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Drought frequency is increasing in many parts of the world and may enhance tree decline and mortality. The underlying physiological mechanisms are poorly understood, however, particularly regarding chronic effects of long-term drought and the response to increasing temperature and vapor pressure deficit (VPD). We combined analyses of radial growth and stable carbon isotope ratios (δ13C) in tree rings in a mature Scots pine (Pinus sylvestris L.) forest over the 20th century to elucidate causes of tree mortality in one of the driest parts of the European Alps (Pfynwald, Switzerland). We further compared trees that have recently died with living trees in a 10-year irrigation experiment, where annual precipitation was doubled. We found a sustained growth increase and immediate depletion of δ13C values for irrigated trees, indicating higher stomatal conductance and thus indeed demonstrating that water is a key limiting factor for growth. Growth of the now-dead trees started declining in the mid-1980s, when both mean temperature and VPD increased strongly. But growth of these trees was reduced to some extent already several decades earlier, while intrinsic water-use efficiency derived from δ13C values was higher. This indicates a more conservative water-use strategy compared with surviving trees, possibly at the cost of low carbon uptake and long-term reduction of the needle mass. We observed reduced climatic sensitivity of raw tree-ring δ13C for the now-dead in contrast to surviving trees, indicating impaired stomatal regulation, although this difference between the tree groups was smaller after detrending the data. Higher autocorrelation and a lower inter-annual δ13C variability of the now-dead trees further indicates a strong dependence on (low) carbon reserves. We conclude that the recent increase in atmospheric moisture demand in combination with insufficient soil water supply was the main trigger for mortality of those trees that were weakened by long-term reduced carbon uptake., Tree Physiology, 37 (8), ISSN:0829-318X, ISSN:1758-4469

Stratospheric volcanic eruptions have had significant impacts on the radiation budget, atmospheric and surface temperatures, precipitation and regional weather patterns, resulting in global climatic changes. The changes associated with such eruptions most commonly result in cooling during several years after events. This study aimed to reveal eco-physiological response of larch trees from northeastern Yakutia (YAK), eastern Taimyr (TAY) and Altai (ALT) regions to climatic anomalies after major volcanic eruptions CE 535, 540, 1257, 1641, 1815 and 1991 using new multiple tree-ring parameters: tree-ring width (TRW), maximum latewood density (MXD), cell wall thicknesses (CWT), δ13C and δ18O in tree-ring cellulose. This investigation showed that TRW, CWT, MXD and δ18O chronologies recorded temperature signal, while information about precipitation and vapor pressure deficit was captured by δ13C chronologies. Sunshine duration was well recorded in δ18O from YAK and ALT. Tree-ring parameters recorded cold, wet and cloudy summer anomalies during the 6th and 13th centuries. However, significant summer anomalies after Tambora (1815) and Pinatubo (1991) eruptions were not captured by any tree-ring parameters

The

Tropical cyclones (TCs) are one of the most devastating storm systems with high socioeconomic impacts around the world. The drivers of long-term changes in TC frequency and intensity, including the recent global climatic changes, are, however, poorly understood due to short instrumental measurements and a lack of accurate proxy records. Here we present the long-term impacts of large volcanic eruptions on TC activity in northeast Asia. For this purpose, we performed a reconstruction of the frequency and intensity of landfalling TCs based on tree-ring oxygen isotope ratios over the past 350 years. Our results revealed markedly depleted δ18O values (P

We aim to achieve a mechanistic understanding of the eco-physiological processes in Larix decidua and Pinus mugo var. uncinata growing on north- and south-facing aspects in the Swiss National Park in order to distinguish the short- and long-term effects of a changing climate. To strengthen the interpretation of the δ(18)O signal in tree rings and its coherence with the main factors and processes driving evaporative δ(18)O needle water enrichment, we analyzed the δ(18)O in needle, xylem and soil water over the growing season in 2013 and applied the mechanistic Craig-Gordon model (1965) for the short-term responses. We found that δ(18)O needle water strongly reflected the variability of relative humidity mainly for larch, while only δ(18)O in pine xylem water showed a strong link to δ(18)O in precipitation. Larger differences in offsets between modeled and measured δ(18)O needle water for both species from the south-facing aspects were detected, which could be explained by the high transpiration rates. Different soil water and needle water responses for the two species indicate different water-use strategies, further modulated by the site conditions. To reveal the long-term physiological response of the studied trees to recent and past climate changes, we analyzed δ(13)C and δ(18)O in wood chronologies from 1900 to 2013. Summer temperatures as well as summer and annual amount of precipitations are important factors for growth of both studied species from both aspects. However, mountain pine trees reduced sensitivity to temperature changes, while precipitation changes come to play an important role for the period from 1980 to 2013. Intrinsic water-use efficiency (WUEi) calculated for larch trees since the 1990s reached a saturation point at elevated CO2 Divergent trends between pine WUEi and δ(18)O are most likely indicative of a decline of mountain pine trees and are also reflected in decoupling mechanisms in the isotope signals between needles and tree-rings.

The analysis of stable carbon and oxygen isotopes in tree-rings is a widely applied tool which allows to retrieve information about past climatic conditions, as well as tree physiological responses to environmental changes. This is based on well-established mechanistic models and firm statistical relationships with climate variables. In contrast, the hydrogen isotopic signature (δ2H) of tree-rings has been reported to be poorly correlated to climate or difficult to explain, and as a consequence, hydrogen isotopes are far less utilized. However, recent plant-physiological experiments have highlighted the role of autotrophic versus heterotrophic processes affecting δ2H values, i.e. use of fresh assimilates versus stored carbohydrates, and have much improved our understanding of the role of post-photosynthetic 2H-fractionation. Using unpublished and literature δ2H data of tree-ring cellulose (δ2HC) of 5 study sites in Europe and Asia, we systematically investigated the relationships between δ2HC and tree-ring width (TRW), which, in contrast to previous research, could now be explained through post-photosynthetic 2H-fractionation. In most cases, these relationships were found to be negative (r2 = 0.23 to 0.51, all P

