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Spanish black pine showed greater resilience and resistance, but generally lower recovery to drought events in managed than in unmanaged forest stands under Mediterranean humid climate. Drought negative effects on forest ecosystems are projected to increase under global warming all over the world. In this context, forest management can be an effective option for reducing drought impacts and increasing tree growth stability to extreme drought events. Here, we aim to evaluate black pine (Pinus nigra subsp. salzmannii) growth response to climatic variability and drought events in managed and unmanaged stands under similar Mediterranean climatic conditions. Drought events were identified using long-term climatic data, and basal area increments were calculated for 100–120-year old trees cored in managed and unmanaged plots. Results showed that tree size, temperature, and the interaction between management treatment and water availability significantly influenced tree growth. Basal area increment was reduced in response to the 1983, 1991, 1994‒1995, 1999–2000 and 2005 drought events. Trees in managed plots showed lower growth reductions in response to drought than those located in unmanaged plots, probably experiencing higher competition for soil water, whereas the reverse happened under wet climate conditions. Black pines showed greater resilience and resistance, but generally lower recovery to drought events in managed than in unmanaged stands. Our results suggest that forest management enhances drought tolerance in black pine stands, which may help to ameliorate the negative impacts of global warming across Mediterranean forest ecosystems.

Significance Forests are experiencing growing risks of drought-induced mortality in a warming world. Yet, ecosystem dynamics following drought mortality remain unknown, representing a major limitation to our understanding of the ecological consequences of climate change. We provide an emerging picture of postdrought ecological trajectories based on field indicators of forest dynamics. Replacement patterns following mortality indicate limited short-term persistence of predrought dominant tree species, highlighting the potential for major ecosystem reorganization in the coming decades. The great variability of the observed dynamics within and among species reinforces the primary influence of drought characteristics and ecosystem legacies, modulated by land use, management, and past disturbances, on ongoing drought-related species turnover and their potential implications for future forest biodiversity and ecosystem services., 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.

Current knowledge of climate change effects on forest ecology and species conservation should be linked to understanding of the past-time. Abies pinsapo forests constitute a model of an endangered ecosystem, highly vulnerable to ongoing warming, whose populations have been declining for centuries, while the drivers of this local depletion trend remain poorly understood. We hypothesized that long-term disturbances, both human- and natural-induced, have shaped A. pinsapo forests, contributing to these decline processes. Until today, studies using fossil pollen record to identify past climate impacts and land-use changes on A. pinsapo populations have not been done. Here, we investigate forests’ dynamics since the late Holocene (1180 cal. AD to present) in Southern Iberian Peninsula from a fossil pollen record by comparing the results obtained with climate fluctuations and land-uses changes. The pollen sequence shows a phase of stability during the Islamic Period (~1180–1400 cal. AD; ‘Medieval Climate Anomaly’), followed by increasing degradation at Christian Period concurrent with ‘Little Ice Age’ (LIA) (ca. 1487–1530 cal. AD). The Modern Period (1530–1800 cal. AD; LIA) is linked to intensive forest management, related to the naval industry. Afterwards, a progressive reduction is recorded during the Contemporary Age period (‘Industrial Period’) until ‘Recent Warming’. In short, historical severe forest management coupled with increasing aridity since LIA appear to influence A. pinsapo forest current species composition and poor structural diversity. These disturbances might be limiting the resilience of A. pinsapo forests under a climate change scenario. A selected forest management could promote a more complex forest structure.

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. climate warming, forest limit, growth model, mountain ecosystems, tree rings acceptedVersion

Global climate change is expected to further raise the frequency and severity of extreme events, such as droughts. The effects of extreme droughts on trees are difficult to disentangle given the inherent complexity of drought events (frequency, severity, duration, and timing during the growing season). Besides, drought effects might be modulated by trees’ phenotypic variability, which is, in turn, affected by long‐term local selective pressures and management legacies. Here we investigated the magnitude and the temporal changes of tree‐level resilience (i.e., resistance, recovery, and resilience) to extreme droughts. Moreover, we assessed the tree‐, site‐, and drought‐related factors and their interactions driving the tree‐level resilience to extreme droughts. We used a tree‐ring network of the widely distributed Scots pine (Pinus sylvestris) along a 2,800 km latitudinal gradient from southern Spain to northern Germany. We found that the resilience to extreme drought decreased in mid‐elevation and low productivity sites from 1980–1999 to 2000–2011 likely due to more frequent and severe droughts in the later period. Our study showed that the impact of drought on tree‐level resilience was not dependent on its latitudinal location, but rather on the type of sites trees were growing at and on their growth performances (i.e., magnitude and variability of growth) during the predrought period. We found significant interactive effects between drought duration and tree growth prior to drought, suggesting that Scots pine trees with higher magnitude and variability of growth in the long term are more vulnerable to long and severe droughts. Moreover, our results indicate that Scots pine trees that experienced more frequent droughts over the long‐term were less resistant to extreme droughts. We, therefore, conclude that the physiological resilience to extreme droughts might be constrained by their growth prior to drought, and that more frequent and longer drought periods may overstrain their potential for acclimation., We examined tree growth resilience of Scots pine along a 2,800 km latitudinal gradient from southern Spain to north‐eastern Germany using 615 adult trees from 30 different sites. We found that the resilience of Scots pine to extreme drought decreased in mid‐elevation and low productivity sites from 1980–1999 to 2000–2011 due to more frequent and severe droughts in the later period. We showed that the impact of drought on tree‐level resilience was not dependent on its latitudinal location, but rather on the type of sites trees were growing at and on their growth performances during the pre‐drought period.

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., Resilience to drought is crucial for tree survival under climate change. Here, DeSoto et al. show that trees that died during drought were less resilient to previous dry events compared to surviving conspecifics, but the resilience strategies differ between angiosperms and gymnosperms.

