'How to adapt forests?'-Exploring the role of leaf trait diversity for long-term forest biomass under new climate normals.

Forests, critical components of global ecosystems, face unprecedented challenges due to climate change. This study investigates the influence of functional diversity-as a component of biodiversity-to enhance long-term biomass of European forests in the context of changing climatic conditions. Using the next-generation flexible trait-based vegetation model, LPJmL-FIT, we explored the impact of functional diversity on long-term forest biomass under three different climate change scenarios (video abstract: https://www.pik-potsdam.de/~billing/video/2023/video_abstract_billing_et_al_LPJmLFIT.mp4). Four model set-ups were tested with varying degrees of functional diversity and best-suited functional traits. Our results show that functional diversity positively influences long-term forest biomass, particularly when climate warming is low (RCP2.6). Under these conditions, high-diversity simulations led to an approximately 18.2% increase in biomass compared to low-diversity experiments. However, as climate change intensity increased, the benefits of functional diversity diminished (RCP8.5). A Bayesian multilevel analysis revealed that both full leaf trait diversity and diversity of plant functional types contributed significantly to biomass enhancement under low warming scenarios in our model simulations. Under strong climate change, the presence of a mixture of different functional groups (e.g. summergreen and evergreen broad-leaved trees) was found more beneficial than the diversity of leaf traits within a functional group (e.g. broad-leaved summergreen trees). Ultimately, this research challenges the notion that planting only the most productive and climate-suited trees guarantees the highest future biomass and carbon sequestration. We underscore the importance of high functional diversity and the potential benefits of fostering a mixture of tree functional types to enhance long-term forest biomass in the face of climate change.

Earth beyond six of nine planetary boundaries.

This planetary boundaries framework update finds that six of the nine boundaries are transgressed, suggesting that Earth is now well outside of the safe operating space for humanity. Ocean acidification is close to being breached, while aerosol loading regionally exceeds the boundary. Stratospheric ozone levels have slightly recovered. The transgression level has increased for all boundaries earlier identified as overstepped. As primary production drives Earth system biosphere functions, human appropriation of net primary production is proposed as a control variable for functional biosphere integrity. This boundary is also transgressed. Earth system modeling of different levels of the transgression of the climate and land system change boundaries illustrates that these anthropogenic impacts on Earth system must be considered in a systemic context.

Future tree survival in European forests depends on understorey tree diversity.

Climate change heavily threatens forest ecosystems worldwide and there is urgent need to understand what controls tree survival and forests stability. There is evidence that biodiversity can enhance ecosystem stability (Loreau and de Mazancourt in Ecol Lett 16:106-115, 2013; McCann in Nature 405:228-233, 2000), however it remains largely unclear whether this also holds for climate change and what aspects of biodiversity might be most important. Here we apply machine learning to outputs of a flexible-trait Dynamic Global Vegetation Model to unravel the effects of enhanced functional tree trait diversity and its sub-components on climate-change resistance of temperate forests ( http://www.pik-potsdam.de/~billing/video/Forest_Resistance_LPJmLFIT.mp4 ). We find that functional tree trait diversity enhances forest resistance. We explain this with 1. stronger complementarity effects (~ 25% importance) especially improving the survival of trees in the understorey of up to + 16.8% (± 1.6%) and 2. environmental and competitive filtering of trees better adapted to future climate (40-87% importance). We conclude that forests containing functionally diverse trees better resist and adapt to future conditions. In this context, we especially highlight the role of functionally diverse understorey trees as they provide the fundament for better survival of young trees and filtering of resistant tree individuals in the future.

Tackling unresolved questions in forest ecology: The past and future role of simulation models.

Understanding the processes that shape forest functioning, structure, and diversity remains challenging, although data on forest systems are being collected at a rapid pace and across scales. Forest models have a long history in bridging data with ecological knowledge and can simulate forest dynamics over spatio-temporal scales unreachable by most empirical investigations.We describe the development that different forest modelling communities have followed to underpin the leverage that simulation models offer for advancing our understanding of forest ecosystems.Using three widely applied but contrasting approaches - species distribution models, individual-based forest models, and dynamic global vegetation models - as examples, we show how scientific and technical advances have led models to transgress their initial objectives and limitations. We provide an overview of recent model applications on current important ecological topics and pinpoint ten key questions that could, and should, be tackled with forest models in the next decade.Synthesis. This overview shows that forest models, due to their complementarity and mutual enrichment, represent an invaluable toolkit to address a wide range of fundamental and applied ecological questions, hence fostering a deeper understanding of forest dynamics in the context of global change.

Constraining modelled global vegetation dynamics and carbon turnover using multiple satellite observations.