Drought responses of mature trees are still poorly understood making it difficult to predict species distributions under a warmer climate. Using mature European beech (Fagus sylvatica L.), a widespread and economically important tree species in Europe, we aimed at developing an empirical stress-level scheme to describe its physiological response to drought. We analysed effects of decreasing soil and leaf water potential on soil water uptake, stem radius, native embolism, early defoliation and crown dieback with comprehensive measurements from overall nine hydrologically distinct beech stands across Switzerland, including records from the exceptional 2018 drought and the 2019/2020 post-drought period. Based on the observed responses to decreasing water potential we derived the following five stress levels: I (predawn leaf water potential-0.4 MPa): no detectable hydraulic limitations; II (-0.4 to -1.3): persistent stem shrinkage begins and growth ceases; III (-1.3 to -2.1): onset of native embolism and defoliation; IV (-2.1 to -2.8): onset of crown dieback; V (-2.8): transpiration ceases and crown dieback is20%. Our scheme provides, for the first time, quantitative thresholds regarding the physiological downregulation of mature European beech trees under drought and therefore synthesises relevant and fundamental information for process-based species distribution models. Moreover, our study revealed that European beech is drought vulnerable, because it still transpires considerably at high levels of embolism and because defoliation occurs rather as a result of embolism than preventing embolism. During the 2018 drought, an exposure to the stress levels III-V of only one month was long enough to trigger substantial crown dieback in beech trees on shallow soils. On deep soils with a high water holding capacity, in contrast, water reserves in deep soil layers prevented drought stress in beech trees. This emphasises the importance to include local data on soil water availability when predicting the future distribution of European beech.

The tree-ring stable carbon and oxygen isotope chronologies from two forest sites located in the Forni Glacier forefield (Italy)—one along the glacier stream (GL) and the other toward the valley slope (SL)—were analyzed with the aim of disentangling the precipitation and glacier meltwater inputs in source water δ18O, as reflected by the tree-ring cellulose δ18O. The cellulose δ18O from the GL trees has a negative correlation with winter and summer temperatures, whereas the cellulose δ18O from the SL trees has a positive correlation with precipitation δ18O. The isotopic signature of the source water at the GL site is also influenced by waters of glacial origin, as confirmed by the 18O-depleted glacier meltwater inputs (GMWI_δ18O) estimated by means of an isotope model. The GMWI_δ18O values are consistent with the mean difference measured between the δ18O in the glacier stream and in the precipitation and the winter and summer temperature explains up to 37 percent of the GMWI_δ18O variance. Our results show an increasing influence of glacier meltwater throughout the past decade for the GL site. Our analysis opens new opportunities to reconstruct changes in water regimes of the glacier streams by means of the tree-ring cellulose δ18O.

Elevated CO2 along with rising temperature and water deficits can lead to changes in tree physiology and leaf biochemistry. These changes can increase heat- and drought-induced tree mortality. We aim to reveal the impacts of climatic drivers on individual compounds at the leaf level among European larch (Larix decidua) and mountain pine (Pinus mugo) trees, which are widely distributed at high elevations. We investigated seasonal carbon isotope composition (δ13C) and concentration patterns of carbohydrates and organic acids in needles of these two different species from a case study in the Swiss National Park (SNP). We found that average and minimum air temperatures were the main climatic drivers of seasonal variation of δ13C in sucrose and glucose as well as in concentrations of carbohydrates and citric acid/citrate in needles of both tree species. The impact of seasonal climatic drivers on larch and mountain pine trees at the needle level is in line with our earlier study in this region for long-term changes at the tree-ring level. We conclude that the species-specific changes in δ13C and concentrations of carbohydrates and organic acids are sensitive indicators of changes in the metabolic pathways occurring as a result of climatic changes.

For the first time we present a multi-proxy data set for the Russian Altai, consisting of Siberian larch tree-ring width (TRW), latewood density (MXD), δ13C and δ18O in cellulose chronologies obtained for the period 1779–2007 and cell wall thickness (CWT) for 1900–2008. All of these parameters agree well between each other in the high-frequency variability, while the low-frequency climate information shows systematic differences. The correlation analysis with temperature and precipitation data from the closest weather station and gridded data revealed that annual TRW, MXD, CWT, and δ13C data contain a strong summer temperature signal, while δ18O in cellulose represents a mixed summer and winter temperature and precipitation signal. The temperature and precipitation reconstructions from the Belukha ice core and Teletskoe lake sediments were used to investigate the correspondence of different independent proxies. Low frequency patterns in TRW and δ13C chronologies are consistent with temperature reconstructions from nearby Belukha ice core and Teletskoe lake sediments showing a pronounced warming trend in the last century. Their combination could be used for the regional temperature reconstruction. The long-term δ18O trend agrees with the precipitation reconstruction from the Teletskoe lake sediment indicating more humid conditions during the twentieth century. Therefore, these two proxies could be combined for the precipitation reconstruction.

Tree-ring width of Larix gmelinii (Rupr.) Rupr., ratios of stable isotopes of C (delta(13)C) and O (delta(18)O) of whole wood and cellulose chronologies were obtained for the northern part of central Siberia (Tura, Russia) for the period 1864-2006. A strong decrease in the isotope ratios of O and C (after atmospheric delta(13)C corrections) and tree-ring width was observed for the period 1967-2005, while weather station data show a decrease in July precipitation, along with increasing July air temperature and vapor pressure deficit (VPD). Temperature at the end of May and the whole month of June mainly determines tree radial growth and marks the beginning of the vegetation period in this region. A positive correlation between tree-ring width and July precipitation was found for the calibration period 1929-2005. Positive significant correlations between C isotope chronologies and temperatures of June and July were found for whole wood and cellulose and negative relationships with July precipitation. These relationships are strengthened when the likely physiological response of trees to increased CO(2) is taken into account (by applying a recently developed delta(13)C correction). For the O isotope ratios, positive relationships with annual temperature, VPD of July and a negative correlation with annual precipitation were observed. The delta(18)O in tree rings may reflect annual rather than summer temperatures, due to the late melting of the winter snow and its contribution to the tree water supply in summer. We observed a clear change in the isotope and climate trends after the 1960s, resulting in a drastic change in the relationship between C and O isotope ratios from a negative to a positive correlation. According to isotope fractionation models, this indicates reduced stomatal conductance at a relatively constant photosynthetic rate, as a response of trees to water deficit for the last half century in this permafrost region.

We investigated seasonal variations in &delta, 13C values and concentrations of carbohydrates and organic acids in needles of declining and healthy mountain pine (Pinus mugo ssp. uncinata (DC.) Domin) trees from the Swiss National Park (SNP), using compound-specific isotopes analysis (CSIA). Our goal was to study the impact of climatic drivers on the individual compounds and understand the reasons of partial tree declines in relation to healthy mountain pine trees under seasonal weather patterns. We found that temperature is the main climatic driver determining the seasonal carbon dynamics at the needle level. Lower seasonal &delta, 13C variability and lower concentration levels of sucrose in needles suggest less photosynthetic activity and sink carbon demand in declining compared to healthy mountain pine trees. Higher concentration levels of hexose (glucose and fructose) can play a reserve function for surviving mechanisms of mountain pine trees. Seasonal patterns of organic acid (malate and citrate) suggest an increasing investment in maintenance and repair mechanisms. The seasonal course of carbohydrates and organic acids can therefore be considered an indicator for a modified carbon metabolism within the leaves and possibly within the other tree tissues, partially explaining the decline of mountain pine trees.

