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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.

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.

Partitioned estimates of the boreal forest carbon (C) sink components are crucial for understanding processes and developing science-driven adaptation and mitigation strategies under climate change. Here, we provide a concise tree-centered overview of the boreal forest C balance and offer a circumpolar perspective on the contribution of trees to boreal forest C dynamics. We combine an ant’s-eye view, based on quantitative in situ observations of C balance, with a bird’s-eye perspective on C dynamics across the circumboreal region using large-scale data sets. We conclude with an outlook addressing the trajectories of the circumboreal C dynamics in response to projected environmental changes.

Drought legacies in radial tree growth are an important feature of variability in biomass accumulation and are widely used to characterize forest resilience to climate change. Defined as a deviation from normal growth, the statistical significance of legacy effects depends on the definition of “normal”—expected growth under average conditions—which has not received sufficient scrutiny. We re-examined legacy effect analyses using the International Tree-Ring Data Bank (ITRDB) and then produced synthetic tree-ring data to disentangle four key variables influencing the magnitude of legacy effects. We hypothesized that legacy effects (i) are mainly influenced by the auto-correlation of the radial growth time series (phi), (ii) depend on climate-growth cross-correlation (rho), (iii) are directly proportional to the inherent variability of the growth time series (standard deviation, SD), and (iv) scale with the chosen extreme event threshold. Using a data simulation approach, we were able to reproduce observed lag patterns, demonstrating that legacy effects are a direct outcome of ubiquitous biological memory. We found that stronger legacy effects for conifers compared to angiosperms is a consequence of their higher auto-correlation, and that the detectability of legacy effects following rare drought events at individual sites is compromised by strong background stochasticity. Synthesis. We propose two pathways forward to improve the assessment and interpretation of legacy effects: First, we highlight the need to account for auto-correlated residuals of climate-growth regression models a posteriori, thereby retrospectively adjusting expectations for “normal” growth variability. Alternatively, we recommend including lagged climate variables in regression models a priori. By doing so, the magnitude of detected legacy effects is greatly reduced and biological memory is directly attributed to antecedent climatic drivers. We argue that future analyses should focus on understanding the functional reasons for how and why key statistical parameters describing this biological memory differ across species and sites. These two pathways should also stimulate improved process-based representation of vegetation carbon dynamics in mechanistic models.

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.

Sapwood characteristics, such as sapwood area as well as thermal and hydraulic conductivity, are linked to species-specific hydraulic function and resource allocation to water transport tissues (xylem). These characteristics are often unknown and thus a major source of uncertainty in sap flow data processing and transpiration estimates because bulk rather than species-specific values are usually applied. Here, we analyzed the sapwood characteristics of fifteen common tree species in eastern North America from different taxa (i.e., angiosperms and gymnosperms) and xylem porosity groups (i.e., tracheid-bearing, diffuse- or ring-porous species). We quantified their sapwood area changes with stem diameter (allometric scaling) and thermal conductivity. We combined these measurements with species-specific values of wood density and hydraulic conductivity found in literature and assessed the role of wood anatomy in orchestrating their covariation. Angiosperms (ring- and diffuse-porous species), with specialized vessels for water transport, showed steeper relation (scaling) between tree size and sapwood area in comparison to gymnosperms (tracheid-bearing species). Despite the variability in thermal conductivity between species, gymnosperms (angiosperms) were characterized by lower (higher) wood density and higher (lower) sapwood moisture content, resulting in non-significant differences in sapwood thermal conductivity between taxa and xylem porosity groups. Clustering of species sapwood characteristics based on taxa or xylem porosity could facilitate more accurate parameterizations of these attributes. When combined with an increasing number of sap flow observations, these findings can improve tree- and landscape-level transpiration estimates, leading to more robust partitioning of terrestrial water fluxes.

