Density-dependent species interactions modulate alpine treeline shifts.

Species interactions such as facilitation and competition play a crucial role in driving species range shifts. However, density dependence as a key feature of these processes has received little attention in both empirical and modelling studies. Herein, we used a novel, individual-based treeline model informed by rich in situ observations to quantify the contribution of density-dependent species interactions to alpine treeline dynamics, an iconic biome boundary recognized as an indicator of global warming. We found that competition and facilitation dominate in dense versus sparse vegetation scenarios respectively. The optimal balance between these two effects was identified at an intermediate vegetation thickness where the treeline elevation was the highest. Furthermore, treeline shift rates decreased sharply with vegetation thickness and the associated transition from positive to negative species interactions. We thus postulate that vegetation density must be considered when modelling species range dynamics to avoid inadequate predictions of its responses to climate warming.

Drought alters aboveground biomass production efficiency: Insights from two European beech forests.

The fraction of photosynthetically assimilated carbon that trees allocate to long-lasting woody biomass pools (biomass production efficiency - BPE), is a key metric of the forest carbon balance. Its apparent simplicity belies the complex interplay between underlying processes of photosynthesis, respiration, litter and fruit production, and tree growth that respond differently to climate variability. Whereas the magnitude of BPE has been routinely quantified in ecological studies, its temporal dynamics and responses to extreme events such as drought remain less well understood. Here, we combine long-term records of aboveground carbon increment (ACI) obtained from tree rings with stand-level gross primary productivity (GPP) from eddy covariance (EC) records to empirically quantify aboveground BPE (= ACI/GPP) and its interannual variability in two European beech forests (Hainich, DE-Hai, Germany; Sorø, DK-Sor, Denmark). We found significant negative correlations between BPE and a daily-resolved drought index at both sites, indicating that woody growth is de-prioritized under water limitation. During identified extreme years, early-season drought reduced same-year BPE by 29 % (Hainich, 2011), 31 % (Sorø, 2006), and 14 % (Sorø, 2013). By contrast, the 2003 late-summer drought resulted in a 17 % reduction of post-drought year BPE at Hainich. Across the entire EC period, the daily-to-seasonal drought response of BPE resembled that of ACI, rather than that of GPP. This indicates that BPE follows sink dynamics more closely than source dynamics, which appear to be decoupled given the distinctive climate response patterns of GPP and ACI. Based on our observations, we caution against estimating the magnitude and variability of the carbon sink in European beech (and likely other temperate forests) based on carbon fluxes alone. We also encourage comparable studies at other long-term EC measurement sites from different ecosystems to further constrain the BPE response to rare climatic events.

Global tree growth resilience to cold extremes following the Tambora volcanic eruption.

Although the global climate is warming, external forcing driven by explosive volcanic eruptions may still cause abrupt cooling. The 1809 and 1815 Tambora eruptions caused lasting cold extremes worldwide, providing a unique lens that allows us to investigate the magnitude of global forest resilience to and recovery from volcanic cooling. Here, we show that growth resilience inferred from tree-ring data was severely impacted by cooling in high latitudes and elevations: the average tree growth decreased substantially (up to 31.8%), especially in larch forests, and regional-scale probabilities of severe growth reduction (below -2σ) increased up to 1390%. The influence of the eruptions extended longer (beyond the year 1824) in mid- than in high-latitudes, presumably due to the combined impacts of cold and drought stress. As Tambora-size eruptions statistically occur every 200-400 years, assessing their influences on ecosystems can help humankind mitigate adverse impacts on natural resources through improved management, especially in high latitude and elevation regions.

Daytime stomatal regulation in mature temperate trees prioritizes stem rehydration at night.

Trees remain sufficiently hydrated during drought by closing stomata and reducing canopy conductance (Gc ) in response to variations in atmospheric water demand and soil water availability. Thresholds that control the reduction of Gc are proposed to optimize hydraulic safety against carbon assimilation efficiency. However, the link between Gc and the ability of stem tissues to rehydrate at night remains unclear. We investigated whether species-specific Gc responses aim to prevent branch embolisms, or enable night-time stem rehydration, which is critical for turgor-dependent growth. For this, we used a unique combination of concurrent dendrometer, sap flow and leaf water potential measurements and collected branch-vulnerability curves of six common European tree species. Species-specific Gc reduction was weakly related to the water potentials at which 50% of branch xylem conductivity is lost (P50 ). Instead, we found a stronger relationship with stem rehydration. Species with a stronger Gc control were less effective at refilling stem-water storage as the soil dries, which appeared related to their xylem architecture. Our findings highlight the importance of stem rehydration for water-use regulation in mature trees, which likely relates to the maintenance of adequate stem turgor. We thus conclude that stem rehydration must complement the widely accepted safety-efficiency stomatal control paradigm.

