Triple-isotope analysis in tree-ring cellulose suggests only moderate effects of tree species mixture on the climate sensitivity of silver fir and Douglas-fir.

Disentangling the factors influencing the climate sensitivity of trees is crucial to understanding the susceptibility of forests to climate change. Reducing tree-to-tree competition and mixing tree species are two strategies often promoted to reduce the drought sensitivity of trees, but it is unclear how effective these measures are in different ecosystems. Here, we studied the growth and physiological responses to climate and severe droughts of silver fir and Douglas-fir growing in pure and mixed conditions at three sites in Switzerland. We used tree-ring width data and carbon (δ13C), oxygen (δ18O) and hydrogen (δ2H) stable isotope ratios from tree-ring cellulose to gain novel information on water relations and the physiology of trees in response to drought and how tree species mixture and competition modulate these responses. We found significant differences in isotope ratios between trees growing in pure and mixed conditions for the two species, although these differences varied between sites, e.g. trees growing in mixed conditions had higher δ13C values and tree-ring width than trees growing in pure conditions for two of the sites. For both species, differences between trees in pure and mixed conditions regarding their sensitivity to temperature, precipitation, climatic water balance and vapor pressure deficit were minor. Furthermore, trees growing in pure and mixed conditions showed similar responses of tree-ring width and isotope ratios to the past severe droughts of 2003, 2015 and 2018. Competition had only a significantly negative effect on δ13C of silver fir, which may suggest a decrease in photosynthesis due to higher competition for light and nutrients. Our study highlights that tree species mixture may have only moderate effects on the radial growth and physiological responses of silver fir and Douglas-fir to climatic conditions and that site condition effects may dominate over mixture effects.

Contrasting water-use strategies to climate warming in white birch and larch in a boreal permafrost region.

The effects of rising atmospheric CO2 concentrations (Ca) with climate warming on intrinsic water-use efficiency and radial growth in boreal forests are still poorly understood. We measured tree-ring cellulose δ13C, δ18O, and tree-ring width in Larix dahurica (larch) and Betula platyphylla (white birch), and analyzed their relationships with climate variables in a boreal permafrost region of northeast China over past 68 years covering a pre-warming period (1951-1984; base period) and a warm period (1985-2018; warm period). We found that white birch but not larch significantly increased their radial growth over the warm period. The increased intrinsic water-use efficiency in both species was mainly driven by elevated Ca but not climate warming. White birch but not larch showed significantly positive correlations between tree-ring δ13C, δ18O and summer maximum temperature as well as vapor pressure deficit in the warm period, suggesting a strong stomatal response in the broad-leaved birch to temperature changes. The climate warming-induced radial growth enhancement in white birch is primarily associated with a conservative water-use strategy. In contrast, larch exhibits a profligate water-use strategy. It implies an advantage for white birch over larch in the warming permafrost regions.

Contrasting physiological strategies explain heterogeneous responses to severe drought conditions within local populations of a widespread conifer.

Understanding how trees prioritize carbon gain at the cost of drought vulnerability under severe drought conditions is crucial for predicting which genetic groups and individuals will be resilient to future climate conditions. In this study, we investigated variations in growth, tree-ring anatomy as well as carbon and oxygen isotope ratios to assess the sensitivity and the xylem formation process in response to an episode of severe drought in 29 mature white spruce (Picea glauca [Moench] Voss) families grown in a common garden trial. During the drought episode, the majority of families displayed decreased growth and exhibited either sustained or increased intrinsic water-use efficiency (iWUE), which was largely influenced by reduced stomatal conductance as revealed by the dual carbon­oxygen isotope approach. Different water-use strategies were detected within white spruce populations in response to drought conditions. Our results revealed intraspecific variation in the prevailing physiological mechanisms underlying drought response within and among populations of Picea glauca. The presence of different genetic groups reflecting diverse water-use strategies within this largely-distributed conifer is likely to lessen the negative effects of drought and decrease the overall forest ecosystems' sensitivity to it.

Unenriched xylem water contribution during cellulose synthesis influenced by atmospheric demand governs the intra-annual tree-ring δ18 O signature.