Drought legacy effects in radial tree growth have been extensively studied over the last decade and are found to critically influence carbon sequestration in woody biomass. Typically quantified as a deviation from “normal” growth, drought legacy magnitude and statistical significance depend on the definition of expected vs. unexpected growth variability under average conditions – a definition that has received insufficient theoretical validation.Here, we revisit popular legacy effect analyses using the International Tree-Ring Data Bank (ITRDB) and employ a synthetic data simulation to disentangle four key variables influencing the magnitude of legacy effects. We show that legacy effects i) are mainly influenced by the overall auto-correlation of radial growth time series, ii) depend on climate-growth cross-correlation, iii) are directly proportional to the year-to-year variability of the growth time series, and iv) scale with the chosen extreme event threshold. Our analysis revealed that legacy effects are a direct outcome of the omnipresent biological memory.We further found that the interpretation of legacy effects following individual drought events at specific sites is challenged by high stochasticity, and show that the commonly perceived stronger legacy effects for conifers are the result of higher auto-correlation compared to deciduous broadleaves. Given that the existing literature has not sufficiently addressed biological memory, we present two pathways to improve future assessment and interpretation of legacy effects. First, we provide a simulation algorithm to a posteriori account for auto-correlated residuals of the initial regression model between growth and climate, i.e. a corrected Null model to determine statistical significance, thereby retrospectively adjusting expectations for “normal” growth variability. The second pathway is to a priori include lagged climate parameters in the regression model. This substantially reduces the magnitude of observed legacy effects and thus challenges us to revisit estimates of drought-induced growth deviations by considering the full spectrum of expected growth behavior.

Sapwood characteristics, such as sapwood area as well as thermal and hydraulic conductivity, are linked to species-specific hydraulic function and resource allocation to water transport tissues (xylem). These characteristics are often unknown and thus a major source of uncertainty in sap flow data processing and transpiration estimates because bulk rather than species-specific values are usually applied. Here, we analyzed the sapwood characteristics of fifteen common tree species in eastern North America from different taxonomic (i.e., angiosperms and gymnosperms) and xylem porosity groups (i.e., tracheid-bearing, diffuse- or ring-porous species) and we assessed how uncertainties in sapwood characteristics involved in sap flow calculations are propagated in tree water use estimates. We quantified their sapwood area changes with stem diameter (allometric scaling) and thermal conductivity. We combined these measurements with species-specific values of wood density and hydraulic conductivity found in the literature and assessed the role of wood anatomy in orchestrating their covariation. Using an example sap flow dataset from tree species with different xylem porosity, we assessed the sensitivity of tree water use estimates to sapwood characteristics and their interactions. Angiosperms (ring- and diffuse-porous species), with specialized vessels for water transport, showed a steeper relationship (scaling) between tree stem diameter and sapwood area in comparison to gymnosperms (tracheid-bearing species). Gymnosperms (angiosperms) were characterized by lower (higher) wood density and higher (lower) sapwood moisture content, resulting in non-significant differences in sapwood thermal conductivity between taxonomic and xylem porosity groups. Clustering of species sapwood characteristics based on taxonomic or xylem porosity groups and constraining these parameters could facilitate more accurate sap flow calculations and tree water use estimates. When combined with an increasing number of sap flow observations, these findings should improve tree- and landscape-level transpiration estimates, leading to more robust partitioning of terrestrial water fluxes.

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.

Since the 1990s the invasive fungus Hymenoscyphus fraxineus has led to 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 high landscape variability in the severity of dieback symptoms. However, apart from heat and humidity-related climate conditions favoring fungal development 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 eight years of intra-annual diameter growth and annual crown health assessments, together with ring-width and quantitative wood anatomical measurements preceding the monitoring, to investigate if wood anatomical adjustments can help better explaining 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. We show that defoliation directly reduces growth as well as maximum earlywood vessel size, and that the positive relationship between vessel size and growth rate causes a positive feedback amplifying crown dieback. Because leaf necrosis happens during late summer when ring formation has already finished, photosynthesis is heavily reduced during a time when non-structural carbohydrates (NSCs, sugars and starch) are stored. Thus, we hypothesize that a lack of NSCs (mainly sugars) leads to lower turgor pressure and smaller earlywood vessels in the next year impeding efficient water transport and photosynthesis, and is responsible why smaller and slower growing trees show stronger symptoms of dieback and higher mortality rates.