Networking the forest infrastructure towards near real-time monitoring - A white paper.

Forests account for nearly 90 % of the world's terrestrial biomass in the form of carbon and they support 80 % of the global biodiversity. To understand the underlying forest dynamics, we need a long-term but also relatively high-frequency, networked monitoring system, as traditionally used in meteorology or hydrology. While there are numerous existing forest monitoring sites, particularly in temperate regions, the resulting data streams are rarely connected and do not provide information promptly, which hampers real-time assessments of forest responses to extreme climate events. The technology to build a better global forest monitoring network now exists. This white paper addresses the key structural components needed to achieve a novel meta-network. We propose to complement - rather than replace or unify - the existing heterogeneous infrastructure with standardized, quality-assured linking methods and interacting data processing centers to create an integrated forest monitoring network. These automated (research topic-dependent) linking methods in atmosphere, biosphere, and pedosphere play a key role in scaling site-specific results and processing them in a timely manner. To ensure broad participation from existing monitoring sites and to establish new sites, these linking methods must be as informative, reliable, affordable, and maintainable as possible, and should be supplemented by near real-time remote sensing data. The proposed novel meta-network will enable the detection of emergent patterns that would not be visible from isolated analyses of individual sites. In addition, the near real-time availability of data will facilitate predictions of current forest conditions (nowcasts), which are urgently needed for research and decision making in the face of rapid climate change. We call for international and interdisciplinary efforts in this direction.

Cross-biome synthesis of source versus sink limits to tree growth.

Uncertainties surrounding tree carbon allocation to growth are a major limitation to projections of forest carbon sequestration and response to climate change. The prevalence and extent to which carbon assimilation (source) or cambial activity (sink) mediate wood production are fundamentally important and remain elusive. We quantified source-sink relations across biomes by combining eddy-covariance gross primary production with extensive on-site and regional tree ring observations. We found widespread temporal decoupling between carbon assimilation and tree growth, underpinned by contrasting climatic sensitivities of these two processes. Substantial differences in assimilation-growth decoupling between angiosperms and gymnosperms were determined, as well as stronger decoupling with canopy closure, aridity, and decreasing temperatures. Our results reveal pervasive sink control over tree growth that is likely to be increasingly prominent under global climate change.

Jet stream position explains regional anomalies in European beech forest productivity and tree growth.

The mechanistic pathways connecting ocean-atmosphere variability and terrestrial productivity are well-established theoretically, but remain challenging to quantify empirically. Such quantification will greatly improve the assessment and prediction of changes in terrestrial carbon sequestration in response to dynamically induced climatic extremes. The jet stream latitude (JSL) over the North Atlantic-European domain provides a synthetic and robust physical framework that integrates climate variability not accounted for by atmospheric circulation patterns alone. Surface climate impacts of north-south summer JSL displacements are not uniform across Europe, but rather create a northwestern-southeastern dipole in forest productivity and radial-growth anomalies. Summer JSL variability over the eastern North Atlantic-European domain (5-40E) exerts the strongest impact on European beech, inducing anomalies of up to 30% in modelled gross primary productivity and 50% in radial tree growth. The net effects of JSL movements on terrestrial carbon fluxes depend on forest density, carbon stocks, and productivity imbalances across biogeographic regions.

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.

An earlier start of the thermal growing season enhances tree growth in cold humid areas but not in dry areas.