The oxygen isotope composition (δ18 O) of tree-ring cellulose is used to evaluate tree physiological responses to climate, but their interpretation is still limited due to the complexity of the isotope fractionation pathways. We assessed the relative contribution of seasonal needle and xylem water δ18 O variations to the intra-annual tree-ring cellulose δ18 O signature of larch trees at two sites with contrasting soil water availability in the Swiss Alps. We combined biweekly δ18 O measurements of soil water, needle water, and twig xylem water with intra-annual δ18 O measurements of tree-ring cellulose, xylogenesis analysis, and mechanistic and structural equation modeling. Intra-annual cellulose δ18 O values resembled source water δ18 O mean levels better than needle water δ18 O. Large parts of the rings were formed under high proportional exchange with unenriched xylem water (pex ). Maximum pex values were achieved in August and imprinted on sections at 50-75% of the ring. High pex values were associated with periods of high atmospheric evaporative demand (VPD). While VPD governed needle water δ18 O variability, we estimated a limited Péclet effect at both sites. Due to a variable pex , source water has a strong influence over large parts of the intra-annual tree-ring cellulose δ18 O variations, potentially masking signals coming from needle-level processes.

Tree-ring isotopes from the Swiss Alps reveal non-climatic fingerprints of cyclic insect population outbreaks over the past 700 years.

Recent experiments have underlined the potential of δ2H in tree-ring cellulose as a physiological indicator of shifts in autotrophic versus heterotrophic processes (i.e., the use of fresh versus stored non-structural carbohydrates). However, the impact of these processes has not yet been quantified under natural conditions. Defoliator outbreaks disrupt tree functioning and carbon assimilation, stimulating remobilization, therefore providing a unique opportunity to improve our understanding of changes in δ2H. By exploring a 700-year tree-ring isotope chronology from Switzerland, we assessed the impact of 79 larch budmoth (LBM, Zeiraphera griseana [Hübner]) outbreaks on the growth of its host tree species, Larix decidua [Mill]. The LBM outbreaks significantly altered the tree-ring isotopic signature, creating a 2H-enrichment and an 18O- and 13C-depletion. Changes in tree physiological functioning in outbreak years are shown by the decoupling of δ2H and δ18O (O-H relationship), in contrast to the positive correlation in non-outbreak years. Across the centuries, the O-H relationship in outbreak years was not significantly affected by temperature, indicating that non-climatic physiological processes dominate over climate in determining δ2H. We conclude that the combination of these isotopic parameters can serve as a metric for assessing changes in physiological mechanisms over time.

Progress in high-resolution isotope-ratio analysis of tree rings using laser ablation.

Stable isotope ratio analysis of tree rings has been widely and successfully applied in recent decades for climatic and environmental reconstructions. These studies were mostly conducted at an annual resolution, considering one measurement per tree ring, often focusing on latewood. However, much more information could be retrieved with high-resolution intra-annual isotope studies, based on the fact that the wood cells and the corresponding organic matter are continuously laid down during the growing season. Such studies are still relatively rare, but have a unique potential for reconstructing seasonal climate variations or short-term changes in physiological plant properties, like water-use efficiency. The reason for this research gap is mostly technical, as on the one hand sub-annual, manual splitting of rings is very tedious, while on the other hand automated laser ablation for high-resolution analyses is not yet well established and available. Here, we give an update on the current status of laser ablation research for analysis of the carbon isotope ratio (δ13C) of wood, describe an easy-to-use laser ablation system, its operation and discuss practical issues related to tree core preparation, including cellulose extraction. The results show that routine analysis with up to 100 laser shot-derived δ13C-values daily and good precision and accuracy (ca. 0.1‰) comparable to conventional combustion in an elemental analyzer are possible. Measurements on resin-extracted wood is recommended as most efficient, but laser ablation is also possible on cellulose extracted wood pieces. Considering the straightforward sample preparation, the technique is therefore ripe for wide-spread application. With this work, we hope to stimulate future progress in the promising field of high-resolution environmental reconstruction using laser ablation.

Permafrost degradation alters the environmental signals recorded in tree-ring lignin methoxy group δ2H in northeastern China.

Climate warming has profoundly altered the status of permafrost and has caused extensive permafrost degradation in the Northern Hemisphere. However, long-term observations investigating the hydrological dynamics of permafrost and its ecological effects on plant growth are lacking. Previous studies have reported tree-ring stable hydrogen isotope ratios of lignin methoxy groups (δ2HLM) as an archive of hydrological signals. This study sampled tree-ring cores from a Larix gmelinii forest in Nanwenghe Forest Park, Northeastern China, and separately measured the tree-ring δ2HLM for earlywood and latewood from 1900 to 2020. Earlywood and latewood δ2HLM values, as well as the difference between them, showed no significant long-term trend from 1900 to 1987; however, they both exhibited significant increasing trends since 1988 at rates of 2.6 ‰ and 4.9 ‰ per decade, respectively. This variance changes the magnitude of the difference between the two chronologies and can be explained by the shift in source water δ2H values during tree growth. Based on a structural equation model analysis, when the influence of permafrost melting weakened due to permafrost degradation, the growing season temperature was better recorded in latewood δ2HLM through the effects of precipitation δ2H from July to September. Based on the environmental response of tree-ring δ2HLM in the permafrost region, permafrost degradation influences the source water δ2H values of trees, thereby affecting the expression of temperature signals in tree-ring δ2HLM. The novel results in this study provide a new perspective on permafrost degradation based on the dynamic responses of tree-ring δ2HLM to source water δ2H during permafrost degradation.

