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Tree species have good tolerance to a range of environmental conditions, though their ability to respond and persist to environmental changes is dramatically reduced at the rear-edge distribution limits. At those edges, gene flow conferring adaptation is impaired due to lack of populations at lower latitudes. Thus, trees mainly rely on phenotypic changes to buffer against long-term environmental changes. Interspecific hybridization may offer an alternative mechanism in the generation of novel genetic recombinants that could be particularly valuable to ensure persistence in geographically isolated forests. In this paper, we take advantage of the longevity of a temperate-submediterranean mixed-oak forest to explore the long-term impact of environmental changes on two different oak species and their hybrid. Individual trees were genetically characterized and classified into three groups: pure Quercus petraea (Matt.), Liebl, pure Q. pyrenaica Willd, and hybrids. We calculated basal area increment and intrinsic water-use efficiency (iWUE) from tree-ring width and δ13C per genetic group, respectively. Tree-growth drivers were assessed using correlation analyses and generalized linear mixed models for two contrasting climatic periods: (1880–1915, colder with [CO2] < 303 ppm; and 1980–2015, warmer with [CO2] > 338 ppm). The three genetic groups have increased radial growth and iWUE during the last decades, being the least drought-tolerant QuPe the most sensitive species to water stress. However, no significant differences were found among genetic groups neither in mean growth rate nor in mean iWUE. Furthermore, little differences were found in the response to climate among groups. Genetic groups only differed in the relationship between δ13C and temperature and precipitation during the earlier period, but such a difference disappeared during the recent decades. Climate change may have promoted species-level convergence as a response to environment-induced growth limitations, which translated in synchronized growth and response to climate as well as a tighter stomatal control and increased iWUE across coexisting oak species. [ABSTRACT FROM AUTHOR]

Aim: The aim was to decipher Europe‐wide spatio‐temporal patterns of forest growth dynamics and their associations with carbon isotope fractionation processes inferred from tree rings as modulated by climate warming. Location: Europe and North Africa (30‒70° N, 10° W‒35° E). Time period: 1901‒2003. Major taxa studied: Temperate and Euro‐Siberian trees. Methods: We characterize changes in the relationship between tree growth and carbon isotope fractionation over the 20th century using a European network consisting of 20 site chronologies. Using indexed tree‐ring widths (TRWi), we assess shifts in the temporal coherence of radial growth across sites (synchrony) for five forest ecosystems (Atlantic, boreal, cold continental, Mediterranean and temperate). We also examine whether TRWi shows variable coupling with leaf‐level gas exchange, inferred from indexed carbon isotope discrimination of tree‐ring cellulose (Δ13Ci). Results: We find spatial autocorrelation for TRWi and Δ13Ci extending over a maximum of 1,000 km among forest stands. However, growth synchrony is not uniform across Europe, but increases along a latitudinal gradient concurrent with decreasing temperature and evapotranspiration. Latitudinal relationships between TRWi and Δ13Ci (changing from negative to positive southwards) point to drought impairing carbon uptake via stomatal regulation for water saving occurring at forests below 60° N in continental Europe. An increase in forest growth synchrony over the 20th century together with increasingly positive relationships between TRWi and Δ13Ci indicate intensifying impacts of drought on tree performance. These effects are noticeable in drought‐prone biomes (Mediterranean, temperate and cold continental). Main conclusions: At the turn of this century, convergence in growth synchrony across European forest ecosystems is coupled with coordinated warming‐induced effects of drought on leaf physiology and tree growth spreading northwards. Such a tendency towards exacerbated moisture‐sensitive growth and physiology could override positive effects of enhanced leaf intercellular CO2 concentrations, possibly resulting in Europe‐wide declines of forest carbon gain in the coming decades. [ABSTRACT FROM AUTHOR]