12 Pags.- 34Figs.- 3 Tabls., 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 duration of drought legacies on forest growth and productivity. Despite the widespread occurrence of growth reductions in the years during and after drought occurrence, significant legacies were not very common, mostly lasted one year, and were more widespread in gymnosperms. These are relevant factors to be considered in the future when studying the consequences of drought on forest productivity and tree growth., This study was financially supported by: Xunta de Galicia, Grant/Award Number: PGIDIT06PXIB502262PR, GRC GI‐1809; INIA, Grant/Award Number: RTA2006‐00117; CANOPEE, 2014‐2020‐FEDER funds; and Spanish Science Ministry: RTI2018‐096884‐B‐C31, RTI2018‐096884‐B‐C33, AGL2017‐83828‐C2‐2R and ENV4‐CT97‐0641 projects. GSB was supported by a ‘Juan de la Cierva Formación’ grant from MINECO (FJCI 2016‐30121). This research was supported by the BERC 2018‐2021 program, and by the Spanish Ministry of Science, Innovation and Universities through the BC3 María de Maeztu excellence accreditation (MDM‐2017‐0714) and the IBERYCA (CGL2017‐84723‐P) project. R.S.S. was supported by VULBOS project (UPO‐1263216, FEDER Funds, Andalusia Regional Government, Consejería de Economía, Conocimiento, Empresas y Universidad 2014‐2020), and A.H. by PinCaR project (UHU‐1266324, FEDER Funds, Andalusia Regional Government, Consejería de Economía, Conocimiento, Empresas y Universidad 2014‐2020). K.N. was supported by the Ministry of Education, Science and Sports (Republic of Slovenia, Raziskovalci‐2.1‐UL‐BF‐ C3330‐19‐952011).

21 Pags.- 5 Figs.- 1 Appendix. - Suppl. Data., 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., This work was supported by the research projects CGL2014-52135-C03-01, CGL2015-69186-C2-1-R, CGL2015-69985-R, CGL2013-48843-C2-1-R, AGL2014-53822-C2-1-R, (BFU2010-21451), PCIN-2015-220, CGL2016-81706-REDT and CGL2017-82216-R financed by the Spanish Commission of Science and Technology and FEDER, 1560/2015: Herramientas de monitorización de la vegetación mediante modelización ecohidrológica en parques continentales financed by the Red de Parques Nacionales, IMDROFLOOD financed by the Water Works 2014 co-funded call of the European Commission, CROSSDRO financed by the AXIS (Assessment of Cross(X) – sectorial climate Impacts and pathways for Sustainable transformation) JPI-Climate co-funded call of the European Commission and INDECIS, which is part of ERA4CS, an ERA-NET initiated by JPI Climate, and funded by FORMAS (SE), DLR (DE), BMWFW (AT), IFD (DK), MINECO (ES), ANR (FR) with co-funding by the European Union (Grant 690462). Financed by the Spanish Commission of Science and Technology and FEDER, and INDECIS, which is part of ERA4CS, an ERA-NET initiated by JPI Climate, and funded by FORMAS (SE), DLR (DE), BMWFW (AT), IFD (DK), MINECO (ES), ANR (FR) with co-funding by the European Union (Grant 690462). Natalia Martin-Hernandez was supported by a doctoral grant by the Aragon Regional Government, Marina Peña-Gallardo was granted by the Spanish Ministry of Economy and Competitiveness and Miquel Tomas-Burguera was supported by a doctoral grant by the Spanish Ministry of Education, Culture and Sport. Raúl Sánchez-Salguero, Antonio Gazol and Gabriel Sangüesa-Barreda were supported by Postdoctoral grants (IJCI-2015-25845, MINECO-FPDI 2013-16600 and FJCI 2016-30121, respectively; FEDER funds). This work also benefited from funding from Xunta de Galicia (PGIDIT06PXIB502262PR, GRC GI-1809, ROCLIGAL-10MDS291009PR), INIA (RTA2006-00117), and Interreg V-A POCTEFA (CANOPEE, 2014-2020-FEDER funds) projects.

Plants in Mediterranean mountains are particularly vulnerable to climatic change. In these environments, low temperature is combined with water shortage during summer, and as a result, the positive effect of global warming theoretically expanding the growing season length may be counterbalanced by rising drought stress. These circumstances may be exacerbated in the rear edge of species distribution, where warmer conditions occur. Here, we examined the climate-growth relationships of Juniperus sabina, a major prostrate shrub above the treeline in Mediterranean mountains, to investigate climate sensitivity and long-term signals stability in four rear-edge populations from southern Spain. We demonstrate that, over recent decades, local climatic conditions have modulated the response of J. sabina secondary growth to the ongoing climate change. We observed a negative effect of winter-spring moisture on secondary growth that suggests a limitation for earlier growth activity at higher elevation, potentially hindering the ability of J. sabina to compensate forthcoming increases in summer drought. At the driest site, we also detected a positive effect of October precipitation, suggesting a second growth pulse by early autumn. Our results provide an example of how local climatic conditions may limit plastic responses of secondary growth to climatic variability. The implications of growth limitation in J. sabina exceed the species-scale level, since these prostrate shrubs play a critical role as nurse plants and local biodiversity foci in Mediterranean mountains.

