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.

Forests play a key role in the carbon balance of terrestrial ecosystems. One of the main uncertainties in global change predictions lies in how the spatiotemporal dynamics of forest productivity will be affected by climate warming. Here we show an increasing influence of climate on the spatial variability of tree growth during the last 120 y, ultimately leading to unprecedented temporal coherence in ring-width records over wide geographical scales (spatial synchrony). Synchrony in growth patterns across cold-constrained (central Siberia) and droughtconstrained (Spain) Eurasian conifer forests have peaked in the early 21st century at subcontinental scales (∼1,000 km). Such enhanced synchrony is similar to that observed in trees co-occurring within a stand. In boreal forests, the combined effects of recent warming and increasing intensity of climate extremes are enhancing synchrony through an earlier start of wood formation and a stronger impact of year-to-year fluctuations of growing-season temperatures on growth. In Mediterranean forests, the impact of warming on synchrony is related mainly to an advanced onset of growth and the strengthening of droughtinduced growth limitations. Spatial patterns of enhanced synchrony represent early warning signals of climate change impacts on forest ecosystems at subcontinental scales. U nderstanding how climate change affects forests across multiple spatiotemporal scales is important for anticipating its impacts on terrestrial ecosystems. Increases in atmospheric CO 2 concentration and shifts in phenology (1-3) could favor tree growth by enhancing photosynthesis and extending the effective growing period, respectively (4). Conversely, recent warming could increase respiration rates and, together with increasing heat and drought stresses, exert negative impacts on forest productivity (5, 6). Given the uncertainty as to what extent enhanced carbon uptake could be offset by the detrimental effects of warming on tree performance, the actual consequences of climate change on forest carbon cycling remain under debate. Notably, climate change has a stronger impact on forests constrained by climatic stressors, such as suboptimal temperatures or water shortage (7). As high-resolution repositories of biological responses to the environment, dendrochronological archives can be used to monitor this impact (8).The concept of spatial synchrony in tree growth refers to the extent of coincident changes in ring-width patterns among geographically disjunct tree populations (9). Climatic restrictions tend to strengthen growth-climate relationships, resulting in enhanced common ringwidth signals (i.e., more synchronous tree growth). Thus, regional bioclimatic patterns can be delineated by identifying groups of trees whose growth is synchronously driven by certain climatic constraints (10, 11). Previous synthesis studies have provided evidence for globally coherent multispecies responses to climate change in natural systems, including forests, with a focus on the role of increasingly warmer tempe...

Drought-triggered declines in forest productivity and associated die-off events have increased considerably due to climate warming in the last decades. There is an increasing interest in quantifying the resilience capacity of forests against climate warming and drought to uncover how different stands and tree species will resist and recover after more frequent and intense droughts. Trees form annual growth rings that represent an accurate record of how forest growth responded to past droughts. Here we use dendrochronology to quantify the radial growth of different forests subjected to contrasting climatic conditions in Spain during the last half century. Particularly, we considered four climatically contrasting areas where dominant forests showed clear signs of drought-induced dieback. Studied forests included wet sites dominated by silver fir (Abies alba) in the Pyrenees and beech (Fagus sylvatica) stands in northern Spain, and drought-prone sites dominated by Scots pine (Pinus sylvestris) in eastern Spain and black pine (Pinus nigra) in the semi-arid south-eastern Spain. We quantified the growth reduction caused by different droughts and assessed the short-and long-term resilience capacity of declining vs. non-declining trees in each forest. In all cases, drought induced a marked growth reduction regardless tree vigor. However, the capacity to recover after drought (resilience) at short-and long-term scales varied greatly between declining and non-declining individuals. In the case of beech and silver fir, non-declining individuals presented greater growth rates and capacity to recover after drought than declining individuals. For Scots pine, the resilience to drought was found to be lower in recent years regardless the tree vigor, but the growth reduction caused by successive droughts was more pronounced in declining than in non-declining individuals. In the black pine forest an extreme drought induced a marked growth reduction in declining individuals when accounting for age effects on growth rates. We demonstrate the potential of tree-ring data to record short-and long-term impacts of drought on forest growth and to quantify the resilience capacity of trees.

The negative impacts of severe drought on the growth and vigor of tree species and their relationship with forest decline have not been properly evaluated taking into account the differential responses to such stress of trees, sites and species. We evaluated these responses by quantifying the changes in radial growth of plantations of four pine species (Pinus sylvestris, Pinus nigra, Pinus pinaster, Pinus halepensis) which showed distinct decline and defoliation levels in southeastern Spain. We used dendrochronological methods, defoliation records, linear mixed models of basal area increment and dynamic factor analysis to quantify the responses of trees at the species and individual scales to site conditions and drought stress. In the region a temperature rise and a decrease in spring precipitation have led to drier conditions during the late twentieth century characterized by severe droughts in the 1990s and 2000s. As expected, the defoliation levels and the reductions in basal area increment were higher in those species more vulnerable to drought-induced xylem embolism (P. sylvestris) than in those more resistant (P. halepensis). Species adapted to xeric conditions but with high growth rates, such as P. pinaster, were also vulnerable to drought-induced decline. The reduction in basal area increment and the Climatic Change (2012) 113:767-785 defoliation events occurred after consecutive severe droughts. A decrease in spring precipitation, which is the main driver of radial growth, is the most plausible cause of recent forest decline. The sharp growth reduction and widespread defoliation of the most affected pine plantations of Scots pine make their future persistence in drought-prone sites unlikely under the forecasted warmer and drier conditions.