Records of stable oxygen isotope ratios in tree rings are valuable tools to reconstruct past climatic conditions and investigate the response of trees to those conditions. So far the use of stable oxygen isotope signatures of tree rings has not been systematically evaluated in dynamic global vegetation models (DGVMs). DGVMs integrate many hydrological and physiological processes and their application could improve proxy-model comparisons and the interpretation of oxygen isotope records. Here we present an approach to simulate leaf water and stem cellulose δ18O of trees using the LPX-Bern DGVM (LPX-Bern). Our results lie within a few per mil of measured tree ring δ18O of 31 different forest stands mainly located in Europe. Temporal means over the last 5 decades as well as interannual variations for a subset of sites in Switzerland are captured. A sensitivity analysis reveals that relative humidity, temperature, and the water isotope boundary conditions have the largest influence on simulated stem cellulose δ18O, followed by all climatic factors combined, whereas increasing atmospheric CO2 and nitrogen deposition exert no impact. We conclude that simulations with LPX-Bern are useful for investigating large-scale oxygen isotope patterns of tree ring cellulose to elucidate the importance of different environmental factors on isotope variations and therefore help to reduce uncertainties in the interpretation of δ18O of tree rings.

RATIONALE The oxygen isotope ratio (δ(18)O) of carbohydrates derived from animals, plants, sediments, and soils provides important information about biochemical and physiological processes, past environmental conditions, and geographical origins, which are otherwise not available. Nowadays, δ(18)O analyses are often performed on carbohydrate bulk material, while compound-specific δ(18)O analyses remain challenging and methods for a wide range of individual carbohydrates are rare. METHODS To improve the δ(18)O analysis of individual carbohydrates by gas chromatography/pyrolysis-isotope ratio mass spectrometry (GC/Pyr-IRMS) we developed a new methylation derivatization method. Carbohydrates were fully methylated within 24 h in an easy-to-handle one-pot reaction in acetonitrile, using silver oxide as proton acceptor, methyl iodide as methyl group carrier, and dimethyl sulfide as catalyst. RESULTS The precision of the method ranged between 0.12 and 1.09‰ for the δ(18)O values of various individual carbohydrates of different classes (mono-, di-, and trisaccharides, alditols), with an accuracy of a similar order of magnitude, despite high variation in peak areas. Based on the δ(18)O values of the main isomers, important monosaccharides such as glucose and fructose could also be precisely analyzed for the first time. We tested the method on standard mixtures, honey samples, and leaf carbohydrates extracted from Pinus sylvestris, showing that the method is also applicable to different carbohydrate mixtures. CONCLUSIONS The new methylation method shows unrivalled accuracy and precision for δ(18)O analysis of various individual carbohydrates; it is fast and easy-to-handle, and may therefore find wide-spread application.

Stable oxygen isotope ratios (δ(18) O) in trees from high latitude ecosystems are valuable sources of information for recent and past environmental changes, but the interpretation is hampered by the complex hydrology of forests growing under permafrost conditions, where only a shallow layer of soil thaws in summer. We investigated larch trees (Larix gmelinii) at two sites with contrasting soil conditions in Siberia and determined δ(18) O of water from different soil depths, roots, twigs, and needles as well as δ(18) O of soluble carbohydrates regularly over two growing seasons. A comparison of results from the 2 yrs revealed an unexpected 'inverse' climate-isotope relationship, as dry and warm summer conditions resulted in lower soil and root δ(18) O values. This was due to a stronger uptake of isotopically depleted water pools originating from melted permafrost or previous winter snow. We developed a conceptual framework that considers the dependence of soil water profiles on climatic conditions for explaining δ(18) O in needle water, needle soluble carbohydrates and stem cellulose. The negative feedback of drought conditions on the source isotope value could explain decreasing tree-ring δ(18) O trends in a warming climate and is likely relevant in many ecosystems, where a soil isotope gradient with depth is observed.

RATIONALE We investigated the applicability of tree-ring whole-wood material for δ(18)O and δ(13)C analysis in comparison with the more time- and resource-intensive use of cellulose, by considering possible variability between (i) five different tree species (Fagus sylvatica, Quercus robur, Picea abies, Abies alba, Pseudotsuga menziesii), (ii) two sites that differ in soil moisture, and (iii) climate conditions within a 10-year period. METHODS Stem cores of 30 individual trees (n = 3 trees per each species and site) were sampled from two sites in south Germany (Bavaria), and tree rings within sapwood of the years 2001-2010 were separated. The δ(18)O and δ(13)C values from homogenized tree-ring whole wood and from extracted cellulose were measured by mass spectrometry. Species-specific offsets in isotope values were analyzed and the responses in isotopic signature to climate variability including a single drought event were compared between whole-wood and cellulose. RESULTS A constant offset in δ(18)O values of ca 5‰ between wood and cellulose was observed for most species independent of site conditions, with a significant difference between beech and Douglas-fir, while inter-annual variability was only observed in oak. The offset in δ(13)C values ranged between 1.45 and 1.84‰ across species, sites and years. Both materials generally showed similar strength in responses to temperature, precipitation and soil water availability, particularly for conifers. Resistance to severe drought stress--partly more strongly reflected in the δ(13)C values of cellulose--was lower for conifers than for the deciduous species. CONCLUSIONS Wood material from the sapwood of the studied tree species is as useful as cellulose for studying environmental effects on tree-ring δ(18)O and δ(13)C values at a short-term scale as considered in most ecophysiological studies. The more variable response of oak may require further investigations.

Little is known about the dynamics of concentrations and carbon isotope ratios of individual carbohydrates in leaves in response to climatic and physiological factors. Improved knowledge of the isotopic ratio in sugars will enhance our understanding of the tree ring isotope ratio and will help to decipher environmental conditions in retrospect more reliably. Carbohydrate samples from larch (Larix gmelinii) needles of two sites in the continuous permafrost zone of Siberia with differing growth conditions were analysed with the Compound-Specific Isotope Analysis (CSIA). We compared concentrations and carbon isotope values (δ13C) of sucrose, fructose, glucose and pinitol combined with phenological data. The results for the variability of the needle carbohydrates show high dynamics with distinct seasonal characteristics between and within the studied years with a clear link to the climatic conditions, particularly vapour pressure deficit. Compound-specific differences in δ13C values as a response to climate were detected. The δ13C of pinitol, which contributes up to 50% of total soluble carbohydrates, was almost invariant during the whole growing season. Our study provides the first in-depth characterization of compound-specific needle carbohydrate isotope variability, identifies involved mechanisms and shows the potential of such results for linking tree physiological responses to different climatic conditions.