Intraspecific variability in morphological and ecophysiological leaf traits might be theorized to be present in declining populations, since they seem to be exposed to stress and plasticity could be advantageous. Here we focused on declining Persian oaks (Quercus brantii Lindl. var. persica (Jaub and Spach) Zohary) in the Zagros Mountains of western Iran, representing the most important tree species of this region. We selected trees with contrasting crown dieback, from healthy to severely defoliated, to investigate the relationships between canopy dieback and leaf morphology, water content and pigments. We also measured esterase and peroxidase, as enzymatic antioxidants and indicators of contrasting genotypes. Trees showing moderate to severe defoliation showed higher leaf mass area (LMA), reduced relative water content (RWC), and lower stomatal density (SD). Increasing LMA indicates a more sclerophyllic structure, according to drier conditions. We did not find significant differences in leaf pigments (chlorophyll a and b, and carotenoids) among crown dieback classes, suggesting that Persian oak trees are able to maintain accurate photochemical efficiency, while reduced RWC and SD suggest hydraulic limitations. Our results do not provide a consistent pattern as regards enzymatic antioxidant defense in Persian oak. Morphological leaf traits would be important drivers of future adaptive evolution in Persian oak, leading to smaller and thicker leaves, which have fitness benefits in dry environments. Nonetheless, drought responses may be critically affecting carbon uptake, as photosynthetic compounds are less effectively used in leaves with higher sclerophylly.

Stand-level competition and local climate influence tree responses to increased drought at the regional scale. To evaluate stand density and elevation effects on tree carbon and water balances, we monitored seasonal changes in sap-flow density (SFD), gas exchange, xylem water potential, secondary growth, and non-structural carbohydrates (NSCs) in Abies pinsapo. Trees were subjected to experimental thinning within a low-elevation stand (1200 m), and carbon and water balances were compared to control plots at low and high elevation (1700 m). The hydraulic conductivity and the resistance to cavitation were also characterized, showing relatively high values and no significant differences among treatments. Trees growing at higher elevations presented the highest SFD, photosynthetic rates, and secondary growth, mainly because their growing season was extended until summer. Trees growing at low elevation reduced SFD during late spring and summer while SFD and secondary growth were significantly higher in the thinned stands. Declining NSC concentrations in needles, branches, and sapwood suggest drought-induced control of the carbon supply status. Our results might indicate potential altitudinal shifts, as better performance occurs at higher elevations, while thinning may be suitable as adaptive management to mitigate drought effects in endangered Mediterranean trees.

AIM: We investigate the effects of the environmental and geographical processes driving growth resilience and recovery in response to drought in Mediterranean Pinus pinaster forests. We explicitly consider how intraspecific variability modulates growth resilience to drought. LOCATION: Western Mediterranean basin. METHODS: We analysed tree rings from a large network of 48 forests (836 trees) encompassing wide ecological and climatic gradients, including six provenances. To characterize the major constraints of P. pinaster growth under extremely dry conditions, we simulated growth responses to temperature and soil moisture using a process‐based growth model coupled with the quantification of climate–growth relationships. Then, we related growth–resilience indices to provenance and site variables considering different drought events. RESULTS: Pinus pinaster displayed strong variation in growth resilience across its distributional range, but common patterns were found within each provenance. Post‐drought resilience increased with elevation and drier conditions but decreased with spring precipitation. Trees from dry sites were less resistant to drought but recovered faster than trees from wet sites. MAIN CONCLUSIONS: Resilience strategies differed among tree provenances: wet forests showed higher growth resistance to drought, while dry forests presented faster growth recovery, suggesting different impacts of climate warming on forest productivity. We detected geographically structured resilience patterns corresponding to different provenances, confirming high intraspecific variability in response to drought. This information should be included in species distribution models to simulate forest responses to climate warming and forecasted aridification.

Climate extremes are expected to increase, which will affect oak forest ecosystems in Central European areas. Intensively managed forests, such as sessile oak stands, may alter their structure and function under a warming scenario. Here we analyse and project the climate-growth relationships of sessile oak ( Quercus petraea ) from high forests, originated from seed, and coppice forests, originated from vegetative reproduction in the Czech Republic. Dendrochronological data and linear mixed-effects models show similar effects of precipitation for April–May (positive) for both seedling- and coppice-origin trees. Previous autumn and current June temperature show significant negative correlations to growth in coppice forests. Nonetheless, trees from high forest stands showed increasing drought sensitivity and tended to show a stronger response to the previous autumn’s temperature during the 20th century. The positive effect of warmer autumns, only found on high stands, might be related to the extended growing season, suggesting improved adaptive capacity to cope with impending warmer conditions. In contrast, coppice trees might be able to buffer soil water shortage during spring and summer by higher root/shoot ratio. The obtained models were used to estimate the impact of 21st century-emission scenarios on tree basal area increments and stand basal area dynamics under different stand structures. Our results support that growth responses to climate warming are age and/or structure-dependent in sessile oak, particularly with regards to coppiced stands. Converting coppice forests to high stand structures, as well as thinning high stands, would enhance sessile oak forest adaptive capacity to cope with warming in Central Europe.

Understanding forest responses to the current climate change requires to investigate the effects of competition, buffering or enhancing process of forests decline. Here we attempt to place intra-specific competition in a climate change context, using as experimental system the drought-sensitive fir Abies pinsapo. We conducted a decade-long (2004–2015) investigation at four permanent plots (two thinned plots and two controls), where all trees were identified, tagged and mapped. Experimental thinning was performed attempting to enhance stand structure diversity by (1) reducing tree competition, (2) increasing the equitability of size classes (i.e. diversity) and (3) randomizing tree spatial pattern. We focus on tree basal area increment and sap flow density, as regards microclimate and competition, to quantify the extent to that thinning alleviates drought in the remaining trees. The results support that lower competition fosters short- to long-term tree-level physiological responses, specifically, improved growth and water uptake in the remaining trees. Density reduction provides a promising strategy for minimizing climate change effects on drought-sensitive tree species by improving resources availability to the remaining trees. Nevertheless, our results also support a higher air temperature and soil water depletion according to density reduction, suggesting that lower competition might also increase soil and understory evapotranspiration. Adaptive management, as the experimental thinning reported here, may increase tree-level resources availability, but the long-term stand-level effectiveness of this approach at sustaining forest ecosystem functioning remains uncertain.

