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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.

Warming alters the variability and trajectories of tree growth around the world by intensifying or alleviating energy and water limitation. This insight from regional to global-scale research emphasizes the susceptibility of forest ecosystems and resources to climate change. However, globally-derived trends are not necessarily meaningful for local nature conservation or management considerations, if they lack specific information on present or prospective tree species. This is particularly the case towards the edge of their distribution, where shifts in growth trajectories may be imminent or already occurring.Importantly, the geographic and bioclimatic space (or “niche”) occupied by a tree species is not only constrained by climate, but often reflects biotic pressure such as competition for resources with other species. This aspect is underrepresented in many species distribution models that define the niche as a climatic envelope, which is then allowed to shift in response to changes in ambient conditions. Hence, distinguishing climatic from competitive niche boundaries becomes a central challenge to identifying areas where tree species are most susceptible to climate change.Here we employ a novel concept to characterize each position within a species’ bioclimatic niche based on two criteria: a climate sensitivity index (CSI) and a habitat suitability index (HSI). The CSI is derived from step-wise multiple linear regression models that explain variability in annual radial tree growth as a function of monthly climate anomalies. The HSI is based on an ensemble of five species distribution models calculated from a combination of observed species occurrences and twenty-five bioclimatic variables. We calculated these two indices for 11 major tree species across the Northern Hemisphere.The combination of climate sensitivity and habitat suitability indicated hotspots of change, where tree growth is mainly limited by competition (low HSI and low CSI), as well as areas that are particularly sensitive to climate variability (low HSI and high CSI). In the former, we expect that forest management geared towards adjusting the competitive balance between several candidate species will be most effective under changing environmental conditions. In the latter areas, selecting particularly drought-tolerant accessions of a given species may reduce forest susceptibility to the predicted warming and drying.

Picea abies and Fagus sylvatica, are two of the most important tree species in Europe, and their responses to climate are being extensively investigated, especially at the limits of their distribution. However, their physiology at temperate sites is not yet fully understood. In a European tree-ring network, 10 sites along a climate gradient were sampled throughout Central Europe, and tree-ring width and stable isotope chronologies (C and O) were measured. The year-to-year variability of the isotopes time series for the last 100 years was analyzed in relation to tree-ring growth, spatial distribution, and seasonal climate.Climate sensitivity of radial growth of both species was rather variable and site-dependent, and was strongest at the driest sites. On the contrary, variability in the isotopic ratios consistently responded to summer climate, particularly to vapor pressure deficit. The high δ18O coherence of the short-term variability between sites and species highlights the strength of the environmental signal in the O chronology also across long distances. On the contrary, δ13C shows lower correlations between sites and species, showing a stronger site-dependency, and a lower intra-annual variability. The generally positive correlation between the year-to-year differences in δ13C and δ18O across most sites demonstrates the strong role of stomatal conductance in controlling leaf gas exchange for these species. However, in the last decades, sites showed a dissimilar shift in the isotopes relationships, with the warmer sites showing an increase of either or both δ13C and δ18O and consequent decrease of photosynthetic rates and stomatal conductance, highlighting their dependency to atmospheric moisture demand and soil water availability.Understanding the underlying physiological mechanisms controlling the short-term variation in tree-ring records will help with defining the performance of these ecologically and economically important tree species under future climate conditions.

Climate−tree growth relationships recorded in annual growth rings have recently been the basis for projecting climate change impacts on forests. However, most trees and sample sites represented in the International Tree-Ring Data Bank (ITRDB) were chosen to maximize climate signal and are characterized by marginal growing conditions not representative of the larger forest ecosystem. We evaluate the magnitude of this potential bias using a spatially unbiased tree-ring network collected by the USFS Forest Inventory and Analysis (FIA) program. We show that U.S. Southwest ITRDB samples overestimate regional forest climate sensitivity by 41–59%, because ITRDB trees were sampled at warmer and drier locations, both at the macro- and micro-site scale, and are systematically older compared to the FIA collection. Although there are uncertainties associated with our statistical approach, projection based on representative FIA samples suggests 29% less of a climate change-induced growth decrease compared to projection based on climate-sensitive ITRDB samples., Sampling strategies may bias tree-ring datasets to not accurately represent the regional response to climate change. Here, Klesse et al. use a new representative dataset to show that the International Tree-Ring Data Bank in the U.S. Southwest overestimates climate sensitivity of forests by 41–59%