Climate change may reduce forest growth and increase forest mortality, which is connected to high carbon costs through reductions in gross primary production and net ecosystem exchange. Yet, the spatiotemporal patterns of vulnerability to both short‐term extreme events and gradual environmental changes are quite uncertain across the species' limits of tolerance to dryness. Such information is fundamental for defining ecologically relevant upper limits of species tolerance to drought and, hence, to predict the risk of increased forest mortality and shifts in species composition. We investigate here to what extent the impact of short‐ and long‐term environmental changes determines vulnerability to climate change of three evergreen conifers (Scots pine, silver fir, Norway spruce) and two deciduous hardwoods (European beech, sessile oak) tree species at their southernmost limits of distribution in the Mediterranean Basin. Finally, we simulated future forest growth under RCP 2.6 and 8.5 emission scenarios using a multispecies generalized linear mixed model. Our analysis provides four key insights into the patterns of species' vulnerability to climate change. First, site climatic marginality was significantly linked to the growth trends: increasing growth was related to less climatically limited sites. Second, estimated species‐specific vulnerability did not match their a priori rank in drought tolerance: Scots pine and beech seem to be the most vulnerable species among those studied despite their contrasting physiologies. Third, adaptation to site conditions prevails over species‐specific determinism in forest response to climate change. And fourth, regional differences in forests vulnerability to climate change across the Mediterranean Basin are linked to the influence of summer atmospheric circulation patterns, which are not correctly represented in global climate models. Thus, projections of forest performance should reconsider the traditional classification of tree species in functional types and critically evaluate the fine‐scale limitations of the climate data generated by global climate models. Current and forecasted trends in tree growth from temperate and boreal tree species growing at the southernmost limit of their natural distribution in Europe. The tree growth projections by 2100 revealed a generalized decrease in growth under the climatic conditions derived from the RCP 8.5 emission scenario. [ABSTRACT FROM AUTHOR]

Understanding how climate has modulated forest growth and composition in the past is necessary to predict the influence of the ongoing climate warming on the dynamics of mountain forests. We studied the past dynamics of subalpine Pyrenean forests during the last 700 years by assessing the relationships between sedimentary pollen and tree-ring records, and their link with climatic drivers. We compared the pollen record and the montane pollen ratio, an integrative index obtained from sedimentary pollen that allows inferring past altitudinal variations in the montane-subalpine ecotone, with tree-ring width from mountain pine (Pinus uncinata) subalpine forests located in Central Pyrenees. To assess climate-growth associations, we related the dendrochronological data with instrumental meteorological records (1901-2010) and temperature reconstructions from the Pyrenees and Northern Hemisphere. Few robust associations were found between arboreal pollen taxa and tree-ring width series of the surrounding forests. However, significant correlations were found between the montane pollen ratio and tree-ring width series from nearby forests (located less than 10 km apart). This relationship could be potentially useful to infer long-term forest growth changes at decadal to centennial scales using the montane pollen ratio. On the contrary, our results show that tree radial growth has mainly been constrained by low temperatures although the growth sensitivity to climate has considerably varied over the last 700 years. Similar results were obtained for the last century as growth variability of these high-elevation forests is still driven by low temperatures, but a relaxation of this constrain in recent decades has been detected. [ABSTRACT FROM AUTHOR]

Scots pine (Pinus sylvestris L.) is a widely used tree species in European dendroclimatology studies due to its common distribution across much of the continent. Almost all studies find radial growth strongly related to summer temperature, a result reflecting site selection at high elevation/latitude environments where trees grow at their ecophysiological limits. Due to the amount of attention spent on these sites there is a geographical and seasonal bias in temperature reconstructions based upon tree‐ring proxies in Europe. To overcome the limited availability of tree‐ring data in temperate lowlands, we present a northern Poland ring‐width chronology developed from living and historic Scots pine material with a strong common growth signal going back to AD 1200. Investigations into climate‐growth relationships found year‐to‐year ring‐width variability to be more strongly correlated to cold season temperature (November to April) prior to the growing season than summer temperatures during tree‐ring formation. Based on this relationship it was possible to reconstruct cold season temperature conditions for the last 810 years. Spatial field correlations with gridded instrumental records indicated that the reconstruction provides relevant cold season temperature information across the land regions bordering the North Atlantic Ocean and Baltic Sea, lowlands and uplands of western and central Europe, and the eastern and central interior of Russia. Despite an unsuccessful attempt to find a stationary relationship with the North Atlantic Oscillation, comparisons with several cold season temperature reconstructions confirmed the long‐term connection between our reconstructed temperature series for northern Poland and the wider area. [ABSTRACT FROM AUTHOR]