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 CO2 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. This article is protected by copyright. All rights reserved.

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.

Wood constitutes the unique source of DNA in dead trees, but extraction of adequate quality DNA from dry wood is usually challenging. However, many different molecular studies require the use of such DNA. We have standardized and validated a modified CTAB protocol to isolate DNA from dry wood from Abies pinsapo and Cedrus atlantica species. Due to the degradation and very little DNA that is normally present in the wood from dead trees we have developed a PCR based test to certify the quality of the extracted samples. In the present study, we have proved too the effectiveness of this methodology to isolate DNA from conifer dry wood samples of sufficient quality to perform further molecular genetic experiments.

Alpine treeline ecotones are considered early-warning monitors of the effects of climate change on terrestrial ecosystems, but it is still unclear how accurately treeline dynamics may track the expected temperature rises. Site-specific abiotic constraints, such as topography and demographic trends may make treelines less responsive to environmental fluctuations. A better understanding on how local processes modulate treelines’ response to warming is thus required. We developed a model of treeline dynamics based on individual data of growth, mortality and reproduction. Specifically, we modeled growth patterns, mortality rates and reproductive size thresholds as a function of temperature and stand structure to evaluate the influence of climate- and stand-related processes on treeline dynamics. In this study, we analyze the dynamics of four Pyrenean mountain pine treeline sites with contrasting stand structures, and subjected to differing rates of climate warming. Our models indicate that Pyrenean treelines could reach basal areas and reproductive potentials similar to those currently observed in high-elevation subalpine forest by the mid twenty-first century. The fastest paces of treeline densification are forecasted by the late twenty-first century and are associated with higher warming rates. We found a common densification response of Pyrenean treelines to climate warming, but contrasting paces arise due to current size structures. Treelines characterized by a multistratified stand structure and subjected to lower mean annual temperatures were the most responsive to climate warming. In monostratified stands, tree growth was less sensitive to temperature than in multistratified stands and trees reached their reproductive size threshold later. Therefore, our simulations highlight that stand structure is paramount in modulating treeline responsiveness to ongoing climate warming. Synthesis. Treeline densification over the twenty-first century is likely to occur at different rates contingent on current stand structure and its effects on individual-level tree growth responses to warming. Accurate projections of future treeline dynamics must thus incorporate site-specific factors other than climate, specifically those related to stand structure and its influence on tree growth.

Aim Stressful environments, like those at the limits of species ranges, are closely associated with the selective establishment of particular traits, yet the mechanisms behind this relationship are poorly understood. Our main aim was to investigate spatial and temporal phenotypic plasticity related to environmental conditions and biogeographical origin. Location A latitudinal gradient along Iberian Peninsula and Morocco, western Mediterranean Basin. Methods We investigate the intra-specific variability in the Mediterranean oak Quercus ilex subsp. ballota, covering most of its latitudinal range. Acorns were collected in the years 2009, 2011, 2012 and 2013 from natural populations and were grown under controlled conditions. We measured acorn mass and germination dynamics, as well as seedling morphology. Results Field data showed significant differences in stand structure among populations. The southernmost population presented the smallest trees, followed by the northernmost population. Core-range trees registered higher values for leaf-nitrogen concentration. Leaf size showed latitudinal variation, with the lowest values at the northern range limit. In addition, the germination pattern also significantly differed among populations, showing a latitudinal pattern of variation (i.e. the southern populations germinated earlier). Fresh acorn mass showed significant within- and among-population differences, displaying a range margin pattern (i.e. similarities between the edge populations). Furthermore, seedling total weight and biomass allocation also followed the same pattern. The models explaining the most part of the trait variance were related mainly to climatic variables (reflecting temporal variability), while the random effect of the models (population) showed a range margin pattern with lower values in edge populations compared with core populations. Main conclusions Similar phenotypic traits among individuals native to the periphery of the distribution range were found despite the temporal variability, suggesting adaptation to stressful environments at the range edges.

Determining the increase in maintenance respiration in response to stress is critical for understanding the cost of adaptation, in terms of expenditure of assimilated carbon. Here, we ask how maintenance costs vary for populations native to contrasting habitats and whether maintenance cost remains constitutive or induced in response to stress. Two populations of Quercus ilex were selected in southern Iberian Peninsula, one growing close to the altitudinal limit and the other growing at mean elevations for the species. Maintenance respiration, growth, and structural variables were measured in leaves and fine roots. We modelled the results found here and those published for populations native to stressed (both soil and latitudinal) habitats. The maintenance respiration measured at 20 °C was higher in expanding (90 %) and in mature (35 %) leaves and in roots (78 %) of individuals growing at higher elevations. Furthermore, our meta-analysis supports that the cost of organ maintenance is higher in stressed habitats, irrespective of the stress factor. As regards the whole plant metabolic design, Q. ilex seems to combine high phenotypic plasticity according to current growth conditions, and high adaptive capacity for local differentiation in response to average stress conditions in the provenance habitat.