Climatic warming alters the onset, duration and cessation of the vegetative season. While previous studies have shown a tight link between thermal conditions and leaf phenology, less is known about the impacts of phenological changes on tree growth. Here, we assessed the relationships between the start of the thermal growing season and tree growth across the extratropical Northern Hemisphere using 3,451 tree-ring chronologies and daily climatic data for 1948-2014. An earlier start of the thermal growing season promoted growth in regions with high ratios of precipitation to temperature but limited growth in cold-dry regions. Path analyses indicated that an earlier start of the thermal growing season enhanced growth primarily by alleviating thermal limitations on wood formation in boreal forests and by lengthening the period of growth in temperate and Mediterranean forests. Semi-arid and dry subalpine forests, however, did not benefit from an earlier onset of growth and a longer growing season, presumably due to associated water loss and/or more frequent early spring frosts. These emergent patterns of how climatic impacts on wood phenology affect tree growth at regional to hemispheric scales hint at how future phenological changes may affect the carbon sequestration capacity of extratropical forest ecosystems.

Warming-induced tipping points of Arctic and alpine shrub recruitment.

Shrub recruitment, a key component of vegetation dynamics beyond forests, is a highly sensitive indicator of climate and environmental change. Warming-induced tipping points in Arctic and alpine treeless ecosystems are, however, little understood. Here, we compare two long-term recruitment datasets of 2,770 shrubs from coastal East Greenland and from the Tibetan Plateau against atmospheric circulation patterns between 1871 and 2010 Common Era. Increasing rates of shrub recruitment since 1871 reached critical tipping points in the 1930s and 1960s on the Tibetan Plateau and in East Greenland, respectively. A recent decline in shrub recruitment in both datasets was likely related to warmer and drier climates, with a stronger May to July El Niño Southern Oscillation over the Tibetan Plateau and a stronger June to July Atlantic Multidecadal Oscillation over Greenland. Exceeding the thermal optimum of shrub recruitment, the recent warming trend may cause soil moisture deficit. Our findings suggest that changes in atmospheric circulation explain regional climate dynamics and associated response patterns in Arctic and alpine shrub communities, knowledge that should be considered to protect vulnerable high-elevation and high-latitude ecosystems from the cascading effects of anthropogenic warming.

The 2018 European heatwave led to stem dehydration but not to consistent growth reductions in forests.

Heatwaves exert disproportionately strong and sometimes irreversible impacts on forest ecosystems. These impacts remain poorly understood at the tree and species level and across large spatial scales. Here, we investigate the effects of the record-breaking 2018 European heatwave on tree growth and tree water status using a collection of high-temporal resolution dendrometer data from 21 species across 53 sites. Relative to the two preceding years, annual stem growth was not consistently reduced by the 2018 heatwave but stems experienced twice the temporary shrinkage due to depletion of water reserves. Conifer species were less capable of rehydrating overnight than broadleaves across gradients of soil and atmospheric drought, suggesting less resilience toward transient stress. In particular, Norway spruce and Scots pine experienced extensive stem dehydration. Our high-resolution dendrometer network was suitable to disentangle the effects of a severe heatwave on tree growth and desiccation at large-spatial scales in situ, and provided insights on which species may be more vulnerable to climate extremes.

Climate sensitivity and drought seasonality determine post-drought growth recovery of Quercus petraea and Quercus robur in Europe.

Recent studies have identified strong relationships between delayed recovery of tree growth after drought and tree mortality caused by subsequent droughts. These observations raise concerns about forest ecosystem services and post-drought growth recovery given the projected increase in drought frequency and extremes. For quantifying the impact of extreme droughts on tree radial growth, we used a network of tree-ring width data of 1689 trees from 100 sites representing most of the distribution of two drought tolerant, deciduous oak species (Quercus petraea and Quercus robur). We first examined which climatic factors and seasons control growth of the two species and if there is any latitudinal, longitudinal or elevational trend. We then quantified the relative departure from pre-drought growth during droughts, and how fast trees were able to recover the pre-drought growth level. Our results showed that growth was more related to precipitation and climatic water balance (precipitation minus potential evapotranspiration) than to temperature. However, we did not detect any clear latitudinal, longitudinal or elevational trends except a decreasing influence of summer water balance on growth of Q. petraea with latitude. Neither species was able to maintain the pre-drought growth level during droughts. However, both species showed rapid recovery or even growth compensation after summer droughts but displayed slow recovery in response to spring droughts where none of the two species was able to fully recover the pre-drought growth-level over the three post-drought years. Collectively, our results indicate that oaks which are considered resilient to extreme droughts have also shown vulnerability when droughts occurred in spring especially at sites where long-term growth is not significantly correlated with climatic factors. This improved understanding of the role of drought seasonality and climate sensitivity of sites is key to better predict trajectories of post-drought growth recovery in response to the drier climate projected for Europe.