Abstract: In forests, the increase in atmospheric CO2 concentrations (Ca) has been related to enhanced tree growth and intrinsic water‐use efficiency (iWUE). However, in drought‐prone areas such as the Mediterranean Basin, it is not yet clear to what extent this “fertilizing” effect may compensate for drought‐induced growth reduction. We investigated tree growth and physiological responses at five Scots pine (Pinus sylvestris L.) and five sessile oak (Quercus petraea (Matt.) Liebl.) sites located at their southernmost distribution limits in Europe for the period 1960–2012 using annually resolved tree‐ring width and δ13C data to track ecophysiological processes. Results indicated that all 10 natural stands significantly increased their leaf intercellular CO2 concentration (Ci), and consequently iWUE. Different trends in the theoretical gas‐exchange scenarios as a response to increasing Ca were found: generally, Ci tended to increase proportionally to Ca, except for trees at the driest sites in which Ci remained constant. Ci from the oak sites displaying higher water availability tended to increase at a comparable rate to Ca. Multiple linear models fitted at site level to predict basal area increment (BAI) using iWUE and climatic variables better explained tree growth in pines (31.9%–71.4%) than in oak stands (15.8%–46.8%). iWUE was negatively linked to pine growth, whereas its effect on growth of oak differed across sites. Tree growth in the western and central oak stands was negatively related to iWUE, whereas BAI from the easternmost stand was positively associated with iWUE. Thus, some Q. petraea stands might have partially benefited from the “fertilizing” effect of rising Ca, whereas P. sylvestris stands due to their strict closure of stomata did not profit from increased iWUE and consequently showed in general growth reductions across sites. Additionally, the inter‐annual variability of BAI and iWUE displayed a geographical polarity in the Mediterranean. [ABSTRACT FROM AUTHOR]

Climate change-induced rise of air temperatures and the increase of extreme climatic events, such as droughts, will largely affect plant growth and hydraulics, leading to mortality events all over the globe. In this study, we investigated the growth and hydraulic responses of seedlings of contrasting functional types. Pinus sylvestris, Quercus spp. and Castanea sativa seedlings were grown in a common garden experiment under four treatments: control, air warming, drought and their combination during two consecutive growing periods. Height and diameter increments, stomatal conductance and stem water potentials were measured during both growing seasons. Additionally, hydraulic parameters such as xylem-specific native and maximum hydraulic conductivities, and native percentage of loss of conductivity were measured at the end of the entire experiment. Our results clearly pointed to different adaptive strategies of the studied species. Scots pine displayed a relatively isohydric behavior with a strict stomata control prohibiting native embolism whereas sweet chestnut and oak as relatively anisohydric species displayed an increased loss of native conductivity as a results of low water potentials. Seasonal timing of shoot and diameter growth also differed among functional types influencing drought impacts. Additionally, the possibility of embolism reversal seemed to be limited under the study conditions. [ABSTRACT FROM AUTHOR]

Drought is a key limiting factor for tree growth in the Mediterranean Basin. However, the variability in acclimation via xylem traits is largely unknown. We studied tree growth and vessel features of Quercus petraea (Matt.) Lieb. in five marginal stands across southern Europe. Tree-ring width (TRW), mean earlywood vessel area (MVA) and number of earlywood vessels (NV) as well as theoretical hydraulic conductivity (Kh) chronologies were developed for the period 1963-2012. Summer drought signals were consistent among TRW chronologies; however, climatic responses of vessel features differed considerably among sites. At the three xeric sites, previous year's summer drought had a negative effect on MVA and a positive effect on NV. In contrast, at the two mesic sites, current year's spring drought negatively affected NV, while exerting a positive influence on MVA. In both cases, Kh was not altered by this xylem adjustment. All variables revealed identical east-west geographical patterns in growth and anatomical features. Sessile oak copes with drought in different ways: at xeric sites and after unfavourable previous summer conditions more but smaller vessels are built, lowering vulnerability to cavitation, whereas at mesic sites, dry springs partly lead to tree-rings with wider but fewer vessels. The variability of vessel-related features displays a similar geographical dipole in the Mediterranean Basin previously described for tree growth by other studies. [ABSTRACT FROM AUTHOR]