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%

The position of the northern boundary of the tropical belt affects the hydroclimate of many arid and semi-arid regions in the Northern Hemisphere. Widening of the tropical belt since the 1970s has largely been attributed to anthropogenic forcing. However, the relative influence of natural drivers of tropical belt expansion and contraction before this time is poorly understood. Here we use data on tree-ring widths from five mid-latitude regions in the Northern Hemisphere to reconstruct the movement of the northern boundary of the early spring tropical belt over the past 800 years (ad 1203–2003). Our reconstruction explains 45% of the interannual variance in the latitudinal extent of the Hadley circulation, a metric of the position of the tropical belt boundary. We find that the tropical belt contracted (expanded) during positive (negative) phases of the El Niño Southern Oscillation and Pacific North American teleconnection patterns. The tropical belt also contracted significantly following major volcanic events that injected sulfur into the stratosphere. The longest period of persistent tropical belt expansion occurred in the late sixteenth century, during one of the coldest periods of the Little Ice Age. Our results warn of potential socio-economic consequences of future variations in tropical belt width driven by natural climate variability or stratospheric aerosol injections, whether volcanic or artificial. Fil: Alfaro Sánchez, R.. Consejo Superior de Investigaciones Científicas. Centre de Recerca Ecológica I Aplicacions Forestals; España. University of Arizona; Estados Unidos Fil: Nguyen, H.. Bureau of Meteorology; Australia Fil: Klesse, S.. University of Arizona; Estados Unidos Fil: Hudson, A.. University of Arizona; Estados Unidos Fil: Belmecheri, S.. University of Arizona; Estados Unidos Fil: Köse, N.. Istanbul University; Turquía Fil: Diaz, H. F.. University of Hawaii at Manoa; Estados Unidos Fil: Monson, R. K.. University of Arizona; Estados Unidos Fil: Villalba, Ricardo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mendoza. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales. Provincia de Mendoza. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales. Universidad Nacional de Cuyo. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales; Argentina Fil: Trouet, V.. University of Arizona; Estados Unidos

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.

Forest growth and biomass response to environmental change depends upon climatic, but also upon interactions with biotic drivers, such as insect outbreak activity. In this study we use tree-rings along a temperature gradient to assess the relative importance of climate versus altered larch budmoth (Zeiraphera diniana) outbreak cycles for forest biomass accumulation at high elevations. We established climate-growth relationships and performed outbreak-growth response analysis for >500 individuals from host (Larix decidua) and non-host trees (Picea abies) at different elevations (from 1300 to 2200 m a.s.l.) in the Swiss Alps. We quantified outbreak-induced reductions of absolute biomass increment and modelled effects of the recent absence of outbreaks. Our results reveal that average outbreaks reduced biomass accumulation by 1130 kg ha−1 y−1 during the four years after the event, having an equal or even greater impact on carbon sequestration than climate. Recent growth increases previously observed at the study sites are largely attributable to the outbreaks absence since 1981, suggesting that regular outbreaks have hampered host-trees from realizing their growth potential for centuries. The presented impact analysis quantifies the importance of non-lethal insect activity on forest biomass dynamics, revealing the relevance of including such biotic drivers and their interactions with climate in models assessing the future productivity and carbon sink capacity of forests.

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.

The “two-third spline” (2/3S) is a frequently applied method to detrend tree-ring series. It fits a spline with a 50% frequency cutoff at a frequency equal to two-thirds of each sample length in a dataset. It was introduced to ensure a minimum loss of low-frequency variance, which is resolvable during the detrending of ring-width series. In this paper I show potential problems that arise when rusing this method. The 2/3S runs counter the strengths of using a digital filter to detrend – i.e. one is giving up full control over the frequency-removing characteristics of the growth curve and each individual time series retains a different amount of low frequency. Thus, the 2/3S is less suitable for reconstructing climate or to compare environmental impacts on tree growth between groups – both of which comprise the majority of dendrochronological analyses – as it will likely introduce a temporal frequency bias. Within a long chronology it will result in decreasing power to resolve low frequencies towards present in a living-only trees setting, especially when the youngest segment lengths are 100 years and shorter, and more generally during the period where the chronology is constructed from samples with shorter segment lengths compared to the period with longer segment lengths. The frequency bias will also significantly impact regression slopes and correlation coefficients, possibly distorting analyses investigating multiple groups with different mean segment lengths. Highlighting these potential biases, I recommend the community to not use this method on an individual basis but rather to use a fixed spline stiffness for all samples based on the n% criterion (n = 67) of e.g. the mean segment length of the entire dataset.

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.