Stable isotopes of tree rings reveal seasonal-to-decadal patterns during the emergence of a megadrought in the Southwestern US.

Recent evidence has revealed the emergence of a megadrought in southwestern North America since 2000. Megadroughts extend for at least 2 decades, making it challenging to identify such events until they are well established. Here, we examined tree-ring growth and stable isotope ratios in Pinus ponderosa at its driest niche edge to investigate whether trees growing near their aridity limit were sensitive to the megadrought climatic pre-conditions, and were capable of informing predictive efforts. During the decade before the megadrought, trees in four populations revealed increases in the cellulose δ13C content of earlywood, latewood, and false latewood, which, based on past studies are correlated with increased intrinsic water-use efficiency. However, radial growth and cellulose δ18O were not sensitive to pre-megadrought conditions. During the 2 decades preceding the megadrought, at all four sites, the changes in δ13C were caused by the high sensitivity of needle carbon and water exchange to drought trends in key winter months, and for three of the four sites during crucial summer months. Such pre-megadrought physiological sensitivity appears to be unique for trees near their arid range limit, as similar patterns were not observed in trees in ten reference sites located along a latitudinal gradient in the same megadrought domain, despite similar drying trends. Our results reveal the utility of tree-ring δ13C to reconstruct spatiotemporal patterns during the organizational phase of a megadrought, demonstrating that trees near the arid boundaries of a species' distribution might be useful in the early detection of long-lasting droughts.

Modeling Ambitions Outpace Observations of Forest Carbon Allocation.

There have been vociferous calls for 'tree-centered' vegetation models to refine predictions of forest carbon (C) cycling. Unfortunately, our global survey at flux-tower sites indicates insufficient empirical data support for this much-needed model development. We urge for a new generation of studies across large environmental gradients that strategically pair long-term ecosystem monitoring with manipulative experiments on mature trees. For this, we outline a versatile experimental framework to build cross-scale data archives of C uptake and allocation to structural, non-structural, and respiratory sinks. Community-wide efforts and discussions are needed to implement this framework, especially in hitherto underrepresented tropical forests. Global coordination and realistic priorities for data collection will thereby be key to achieve and maintain adequate empirical support for tree-centered vegetation modeling.

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.

Axial changes in wood functional traits have limited net effects on stem biomass increment in European beech (Fagus sylvatica).

During the growing season, trees allocate photoassimilates to increase their aboveground woody biomass in the stem (ABIstem). This 'carbon allocation' to structural growth is a dynamic process influenced by internal and external (e.g., climatic) drivers. While radial variability in wood formation and its resulting structure have been intensively studied, their variability along tree stems and subsequent impacts on ABIstem remain poorly understood. We collected wood cores from mature trees within a fixed plot in a well-studied temperate Fagus sylvatica L. forest. For a subset of trees, we performed regular interval sampling along the stem to elucidate axial variability in ring width (RW) and wood density (ρ), and the resulting effects on tree- and plot-level ABIstem. Moreover, we measured wood anatomical traits to understand the anatomical basis of ρ and the coupling between changes in RW and ρ during drought. We found no significant axial variability in ρ because an increase in the vessel-to-fiber ratio with smaller RW compensated for vessel tapering towards the apex. By contrast, temporal variability in RW varied significantly along the stem axis, depending on the growing conditions. Drought caused a more severe growth decrease, and wetter summers caused a disproportionate growth increase at the stem base compared with the top. Discarding this axial variability resulted in a significant overestimation of tree-level ABIstem in wetter and cooler summers, but this bias was reduced to ~2% when scaling ABIstem to the plot level. These results suggest that F. sylvatica prioritizes structural carbon sinks close to the canopy when conditions are unfavorable. The different axial variability in RW and ρ thereby indicates some independence of the processes that drive volume growth and wood structure along the stem. This refines our knowledge of carbon allocation dynamics in temperate diffuse-porous species and contributes to reducing uncertainties in determining forest carbon fixation.

Assessing the response of forest productivity to climate extremes in Switzerland using model-data fusion.

