Tree rings reveal the transient risk of extinction hidden inside climate envelope forecasts.

Given the importance of climate in shaping species' geographic distributions, climate change poses an existential threat to biodiversity. Climate envelope modeling, the predominant approach used to quantify this threat, presumes that individuals in populations respond to climate variability and change according to species-level responses inferred from spatial occurrence data-such that individuals at the cool edge of a species' distribution should benefit from warming (the "leading edge"), whereas individuals at the warm edge should suffer (the "trailing edge"). Using 1,558 tree-ring time series of an aridland pine (Pinus edulis) collected at 977 locations across the species' distribution, we found that trees everywhere grow less in warmer-than-average and drier-than-average years. Ubiquitous negative temperature sensitivity indicates that individuals across the entire distribution should suffer with warming-the entire distribution is a trailing edge. Species-level responses to spatial climate variation are opposite in sign to individual-scale responses to time-varying climate for approximately half the species' distribution with respect to temperature and the majority of the species' distribution with respect to precipitation. These findings, added to evidence from the literature for scale-dependent climate responses in hundreds of species, suggest that correlative, equilibrium-based range forecasts may fail to accurately represent how individuals in populations will be impacted by changing climate. A scale-dependent view of the impact of climate change on biodiversity highlights the transient risk of extinction hidden inside climate envelope forecasts and the importance of evolution in rescuing species from extinction whenever local climate variability and change exceeds individual-scale climate tolerances.

Finding balance: Tree-ring isotopes differentiate between acclimation and stress-induced imbalance in a long-term irrigation experiment.

Scots pine (Pinus sylvestris L.) is a common European tree species, and understanding its acclimation to the rapidly changing climate through physiological, biochemical or structural adjustments is vital for predicting future growth. We investigated a long-term irrigation experiment at a naturally dry forest in Switzerland, comparing Scots pine trees that have been continuously irrigated for 17 years (irrigated) with those for which irrigation was interrupted after 10 years (stop) and non-irrigated trees (control), using tree growth, xylogenesis, wood anatomy, and carbon, oxygen and hydrogen stable isotope measurements in the water, sugars and cellulose of plant tissues. The dendrochronological analyses highlighted three distinct acclimation phases to the treatments: irrigated trees experienced (i) a significant growth increase in the first 4 years of treatment, (ii) high growth rates but with a declining trend in the following 8 years and finally (iii) a regression to pre-irrigation growth rates, suggesting the development of a new growth limitation (i.e. acclimation). The introduction of the stop treatment resulted in further growth reductions to below-control levels during the third phase. Irrigated trees showed longer growth periods and lower tree-ring δ13 C values, reflecting lower stomatal restrictions than control trees. Their strong tree-ring δ18 O and δ2 H (O-H) relationship reflected the hydrological signature similarly to the control. On the contrary, the stop trees had lower growth rates, conservative wood anatomical traits, and a weak O-H relationship, indicating a physiological imbalance. Tree vitality (identified by crown transparency) significantly modulated growth, wood anatomical traits and tree-ring δ13 C, with low-vitality trees of all treatments performing similarly regardless of water availability. We thus provide quantitative indicators for assessing physiological imbalance and tree acclimation after environmental stresses. We also show that tree vitality is crucial in shaping such responses. These findings are fundamental for the early assessment of ecosystem imbalances and decline under climate change.

No Future Growth Enhancement Expected at the Northern Edge for European Beech due to Continued Water Limitation.

With ongoing global warming, increasing water deficits promote physiological stress on forest ecosystems with negative impacts on tree growth, vitality, and survival. How individual tree species will react to increased drought stress is therefore a key research question to address for carbon accounting and the development of climate change mitigation strategies. Recent tree-ring studies have shown that trees at higher latitudes will benefit from warmer temperatures, yet this is likely highly species-dependent and less well-known for more temperate tree species. Using a unique pan-European tree-ring network of 26,430 European beech (Fagus sylvatica L.) trees from 2118 sites, we applied a linear mixed-effects modeling framework to (i) explain variation in climate-dependent growth and (ii) project growth for the near future (2021-2050) across the entire distribution of beech. We modeled the spatial pattern of radial growth responses to annually varying climate as a function of mean climate conditions (mean annual temperature, mean annual climatic water balance, and continentality). Over the calibration period (1952-2011), the model yielded high regional explanatory power (R2 = 0.38-0.72). Considering a moderate climate change scenario (CMIP6 SSP2-4.5), beech growth is projected to decrease in the future across most of its distribution range. In particular, projected growth decreases by 12%-18% (interquartile range) in northwestern Central Europe and by 11%-21% in the Mediterranean region. In contrast, climate-driven growth increases are limited to around 13% of the current occurrence, where the historical mean annual temperature was below ~6°C. More specifically, the model predicts a 3%-24% growth increase in the high-elevation clusters of the Alps and Carpathian Arc. Notably, we find little potential for future growth increases (-10 to +2%) at the poleward leading edge in southern Scandinavia. Because in this region beech growth is found to be primarily water-limited, a northward shift in its distributional range will be constrained by water availability.