Forest performance is challenged by climate change but higher atmospheric [CO2] ( ca) could help trees mitigate the negative effect of enhanced water stress. Forest projections using data assimilation with mechanistic models are a valuable tool to assess forest performance. Firstly, we used dendrochronological data from 12 Mediterranean tree species (six conifers and six broadleaves) to calibrate a process-based vegetation model at 77 sites. Secondly, we conducted simulations of gross primary production (GPP) and radial growth using an ensemble of climate projections for the period 2010-2100 for the high-emission RCP8.5 and low-emission RCP2.6 scenarios. GPP and growth projections were simulated using climatic data from the two RCPs combined with (i) expected ca; (ii) constant ca = 390 ppm, to test a purely climate-driven performance excluding compensation from carbon fertilization. The model accurately mimicked the growth trends since the 1950s when, despite increasing ca, enhanced evaporative demands precluded a global net positive effect on growth. Modeled annual growth and GPP showed similar long-term trends. Under RCP2.6 (i.e., temperatures below +2 °C with respect to preindustrial values), the forests showed resistance to future climate (as expressed by non-negative trends in growth and GPP) except for some coniferous sites. Using exponentially growing ca and climate as from RCP8.5, carbon fertilization overrode the negative effect of the highly constraining climatic conditions under that scenario. This effect was particularly evident above 500 ppm (which is already over +2 °C), which seems unrealistic and likely reflects model miss-performance at high ca above the calibration range. Thus, forest projections under RCP8.5 preventing carbon fertilization displayed very negative forest performance at the regional scale. This suggests that most of western Mediterranean forests would successfully acclimate to the coldest climate change scenario but be vulnerable to a climate warmer than +2 °C unless the trees developed an exaggerated fertilization response to [CO2]. [ABSTRACT FROM AUTHOR]

Many temperate European tree species have their southernmost distribution limits in the Mediterranean Basin. The projected climatic conditions, particularly an increase in dryness, might induce an altitudinal and latitudinal retreat at their southernmost distribution limit. Therefore, characterizing the morphological and physiological variability of temperate tree species under dry conditions is essential to understand species’ responses to expected climate change. In this study, we compared branch-level hydraulic traits of four Scots pine and four sessile oak natural stands located at the western and central Mediterranean Basin to assess their adjustment to water limiting conditions. Hydraulic traits such as xylem- and leaf-specific maximum hydraulic conductivity (KS-MAX and KL-MAX), leaf-to-xylem area ratio (AL:AX) and functional xylem fraction (FX) were measured in July 2015 during a long and exceptionally dry summer. Additionally, xylem-specific native hydraulic conductivity (KS-N) and native percentage of loss of hydraulic conductivity (PLC) were measured for Scots pine. Interspecific differences in these hydraulic traits as well as intraspecific variability between sites were assessed. The influence of annual, summer and growing season site climatic aridity (P/PET) on intraspecific variability was investigated. Sessile oak displayed higher values of KS-MAX, KL-MAX, AL:AX but a smaller percentage of FX than Scots pines. Scots pine did not vary in any of the measured hydraulic traits across the sites, and PLC values were low for all sites, even during one of the warmest summers in the region. In contrast, sessile oak showed significant differences in KS-MAX, KL-MAX, and FX across sites, which were significantly related to site aridity. The striking similarity in the hydraulic traits across Scots pine sites suggests that no adjustment in hydraulic architecture was needed, likely as a consequence of a drought-avoidance strategy. In contrast, sessile oak displayed adjustments in the hydraulic architecture along an aridity gradient, pointing to a drought-tolerance strategy. [ABSTRACT FROM AUTHOR]