Changes in climate, land use and management have led to an increase in woody species encroachment into grasslands, threatening key ecosystem services and resilience. This study uses dendroecological techniques to investigate acclimation strategies of two native woody species, Juniperus virginiana and Pinus ponderosa, to spatial and temporal variability in precipitation and stand density in Nebraska (NE). Two locations in Eastern NE and one in Western NE, were selected and tree cores for ring-width measurements were collected. Stable Isotope ratios of carbon were analysed to investigate water use efficiency (WUE), and stable isotope ratios of oxygen were used to understand stomatal conductance and control. Our results showed trees in dense stands exhibited less intra-specific variability in response to climate and narrower tree rings than sparse open stands, with P. ponderosa being more responsive to density and climate variability than J. virginiana. Populations in the drier Sandhills grasslands of Western NE were more dependent on intra- and inter-annual precipitation and generally more impacted by drought events than those in the East. Eastern NE trees were less limited by drought and displayed lower WIT relative to Western location, even though temperatures were slightly higher, but this was compensated for by the overall higher precipitation levels in the East. Generally, above average winter temperatures were positively correlated with ring widths, while temperature extremes during the growing season were negatively correlated. Although years of extreme drought events were visible in the tree-ring width patterns for both species independent of stand density and location, it seems that once the trees are established and have access to soil moisture, the current climate variability and extremes in NE do not limit or cause a permanent it decline in growth of either species.

Global warming is most pronounced in high-latitude regions by altering habitat conditions and affecting permafrost degradation, which may significantly influence tree productivity and vegetation changes. In this study, by applying a “space-for-time” approach, we selected three plots of Larix gmelinii forest from a continuous permafrost zone in Siberia with different thermo-hydrological soil regimes and ground cover vegetation with the objective of assessing how tree growth and productivity will change under different stages of permafrost degradation. A tree-ring multi-proxy characterization of mature trees was used to identify shift in ecophysiological responses related to the modified plant-soil system. Variability of tree-ring width (1975–2009), stable isotope ratios (oxygen and carbon, 2000–2009) and xylem structural characteristics (2000–2009) under climatic conditions of particular years indicated that an increased depth of the soil active layer will initially lead to increase of tree productivity. However, due to an expected water use increase through transpiration, the system might progressively shift from a temperature to a moisture-limited environment.

Stable isotope ratios in tree rings have become an important proxy for palaeoclimatology, particularly in temperate regions. Yet temperate forests are often characterized by heterogeneous stand structures, and the effects of stand dynamics on carbon (δ

Rationale Levoglucosan is formed from cellulose during biomass burning. It is therefore often used as a specific tracer to quantify the contribution of wood burning to the aerosol loading. The stable oxygen isotope composition (δ18 O value) of biomass is determined by the water cycle and varies regionally, and hence the δ18 O value of levoglucosan could help to identify source regions of organic aerosols. Methods After solvent extraction of the organic fraction and concentration steps, a recently developed methylation derivatisation technique was applied on experimental (i.e. controlled wood-burning experiments) and on ambient aerosol samples from Switzerland and Lithuania. The method achieves sufficient compound separation for isotope analysis in atmospheric particulate matter, enabling δ18 O analysis of levoglucosan by gas chromatography/pyrolysis-isotope ratio mass spectrometry (GC/Pyr-IRMS), with a precision better than 1.0 ‰ and an accuracy of 0.3 ‰. Results The δ18 O value of the levoglucosan released during controlled wood-burning experiments was not significantly different from the cellulose δ18 O values, which implies very little or no isotope fractionation during wood burning under the given conditions. While the δ18 O values of levoglucosan in Swiss samples were as expected for the source region, those in Lithuania were 1-4 ‰ lower than expected. This may be due to differences in vegetation (grass vs wood) or burning conditions (high vs low temperatures). Conclusions Low oxygen isotope fractionation between cellulose and levoglucosan and clear differences in levoglucosan δ18 O values between the Swiss and Lithuanian ambient samples demonstrate that our new method is useful for source appointment studies on wood-burning-derived aerosols.

Our understanding of how temporal variations of atmospheric water vapour and its isotopic composition (δ

Recently published, improved chronologies for volcanic sulfate in Greenland and Antarctic ice permit a comparison of the growth responses of absolutely annually dated tree rings at three locations in Siberia with annual ice-core records of volcanic eruptions centered on AD 536. For the first time for this region and period, we present unique data sets for tree-ring width, cell-wall thickness, δ13C and δ18O in cellulose. These were based on multiple samples from relict wood of larch obtained from two sites close to the northern limit of tree growth on the Taimyr Peninsula and in northeastern Yakutia, and at a high-elevation, location 20° further South in the Altai Mts. An event in AD 536 was associated with different, but marked, changes in tree-ring parameters at the high-latitude sites compared with the high elevation site. An AD 541 event was associated with its own distinctive tree-ring responses across the three sites and multiple variables. The years after AD 532 were marked by a strong and sustained decrease in growth at the high-elevation, more southerly, site. The combination of improved ice-core chronology for the climatically effective volcanic eruptions of this part of the 6th century AD, and an array of tree-ring sites with different climates and multiple tree-ring variables permits a richer description of tree responses to this cluster of events. The pattern of tree-ring parameter responses at the three locations in AD 536, AD 541, and perhaps AD 532 is consistent with responses to climatically effective volcanic eruptions influencing tree response in those and subsequent years.

We present a 1200 year drought reconstruction for the European Alpine region based on carbon isotope variations of tree rings from living larch trees and historic timber. The carbon isotope fractionation at the study site is sensitive to summer precipitation, temperature, and irradiance, resulting in a stable and high correlation with a drought index for interannual to decadal frequencies and possibly beyond (r(2)=0.58 for 1901-2004, July/August). When combining this information with maximum latewood density-derived summer temperature, a strongly reduced occurrence of summer droughts during the warm A.D. 900-1200 period is evident, coinciding with the Medieval Climate Anomaly (MCA), with a shift to colder and drier conditions for the subsequent centuries. The warm-wet MCA contrasts strongly with the climate of the drought-prone warm phase of the recent decades, indicating different forcing mechanism for these two warm periods and pointing to beneficial conditions for agriculture and human well-being during the MCA in this region.