Increasing water-use efficiency mediates effects of atmospheric carbon, sulfur, and nitrogen on growth variability of central European conifers.

Climate controls forest biomass production through direct effects on cambial activity and indirectly through interactions with CO2, air pollution, and nutrient availability. The atmospheric concentration of CO2, sulfur and nitrogen deposition can also exert a significant indirect control on wood formation since these factors influence the stomatal regulation of transpiration and carbon uptake, that is, intrinsic water use efficiency (iWUE). Here we provide 120-year long tree-ring time series of iWUE, stem growth, climatic and combined sulfur and nitrogen (SN) deposition trends for two common tree species, Pinus sylvestris (PISY) and Picea abies (PCAB), at their lower and upper distribution margins in Central Europe. The main goals were to explain iWUE trends using theoretical scenarios including climatic and SN deposition data, and to assess the contribution of climate and iWUE to the observed growth trends. Our results showed that after a notable increase in iWUE between the 1950s and 1980s, this positive trend subsequently slowed down. The substantial rise of iWUE since the 1950s resulted from a combination of an accelerated increase in atmospheric CO2 concentrations (Ca) and a stable level of leaf intercellular CO2 (Ci). The offset of observed iWUE values above the trajectory of a constant Ci/Ca scenario was explained by trends in SN deposition (all sites) together with the variation of drought conditions (low-elevation sites only). Increasing iWUE over the 20th and 21st centuries improved tree growth at low-elevation drought-sensitive sites. In contrast, at high-elevation PCAB sites, growth was mainly stimulated by recent warming. We propose that SN pollution should be considered in order to explain the steep increase in iWUE of conifers in the 20th century throughout Central Europe and other regions with a significant SN deposition history.

Drought impacts on tree carbon sequestration and water use - evidence from intra-annual tree-ring characteristics.

The impact of climate extremes on forest ecosystems is poorly understood but important for predicting carbon and water cycle feedbacks to climate. Some knowledge gaps still remain regarding how drought-related adjustments in intra-annual tree-ring characteristics directly impact tree carbon and water use. In this study we quantified the impact of an extreme summer drought on the water-use efficiency and carbon sequestration of four mature Norway spruce trees. We used detailed observations of wood formation (xylogenesis) and intra-annual tree-ring properties (quantitative wood anatomy and stable carbon isotopes) combined with physiological water-stress monitoring. During 41 d of tree water deficit, we observed an enrichment in 13 C but a reduction in cell enlargement and wall-thickening processes, which impacted the anatomical characteristics. These adjustments diminished carbon sequestration by 67% despite an 11% increase in water-use efficiency during drought. However, with the resumption of a positive hydric state in the stem, we observed a fast recovery of cell formation rates based on the accumulated assimilates produced during drought. Our findings enhance our understanding of carbon and water fluxes between the atmosphere and forest ecosystems, providing observational evidence on the tree intra-annual carbon sequestration and water-use efficiency dynamics to improve future generations of vegetation models.

Drought reduces water uptake in beech from the drying topsoil, but no compensatory uptake occurs from deeper soil layers.

The intensity and frequency of droughts events are projected to increase in future with expected adverse effects for forests. Thus, information on the dynamics of tree water uptake from different soil layers during and after drought is crucial. We applied an in situ water isotopologue monitoring system to determine the oxygen isotope composition in soil and xylem water of European beech with a 2-h resolution together with measurements of soil water content, transpiration and tree water deficit. Using a Bayesian isotope mixing model, we inferred the relative and absolute contribution of water from four different soil layers to tree water use. Beech took up more than 50% of its water from the uppermost 5 cm soil layer at the beginning of the 2018 drought, but then reduced absolute water uptake from the drying topsoil by 84%. The trees were not able to quantitatively compensate for restricted topsoil water availability by additional uptake from deeper soil layers, which is related to the fine root depth distribution. Absolute water uptake from the topsoil was restored to pre-drought levels within 3 wk after rewetting. These uptake patterns help to explain both the drought sensitivity of beech and its high recovery potential after drought release.