Summary The growth limitation hypothesis (GLH) is the most accepted explanation for treeline formation, but it has been scarcely examined in Mediterranean regions, where treelines are located at lower elevations than in temperate regions. The GLH states that low temperature is the ultimate environmental driver for treeline formation, constraining C-sinks (i.e. tissue formation) more than C-sources. The GLH predicts similar or increasing (but not decreasing) non-structural carbohydrate (NSC) concentrations with elevation throughout the course of the growing season. We hypothesized that elevational trends in growth and NSC in Mediterranean regions are not determined by low temperature alone. We tested the GLH by examining the mean annual basal area and NSC concentrations in developing (new twigs) and ripened tissues (branch, stem) of trees growing at three elevations in three Mediterranean and three temperate treelines in the Chilean southern Andes (33° S, Kageneckia angustifolia; 36, 40 and 46° S, Nothofagus pumilio) and in Spain (36° N, Pinus sylvestris; 42° N, Pinus uncinata). Samples for NSC were taken at the onset of summer and autumn, which represent periods of contrasting drought intensities in Mediterranean regions. Tree growth decreased significantly with elevation in temperate treelines but not in Mediterranean treelines. In Mediterranean treelines, new twig NSC concentrations increased significantly with elevation in the early summer but not in the early autumn. In temperate treelines, in contrast, no elevational or seasonal variation (or interaction between them) was observed in new twig NSC concentrations. The NSC concentrations of the branches and stems from both climates showed no elevational trends. The soluble sugars' NSC fraction increased over the season in Mediterranean treelines and decreased in temperate treelines. Synthesis. Although we found support for the growth limitation hypothesis (GLH) in temperate and Mediterranean treelines, our study shows that the tree growth and C balance in Mediterranean treelines are not controlled by low temperature alone. We suggest that environmental factors other than temperature explain the lower global elevation of Mediterranean treelines when compared to temperate treelines.

Aim of study: Understanding small-scale patterns caused by stochastic factors or community interactions driving forest structure and diversity of Moroccan fir Abies marocana Trab.Area of study: Talassemtane fir forest, Talassemtane National Park, Rif Mountains, northern Morocco.Material and methods: Eight plots representative of the structural variability of A. marocana forests were selected, and all tree individuals with diameter at breast height (dbh) ≥2 cm were mapped and measured. We performed four types of spatial point pattern analyses: (1) Univariate analyses to explore the overall trees spatial pattern, (2) bivariate analyses to assess the spatial relationship between juveniles and A. marocana adults, (3) correlation between tree sizes (dbh) and distance between points pairs using the univariate mark correlation function, and (4) random labeling analyses between dominant and suppressed Moroccan fir individuals to assess competition patterns.Main results: We found a strong spatial aggregation of fir individuals and a positive intraspecific association between juveniles and adult trees. However, there were weak but significant distance-dependent effect on tree size and density-dependent effect on suppression pattern.Research highlights: Shade-tolerance, seed dispersal and/or microsite heterogeneity might play important roles in the observed fir patterns. Our results provide a basic knowledge on within-stand Moroccan fir spatial distribution, with implications for adaptive management of these relic forests, and prompting to further research to test advanced hypotheses.Additional key words: Moroccan fir; Talassemtane National Park; Rif forest; Spatial analysis; summary statistics; facilitation; aggregation.Abbreviations used: MF (Moroccan fir); Talassemtane National Park (TNP); dbh (diameter at breast height); Univariate pair-correlation function (g(r)); Heterogeneous Poisson (HP); Bivariate pair-correlation function (g12(r)); Univariate mark correlation function (kmm(r)), Mark connection function (p12(r)).

Extreme drought events are becoming increasingly frequent and extended, particularly in Mediterranean drought-prone regions. In this sense, atmospheric oscillations patterns, such as those represented by the North Atlantic Oscillation (NAO) index and the Westerly Index (WI) have been widely proven as reliable proxies of drought trends. Here, we used the Standardized Precipitation&ndash, Evapotranspiration Index (SPEI), as a reliable indicator of drought, to investigate the drought sensitivity of tree-ring width data (TRW) from several long-lived tree species (Abies borisii-regis, Abies cilicica, Abies pinsapo, Cedrus atlantica, Cedrus libanii, Pinus nigra, Pinus heldreichii). NAO and WI relations with TRW were also investigated in order to identify potential non-stationary responses among those drought proxies. Our temporal and spatial analyses support contrasting Mediterranean dipole patterns regarding the drought sensitivity of tree growth for each tree species. The spatial assessment of NAO and WI relationships regarding SPEI and TRW showed on average stronger correlations westward with non-stationary correlations between annual WI index and TRW in all species. The results indicate that the drought variability and the inferred drought-sensitive trees species (e.g., C. atlantica) are related to the NAO and the WI, showing that TRW is a feasible proxy to long-term reconstructions of Westerly Index (WI) variability in the Western Mediterranean region. Spatial variability of drought severity suggests a complex association between NAO and WI, likely modulated by an east&ndash, west Mediterranean climate dipole.

Increasing variability and uncertainty regarding future climate provide new challenges for the conservation of endangered tree species. For example, threat status can be impacted by genetic diversity, where forest trees show reduced geographic range size, isolated populations and fragmented distribution. We place the conservation insights of population genetic structure in a climate change context, using as experimental system a relict drought-sensitive fir (Abies pinsapo Boiss.). Nuclear (nSSR, ISSR) and chloroplast (cpSSR) markers were analysed to investigate the extent to that A. pinsapo evidences ongoing genetic erosion, isolation and divergent genetic diversity, among populations, elevations and cohorts (young, adult and old trees). We obtained contrasting patterns among chloroplast and nuclear markers. Based on cpSSRs, the highest genetic distances were found in the western portion of the distribution, while based on both nSSRs and ISSRs, differentiation appeared in the eastern portion of the distribution. Evidence for bottlenecks and genetic drift were found in all the studied populations, as well as low among-population genetic differentiation. Land use legacies e.g. impacting current forest structural diversity might be related to observed genetic diversity. No evidence of demographic genetic erosion among cohorts were found. Conservation efforts should focus on reducing the probability of occurrence of stochastic events such as fires and habitat loss due to human impacts or climate change to maximise A. pinsapo population sizes. Further research on adaptive potential should focus on identifying active genetic management strategies that might improve adaptation to future climates in such endangered relict species.