Regulation of stomatal (gs ) and mesophyll conductance (gm ) is an efficient means for optimizing the relationship between water loss and carbon uptake in plants. We assessed water-use efficiency (WUE)-based drought adaptation strategies with respect to mesophyll conductance of different functional plant groups of the forest understory. Moreover we aimed at assessing the mechanisms of and interactions between water and CO2 conductance in the mesophyll. The facts that an increase in WUE was observed only in the two species that increased gm in response to moderate drought, and that over all five species examined, changes in mesophyll conductance were significantly correlated with the drought-induced change in WUE, proves the importance of gm in optimizing resource use under water restriction. There was no clear correlation of mesophyll CO2 conductance and the tortuosity of water movement in the leaf across the five species in the control and drought treatments. This points either to different main pathways for CO2 and water in the mesophyll either to different regulation of a common pathway.

Will warming lead to an increased use of older soil organic carbon (SOC) by microbial communities, thereby inducing C losses from C-rich alpine soils? We studied soil microbial community composition, activity, and substrate use after 3 and 4 years of soil warming (+4 °C, 2007-2010) at the alpine treeline in Switzerland. The warming experiment was nested in a free air CO2 enrichment experiment using depleted (13)CO2 (δ(13)C = -30‰, 2001-2009). We traced this depleted (13)C label in phospholipid fatty acids (PLFA) of the organic layer (0-5 cm soil depth) and in C mineralized from root-free soils to distinguish substrate ages used by soil microorganisms: fixed before 2001 ('old'), from 2001 to 2009 ('new') or in 2010 ('recent'). Warming induced a sustained stimulation of soil respiration (+38%) without decline in mineralizable SOC. PLFA concentrations did not reveal changes in microbial community composition due to soil warming, but soil microbial metabolic activity was stimulated (+66%). Warming decreased the amount of new and recent C in the fungal biomarker 18:2ω6,9 and the amount of new C mineralized from root-free soils, implying a shift in microbial substrate use toward a greater use of old SOC. This shift in substrate use could indicate an imbalance between C inputs and outputs, which could eventually decrease SOC storage in this alpine ecosystem.

Relevance of spring temperatures for tree-ring growth steadily increased since 1950s. Closely linked tree-ring growth and net CO 2 exchange driven by spring temperatures. We investigated long-term (over 100 years) tree-ring width (TRW) variabilities as well as short-term (10 years) variations in net ecosystem productivity (NEP) in response to climate to assess the driving factors for stem growth of Norway spruce in a subalpine forest at Davos in Switzerland. A tree-ring width index (TRWi) chronology for the period from 1750 to 2006 was constructed and linked with climate data from 1876 to 2006, and with NEP available for the period from 1997 to 2006. Based on TRWi, we found that only two out of the 257 years exhibited extreme negative TRWi, compared to 29 years with extreme positive anomalies, observed mainly in recent decades. Annual temperature, annual precipitation, as well as autumn and winter temperature signals were well preserved in the TRWi chronology over the last 130 years. Spring temperatures became increasingly relevant for TRWi, explaining less than 1 % of the variation in TRWi for the period from 1876 to 2006, but 8 % for the period from 1950 to 2006 (p = 0.032), and even 47 % for 1997–2006 (p = 0.028). We also observed a strong positive relationship between annual TRWi and annual NEP (r = 0.661; p = 0.037), both strongly related to spring temperatures (r = 0.687 and r = 0.678 for TRWi and NEP, respectively; p = 0.028; p = 0.032). Moreover, we found strong links between monthly NEP of March and annual TRWi (r = 0.912; p = 0.0001), both related to March temperatures (r = 0.767, p = 0.010 and r = 0.724, p = 0.018, respectively). Thus, under future climate warming, we expect stem growth of these subalpine trees and also ecosystem carbon (C) sequestration to increase, as long as water does not become a limiting factor.

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 19012000. The results were compared with simulations of a dynamic vegetation model (LPX-Bern 1.0) that integrates numerous ecosystem and landatmosphere 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 of iWUE. These findings will help to reduce uncertainties in the land surface schemes of global climate models, where vegetationclimate feedbacks are currently still poorly constrained by observational data.

With the aim of evaluating the influence of glacial meltwater signature on tree-ring stable isotopes, we analysed δ18O and δ13C in the tree rings of Larix decidua Mill. specimens growing in the area of an ice-contact lake (Lago Verde, at Miage Glacier, European Alps). Additionally, we analysed δ18O in the glacial meltwater of the lake and of the glacier stream and compared it with the δ18O of precipitation predicted by a spatial model. We found that tree-ring cellulose of trees fed by glacial meltwaters (LVW site) is significantly more depleted in δ18O than at a control site LVM (−0.91‰) fed only by precipitation, thus reflecting the measured higher depletion of glacial meltwaters with respect to local precipitation. δ13C values did not show significant differences in mean values between the two sites but an anomalous correlation with summer temperature was found at the LVW site, probably due to the different responses of trees stomatal conductance. Over the 30-year period of analysis, four years at LVW (1992, 1995, 2003 and 2009) were markedly depleted in δ18O. These years are those when the highest summer temperatures were recorded in the area (the ones during which glacier ablation usually increases and more depleted meltwaters fill the lake), with the exception of 1995 during which high water levels occurred following the year with the second highest summer temperature (1994). Overall, our analysis demonstrates that tree-ring δ18O, driven by the glacial meltwater signature in the lake, can be used for detecting past major glacier runoff events. The proposed approach could also be used for quantifying past glacier runoff and for defining past distribution areas of glacial meltwaters in glacier forefields, thus contributing to past environmental reconstructions and to hazard assessment.

Erica arborea (L) is a widespread Mediterranean species, able to cope with water stress and colonize semiarid environ- ments. The eco-physiological plasticity of this species was evaluated by studying plants growing at two sites with differ- ent soil moistures on the island of Elba (Italy), through dendrochronological, wood-anatomical analyses and stable isotopes measurements. Intra-annual density fluctuations (IADFs) were abundant in tree rings, and were identified as the key parameter to understand site-specific plant responses to water stress. Our findings showed that the formation of IADFs is mainly related to the high temperature, precipita- tion patterns and probably to soil water availability, which differs at the selected study sites. The recorded increase in the 13 C-derived intrinsic water use efficiency at the IADFs level was linked to reduced water loss rather than to increas- ing C assimilation.The variation in vessel size and the differ- ent absolute values of δ 18 O among trees growing at the two study sites underlined possible differences in stomatal control of water loss and possible differences in sources of water uptake. This approach not only helped monitor sea- sonal environmental differences through tree-ring width, but also added valuable information on E. arborea responses to drought and their ecological implications for Mediterranean vegetation dynamics.