FB acknowledges funding from the Swiss National Science Foundation (P300P2_154543) and the EU Horizon-2020 project “BACI” (grant 640176). SK, FJ, RS and DCF are supported by the SNF iTREE sinergia project 136295. FJ, SL, and RS acknowledge support by the Swiss National Science Foundation (#200020_172476), OB acknowledges funding from UEFISCDI project PN-II-ID-PCE-2011-3-07 and VT is supported by the GACR 15-14840S and CIGA 20154316. The response of forest growth to climate variability varies along environmental gradients. A growth increase and decrease with warming is usually observed in cold‐humid and warm‐dry regions, respectively. However, it remains poorly known where the sign of these temperature effects switches. Here we introduce a newly developed European tree‐ring network that has been specifically collected to reconstruct forest aboveground biomass increment (ABI). We quantify, how the long‐term (1910‐2009) inter‐annual variability of ABI depends on local mean May‐August temperature and test, if a dynamic global vegetation model (DGVM) ensemble reflects the resulting patterns. We find that sites at 8°C mean May‐August temperature increase ABI on average by 5.7 ± 1.3 %, whereas sites at 20°C decrease ABI by 3.0±1.8 % m‐2 y‐1 ∆°C‐1. A threshold temperature between beneficial and detrimental effects of warming and the associated increase in water demand on tree growth emerged at 15.9 ± 1.4°C mean May‐August temperature. Because inter‐annual variability increases proportionally with mean growth rate – i.e. the coefficient of variation stays constant – we were able to validate these findings with a much larger tree‐ring dataset that had been established following classic dendrochronological sampling schemes. While the observed climate sensitivity pattern is well reflected in the DGVM ensemble, there is a large spread of threshold temperatures between the individual models. Also, individual models disagree strongly on the magnitude of climate impact at the coldest and warmest locations, suggesting where model improvement is most needed to more accurately predict forest growth and effectively guide silvicultural practices. Publisher PDF

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.

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

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.

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.

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

Текст статьи не публикуется в открытом доступе в соответствии с политикой журнала. Large-scale millennial length Northern Hemisphere (NH) temperature reconstructions have been progressively improved over the last 20 years as new datasets have been developed. This paper, and its companion (Part II, Anchukaitis et al. in prep), details the latest tree-ring (TR) based NH land air temperature reconstruction from a temporal and spatial perspective. This work is the first product of a consortium called N-TREND (Northern Hemisphere Tree-Ring Network Development) which brings together dendroclimatologists to identify a collective strategy for improving large-scale summer temperature reconstructions. The new reconstruction, N-TREND2015, utilises 54 records, a significant expansion compared with previous TR studies, and yields an improved reconstruction with stronger statistical calibration metrics. N-TREND2015 is relatively insensitive to the compositing method and spatial weighting used and validation metrics indicate that the new record portrays reasonable coherence with large scale summer temperatures and is robust at all time-scales from 918 to 2004 where at least 3 TR records exist from each major continental mass. N-TREND2015 indicates a longer and warmer medieval period (~900 e1170) than portrayed by previous TR NH reconstructions and by the CMIP5 model ensemble, but with better overall agreement between records for the last 600 years. Future dendroclimatic projects should focus on developing new long records from data-sparse regions such as North America and eastern Eurasia as well as ensuring the measurement of parameters related to latewood density to complement ring-width records which can improve local based calibration substantially.

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.

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.

The demand for large-scale and long-term information on tree growth is increasing rapidly as environmental change research strives to quantify and forecast the impacts of continued warming on forest ecosystems. This demand, combined with the now quasi-global availability of tree-ring observations, has inspired researchers to compile large tree-ring networks to address continental or even global-scale research questions. However, these emergent spatial objectives contrast with paleo-oriented research ideas that have guided the development of many existing records. A series of challenges related to how, where, and when samples have been collected is complicating the transition of tree rings from a local to a global resource on the question of tree growth. Herein, we review possibilities to scale tree-ring data (A) from the sample to the whole tree, (B) from the tree to the site, and (C) from the site to larger spatial domains. Representative tree-ring sampling supported by creative statistical approaches is thereby key to robustly capture the heterogeneity of climate-growth responses across forested landscapes. We highlight the benefits of combining the temporal information embedded in tree rings with the spatial information offered by forest inventories and earth observations to quantify tree growth and its drivers. In addition, we show how the continued development of mechanistic tree-ring models can help address some of the non-linearities and feedbacks that complicate making inference from tree-ring data. By embracing scaling issues, the discipline of dendrochronology will greatly increase its contributions to assessing climate impacts on forests and support the development of adaptation strategies.