Radial and axial water movement in adult trees recorded by stable isotope tracing.

The capacity of trees to release water from storage compartments into the transpiration stream can mitigate damage to hydraulic functioning. However, the location of these 'transient' water sources and also the pathways of water movement other than vertical through tree stems still remain poorly understood. We conducted an experiment on two tree species in a common garden in eastern Australia that naturally grow in regions of high (Eucalyptus tereticornis, 'Red Gum') and low (Eucalyptus sideroxylon, 'Ironbark') annual precipitation rates. Deuterium-enriched water (1350% label strength) was directly introduced into the transpiration stream of three trees per species for four consecutive days. Subsequently, the trees were felled, woody tissue samples were collected from different heights and azimuthal positions of the stems, and stable isotope ratios were determined on the water extracted from all samples. The presence/absence of the tracer along the radial and vertical stem axes in combination with xylem hydraulic properties inferred from sapflow, leaf and stem water potentials, wood moisture contents and anatomical sapwood characteristics elucidated species-specific patterns of short-term stem water storage and movement. The distribution of water isotopes at natural abundance among woody tissues indicated systematic differences with highest values of sapwood water and lower values in inner bark and heartwood. Presence of tracer in water of the inner bark highlighted the importance of this tissue as capacitor. Although injected at the northern side of stems, tracer was also discovered at the southern side, providing empirical evidence for circumferential flow in sapwood, particularly of Ironbark. Greater vertical water transport in Red Gum compared with more radial and circumferential water transport in Ironbark were associated with species-specific sapwood anatomy. Our study highlights the value of combining information from stable isotope tracers and wood anatomy to investigate patterns of water transport and storage of tall trees in situ.

High-frequency stable isotope signals in uneven-aged forests as proxy for physiological responses to climate in Central Europe.

Picea abies (L.) Karst. and Fagus sylvatica (L.) are important tree species in Europe, and the foreseen increase in temperature and vapour pressure deficit (VPD) could increase the vulnerability of these species. However, their physiological performance under climate change at temperate and productive sites is not yet fully understood, especially in uneven-aged stands. Therefore, we investigated tree-ring width and stable isotope chronologies (δ13C/δ18O) of these two species at 10 sites along a climate gradient in Central Europe. In these uneven-aged stands, we compared the year-to-year variability of dominant and suppressed trees for the last 80 years in relation to the sites' spatial distribution and climate. δ18O and δ13C were generally consistent across sites and species, showing high sensitivity to summer VPD, whereas climate correlations with radial growth varied much more and depended on mean local climate. We found no significant differences between dominant and suppressed trees in the response of stable isotope ratios to climate variability, especially within the annual high-frequency signals. In addition, we observed a strikingly high coherence of the high-frequency δ18O variations across long distances with significant correlations above 1500 km, whereas the spatial agreement of δ13C variations was weaker (~700 km). We applied a dual-isotope approach that is based on known theoretical understanding of isotope fractionations to translate the observed changes into physiological components, mainly photosynthetic assimilation rate and stomatal conductance. When separating the chronologies in two time windows and investigating the shifts in isotopes ratios, a significant enrichment of either or both isotope ratios over the last decades can be observed. These results, translated by the dual-isotope approach, indicate a general climate-driven decrease in stomatal conductance. This improved understanding of the physiological mechanisms controlling the short-term variation of the isotopic signature will help to define the performance of these tree species under future climate.

Insight into Canary Island pine physiology provided by stable isotope patterns of water and plant tissues along an altitudinal gradient.