The European Dendroecological Fieldweek (EDF) is a one-week course that takes place every year at varying locations in Europe according to the principle "Bring tree-ring research to the people". The EDF welcomes early-career to advanced researchers, but also forest service and other federal agency employees and private people interested in tree-ring sciences from all over the world. It encompasses a large spectrum of dendrochronological field, laboratory and data analysis methods and scientific fields including climatology, ecology, physiology, geomorphology and archaeology. Multiple scales of observations from the individual cell to the ecosystem level and from seasonal to multi-centennial periods are covered. Work on mini research projects in topic groups alternates with keynote lectures and individual participants' presentations. As one of the first in-person tree-ring meetings since the start of the COVID-19 pandemic the 31st EDF was held in summer 2021 in Val Müstair, Switzerland. Topics included i) Tree age and climate sensitivity of a relict, old-growth Scots pine stand, ii) Blue intensity-based climate sensitivity of Norway spruce growth, iii) Tree rings as indicators of grey larch budmoth outbreaks, iv) Growth of larch trees along an abandoned irrigation channel, v) Wood anatomical characteristics of two alpine creeping shrub species, and vi) Historical dating of a stable and a residential house. Alongside with their educational value these projects allowed novel insight into the age structure and growth dynamics of the sub-alpine forests and beyond in the valley and provided valuable outcome to the local stakeholders such as the Nature Park Biosfera Val Müstair, the local forest service and the public of Val Müstair. Under hindered conditions due to the pandemic, the 31st EDF still demonstrated its strength as an international educational and interdisciplinary scientific field and lab course, combining teaching with the application of cutting-edge technologies. [ABSTRACT FROM AUTHOR]

July-to-October temperature variations are reconstructed for the last 800 years based on tree-ring widths from the Cazorla Range. Annual tree-ring width at this site has been found to be negatively correlated with temperature of the previous summer. This relationship is genuine, metabolically plausible, and cannot be explained as an indirect correlation mediated by hydroclimate. The resulting reconstruction (NCZ) represents the southernmost annually resolved temperature record based on tree-rings in Europe and provides detailed information on the regional climate evolution during the Late Holocene in the southeast of the Iberian Peninsula. The tree-ring based temperature reconstruction of Cazorla Range reveals predominantly warm summer temperatures during the transition between the Medieval Climate Anomaly (MCA) and the Little Ice Age (LIA) from the 13th to the mid of the sixteenth century. The LIA spanned a slightly longer time (1500-1930 CE) than in other European summer temperature reconstructions from the Alps and Pyrenees. The twentieth century, though warmer than the preceding centuries, does not show unprecedented warmth in the last 800 years. Three ensembles of climate simulations conducted with two global atmosphere-ocean general circulation climate models (GCMs), considering different external forcings, were used for comparison: ECHO-G (Erik) and MPI-ESM (E1 and E2). Additionally, individual simulations were available from GCM included in the fifth Coupled Model Intercomparison Project, as well as single-forcing simulations performed with MPI-ESM. The comparison of the reconstructed and simulated temperatures revealed a close agreement of NCZ with the simulations performed with total solar irradiance forcing with wider amplitude. Furthermore, the correlations with single-forcing simulations suggest volcanism as the main factor controlling preindustrial summer temperature variations in the Cazorla Range over the last five centuries. The persistent anti-correlation between NCZ and simulated temperatures during the MCA-LIA transitional period underlines the current limitations for attributing temperature variations during that period to internal variability or external forcing. [ABSTRACT FROM AUTHOR]

The increasing carbon dioxide (CO2) concentration in the atmosphere in combination with climatic changes throughout the last century are likely to have had a profound effect on the physiology of trees: altering the carbon and water fluxes passing through the stomatal pores. However, the magnitude and spatial patterns of such changes in natural forests remain highly uncertain. Here, stable carbon isotope ratios from a network of 35 tree-ring sites located across Europe are investigated to determine the intrinsic water-use efficiency ( iWUE), the ratio of photosynthesis to stomatal conductance from 1901 to 2000. The results were compared with simulations of a dynamic vegetation model (LPX-Bern 1.0) that integrates numerous ecosystem and land-atmosphere exchange processes in a theoretical framework. The spatial pattern of tree-ring derived iWUE of the investigated coniferous and deciduous species and the model results agreed significantly with a clear south-to-north gradient, as well as a general increase in iWUE over the 20th century. The magnitude of the iWUE increase was not spatially uniform, with the strongest increase observed and modelled for temperate forests in Central Europe, a region where summer soil-water availability decreased over the last century. We were able to demonstrate that the combined effects of increasing CO2 and climate change leading to soil drying have resulted in an accelerated increase in iWUE. These findings will help to reduce uncertainties in the land surface schemes of global climate models, where vegetation-climate feedbacks are currently still poorly constrained by observational data. [ABSTRACT FROM AUTHOR]