46Scots pine forests subjected to continental Mediterranean climates undergo cold winter 47 temperatures and drought stress. Recent climatic trends towards warmer and drier 48 conditions across the Mediterranean Basin might render some of these pine populations 49 more vulnerable to drought-induced growth decline at the southernmost limit of the 50 species distribution. We investigated how cold winters and dry growing seasons drive 51 the radial growth of Scots pine subject to continental Mediterranean climates by relating 52 growth to climate variables at local (elevational gradient) and regional (latitudinal 53 gradient) scales. Local climate-growth relationships were quantified on different time 54 scales (5-, 10-and 15-days) to evaluate the relative role of elevation and specific site 55 characteristics. A negative water balance driven by high maximum temperatures in June 56 (low-elevation sites) and July (high-elevation sites) was the major constraint on growth, 57 particularly on a 5-to 10-day time scale. Warm nocturnal conditions in January were 58 associated with wider rings at the high-elevation sites. At the regional scale, Scots pine 59 growth mainly responded positively to July precipitation, with a stronger association at 60 lower elevations and higher latitudes. January minimum temperatures showed similar 61 patterns but played a secondary role as a driver of tree growth. The balance between 62 positive and negative effects of summer precipitation and winter temperature on radial 63 growth depends on elevation and latitude, with low-elevation populations being more 64 prone to suffer drought and heat stress, whereas high-elevation populations may be 65 favoured by warmer winter conditions. This negative impact of summer heat and 66 drought has increased during the past decades. This interaction between climate and site 67 conditions and local adaptations is therefore decisive for the future performance and 68 persistence of Scots pine populations in continental Mediterranean climates.

Growth models can be used to assess forest vulnerability to climate warming. If global warming amplifies water deficit in drought-prone areas, tree populations located at the driest and southernmost distribution limits (rear-edges) should be particularly threatened. Here, we address these statements by analyzing and projecting growth responses to climate of three major tree species (silver fir, Abies alba; Scots pine, Pinus sylvestris; and mountain pine, Pinus uncinata) in mountainous areas of NE Spain. This region is subjected to Mediterranean continental conditions, it encompasses wide climatic, topographic and environmental gradients, and, more importantly, it includes rear-edges of the continuous distributions of these tree species. We used tree-ring width data from a network of 110 forests in combination with the process-based Vaganov-Shashkin-Lite growth model and climate-growth analyses to forecast changes in tree growth during the 21st century. Climatic projections were based on four ensembles CO emission scenarios. Warm and dry conditions during the growing season constrain silver fir and Scots pine growth, particularly at the species rear-edge. By contrast, growth of high-elevation mountain pine forests is enhanced by climate warming. The emission scenario (RCP 8.5) corresponding to the most pronounced warming (+1.4 to 4.8 °C) forecasted mean growth reductions of -10.7% and -16.4% in silver fir and Scots pine, respectively, after 2050. This indicates that rising temperatures could amplify drought stress and thus constrain the growth of silver fir and Scots pine rear-edge populations growing at xeric sites. Contrastingly, mountain pine growth is expected to increase by +12.5% due to a longer and warmer growing season. The projections of growth reduction in silver fir and Scots pine portend dieback and a contraction of their species distribution areas through potential local extinctions of the most vulnerable driest rear-edge stands. Our modeling approach provides accessible tools to evaluate forest vulnerability to warmer conditions.

Climate, stand structure and local site conditions are potentially important determinants of forest dynamics. Understanding the relative contributions of competition and climate to tree growth is critical to project likely stand development under different climate change scenarios. Further, current competition levels and stand structure may reflect legacies of past forest management. Here, we analyze the effects of climate, site conditions and competition on radial growth in three Scots pine plots located along an altitudinal gradient. These stands are subjected to Mediterranean climate with continental influence, i.e., growth is limited by low winter temperatures and dry summer conditions. Current stand structure and retrospective analyses of radial growth (basal area increment, BAI) were used to model changes in tree growth as a function of competition (CI) and climate at an annual resolution. Negative exponential functions characterized the CI-BAI associations, whereas linear mixed-effects models were used to model BAI and to quantify the growth response to climate of trees under low and high competition. Competition effects on growth were steady over time regardless of the elevation and tree age. High competition levels negatively affected growth, with a proportionally stronger influence in suppressed trees than in dominant trees. Sensitivity of tree growth to climate increased with decreasing competition. Altitudinal gradientrelated growth responses to climate were found only for temperature variables. Specifically, growth at high elevations was mainly limited by low winter temperatures, whilst warm spring enhanced growth at middle elevations and late summer temperatures did it at low elevations. Since growth and its sensitivity to climate is more pronounced in low competition trees, we argue that the past management of the forest overrides site conditions and climate effects through the legacies on stand structure and competition. Pro-active forest management practices should be adopted to reduce the vulnerability of previously managed Scots pine forests currently threatened by the predicted warmer and drier conditions.

Drought-and heat-driven tree mortality, along with associated insect outbreaks, have been observed globally in recent decades and are expected to increase in future climates. Despite its potential to profoundly alter ecosystem carbon and water cycles, how tree mortality scales up to ecosystem functions and fluxes is uncertain. We describe a framework for this scaling where the effects of mortality are a function of the mortality attributes, such as spatial clustering and functional role of the trees killed, and ecosystem properties, such as productivity and diversity. We draw upon remote-sensing data and ecosystem flux data to illustrate this framework and place climate-driven tree mortality in the context of other major disturbances. We find that emerging evidence suggests that climate-driven tree mortality impacts may be relatively small and recovery times are remarkably fast (4 years for net ecosystem production). We review the key processes in ecosystem models necessary to simulate the effects of mortality on ecosystem fluxes and highlight key research gaps in modeling. Overall, our results highlight the key axes of variation needed for better monitoring and modeling of the impacts of tree mortality and provide a founda- 1133tion for including climate-driven tree mortality in a disturbance framework.