The Canary Islands, an archipelago east of Morocco's Atlantic coast, present steep altitudinal gradients covering various climatic zones from hot deserts to subalpine Mediterranean, passing through fog-influenced cloud forests. Unlike the majority of the Canarian flora, Pinus canariensis C. Sm. ex DC. in Buch grow along most of these gradients, allowing the study of plant functioning in contrasting ecosystems. Here we assess the water sources (precipitation, fog) of P. canariensis and its physiological behavior in its different natural environments. We analyzed carbon and oxygen isotope ratios of water and organics from atmosphere, soil and different plant organs and tissues (including 10-year annual time series of tree-ring cellulose) of six sites from 480 to 1990 m above sea level on the Canary Island La Palma. We found a decreasing δ18O trend in source water that was overridden by an increasing δ18O trend in needle water, leaf assimilates and tree-ring cellulose with increasing altitude, suggesting site-specific tree physiological responses to relative humidity. Fog-influenced and fog-free sites showed similar δ13C values, suggesting photosynthetic activity to be limited by stomatal closure and irradiance at certain periods. In addition, we observed an 18O-depletion (fog-free and timberline sites) and 13C-depletion (fog-influenced and fog-free sites) in latewood compared with earlywood caused by seasonal differences in: (i) water uptake (i.e., deeper ground water during summer drought, fog water frequency and interception) and (ii) meteorological conditions (stem radial growth and latewood δ18O correlated with winter precipitation). In addition, we found evidence for foliar water uptake and strong isotopic gradients along the pine needle axis in water and assimilates. These gradients are likely the reason for an unexpected underestimation of pine needle water δ18O when applying standard leaf water δ18O models. Our results indicate that soil water availability and air humidity conditions are the main drivers of the physiological behavior of pine along the Canary Island's altitudinal gradients.

How does varying water supply affect oxygen isotope variations in needles and tree rings of Scots pine?

In many regions, drought is suspected to be a cause of Scots pine decline and mortality, but the underlying physiological mechanisms remain unclear. Because of their relationship to ecohydrological processes, δ18O values in tree rings are potentially useful for deciphering long-term physiological responses and tree adaptation to increasing drought. We therefore analyzed both needle- and stem-level isotope fractionations in mature trees exposed to varying water supply. In a first experiment, we investigated seasonal δ18O variations in soil and needle water of Scots pine in a dry inner Alpine valley in Switzerland, comparing drought-stressed trees with trees that were irrigated for more than 10 years. In a second experiment, we analyzed twentieth-century δ18O variations in tree rings of the same forest, including a group of trees that had recently died. We observed less 18O enrichment in needle water of drought-stressed compared with irrigated trees. We applied different isotope fractionation models to explain these results, including the Péclet and the two-pool correction, which considers the ratio of unenriched xylem water in the needles to total needle water. Based on anatomical measurements, we found this ratio to be unchanged in drought-stressed needles, although they were shorter. The observed lower 18O enrichment in needles of stressed trees was therefore likely caused by increased effective path length for water movement within the leaf lamina. In the tree-ring study, we observed lower δ18O values in tree rings of dead trees compared with survivors during several decades prior to their death. These lower values in declining trees are consistent with the lower needle water 18O enrichment observed for drought-stressed compared with irrigated trees, suggesting that this needle-level signal is reflected in the tree rings, although changes in rooting depth could also play a role. Our study demonstrates that long-term effects of drought are reflected in the tree-ring δ18O values, which helps to provide a better understanding of past tree physiological changes of Scots pine.

Compression wood has a minor effect on the climate signal in tree-ring stable isotope records of montane Norway spruce.

Compression wood (CW) is a common tissue present in the trunk, branches and roots of mechanically stressed coniferous trees. Its main role is to increase the mechanical strength and regain the vertical orientation of a leaning stem. Compression wood is thought to influence the climate signal in different tree-ring measures. Hence trees containing CW are mostly excluded from tree-ring studies reconstructing past climate variability. There is a large gap of systematic work testing the potential effect of CW on the strength of the climate signal in different tree-ring parameters, especially stable isotope records. Here we test for the first time the effect of CW contained in montane Norway spruce (Picea abies L. Karst) on both δ13C and δ18O tree-ring cellulose records by analyzing compression and opposite wood radii from several disturbed trees together with samples from undisturbed reference trees. We selected four trees tilted by geomorphic processes that were felled by wind and four undisturbed reference trees in the Tatra Mountains, Poland. We qualitatively classified the strength of CW using wood cell anatomical characteristics (tracheid shape, cell wall thickness and presence of intercellular spaces). Then we developed tree-ring width and δ13C and δ18O chronologies from the CW radii, from the opposite radii of the tilted trees and from the reference radii. We tested the effect of CW on tree-ring cellulose δ13C and δ18O variability and on the climate signal strength. We found only minor differences in the means of δ13C and δ18O of compression (δ13C: -22.81‰, δ18O: 28.29‰), opposite (δ13C: -23.02‰; δ18O: 28.05‰) and reference (δ13C: -22.78‰; δ18O: 27.61‰) radii. The statistical relationships between climate variables, δ13C and δ18O, remained consistent among all chronologies. Our findings suggest that moderately tilted trees containing CW can be used to reconstruct past geomorphic activity and for stable isotope-based dendroclimatology.