To answer the question “Has the recent warming no analogues in the Siberian north?” we analyzed larch tree samples ( Larix gmelinii Rupr.) from permafrost zone in the eastern Taimyr (TAY) (72°N, 102°E) using tree-ring and stable isotope analyses for the Climatic Optimum Period (COP) 4111–3806 BC and Medieval Warm Period (MWP) 917–1150 AD, in comparison to the recent period (RP) 1791–2008 AD. We developed a description of the climatic and environmental changes in the eastern Taimyr using tree-ring width and stable isotope (δ 13 C, δ 18 O) data based on statistical verification of the relationships to climatic parameters (temperature and precipitation). Additionally, we compared our new tree-ring and stable isotope data sets with earlier published July temperature and precipitation reconstructions inferred from pollen data of the Lama Lake, Taimyr Peninsula, δ 18 O ice core data from Akademii Nauk ice cap on Severnaya Zemlya (SZ) and δ 18 O ice core data from Greenland (GISP2), as well as tree-ring width and stable carbon and oxygen isotope data from northeastern Yakutia (YAK). We found that the COP in TAY was warmer and drier compared to the MWP but rather similar to the RP. Our results indicate that the MWP in TAY started earlier and was wetter than in YAK. July precipitation reconstructions obtained from pollen data of the Lama Lake, oxygen isotope data from SZ and our carbon isotopes in tree cellulose agree well and indicate wetter climate conditions during the MWP. Consistent large-scale patterns were reflected in significant links between oxygen isotope data in tree cellulose from TAY and YAK, and oxygen isotope data from SZ and GISP2 during the MWP and the RP. Finally, we showed that the recent warming is not unprecedented in the Siberian north. Similar climate conditions were recorded by tree-rings, stable isotopes, pollen, and ice core data 6000 years ago.

Carbon reserves are important for maintaining tree function during and after stress. Increasing tree mortality driven by drought globally has renewed the interest in how plants regulate allocation of recently fixed C to reserve formation. Three-year-old seedlings of two species (Tilia platyphyllos and Pinus sylvestris) were exposed to two intensities of experimental drought during ~10 weeks, and

International audience; We analyzed the chronologies of cellulose stable isotopes (δ13C and δ18O) and tree-ring widths from European larch (Larix decidua) in a high-altitude site (2190ma.s.l.) at the bottom of a glacial valley in the Italian Alps, and investigated their dependence on monthly meteorological variables and δ18O precipitation values. The δ18O of tree-ring cellulose appears to be strongly driven by the δ18O of winter snowfall (November to March), which suggests that larch trees mostly use the snow-melt water of the previous winter during the growing season. This water, which also comes from the slope streams and from the underground flow of nearby steep slopes, infiltrates the soil in the valley bottom. The tree-ring cellulose δ18O values were also found to be influenced by the August precipitation δ18O and mean temperature. The associated regression model shows that the δ18O chronology from the tree rings explains up to 34% of the variance in the winter precipitation δ18O record, demonstrating the potential for reconstructing the δ18O isotopic composition of past winter precipitation in the study region. Unlike most other tree-ring studies that focus on growing season signals, in our study the summer signal was small and the winter signal dominant due to the special conditions of the glacial valley. Site topography, geomorphology and soil characteristics in particular influence the stable isotope signal in tree-ring cellulose.

Understanding climate change impacts on drought-prone forests is a critical issue. We investigated ring-width and stable isotopes (Δ13C and δ18O) in two Pinus sylvestris stands of the cold-dry Siberian forest-steppe growing under contrasting climatic trends over the last 75 years. Despite regional warming, there was increasing precipitation during the growing period at the southern site (MIN) but increasing water deficit (WD) at the northern site (BER). Intrinsic water use efficiency (WUEi) increased similarly (ca. 22%) in response to warming and rising atmospheric CO2. However, the steady increase in WUEi was accompanied by divergent growth patterns since 1980: increasing basal area increment (BAI) in MIN (slope = 0.102 cm2 year−2) and decreasing BAI in BER (slope = −0.129 cm2 year−2). This suggests that increased precipitation, mediated by CO2 effects, promoted growth in MIN, whereas intensified drought stress led to decreased carbon gain and productivity in BER. When compared to warm-dry stands of eastern Spain, the WUEi dependence on WD was three-fold greater in Siberia. Conversely, BAI was more affected by the relative impact of water stress within each region. These results indicate contrasting future trajectories of P. sylvestris forests, which challenge forecasting growth and carbon sequestration in cold-dry areas. This study was funded by the Spanish Government (grant number AGL2015-68274-C3-3-R) and the Russian Science Foundation (project numbers 14-14-00295, sampling and tree-ring data obtainment and 14-14-00219-P, mathematical approach). We acknowledge P. Sopeña and M.J. Pau for technical assistance.

The 2003 and 2012 summer seasons were among the warmest and driest of the last 200 years over southeastern Europe, and in particular in the Karst region (northeastern Italy). Starting from winter-spring 2013, several black pines (Pinus nigra J.F. Arnold) suffered crown die-back. Declining trees occurred nearby individuals with no signs of die-back, raising hypotheses about the occurrence of individual-specific hydraulic strategies underlying different responses to extreme drought. We investigated possible processes driving black pine decline by dendrochronological and wood anatomical measurements, coupled with analysis of tree-ring carbon (δ13C) and oxygen (δ18O) isotopic composition in healthy trees (H) and trees suffering die-back (D). Die-back trees showed higher growth rates than H trees at the beginning of the last century, but suffered important growth reduction following the dry summers in 2003 and 2012. After the 2012 drought, D trees produced tracheids with larger diameter and greater vulnerability to implosion than H ones. Healthy trees had significantly higher wood δ13C than D trees, reflecting higher water-use efficiency for the surviving trees, i.e., less water transpired per unit carbon gain, which could be related to lower stomatal conductance and a more conservative use of water. Relatively high δ18O for D trees indicates that they were strongly dependent on shallow water sources, or that they sustained higher transpiration rates than H trees. Our results suggest that H trees adopted a more conservative water-use strategy under drought stress compared with D trees. We speculate that this diversity might have a genotypic basis, but other possible explanations, like different rooting depth, cannot be ruled out.