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 (δ13 C) and oxygen isotope ratios (δ18 O) in tree rings are not well explored. In this study, we investigated long-term trends and offsets in δ18 O and δ13 C of Picea abies and Fagus sylvatica in relation to tree age, size, and distance to the upper canopy at seven temperate sites across Europe. We observed strong positive trends in δ13 C that are best explained by the reconstructed dynamics of individual trees below the upper canopy, highlighting the influence of light attenuation on δ13 C in shade-tolerant species. We also detected positive trends in δ18 O with increasing tree size. However, the observed slopes are less steep and consistent between trees of different ages and thus can be more easily addressed. We recommend restricting the use of δ13 C to years when trees are in a dominant canopy position to infer long-term climate signals in δ13 C when relying on material from shade-tolerant species, such as beech and spruce. For such species, δ18 O should be in principle the superior proxy for climate reconstructions.

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 conditions, 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 Land Surface Processes and Exchanges (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 76 tree-ring records, mainly from Europe, boreal Asia, and western North America, suggests on average small 20th century changes in isotopic discrimination and in ci∕ca 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 by transpiration. In contrast, CLM4.5 simulates an increase in discrimination and in turn a change in iWUE that is almost twice as large as that 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 downregulation of ci∕ca and of photosynthesis by nitrogen limitation is possibly too strong in the standard setup of CLM4.5 or that there may be problems associated with the implementation of conductance, assimilation, and related adjustment processes on long-term environmental changes.

A central challenge in global change research is the projection of the future behavior of a system based upon past observations. Tree-ring data have been used increasingly over the last decade to project tree growth and forest ecosystem vulnerability under future climate conditions. But how can the response of tree growth to past climate variation predict the future, when the future does not look like the past? Space-for-time substitution (SFTS) is one way to overcome the problem of extrapolation: the response at a given location in a warmer future is assumed to follow the response at a warmer location today. Here we evaluated an SFTS approach to projecting future growth of Douglas-fir (Pseudotsuga menziesii), a species that occupies an exceptionally large environmental space in North America. We fit a hierarchical mixed-effects model to capture ring-width variability in response to spatial and temporal variation in climate. We found opposing gradients for productivity and climate sensitivity with highest growth rates and weakest response to interannual climate variation in the mesic coastal part of Douglas-fir's range; narrower rings and stronger climate sensitivity occurred across the semi-arid interior. Ring-width response to spatial versus temporal temperature variation was opposite in sign, suggesting that spatial variation in productivity, caused by local adaptation and other slow processes, cannot be used to anticipate changes in productivity caused by rapid climate change. We thus substituted only climate sensitivities when projecting future tree growth. Growth declines were projected across much of Douglas-fir's distribution, with largest relative decreases in the semiarid U.S. Interior West and smallest in the mesic Pacific Northwest. We further highlight the strengths of mixed-effects modeling for reviving a conceptual cornerstone of dendroecology, Cook's 1987 aggregate growth model, and the great potential to use tree-ring networks and results as a calibration target for next-generation vegetation models.

Disturbances, both natural and human induced, influence forest dynamics, ecosystem functioning, and ecosystem services. Here, we aim to evaluate the consequences of natural disturbances on the magnitude and dynamics of tree- and stand-level biomass accumulation from decadal to centennial scales. We use tree-ring data from 2301 trees and biometric data from 4909 trees sampled in 96 plots (each 1000 m 2 ) to quantify the influence of mixed severity disturbance regimes on annual aboveground biomass increment (AGBI) and total aboveground biomass accumulation (AGB) across a mountainous monotypic Norway spruce ( Picea abies (L.) Karst.) primary forest. We hypothesise that the multiple internal and external factors constraining tree growth will cause differences in tree and stand-level biomass trajectories in these natural forests. Although we found that tree-level AGB growth increases with tree size, we also found that tree age and disturbance legacies plays a crucial role for AGB in the investigated Norway spruce forests. Importantly, while younger trees of the same diameter class have an average current AGBI rate that is ∼225% higher than older trees (300–400 years), we find trees that have been suppressed for up to 120 years can respond vigorously when competition is reduced. On average, post disturbance AGBI was ∼400% greater than pre-disturbance AGBI. Growth of suppressed trees, independent of their age, followed similar trajectories after canopy accession. While aboveground biomass generally increased through time, the time since disturbance and disturbance severity are important co-predictors for stand-level AGBI and AGB. These forests regained most of the above ground living biomass over short interval (∼50 years) after low intensity disturbances. The highest stand-level living AGB was observed on plots that experienced >40% canopy removal 160–190 years ago, whereas the highest AGBI occurred in plots disturbed recently within the past 40–50 years. Our results emphasize the importance of including both individual tree age and disturbance legacies to accurately characterize biomass dynamics and trajectories in forest ecosystems. Importantly, the period of time that a tree is in the canopy, and not tree age, modulates the trajectory of tree level AGBI. Growth rates begin to decline after ∼30 years (tree-rings width) and ∼100 years (AGBI) in the canopy. We demonstrate that even late-seral forests can rapidly regain biomass lost to low intensity disturbance.