The response of forest productivity to climate extremes strongly depends on ambient environmental and site conditions. To better understand these relationships at a regional scale, we used nearly 800 observation years from 271 permanent long-term forest monitoring plots across Switzerland, obtained between 1980 and 2017. We assimilated these data into the 3-PG forest ecosystem model using Bayesian inference, reducing the bias of model predictions from 14% to 5% for forest stem carbon stocks and from 45% to 9% for stem carbon stock changes. We then estimated the productivity of forests dominated by Picea abies and Fagus sylvatica for the period of 1960-2018, and tested for productivity shifts in response to climate along elevational gradient and in extreme years. Simulated net primary productivity (NPP) decreased with elevation (2.86 ± 0.006 Mg C ha-1  year-1  km-1 for P. abies and 0.93 ± 0.010 Mg C ha-1  year-1  km-1 for F. sylvatica). During warm-dry extremes, simulated NPP for both species increased at higher and decreased at lower elevations, with reductions in NPP of more than 25% for up to 21% of the potential species distribution range in Switzerland. Reduced plant water availability had a stronger effect on NPP than temperature during warm-dry extremes. Importantly, cold-dry extremes had negative impacts on regional forest NPP comparable to warm-dry extremes. Overall, our calibrated model suggests that the response of forest productivity to climate extremes is more complex than simple shift toward higher elevation. Such robust estimates of NPP are key for increasing our understanding of forests ecosystems carbon dynamics under climate extremes.

The climatic drivers of primary Picea forest growth along the Carpathian arc are changing under rising temperatures.

Climatic constraints on tree growth mediate an important link between terrestrial and atmospheric carbon pools. Tree rings provide valuable information on climate-driven growth patterns, but existing data tend to be biased toward older trees on climatically extreme sites. Understanding climate change responses of biogeographic regions requires data that integrate spatial variability in growing conditions and forest structure. We analyzed both temporal (c. 1901-2010) and spatial variation in radial growth patterns in 9,876 trees from fragments of primary Picea abies forests spanning the latitudinal and altitudinal extent of the Carpathian arc. Growth was positively correlated with summer temperatures and spring moisture availability throughout the entire region. However, important seasonal variation in climate responses occurred along geospatial gradients. At northern sites, winter precipitation and October temperatures of the year preceding ring formation were positively correlated with ring width. In contrast, trees at the southern extent of the Carpathians responded negatively to warm and dry conditions in autumn of the year preceding ring formation. An assessment of regional synchronization in radial growth variability showed temporal fluctuations throughout the 20th century linked to the onset of moisture limitation in southern landscapes. Since the beginning of the study period, differences between high and low elevations in the temperature sensitivity of tree growth generally declined, while moisture sensitivity increased at lower elevations. Growth trend analyses demonstrated changes in absolute tree growth rates linked to climatic change, with basal area increments in northern landscapes and lower altitudes responding positively to recent warming. Tree growth has predominantly increased with rising temperatures in the Carpathians, accompanied by early indicators that portions of the mountain range are transitioning from temperature to moisture limitation. Continued warming will alleviate large-scale temperature constraints on tree growth, giving increasing weight to local drivers that are more challenging to predict.

Twentieth century redistribution in climatic drivers of global tree growth.

Energy and water limitations of tree growth remain insufficiently understood at large spatiotemporal scales, hindering model representation of interannual or longer-term ecosystem processes. By assessing and statistically scaling the climatic drivers from 2710 tree-ring sites, we identified the boreal and temperate land areas where tree growth during 1930-1960 CE responded positively to temperature (20.8 ± 3.7 Mio km2; 25.9 ± 4.6%), precipitation (77.5 ± 3.3 Mio km2; 96.4 ± 4.1%), and other parameters. The spatial manifestation of this climate response is determined by latitudinal and altitudinal temperature gradients, indicating that warming leads to geographic shifts in growth limitations. We observed a significant (P < 0.001) decrease in temperature response at cold-dry sites between 1930-1960 and 1960-1990 CE, and the total temperature-limited area shrunk by -8.7 ± 0.6 Mio km2. Simultaneously, trees became more limited by atmospheric water demand almost worldwide. These changes occurred under mild warming, and we expect that continued climate change will trigger a major redistribution in growth responses to climate.