Although the instrumental coupling of gas chromatography pyrolysis isotope ratio mass spectrometry (GC Py IRMS) for compound specific O 18 analysis has been commercially available for more than a decade this method has been hardly applied so far. Here we present the first GC Py IRMS O 18 results for trimethylsilyl derivatives of plant sap relevant sugars and a polyalcohol (glucose fructose sucrose raffinose and pinitol). Particularly we focus on sucrose which is assimilated in leaves and which is the most important transport sugar in plants and hence of utmost relevance in plant physiology and paleoclimate studies. Replication measurements of sucrose standards and concentration series indicate that the GC Py IRMS O 18 measurements are not stable over time and that they are amount (area) dependent. We therefore suggest running sample batch replication measurements in alternation with standard concentration series of reference material. This allows for carrying out (i) a drift correction (ii) a calibration against reference material and (iii) an amount (area) correction. Tests with O 18 enriched water do not provide any evidence for oxygen isotope exchange reactions affecting sucrose and raffinose. We present the first application of GC Py IRMS O 18 analysis for sucrose from needle extract (soluble carbohydrate) samples. The obtained O 18(sucrose/ Vienna Standard Mean Ocean Water (VSMOW)) values are more positive and vary in a wider range (32.1 40.1 parts per thousand) than the O 18(bulk/ VSMOW) values (24.6 27.2 parts per thousand). Furthermore they are shown to depend on the climate parameters maximum day temperature relative air humidity and cloud cover. These findings suggest that O 18(sucrose) of the investigated needles very sensitively reflects the climatically controlled evaporative O 18 enrichment of leaf water and thus highlights the great potential of GC Py IRMS O 18(sucrose) analysis for plant physiology and paleoclimate studies.

Time series of tree-ring growth show significant increases since the early 1970s at the alpine tree line, with simultaneously increasing temperatures and atmospheric CO2 concentration. For a comprehensive understanding of this growth change, the physiological response patterns at both the leaf and stem level need to be separately analyzed and identified, and can be retrieved from tree-ring growth and isotope (δ13C, δ18O) series. In this study, we assessed the relative contribution of environmental factors to interannual tree-ring variability by multivariate linear mixed-effects models and the dual isotope approach on a dataset of tree-ring records of ~400-year-old larch (Larix decidua Mill.) from a non-water-limited high-elevation site in the Swiss Alps. The models suggest that summer temperatures and the recent lack of larch budmoth outbreaks were most important for explaining growth variations and trends, while a significant direct effect of the continuously increasing CO2 concentration could not be confirmed. In contrast, δ13C and δ18O, which are strongly influenced by fractionation changes in the leaf, clearly reflected the impact of air humidity (precipitation and vapor pressure deficit) and CO2 concentration: the increase in (δ13C-derived) intrinsic water-use efficiency over the second half of the 20th century suggests an increase in carbon assimilation as a result of enhanced CO2 concentration. The tree-ring δ18O largely reflected recent precipitation as source water, thus indicating a low variability in stomatal conductance, which was confirmed by the dual isotope approach. These leaf-level effects were not reflected in stem growth as they may have been masked by the temperature-caused growth limitation controlling the allocation of increased amounts of photosynthates into wood growth. Our approach demonstrates that the identification of different roles of environmental factors on leaf and stem processes helps to improve the assessment of site-specific changes of carbon fluxes and growth performance under future environmental conditions.

Measurements of the stable carbon isotope ratio (δ13C) on annual tree rings offer new opportunities to evaluate mechanisms of variations in photosynthesis and stomatal conductance under changing CO2 and climate, especially in conjunction with process-based biogeochemical model simulations. The isotopic discrimination is indicative of the ratio between the CO2 partial pressure in the intercellular cavities and the atmosphere (ci / ca) and of the ratio of assimilation to stomatal conductance, termed intrinsic water-use efficiency (iWUE). We performed isotope-enabled simulations over the industrial period with the land biosphere module (CLM4.5) of the Community Earth System Model and the LPX-Bern dynamic global vegetation model. Results for C3 tree species show good agreement with a global compilation of δ13C measurements on leaves, though modeled 13C discrimination by C3 trees is smaller in arid regions than measured. A compilation of seventy-six tree-ring records, mainly from Europe, boreal Asia, and western North America, suggest on average small 20th-century changes in isotopic discrimination and an increase in iWUE of about 27 % since 1900. LPX-Bern results match these century-scale reconstructions, supporting the idea that the physiology of stomata has evolved to optimize trade-offs between carbon gain by assimilation and water loss. In contrast, CLM4.5 simulates an increase in discrimination and in turn a change in iWUE that is almost twice as large as revealed by the tree-ring data. Factorial simulations show that these changes are mainly in response to rising atmospheric CO2. The results suggest that the down-regulation of ci / ca and of photosynthesis by nitrogen limitation is possibly too strong in the standard setup of CLM4.5 or there may be more fundamental problems associated with the prescribed relationship between conductance and assimilation.

We analyzed the chronologies of cellulose stable isotopes (δ

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Summary How will carbon source–sink relations of 35-yr-old larch trees (Larix decidua) at the alpine treeline respond to changes in atmospheric CO2 and climate? We evaluated the effects of previously elevated CO2 concentrations (9 yr, 580 ppm, ended the previous season) and ongoing soil warming (4 yr, + 4°C). Larch branches were pulse labeled (50 at% 13CO2) in July 2010 to trace fresh assimilates through tissues (buds, needles, bark and wood) and non-structural carbon compounds (NCC; starch, lipids, individual sugars) using compound-specific isotope analysis. Nine years of elevated CO2 did not lead to increased NCC concentrations, nor did soil warming increase NCC transfer velocities. By contrast, we found slower transfer velocities and higher NCC concentrations than reported in the literature for lowland larch. As a result of low dilution with older carbon, sucrose and glucose showed the highest maximum 13C labels, whereas labels were lower for starch, lipids and pinitol. Label residence times in needles were shorter for sucrose and starch (c. 2 d) than for glucose (c. 6 d). Although our treatments showed no persistent effect on larch carbon relations, low temperature at high altitudes clearly induced a limitation of sink activities (growth, respiration, root exudation), expressed in slower carbon transfer and higher NCC concentrations.