Ecological forecasting of tree growth: Regional fusion of tree-ring and forest inventory data to quantify drivers and characterize uncertainty.

Robust ecological forecasting of tree growth under future climate conditions is critical to anticipate future forest carbon storage and flux. Here, we apply three ingredients of ecological forecasting that are key to improving forecast skill: data fusion, confronting model predictions with new data, and partitioning forecast uncertainty. Specifically, we present the first fusion of tree-ring and forest inventory data within a Bayesian state-space model at a multi-site, regional scale, focusing on Pinus ponderosa var. brachyptera in the southwestern US. Leveraging the complementarity of these two data sources, we parsed the ecological complexity of tree growth into the effects of climate, tree size, stand density, site quality, and their interactions, and quantified uncertainties associated with these effects. New measurements of trees, an ongoing process in forest inventories, were used to confront forecasts of tree diameter with observations, and evaluate alternative tree growth models. We forecasted tree diameter and increment in response to an ensemble of climate change projections, and separated forecast uncertainty into four different causes: initial conditions, parameters, climate drivers, and process error. We found a strong negative effect of fall-spring maximum temperature, and a positive effect of water-year precipitation on tree growth. Furthermore, tree vulnerability to climate stress increases with greater competition, with tree size, and at poor sites. Under future climate scenarios, we forecast increment declines of 22%-117%, while the combined effect of climate and size-related trends results in a 56%-91% decline. Partitioning of forecast uncertainty showed that diameter forecast uncertainty is primarily caused by parameter and initial conditions uncertainty, but increment forecast uncertainty is mostly caused by process error and climate driver uncertainty. This fusion of tree-ring and forest inventory data lays the foundation for robust ecological forecasting of aboveground biomass and carbon accounting at tree, plot, and regional scales, including iterative improvement of model skill.

Adding Tree Rings to North America's National Forest Inventories: An Essential Tool to Guide Drawdown of Atmospheric CO2.

Tree-ring time series provide long-term, annually resolved information on the growth of trees. When sampled in a systematic context, tree-ring data can be scaled to estimate the forest carbon capture and storage of landscapes, biomes, and-ultimately-the globe. A systematic effort to sample tree rings in national forest inventories would yield unprecedented temporal and spatial resolution of forest carbon dynamics and help resolve key scientific uncertainties, which we highlight in terms of evidence for forest greening (enhanced growth) versus browning (reduced growth, increased mortality). We describe jump-starting a tree-ring collection across the continent of North America, given the commitments of Canada, the United States, and Mexico to visit forest inventory plots, along with existing legacy collections. Failing to do so would be a missed opportunity to help chart an evidence-based path toward meeting national commitments to reduce net greenhouse gas emissions, urgently needed for climate stabilization and repair.

Continental-scale tree-ring-based projection of Douglas-fir growth: Testing the limits of space-for-time substitution.

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.

Oxygen isotopes in tree rings are less sensitive to changes in tree size and relative canopy position than carbon isotopes.

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.

The influence of sampling design on tree-ring-based quantification of forest growth.

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.

A tree-ring perspective on the terrestrial carbon cycle.

Tree-ring records can provide valuable information to advance our understanding of contemporary terrestrial carbon cycling and to reconstruct key metrics in the decades preceding monitoring data. The growing use of tree rings in carbon-cycle research is being facilitated by increasing recognition of reciprocal benefits among research communities. Yet, basic questions persist regarding what tree rings represent at the ecosystem level, how to optimally integrate them with other data streams, and what related challenges need to be overcome. It is also apparent that considerable unexplored potential exists for tree rings to refine assessments of terrestrial carbon cycling across a range of temporal and spatial domains. Here, we summarize recent advances and highlight promising paths of investigation with respect to (1) growth phenology, (2) forest productivity trends and variability, (3) CO2 fertilization and water-use efficiency, (4) forest disturbances, and (5) comparisons between observational and computational forest productivity estimates. We encourage the integration of tree-ring data: with eddy-covariance measurements to investigate carbon allocation patterns and water-use efficiency; with remotely sensed observations to distinguish the timing of cambial growth and leaf phenology; and with forest inventories to develop continuous, annually-resolved and long-term carbon budgets. In addition, we note the potential of tree-ring records and derivatives thereof to help evaluate the performance of earth system models regarding the simulated magnitude and dynamics of forest carbon uptake, and inform these models about growth responses to (non-)climatic drivers. Such efforts are expected to improve our understanding of forest carbon cycling and place current developments into a long-term perspective.

Slower growth prior to the 2018 drought and a high growth sensitivity to previous year summer conditions predisposed European beech to crown dieback.

