortality and decline of forests are affecting several tree species and biomes. These events of forest decline have been related to the interaction of several global change elements, like the increase in drought stress or a higher incidence of biotic stress factors. However, the roles played by biotic stressors have been scarcely explored as related to drought stress. In this paper we retrospectively characterize the effects of these organisms on radial growth of trees already stressed by drought using dendrochronology. The combined effect of mistletoe and droughts reduce tree growth in Scots pine and increase defoliation, being these effects more noticeable in the apex. The pine processionary moth induces growth losses in the affected stands of black pine, but they usually recover two years after the outbreak. We did not find a determinate role of root pathogens on silver fir dieback. Finally, dead Aleppo pine trees infested by bark beetles showed similar growth patterns than living pine trees. This study allows characterizing some of the complex and multiple effects of different biotic stressors on growth and vulnerability of forests subjected to the negative effects exerted by droughts.

Alpine treelines are considered monitors of the effects of climate on forest growth and dynamics. Treelines are expected to react to current climate warming by showing upslope migrations. However, treeline dynamics are often characterized by lagged responses to rising temperatures, that is, treeline inertia. In addition, encroachment within the treeline ecotone seems to be a more widespread response to climate warming than treeline ascent. We investigate how the treeline responds to climate in a Pyrenean site with an intense Mountain pine (Pinus uncinata) regeneration but also abundant dead trees. We use dendrochronology to reconstruct treeline dynamics (growth, tree recruitment, and death) and to build an age structure of Pyrenean Mountain pine forests, and relate them to temperature reconstructions of the study area. We also describe the spatial structure and estimate the size reproductive threshold of pine recruits. The study treeline showed profuse pine recruitment in the 1980s. These recruits were spatially aggregated and reached the 50% probability of reproduction at 24 years old. Most Pyrenean Mountain pines were recruited in the first half of the 18th century, a warm period when growth was stable, while old treeline trees recruited not only in those decades but also in previous warm periods. Pine deaths concentrated in the cool transition between the mid 17th and the early 18th centuries and mainly from 1820 to 1860, when growth declined as a consequence of temperatures rapidly dropping at the end of the Little Ice Age. Only the amount of dead pines at the treeline was negatively related to temperatures, indicating that cool periods cause high adult mortality rates and trigger long-term treeline decline. But this decline was interrupted by intense regeneration and treeline encroachment, two features that characterize recent treeline dynamics in some mountains. This concurs with the view of a rapid response of alpine treelines to climate during the late 20th

Drought and bark-beetle infestation are major and often interconnected drivers of forest dieback and tree death. These two stressors may interact and accelerate forest mortality, since warmer and drier conditions boost beetle attacks and reduce tree growth. However, the way in which drought and bark-beetle infestation interact and affect declining or dying trees is still poorly understood. To disentangle the long-term interaction between the two stressors, we quantified radial growth (basal area increment), resin production, mortality and intrinsic water-use efficiency (iWUE) inferred from wood-carbon isotope discrimination. We compared trees infested and recently killed by two abundant bark beetle species (. Orthotomicus erosus and Tomicus piniperda) with non-infested living trees in a drought-prone Aleppo pine plantation. Growth and iWUE showed similar values in infested and non-infested trees. Since bark-beetle-infested trees did not grow less than non-infested trees, our results did not support the hypothesis of higher costs of resin production at the expense of stem-wood formation. Radial growth was enhanced by cool and wet winter conditions prior to the growing season. However, infested trees showed lower growth responsiveness than did non-infested trees with respect to this climatic driver of growth. Infested trees also showed a lower resin-duct production two years prior to death than non-infested trees. The growth responsiveness to climate should be characterized in bark-beetle-infested trees, since a weak correlation between climate and growth can be regarded as a predisposing factor of infestation-induced tree death. Such reduced responsiveness to climate stress could be linked to the tree vulnerability to beetle attacks in drought-prone forests. © 2015 Elsevier B.V.