Abstract: δ 13C and δ 18O of tree rings contain time integrated information about the environmental conditions weighted by seasonal growth dynamics and are well established as sources of palaeoclimatic and ecophysiological data. Annually resolved isotope chronologies are frequently produced by pooling dated growth rings from several trees prior to the isotopic analyses. This procedure has the advantage of saving time and resources, but precludes from defining the isotopic error or statistical uncertainty related to the inter-tree variability. Up to now only a few studies have compared isotope series from pooled tree rings with isotopic measurements from individual trees. We tested whether or not the δ 13C and the δ 18O chronologies derived from pooled and from individual tree rings display significant differences at two locations from the Iberian Peninsula to assess advantages and constraints of both methodologies. The comparisons along the period 1900–2003 reveal a good agreement between pooled chronologies and the two mean master series which were created by averaging raw individual values (Mean) or by generating a mass calibrated mean (MassC). In most of the cases, pooled chronologies show high synchronicity with averaged individual samples at interannual scale but some differences also show up especially when comparing δ 18O decadal to multi-decadal variations. Moreover, differences in the first order autocorrelation among individuals may be obscured by pooling strategies. The lack of replication of pooled chronologies prevents detection of a bias due to a higher mass contribution of one sample but uncertainties associated with the analytical process itself, as sample inhomogeneity, seems to account for the observed differences. [Copyright &y& Elsevier]

Temperature and precipitation variability throughout the year control the intra-annual dynamics of tree-ring formation. Physiological adaptation of trees to climate change is among the key issues to better understand and predict future forest performance and composition. In this study, we investigated the species' coexistence and performance of Scots pine and pubescent oak growing in a mixed sub-Mediterranean forest in the northeast of the Iberian Peninsula. We assessed intra-annual cumulative growth patterns derived from band dendrometers during four consecutive growing seasons and long-term changes in basal area increment for the period 1950–2014. Our results revealed that Scots pine followed an intra-annual bimodal growth pattern. Scots pine growth was mainly limited by water availability at intra-annual, interannual and decadal time scales, which resulted in a negative long-term growth trend. Conversely, oak displayed a unimodal growth pattern, which was less climatically constrained. A significant increase in basal area of oak denotes an overall better potential acclimation to prevailing climatic conditions at the expenses of a higher risk of physiological failure during extreme climate events. [ABSTRACT FROM AUTHOR]

There is a lack of knowledge on how tree species respond to climatic constraints like water shortages and related atmospheric patterns across broad spatial and temporal scales. These assessments are needed to project which populations will better tolerate or respond to global warming across the tree species distribution range. Warmer and drier conditions have been forecasted for the Mediterranean Basin, where Aleppo pine (Pinus halepensis Mill.) is the most widely distributed conifer in dry sites. This species shows plastic growth responses to climate, being particularly sensitive to drought. We evaluated how 32 Aleppo pine forests responded to climate during the second half of the 20th century by using dendrochronology. Climatic constraints of radial growth were inferred by fitting the Vaganov–Shashkin (VS-Lite) growth model to ring-width data from our Aleppo pine forest network. Our findings reported that Aleppo pine growth decreased and showed the highest common coherence among trees in dry, continental sites located in southeastern and eastern inland Spain and Algeria. In contrast, growth increased in wetter sites located in northeastern Spain. Overall, across the Aleppo pine network tree growth was enhanced by prior wet winters and cool and wet springs, whilst warm summers were associated with less growth. The relationships between site ring-width chronologies were higher in nearby forests. This explains why Aleppo pine growth was distinctly linked to indices of atmospheric circulation patterns depending on the geographical location of the forests. The western forests were more influenced by moisture and temperature conditions driven by the Western Mediterranean Oscillation (WeMO) and the Northern Atlantic Oscillation (NAO), the southern forests by the East Atlantic (EA) and the august NAO, while the Balearic, Tunisian and northeastern sites by the Arctic Oscillation (AO) and the Scandinavian pattern (SCA). The climatic constraints for Aleppo pine tree growth and its biogeographical variability were well captured by the VS-Lite model. The model performed better in dry and continental sites, showing strong growth coherence between trees and climatic limitations of growth. Further research using similar broad-scale approaches to climate–growth relationships in drought-prone regions deserves more attention. [ABSTRACT FROM AUTHOR]