The record-breaking drought in 2018 caused premature leaf discoloration and shedding (early browning) in many beech (Fagus sylvatica L.) dominated forests in Central Europe. However, a high degree of variability in drought response among individual beech trees was observed. While some trees were severely impacted by the prolonged water deficits and high temperatures, others remained vital with no or only minor signs of crown vitality loss. Why some beech trees were more susceptible to drought-induced crown damage than others and whether growth recovery is possible are poorly understood. Here, we aimed to identify growth characteristics associated with the variability in drought response between individual beech trees based on a sample of 470 trees in northern Switzerland. By combining tree growth measurements and crown condition assessments, we also investigated the possible link between crown dieback and growth recovery after drought. Beech trees with early browning exhibited an overall lower growth vigor before the 2018 drought than co-occurring vital beech trees. This lower vigor is mainly indicated by lower overall growth rates, stronger growth declines in the past decades, and higher growth-climate sensitivity. Particularly, warm previous year summer conditions negatively affected current growth of the early-browning trees. These findings suggest that the affected trees had less access to critical resources and were physiologically limited in their growth predisposing them to early browning. Following the 2018 drought, observed growth recovery potential corresponded to the amount of crown dieback and the local climatic water balance. Overall, our findings emphasize that beech-dominated forests in Central Europe are under increasing pressure from severe droughts, ultimately reducing the competitive ability of this species, especially on lowland sites with shallow soils and low water holding capacity.

Identifying drivers of non-stationary climate-growth relationships of European beech.

The future performance of the widely abundant European beech (Fagus sylvatica L.) across its ecological amplitude is uncertain. Although beech is considered drought-sensitive and thus negatively affected by drought events, scientific evidence indicating increasing drought vulnerability under climate change on a cross-regional scale remains elusive. While evaluating changes in climate sensitivity of secondary growth offers a promising avenue, studies from productive, closed-canopy forests suffer from knowledge gaps, especially regarding the natural variability of climate sensitivity and how it relates to radial growth as an indicator of tree vitality. Since beech is sensitive to drought, we in this study use a drought index as a climate variable to account for the combined effects of temperature and water availability and explore how the drought sensitivity of secondary growth varies temporally in dependence on growth variability, growth trends, and climatic water availability across the species' ecological amplitude. Our results show that drought sensitivity is highly variable and non-stationary, though consistently higher at dry sites compared to moist sites. Increasing drought sensitivity can largely be explained by increasing climatic aridity, especially as it is exacerbated by climate change and trees' rank progression within forest communities, as (co-)dominant trees are more sensitive to extra-canopy climatic conditions than trees embedded in understories. However, during the driest periods of the 20th century, growth showed clear signs of being decoupled from climate. This may indicate fundamental changes in system behavior and be early-warning signals of decreasing drought tolerance. The multiple significant interaction terms in our model elucidate the complexity of European beech's drought sensitivity, which needs to be taken into consideration when assessing this species' response to climate change.

Climate-change-driven growth decline of European beech forests.

The growth of past, present, and future forests was, is and will be affected by climate variability. This multifaceted relationship has been assessed in several regional studies, but spatially resolved, large-scale analyses are largely missing so far. Here we estimate recent changes in growth of 5800 beech trees (Fagus sylvatica L.) from 324 sites, representing the full geographic and climatic range of species. Future growth trends were predicted considering state-of-the-art climate scenarios. The validated models indicate growth declines across large region of the distribution in recent decades, and project severe future growth declines ranging from -20% to more than -50% by 2090, depending on the region and climate change scenario (i.e. CMIP6 SSP1-2.6 and SSP5-8.5). Forecasted forest productivity losses are most striking towards the southern distribution limit of Fagus sylvatica, in regions where persisting atmospheric high-pressure systems are expected to increase drought severity. The projected 21st century growth changes across Europe indicate serious ecological and economic consequences that require immediate forest adaptation.

Amplifying feedback loop between growth and wood anatomical characteristics of Fraxinus excelsior explains size-related susceptibility to ash dieback.

Since the 1990s the invasive fungus Hymenoscyphus fraxineus has caused severe crown dieback and high mortality rates in Fraxinus excelsior in Europe. In addition to a strong genetic control of tolerance to the fungus, previous studies have found landscape heterogeneity to be an additional driver of variability in the severity of dieback symptoms. However, apart from climatic conditions related to heat and humidity influencing fungal infection success, the mechanistic understanding of why smaller or slower-growing trees are more susceptible to dieback remains less well understood. Here, we analyzed three stands in Switzerland with a unique setting of 8 years of data availability of intra-annual diameter growth and annual crown health assessments. We complemented this by ring width and quantitative wood anatomical measurements extending back before the monitoring started to investigate if wood anatomical adjustments can help better explain the size-related dieback phenomenon. We found that slower-growing trees or trees with smaller crowns already before the arrival of the fungus were more susceptible to dieback and mortality. Defoliation directly reduced growth as well as maximum earlywood vessel size, and the positive relationship between vessel size and growth rate caused a positive feedback amplifying and accelerating crown dieback. Measured non-structural carbohydrate (NSC) concentrations in the outermost five rings did not significantly vary between healthy and weakened trees, which translate into large differences in absolute available amount of NSCs. Thus, we hypothesize that a lack of NSCs (mainly sugars) leads to lower turgor pressure and smaller earlywood vessels in the following year. This might impede efficient water transport and photosynthesis, and be responsible for stronger symptoms of dieback and higher mortality rates in smaller and slower-growing trees.

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.

Sampling bias overestimates climate change impacts on forest growth in the southwestern United States.

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.