Aim: Climate change is expected to modify growth trends of forests around the world. However, this modification may vary in strength and intensity across a species' biogeographical range. Here, we study European populations of silver fir (Abies alba) across its southern distribution limits in Spain, Italy and Romania. We hypothesized that growth trends of silver fir will differ across its distribution range, with a marked decline in growth in drought-prone regions near the species' southernmost biogeographical limits. Location: Europe (Spain, Italy, Romania). Methods: We collected tree-ring data from at least 1300 silver fir trees located in 111 sites. The dataset was used to assess and model growth trends, quantified as changes in basal area increment, and to determine how growth responds to climate. Results: We found contrasting patterns of basal area increments among countries and sites. Populations of silver fir located outside the Mediterranean area (e.g. northern Italy, Romania) have shown a clear increase in growth over the last two decades, whereas most populations in Spain and southern Italy have displayed a marked decline in growth since the 1980s. The growth of silver fir forests at the south-western distribution limit is severely constrained by low spring-summer water availability, whereas growth of silver fir forests in non-Mediterranean areas is limited by cold conditions in late winter to early spring. Main conclusions: Climate warming is distinctly modifying growth patterns and responses to climate in silver fir across most of the species' European distribution area. In south-western Europe the reduction in growth of many populations is related to an observed increase in aridity, whereas in more temperate areas warming is enhancing growth. Our results confirm a decline in the growth of silver fir at its south-western distribution limits as a consequence of climate warming. © 2015 John Wiley & Sons Ltd.

Human beings have used fire as an ecosystem management tool for thousands of years. In the context of the scientific and policy debate surrounding potential climate change adaptation and mitigation strategies, the importance of the impact of relatively recent state fire-exclusion policies on fire regimes has been debated. To provide empirical evidence to this ongoing debate we examine the impacts of state fire-exclusion policies in the chestnut forest ecosystems of two geographically neighbouring municipalities in central Spain, Casillas and Rozas de Puerto Real. Extending the concept of 'Traditional Ecological Knowledge' to include the use of fire as a management tool as 'Traditional Fire Knowledge' (TFK), we take a mixed-methods and interdisciplinary approach to argue that currently observed differences between the municipalities are useful for considering the characteristics of >pre-industrial anthropogenic fire regimes> and their impact on chestnut forest ecosystems. We do this by examining how responses from interviews and questionnaire surveys of local inhabitants about TFK in the past and present correspond to the current biophysical landscape state and recent fire activity (based on data from dendrochronological analysis, aerial photography and official fire statistics). We then discuss the broader implications of TFK decline for future fire management policies across Europe particularly in light of the published results of the EU sponsored Fire Paradox research project. In locations where TFK-based >pre-industrial anthropogenic fire regimes> still exist, ecosystem management strategies for adaptation and mitigation to climate change could be conceivably implemented at a minimal economic and political cost to the state by local communities that have both the TFK and the adequate social, economic and cultural incentives to use it. © 2015 Elsevier Ltd.

Scots pine forests subjected to continental Mediterranean climates undergo cold winter temperatures and drought stress. Recent climatic trends towards warmer and drier conditions across the Mediterranean Basin might render some of these pine populations more vulnerable to drought-induced growth decline at the Southernmost limit of the species distribution. We investigated how cold winters and dry growing seasons drive the radial growth of Scots pine subject to continental Mediterranean climates by relating growth to climate variables at local (elevational gradient) and regional (latitudinal gradient) scales. Local climate-growth relationships were quantified on different time scales (5-, 10- and 15-days) to evaluate the relative role of elevation and specific site characteristics. A negative water balance driven by high maximum temperatures in June (low-elevation sites) and July (high-elevation sites) was the major constraint on growth, particularly on a 5- to 10-day time scale. Warm nocturnal conditions in January were associated with wider rings at the high-elevation sites. At the regional scale, Scots pine growth mainly responded positively to July precipitation, with a stronger association at lower elevations and higher latitudes. January minimum temperatures showed similar patterns but played a secondary role as a driver of tree growth. The balance between positive and negative effects of summer precipitation and winter temperature on radial growth depends on elevation and latitude, with low-elevation populations being more prone to suffer drought and heat stress; whereas, high-elevation populations may be favoured by warmer winter conditions. This negative impact of summer heat and drought has increased during the past decades. This interaction between climate and site conditions and local adaptations is therefore decisive for the future performance and persistence of Scots pine populations in continental Mediterranean climates. Forecasting changes in the Scot

The ample distribution of common beech (Fagus sylvatica) across Europe implies that this key tree species occurs under a broad variety of climatic conditions despite its sensitivity to drought stress. Iberian beech rear-edge (southernmost) forests are located along the boundary between the Eurosiberian and Mediterranean biogeographical regions. Therefore, those forests are considered to be sensitive monitors of the effects of warming-induced drought stress on marginal tree populations. We evaluate if the radial growth of Iberian beech populations is mainly constrained by drought. Since previous findings indicated that El Niño-Southern Oscillation (ENSO) teleconnections may influence the rainfall regime in northern Spain, we also assessed if beech response to drought and water availability is modulated by this large-scale climatic pattern. We compared the recent growth patterns and responses to climate across a network of 30 tree-ring site chronologies established throughout northern Spain where beech forests are subjected to contrasting climatic conditions. Iberian beech populations located near or within the Mediterranean biogeographical region were the most sensitive to June water deficit. However, the dependency of beech growth on drought stress near the rear-edge of the species was mitigated where cloudy conditions prevail in summer, namely in mesic stands located in the Eurosiberian region. Drought stress in the latter populations was alleviated by cloudiness, which in turn depended on ENSO, and this effect on growth has been intensifying for the last decades. We prove that the sensitivity of rear-edge populations to drought, in terms of growth reduction, is greatly modulated by local or regional environmental gradients, but also by the influence of large-scale climatic variation.