The Mediterranean region has been identified as a global warming hotspot, where future climate impacts are expected to have significant consequences on societal and ecosystem well-being. To put ongoing trends of summer climate into the context of past natural variability, we reconstructed climate from maximum latewood density (MXD) measurements of Pinus heldreichii (1521–2010) and latewood width (LWW) of Pinus nigra (1617–2010) on Mt. Olympus, Greece. Previous research in the northeastern Mediterranean has primarily focused on inter-annual variability, omitting any low-frequency trends. The present study utilizes methods capable of retaining climatically driven long-term behavior of tree growth. The LWW chronology corresponds closely to early summer moisture variability (May–July, r = 0.65, p < 0.001, 1950–2010), whereas the MXD-chronology relates mainly to late summer warmth (July–September, r = 0.64, p < 0.001; 1899–2010). The chronologies show opposing patterns of decadal variability over the twentieth century (r = −0.68, p < 0.001) and confirm the importance of the summer North Atlantic Oscillation (sNAO) for summer climate in the northeastern Mediterranean, with positive sNAO phases inducing cold anomalies and enhanced cloudiness and precipitation. The combined reconstructions document the late twentieth—early twenty-first century warming and drying trend, but indicate generally drier early summer and cooler late summer conditions in the period ~1700–1900 CE. Our findings suggest a potential decoupling between twentieth century atmospheric circulation patterns and pre-industrial climate variability. Furthermore, the range of natural climate variability stretches beyond summer moisture availabilityobserved in recent decades and thus lends credibility to the significant drying trends projected for this region in current Earth System Model simulations.

The temporal variability of the forest sink is associated with high uncertainties in both its magnitude and the driving ecological and climatic processes. In this study, we assess the inter-annual variability (IAV) of carbon uptake using annually resolved aboveground biomass increment (ABI) estimates from 272 pseudorandomly sampled trees at a long-term monitoring plot in the dry valley of the Valais in Switzerland. Over the 1950–2011 period, the mean ABI is 86.8 g C m−2 year−1 with an IAV of ±31 %. The IAV is largely driven by hydrological conditions throughout the water year from previous August to current August (r SPEI = 0.56; 1st differenced r = 0.75, p

Tree-rings offer one of the few possibilities to empirically quantify and reconstruct forest growth dynamics over years to millennia. Contemporaneously with the growing scientific community employing tree-ring parameters, recent research has suggested that commonly applied sampling designs (i.e. how and which trees are selected for dendrochronological sampling) may introduce considerable biases in quantifications of forest responses to environmental change. To date, a systematic assessment of the consequences of sampling design on dendroecological and-climatological conclusions has not yet been performed. Here, we investigate potential biases by sampling a large population of trees and replicating diverse sampling designs. This is achieved by retroactively subsetting the population and specifically testing for biases emerging for climate reconstruction, growth response to climate variability, long-term growth trends, and quantification of forest productivity. We find that commonly applied sampling designs can impart systematic biases of varying magnitude to any type of tree-ring-based investigations, independent of the total number of samples considered. Quantifications of forest growth and productivity are particularly susceptible to biases, whereas growth responses to short-term climate variability are less affected by the choice of sampling design. The world's most frequently applied sampling design, focusing on dominant trees only, can bias absolute growth rates by up to 459% and trends in excess of 200%. Our findings challenge paradigms, where a subset of samples is typically considered to be representative for the entire population. The only two sampling strategies meeting the requirements for all types of investigations are the (i) sampling of all individuals within a fixed area; and (ii) fully randomized selection of trees. This result advertises the consistent implementation of a widely applicable sampling design to simultaneously reduce uncertainties in tree-ring-based quantifications of forest growth and increase the comparability of datasets beyond individual studies, investigators, laboratories, and geographical boundaries.