Publisher Correction: Northern forest tree populations are physiologically maladapted to drought.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Contrasting stomatal sensitivity to temperature and soil drought in mature alpine conifers.

Conifers growing at high elevations need to optimize their stomatal conductance (gs ) for maximizing photosynthetic yield while minimizing water loss under less favourable thermal conditions. Yet the ability of high-elevation conifers to adjust their gs sensitivity to environmental drivers remains largely unexplored. We used 4 years of sap flow measurements to elucidate intraspecific and interspecific variability of gs in Larix decidua Mill. and Picea abies (L.) Karst along an elevational gradient and contrasting soil moisture conditions. Site- and species-specific gs response to main environmental drivers were examined, including vapour pressure deficit, air temperature, solar irradiance, and soil water potential. Our results indicate that maximum gs of L. decidua is >2 times higher, shows a more plastic response to temperature, and down-regulates gs stronger during atmospheric drought compared to P. abies. These differences allow L. decidua to exert more efficient water use, adjust to site-specific thermal conditions, and reduce water loss during drought episodes. The stronger plasticity of gs sensitivity to temperature and higher conductance of L. decidua compared to P. abies provide new insights into species-specific water use strategies, which affect species' performance and should be considered when predicting terrestrial water dynamics under future climatic change.

Northern forest tree populations are physiologically maladapted to drought.

Northern forests at the leading edge of their distributions may not show increased primary productivity under climate warming, being limited by climatic extremes such as drought. Looking beyond tree growth to underlying physiological mechanisms is fundamental for accurate predictions of forest responses to climate warming and drought stress. Within a 32-year genetic field trial, we analyze relative contributions of xylem plasticity and inferred stomatal response to drought tolerance in regional populations of a widespread conifer. Genetic adaptation leads to varying responses under drought. Trailing-edge tree populations produce fewer tracheids with thicker cell walls, characteristic of drought-tolerance. Stomatal response explains the moderate drought tolerance of tree populations in central areas of the species range. Growth loss of the northern population is linked to low stomatal responsiveness combined with the production of tracheids with thinner cell walls. Forests of the western boreal may therefore lack physiological adaptations necessary to tolerate drier conditions.

Poleward migration of the destructive effects of tropical cyclones during the 20th century.

Determination of long-term tropical cyclone (TC) variability is of enormous importance to society; however, changes in TC activity are poorly understood owing to discrepancies among various datasets and limited span of instrumental records. While the increasing intensity and frequency of TCs have been previously documented on a long-term scale using various proxy records, determination of their poleward migration has been based mostly on short-term instrumental data. Here we present a unique tree-ring-based approach for determination of long-term variability in TC activity via forest disturbance rates in northeast Asia (33-45°N). Our results indicate significant long-term changes in TC activity, with increased rates of disturbances in the northern latitudes over the past century. The disturbance frequency was stable over time in the southern latitudes, however. Our findings of increasing disturbance frequency in the areas formerly situated at the edge of TC activity provide evidence supporting the broad relevance of poleward migration of TCs. Our results significantly enhance our understanding of the effects of climate change on TCs and emphasize the need for determination of long-term variation of past TC activity to improve future TC projections.

Subfossil trees suggest enhanced Mediterranean hydroclimate variability at the onset of the Younger Dryas.

Nearly 13,000 years ago, the warming trend into the Holocene was sharply interrupted by a reversal to near glacial conditions. Climatic causes and ecological consequences of the Younger Dryas (YD) have been extensively studied, however proxy archives from the Mediterranean basin capturing this period are scarce and do not provide annual resolution. Here, we report a hydroclimatic reconstruction from stable isotopes (δ18O, δ13C) in subfossil pines from southern France. Growing before and during the transition period into the YD (12 900-12 600 cal BP), the trees provide an annually resolved, continuous sequence of atmospheric change. Isotopic signature of tree sourcewater (δ18Osw) and estimates of relative air humidity were reconstructed as a proxy for variations in air mass origin and precipitation regime. We find a distinct increase in inter-annual variability of sourcewater isotopes (δ18Osw), with three major downturn phases of increasing magnitude beginning at 12 740 cal BP. The observed variation most likely results from an amplified intensity of North Atlantic (low δ18Osw) versus Mediterranean (high δ18Osw) precipitation. This marked pattern of climate variability is not seen in records from higher latitudes and is likely a consequence of atmospheric circulation oscillations at the margin of the southward moving polar front.