Some disturbances can drive ecological systems to abrupt shifts between alternative stages (tipping points) when critical transitions occur. Drought-induced tree death can be considered as a nonlinear shift in tree vigour and growth. However, at what point do trees become predisposed to drought-related dieback and which factors determine this (tipping) point? We investigated these questions by characterizing the responses of three tree species, silver fir (Abies alba), Scots pine (Pinus sylvestris) and Aleppo pine (Pinus halepensis), to a severe drought event. We compared basal area increment (BAI) trends and responses to climate and drought in declining (very defoliated and dying) vs. non-declining (slightly or not defoliated) trees by using generalized additive mixed models. Defoliation, BAI and sapwood production were related to functional proxies of tree vigour measured at the onset and end of the drought (non-structural carbohydrate concentrations, needle N content and C isotopic discrimination, presence of wood-inhabiting fungi). We evaluated whether early warning signals (increases in synchronicity among trees or in autocorrelation and standard deviation) could be extracted from the BAI series prior to tree death. Declining silver fir and Scots pine trees showed less growth than non-declining trees one to three decades, respectively, before the drought event, whereas Aleppo pines showed growth decline irrespective of tree defoliation. At the end of the drought period, all species showed increased defoliation and a related reduction in the concentration of sapwood soluble sugars. Defoliation was constrained by the BAI of the previous 5 years and sapwood production. No specific wood-inhabiting fungi were found in post-drought declining trees apart from blue-stain fungi, which extensively affected damaged Scots pines. Declining silver firs showed increases in BAI autocorrelation and variability prior to tree death. Synthesis. Early warning signals of drought-tri

The Périgord black truffle is an exclusive culinary delicacy, but its Mediterranean harvests have declined, despite cultivation efforts since the 1970s. The role of long-term irrigation, symbiotic fungus-host interaction, and microbial belowground progression remain poorly understood, because generally too short experimental settings miss the necessary degree of real world complexity and reliable information from truffle orchards is limited. Here, we conduct the first dendrochronological and wood anatomical assessment of 295 holm oaks, which have been growing under different irrigation intensities in the world's largest truffle orchard in Spain. The relationships between different climatic variables (monthly temperature means and precipitation totals) and dendro-parameters (ring width, vessel count and vessel size) of the oak hosts are utilized to disentangle direct and indirect drivers of truffle fruit body production. Irrigation at medium - instead of high - intensity is most beneficial for oak growth. Non-irrigated trees reveal overall lower stem increments. Warmer temperatures from February to April and wetter conditions from May to July enhance host vitality and possibly also its interplay with fungi symbionts via increased fine root production and mycorrhizal colonization. Adequately irrigated Mediterranean orchards may counteract some of the drought-induced natural truffle decline, and help stabilizing rural tourism, regional agriculture and global markets. © 2014 Elsevier B.V.

Warming-induced drought stress has been hypothesized as a major driver of forest net primary production (NPP) reduction, but we lack reliable field data to assess if higher temperatures lead to forest NPP reduction, particularly in humid sites and at basin to landscape spatial scales. The use of a landscape approach would allow considering the feedbacks operating between climate, topography, soil vegetation and water resources. Here we follow that approach by simulating NPP using the regional hydro-ecologic simulation system (RHESSys) model and by comparing the results with radial growth data (tree-ring widths and intrinsic water-use efficiency - iWUE). We evaluate the relationships between climate, growth, NPP, atmospheric CO2 concentrations (ca) and iWUE in xeric and mesic silver fir forests subjected to contrasting water balances. The growth data successfully validated the 11-month NPP cumulated until spring. The main negative climatic driver of growth and NPP was the summer evapotranspiration deficit, which shows a negative association with tree-ring width indices. Sensitivity analyses indicate that rising ca do not compensate the severe NPP reduction associated to warmer and drier conditions. The positive effect of rising ca on NPP is mediated by climatic site conditions being detected only in mesic sites, whereas the negative effects of drought on NPP override any ca-related enhancement of NPP in xeric sites. Future warmer and drier conditions causing a higher evaporative demand by the atmosphere could lead to a NPP decline in temperate conifer forests subjected to episodic droughts. © 2015 Elsevier B.V.