RATIONALE Isotope analysis of carbohydrates is important for improved understanding of plant carbon metabolism and plant physiological response to the environment. High-performance liquid chromatography/isotope ratio mass spectrometry (HPLC/IRMS) for direct compound-specific δ13C measurements of soluble carbohydrates has recently been developed, but the still challenging sample preparation and the fact that no single method is capable of separating all compounds of interest hinder its wide-spread application. Here we tested in detail a chromatography method in alkaline media. METHODS We examined the most suitable chromatographic conditions for HPLC/IRMS analysis of carbohydrates in aqueous conifer needle extracts using a CarboPac PA20 anion-exchange column with NaOH eluent, paying specific attention to compound yields, carbon isotope fractionation processes and the reproducibility of the method. Furthermore, we adapted and calibrated sample preparation methods for HPLC/IRMS analysis. OnGuard II cartridges were used for sample purification. RESULTS Good peak separation and highly linear and reproducible concentration and δ13C measurements were obtained. The alkaline eluent was observed to induce isomerization of hexoses, detected as reduced yields and 13 C fractionation of the affected compounds. A reproducible pre-purification method providing ~100% yield for the carbohydrate compounds of interest was calibrated. CONCLUSIONS The good level of peak separation obtained in this study is reflected in the good precision and linearity of concentration and δ13C results. The data provided crucial information on the behaviour of sugars in LC analysis with alkaline media. The observations highlight the importance for the application of compound-matched standard solution for the detection and correction of instrumental biases in concentration and δ13C analysis performed under identical chromatographic conditions. The calibrated pre-purification method is well suited for studies with complex matrices that disable the use of a spiked internal standard for the detection of procedural losses. Copyright © 2012 John Wiley & Sons, Ltd.

Stable isotopes ratios in tree-rings are increasingly used to assess long-term environmental variation. As with conventional tree-ring investigations, crossdating overlapping sequences of wood material allows chronologies longer than the lifespans of individual trees to be developed. We measured data from four overlapping cohorts of recent (AD 1650–2004) and historic (AD 1100–1198, AD 1150–1325, AD 1256–1700) wood to yield millennial-length δ 13 C , δ ​ D and δ 18 O chronologies. However, we found that offsets in absolute isotopic values between overlapping segments challenge the preservation of low frequency climate signals. In this paper, we describe the development of these records, signal strength assessments, and present the problems associated with overcoming these offsets and preserving low-frequency variance. We then propose and evaluate four different solutions to link the cohorts. These methods include various types of transformations of mean and variance; all present potential advantages and drawbacks. Inflated variance, loss of absolute isotope values, and the loss of low-frequency signals are major problems. Only one method is capable of fully preserving low-frequency trends, but this method may also risk inducing long-term artifacts. While the various solutions we propose may be useful to help overcome such biases, the challenges described here, if typical for other isotopic series, will best be solved by additionally modifying sampling and measurement procedures. We recommend avoiding pooling of tree rings in the overlap period and to increase the replication in order to extract the original signal and to maintain low-frequency signals.

We investigated whether stand structure modulates the long-term physiological performance and growth of Pinus halepensis Mill. in a semiarid Mediterranean ecosystem. Tree radial growth and carbon and oxygen stable isotope composition of latewood (δ(13)C(LW) and δ(18)O(LW), respectively) from 1967 to 2007 were measured in P. halepensis trees from two sharply contrasting stand types: open woodlands with widely scattered trees versus dense afforested stands. In both stand types, tree radial growth, δ(13)C(LW) and δ(18)O(LW) were strongly correlated with annual rainfall, thus indicating that tree performance in this semiarid environment is largely determined by inter-annual changes in water availability. However, trees in dense afforested stands showed consistently higher δ(18)O(LW) and similar δ(13)C(LW) values compared with those in neighbouring open woodlands, indicating lower stomatal conductance and photosynthesis rates in the former, but little difference in water use efficiency between stand types. Trees in dense afforested stands were more water stressed and showed lower radial growth, overall suggesting greater vulnerability to drought and climate aridification compared with trees in open woodlands. In this semiarid ecosystem, the negative impacts of intense inter-tree competition for water on P. halepensis performance clearly outweigh potential benefits derived from enhanced infiltration and reduced run-off losses in dense afforested stands.

Intra-annual density fluctuations (IADFs) are anomalies of tree rings where wood density is abruptly altered after sudden changes in environmental conditions. Their characterisation can provide information about the relationship between environmental factors and eco-physiological processes during tree growth. This paper reports about the variability of anatomical traits and stable carbon isotopic composition along tree rings as resulting from the application of two different methodological approaches: (a) the separation of each ring into different regions (earlywood, latewood and IADF) and the comparison of anatomical and isotopic parameters measured in those specific sectors and (b) the analysis of such features in continuum along ring width. Moreover, different parameters of vessels (i. e. ecd-equivalent circle diameter, elongation, sphericity and convexity of vessel lumen) were considered to identify those more appropriate for the representation of intra-annual anatomical variations. The analysis was conducted on Arbutus unedo L. growing on the Elba Island (Italy); tree rings of this species form IADFs with features clearly responsive to the environmental conditions experienced during plant growth. Results showed that the first approach, although more suitable to obtain data for subsequent statistical comparisons and for the calculation of correlations with environmental parameters, suffers from elements of subjectivity due to the size and position of the selected tree-ring regions. The in continuum method allows a clearer identification of the variation of tree-ring properties along ring width. Regarding anatomical parameters, shape indexes were not suitable indicators of intra-annual variability. The overall analysis suggested that using both methodologies in synergy helps to gain complete information and avoid misleading interpretations of IADFs in tree rings. © 2011 Springer-Verlag.

The idea that photosynthesis-weighted tree canopy leaf temperature (T(canδ)) can be resolved through analysis of oxygen isotope composition in tree wood cellulose (δ(18) O(wc)) has led to the observation of boreal-to-subtropical convergence of T(canδ) to c. 20°C. To further assess the validity of the large-scale convergence of T(canδ), we used the isotope approach to perform calculation of T(canδ) for independent δ(18) O(wc) data sets that have broad coverage of climates. For the boreal-to-subtropical data sets, we found that the deviation of T(canδ) from the growing season temperature systemically increases with the decreasing mean annual temperature. Across the whole data sets we calculated a mean T(canδ) of 19.48°C and an SD of 2.05°C, while for the tropical data set, the mean T(canδ) was 26.40 ± 1.03°C, significantly higher than the boreal-to-subtropical mean. Our study thus offers independent isotopic support for the concept that boreal-to-subtropical trees display conserved T(canδ) near 20°C. The isotopic analysis cannot distinguish between the possibility that leaf temperatures are generally elevated above ambient air temperatures in cooler environments and the possibility that leaf temperature equals air temperature, whereas the leaf/air temperature at which photosynthesis occurs has a weighted average of near 20°C in cooler environments. Future work will separate these potential explanations.