Producción Científica, 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., Ministerio de Economía projects: (CGL2015-69186-C2-1-R (Fundiver)), (CGL2015-69985-R (CLIMED)),( CGL2013-48843-C2-1-R (CoMo-ReAdapt)),( AGL2014-53822-C2-1-R (SATIVA)),( XIRONO (BFU2010-21451)),( CGL2014-52135-C03-01), (PCIN-2015-220), and (CGL2016-81706-REDT (Ecometas Network)), (1560/2015 (ECOHIPRO)), Xunta de Galicia, projects:(PGIDIT06PXIB502262PR, GRC GI-1809, ROCLIGAL-10MDS291009PR), INIA (RTA200600117) e Interreg V-A POCTEFA (CANOPEE, 2014-2020-fondos FEDER), Ministerio de Economía y Empresa, postdoctoral grants (MINECO-FPDI 2013-16600; fondos FEDER), Postdoctoral grants (IJCI-2015-25845)

Aim of study: Drought and stand structure are major and interconnected drivers of forest dynamics. Water shortage and tree-to-tree competition may interact under the current climate change scenario, increasing tree mortality. In this study, we aimed to investigate climate trends, site and stand structure effects on tree mortality, with the main hypothesis that drought-induced mortality is higher as competition increases.Area of study: Persian oak forests from Zagros Range, western Iran.Material and Methods: We split the study area into 20 topographical units (TUs), based on aspect, slope and elevation. In each TU, three 0.1 ha plots were established to quantify site and stand characteristics, namely the diameter of all trees and shrubs, stand density and basal area, canopy dieback and mortality. In addition, soil profiles were analyzed to obtain physical and chemical soil properties. Six transects 100 m length were established per TU to measure tree-to-tree competition for alive and dead trees.Main Results: The highest mortality rates and crown dieback were found at higher elevations and southern and western aspects. Our findings confirm increasing rates of tree mortality in stands with higher tree density and shallow soils. As regard links between climate change and forest decline, our results suggest that changing forest structure may have a significant impact on dust emission.Research highlights: Despite severe dry years occurred recently the study area, they are not significantly different than those recorded in the past. Stand structure appears as a modulating factor of climate change effects, linked to competition-related tree vulnerability to drought.

Warmer and drier climatic conditions are projected for the 21st century; however, the role played by extreme climatic events on forest vulnerability is still little understood. For example, more severe droughts and heat waves could threaten quaternary relict tree refugia such as Circum-Mediterranean fir forests (CMFF). Using tree-ring data and a process-based model, we characterized the major climate constraints of recent (1950–2010) CMFF growth to project their vulnerability to 21st-century climate. Simulations predict a 30% growth reduction in some fir species with the 2050s business-as-usual emission scenario, whereas growth would increase in moist refugia due to a longer and warmer growing season. Fir populations currently subjected to warm and dry conditions will be the most vulnerable in the late 21st century when climatic conditions will be analogous to the most severe dry/heat spells causing dieback in the late 20th century. Quantification of growth trends based on climate scenarios could allow defining vulnerability thresholds in tree populations. The presented predictions call for conservation strategies to safeguard relict tree populations and anticipate how many refugia could be threatened by 21st-century dry spells.

16 Pags.- 3 Tabls.- 5 Figs., 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., This study was financially supported by the Spanish Ministry of Economy projects: CGL2015‐69186‐C2‐1‐R (Fundiver), CGL2015‐69985‐R (CLIMED), CGL2013‐48843‐C2‐1‐R (CoMo‐ReAdapt), AGL2014‐53822‐C2‐1‐R (SATIVA), XIRONO (BFU2010‐21451), CGL2014‐52135‐C03‐01, PCIN‐2015‐220, and CGL2016‐81706‐REDT (Ecometas Network). The study was also funded by IMDROFLOOD (Water Works 2014, EC) and INDECIS (European Research Areas for Climate Services) projects. This work also benefited from funding from Xunta de Galicia (PGIDIT06PXIB502262PR, GRC GI‐1809, ROCLIGAL‐10MDS291009PR), INIA (RTA2006‐00117), and Interreg V‐A POCTEFA (CANOPEE, 2014‐2020‐FEDER funds) projects. RSS and AG were supported by Postdoctoral grants (IJCI‐2015‐25845 and MINECO‐FPDI 2013‐16600; FEDER funds).

11 páginas.--3 figuras.-- 3 tablas.-- 68 referencias.-- Additional Supporting Information may be found in the online version of this article: http://dx.doi.org/10.1111/gcb.13627, Ongoing changes in global climate are altering ecological conditions for many species. The consequences of such changes are typically most evident at the edge of a species’ geographical distribution, where differences in growth or population dynamics may result in range expansions or contractions. Understanding population responses to different climatic drivers along wide latitudinal and altitudinal gradients is necessary in order to gain a better understanding of plant responses to ongoing increases in global temperature and drought severity. We selected Scots pine (Pinus sylvestris L.) as a model species to explore growth responses to climatic variability (seasonal temperature and precipitation) over the last century through dendrochronological methods. We developed linear models based on age, climate and previous growth to forecast growth trends up to year 2100 using climatic predictions. Populations were located at the treeline across a latitudinal gradient covering the northern, central and southernmost populations and across an altitudinal gradient at the southern edge of the distribution (treeline, medium and lower elevations). Radial growth was maximal at medium altitude and treeline of the southernmost populations. Temperature was the main factor controlling growth variability along the gradients, although the timing and strength of climatic variables affecting growth shifted with latitude and altitude. Predictive models forecast a general increase in Scots pine growth at treeline across the latitudinal distribution, with southern populations increasing growth up to year 2050, when it stabilizes. The highest responsiveness appeared at central latitude, and moderate growth increase is projected at the northern limit. Contrastingly, the model forecasted growth declines at lowland-southern populations, suggesting an upslope range displacement over the coming decades. Our results give insight into the geographical responses of tree species to climate change and demonstrate the importance of incorporating biogeographical variability into predictive models for an accurate prediction of species dynamics as climate changes., We thank L. Cavin for methodological advice, and P. González-Diaz and R. Sánchez-Salguero for helping in the tree-ring mea-surements and data processing. This research was funded by EU Marie-Curie (FP7-2011-IEF-300825) and Juan de la Cierva (FPDI-2013-15867) grants to LM. JCL acknowledges financialsupport by CGL2013-48843-C2-2-R (Ministerio de Economíıa y Competitividad) and TRANSHABITAT (European Union FEDER 0087).

12 páginas.-- 4 figuras.-- 3 tablas.-- 80 referencias.-- The online version of this article (doi:10.1007/s10342-017-1064-1) contains supplementary material, which is available to authorized users, Forests worldwide have been recently affected by severe decline and mortality, while our understanding about forest decline across spatial scale is still limited. In this work, we study how Quercus suber trees adjust their physiology, in terms of water use efficiency and secondary growth, to pathogen-induced oak decline at the whole-tree, local and landscape scales. This work was carried out in Mediterranean mixed forests where their dominant key species Q. suber is affected by a severe decline and mortality induced by the exotic soil-borne pathogen Phytophthora cinnamomi. Significant differences were not observed between defoliated and healthy trees, either in terms of water use efficiency or growth at the whole-tree scale. We found that limiting conditions, such as low soil depth and high pathogen abundance, induced trees to higher water use efficiency at local and landscape scales. Overall our findings point out that Q. suber trees subjected to soil drought and root pathogens increase water use efficiency to some extent, while this response might not be enough for the trees to overcome the physiological stress associated with the pathogen-induced dieback. We discuss the complex way by which adult Q. suber trees physiologically respond to P. cinnamomi-induced mortality, improving our understanding of the likely consequences of chronic oak decline in the future., This research was supported by the Ministerio de Ciencia e Innovación (MICINN) projects CGL2011-26877 (RETROBOS) and CGL2014-56739-R (INTERCAPA). J.M.A. was supported by a Formación de Profesorado Universitario FPU-Ministerio de Educación y Ciencia grant (AP2010-0229).

Climate warming is predicted to intensify drought stress in forests by amplifying the severity and frequency of droughts. This drying trend will potentially trigger forest dieback, characterized by tree growth decline and mortality. In drought-prone Spain, forest decline is mainly attributed to severe water shortage. This inciting factor causes a loss in tree vigor and interacts with predisposing or contributing factors that usually act at long time scales. Combined, these factors drive complex patterns of forest decline. Here, we illustrate four case studies on forest decline in Spain, evaluated using dendroecology. These declines were triggered by drought, interacting with other causal factors including past logging in Pyrenean silver fir, increased competition induced by land-use changes in Spanish fir, variation in drought vulnerability of planted pine species, and mistletoe infection in Scots pine. These case studies demonstrate that multiple causes drive forest decline in drought-prone regions. Differentiating the causal factors and inferring their interactions was feasible through tree-ring analyses, which provide robust reconstructions of changes in tree vitality through time.

Treeline ecotones are considered early-warning monitors of the effects of climate warming on terrestrial ecosystems, but it is still unclear how tree growth at treeline will track the forecasted temperature rise in these cold environments. Here, we address this issue by analysing and projecting growth responses to climate on two different cold-limited alpine treelines: Pinus uncinata Ram. in the Spanish Pyrenees and Larix sibirica Ledeb. in the Russian Polar Urals. We assess radial-growth changes as a function of tree age and long-term climate variability using dendrochronology and a process-based model of tree growth. Climate‒growth relationships were compared considering young (age &lt, 50 years) and old trees (age &gt, 75 years) separately. Warm summer conditions enhanced radial growth, particularly after the 1980s, in the Polar Urals sites, whereas growth was positively related to warm spring and winter conditions in the Pyrenees sites. These associations were stronger in young than in old trees for both tree species and regions. Forecasted warm conditions are expected to enhance growth rates in both regions, while the growing season is forecasted to lengthen in the Pyrenees treelines, mostly in young trees. The observed age-related responses to temperature also depend on the forecasted warming rates. Although the temperature sensitivity is overall increasing for young trees, those responses seem more divergent, or even reversed, throughout the contrasting emission scenarios. The RCP 8.5 emission scenario corresponding to the most pronounced warming and drier conditions (+4.8 &deg, C) could also amplify drought stress in young trees from the Pyrenees treelines. Our modelling approach provides accessible tools to evaluate functional thresholds for tree growth in treeline ecotones under warmer conditions.

Facilitation enables plants to improve their fitness in stressful environments. The overall impact of plant-plant interactions on the population dynamics of protégées is the net result of both positive and negative effects that may act simultaneously along the plant life cycle, and depends on the environmental context. This study evaluates the impact of the nurse plant Juniperus sabina on different stages of the life cycle of the forb Helleborus foetidus. Growth, number of leaves, flowers, carpels, and seeds per flower were compared for 240 individuals collected under nurse canopies and in open areas at two sites with contrasting stress levels. Spatial associations with nurse plants and age structures were also checked. A structural equation model was built to test the effect of facilitation on fecundity, accounting for sequential steps from flowering to seed production. The net impact of nurse plants depended on a combination of positive and negative effects on vegetative and reproductive variables. Although nurse plants caused a decrease in flower production at the low-stress site, their net impact there was neutral. In contrast, at the high-stress site the net outcome of plant-plant interactions was positive due to an increase in effective recruitment, plant density, number of viable carpels per flower, and fruit set under nurse canopies. The naturally lower rates of secondary growth and flower production at the high-stress site were compensated by the net positive impact of nurse plants here. Our results emphasize the need to evaluate entire processes and not only final outcomes when studying plant-plant interactions.

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-beetle outbreaks., GGI by the Spanish Ministry of Economy and Competitiveness (Project AGL2014-61175-JIN)., AMH by the Spanish Ministry of Science and Innovation (Projects CGL2007-60120 and CSD2008-0040) and by the Spanish Ministry of Education via a FPU Scholarship., JJC, JCLC, and GSB by the Spanish Ministry of Economy (Projects CGL2015-69186-C2-1-R, CGL2013-48843-C2-2-R, and CGL2012-32965) and the EU (Project FEDER 0087 TRANSHABITAT).

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 drought-constrained (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 drought-induced growth limitations. Spatial patterns of enhanced synchrony represent early warning signals of climate change impacts on forest ecosystems at subcontinental scales. T.A.S. is supported by ERANET-MUNDUS (Euro-Russian Academic Network-Mundus) (European Commission). V.R.d.D. is supported by the Ramón y Cajal Program (Contract RYC-2012-10970). Support for this work was provided by European Union projects ForMAT (Sensitivity of tree growth to climate change and growth modelling from past to future) (Contract ENV4-CT97-0641), ISONET (400 years of annual reconstructions of European climate variability using a high resolution isotopic network) (Contract EV K2-2001-00237), and MILLENNIUM (European climate of the last millennium) (Contract GOCE 017008); the Russian Science Foundation (Project 14-14-00295); the Catalan Government (Project 2014 SGR 1141); Studying Tree Responses to Extreme Events: A Synthesis (STReESS) COST (European Cooperation in Science and Technology) Action FP1106; and Spanish Projects CGL2011-26654 and CGL2013-48843-C2-1-R (Spanish Ministry of Economy and Competitiveness).

Disturbances can affect forest health and are important modulating factors of tree responses to environmental changes. However, standard methods are needed to assess and elucidate the relative effects of disturbance legacies on forest health among species. Here, structural sustainability was used to evaluate and to compare the impacts of contrasting disturbances on the health of Quercus ilex, Cedrus atlantica, and Abies pinsapo forest stands in Morocco and Spain. Disturbance effects on structural sustainability were related to type, severity, and land-use history, and showed inter-regional variability. Cedrus atlantica was structurally sustainable in its core distribution in Morocco, but not at its southern and northernmost geographical range limits. Quercus ilex was structurally sustainable in Morocco. Abies pinsapo was structurally sustainable at optimal elevation sites in Morocco (Rif Mts.), but considering the whole A. pinsapo dataset including Morocco and Spain, the species is structurally unsustainable due to excessive mortality. However, at the lower elevation plots in Spain, unsustainability was due to insufficient mortality. Although some forests were structurally sustainable, none of them were deemed healthy because none met their management objectives. Results also support the key role of disturbance regimes as drivers of forest structural sustainability and adaptive capacity.

International audience; Introduction and statement of the research questions: Gap dynamics have been widely studied in forests of Abies spp. from temperate and boreal regions. The local microclimate and competition for light have been identified as the main factors controlling changes in species composition and canopy structure, however little is known on dynamics of such forests in Mediterranean.* Experimental design and aims: We studied forest structure and dynamics of Abies pinsapo stands in southern Spain, in contrasting habitats and successional status. In addition past regeneration patterns and their relationship to canopy structure, disturbances and forest-use history were investigated.* Results: Stands structure attributes were within the range described for temperate conifer biomes. The age structure revealed two main cohorts comprised of a few > 100 year-old trees and abundant younger trees established in a single recruitment event after the stands were protected in the 1950s. Initial growth-rate analyses indicated that A. pinsapo regenerated mainly in small canopy gaps, while only 15% recruited from the forest understorey. For the last ten years, basal area increment was lower than 10 cm2y−1 in 91% of studied trees and growth rate differences between trees narrowed.* Conclusion: Stand dynamics in A. pinsapo forests maintain general features of temperate fir forests. Tree establishment over time and current stand structure fit to known changes in forest use. Widespread growth decline trends might be linked to stand stagnation and global warming.; Introduction et état des questions de recherche: Les dynamiques des trouées ont été largement étudiées dans les forêts d'Abies spp. à partir des régions tempérées et boréales. Le microclimat local et la compétition pour la lumière ont été identifiés comme les principaux facteurs de contrôle des changements dans la composition des espèces et la structure du couvert, mais on sait peu de choses sur la dynamique de ces forêts dans les écosystèmes méditerranéens.* Plan expérimental et objectifs: Nous avons étudié la structure de la forêt et la dynamique de peuplements d'Abies pinsapo dans le sud de l'Espagne, dans des habitats contrastés et dans des états d'évolution successifs. En outre, les modèles passés de régénération et leurs relations avec la structure du couvert, les perturbations et l'histoire de l'utilisation des forêts ont été étudiés.* Résultats: Les attributs de structure des peuplements se situent dans la gamme décrite pour les biomes de conifères tempérés. La structure des âges a révélé deux cohortes principales composées de quelques vieux arbres d'un âge supérieur à 100 ans, et de nombreux jeunes arbres établis dans un seul évènement de régénération après que les peuplements aient été protégés dans les années 1950. Les premières analyses des taux de croissance ont indiqué que A. pinsapo s'est régénéré principalement dans les petites trouées, tandis que seulement 15 % des arbres ont été recrutés dans le sous-bois. Au cours des dix dernières années, l'augmentation de la surface terrière a été inférieure à 10 cm2 an−1 chez 91 % des arbres étudiés et les différences entre les taux de croissance des arbres étaient réduites.* Conclusion: Les dynamique des peuplements d'A. pinsapo conservent les caractéristiques générales des forêts de sapins tempérés. L'installation des arbres et la structure actuelle des peuplements permettent de connaître les changements dans l'utilisation de la forêt. La baisse généralisée des tendances de croissance pourrait être liée à la stagnation du peuplement et au réchauffement de la planète.