The response of land ecosystems to future climate change is among the largest unknowns in the global climate-carbon cycle feedback. This uncertainty originates from how dynamic global vegetation models (DGVMs) simulate climate impacts on changes in vegetation distribution, productivity, biomass allocation, and carbon turnover. The present-day availability of a multitude of satellite observations can potentially help to constrain DGVM simulations within model-data integration frameworks. Here, we use satellite-derived datasets of the fraction of absorbed photosynthetic active radiation (FAPAR), sun-induced fluorescence (SIF), above-ground biomass of trees (AGB), land cover, and burned area to constrain parameters for phenology, productivity, and vegetation dynamics in the LPJmL4 DGVM. Both the prior and the optimized model accurately reproduce present-day estimates of the land carbon cycle and of temporal dynamics in FAPAR, SIF and gross primary production. However, the optimized model reproduces better the observed spatial patterns of biomass, tree cover, and regional forest carbon turnover. Using a machine learning approach, we found that remaining errors in simulated forest carbon turnover can be explained with bioclimatic variables. This demonstrates the need to improve model formulations for climate effects on vegetation turnover and mortality despite the apparent successful constraint of simulated vegetation dynamics with multiple satellite observations.

The dimensionality of stability depends on disturbance type.

Ecosystems respond in various ways to disturbances. Quantifying ecological stability therefore requires inspecting multiple stability properties, such as resistance, recovery, persistence and invariability. Correlations among these properties can reduce the dimensionality of stability, simplifying the study of environmental effects on ecosystems. A key question is how the kind of disturbance affects these correlations. We here investigated the effect of three disturbance types (random, species-specific, local) applied at four intensity levels, on the dimensionality of stability at the population and community level. We used previously parameterized models that represent five natural communities, varying in species richness and the number of trophic levels. We found that disturbance type but not intensity affected the dimensionality of stability and only at the population level. The dimensionality of stability also varied greatly among species and communities. Therefore, studying stability cannot be simplified to using a single metric and multi-dimensional assessments are still to be recommended.

Alberta wildfire 2016: Apt contribution from anomalous planetary wave dynamics.

In May-June 2016 the Canadian Province of Alberta suffered one of the most devastating wildfires in its history. Here we show that in mid-April to early May 2016 the large-scale circulation in the mid- and high troposphere of the middle and sub-polar latitudes of the northern hemisphere featured a persistent high-amplitude planetary wave structure dominated by the non-dimensional zonal wave number 4. The strongest anticyclonic wing of this structure was located over western Canada. In combination with a very strong El Niño event in winter 2015/2016 this favored highly anomalous, tinder-dry and high-temperature conditions at the surface in that area, entailing an increased fire hazard there. This critically contributed to the ignition of the Alberta Wildfire in May 2016, appearing to be the costliest disaster in Canadian history thus far.

Large-scale impact of climate change vs. land-use change on future biome shifts in Latin America.

Climate change and land-use change are two major drivers of biome shifts causing habitat and biodiversity loss. What is missing is a continental-scale future projection of the estimated relative impacts of both drivers on biome shifts over the course of this century. Here, we provide such a projection for the biodiverse region of Latin America under four socio-economic development scenarios. We find that across all scenarios 5-6% of the total area will undergo biome shifts that can be attributed to climate change until 2099. The relative impact of climate change on biome shifts may overtake land-use change even under an optimistic climate scenario, if land-use expansion is halted by the mid-century. We suggest that constraining land-use change and preserving the remaining natural vegetation early during this century creates opportunities to mitigate climate-change impacts during the second half of this century. Our results may guide the evaluation of socio-economic scenarios in terms of their potential for biome conservation under global change.

Variation in stem mortality rates determines patterns of above-ground biomass in Amazonian forests: implications for dynamic global vegetation models.

Understanding the processes that determine above-ground biomass (AGB) in Amazonian forests is important for predicting the sensitivity of these ecosystems to environmental change and for designing and evaluating dynamic global vegetation models (DGVMs). AGB is determined by inputs from woody productivity [woody net primary productivity (NPP)] and the rate at which carbon is lost through tree mortality. Here, we test whether two direct metrics of tree mortality (the absolute rate of woody biomass loss and the rate of stem mortality) and/or woody NPP, control variation in AGB among 167 plots in intact forest across Amazonia. We then compare these relationships and the observed variation in AGB and woody NPP with the predictions of four DGVMs. The observations show that stem mortality rates, rather than absolute rates of woody biomass loss, are the most important predictor of AGB, which is consistent with the importance of stand size structure for determining spatial variation in AGB. The relationship between stem mortality rates and AGB varies among different regions of Amazonia, indicating that variation in wood density and height/diameter relationships also influences AGB. In contrast to previous findings, we find that woody NPP is not correlated with stem mortality rates and is weakly positively correlated with AGB. Across the four models, basin-wide average AGB is similar to the mean of the observations. However, the models consistently overestimate woody NPP and poorly represent the spatial patterns of both AGB and woody NPP estimated using plot data. In marked contrast to the observations, DGVMs typically show strong positive relationships between woody NPP and AGB. Resolving these differences will require incorporating forest size structure, mechanistic models of stem mortality and variation in functional composition in DGVMs.

Enhanced seasonal CO2 exchange caused by amplified plant productivity in northern ecosystems.

Atmospheric monitoring of high northern latitudes (above 40°N) has shown an enhanced seasonal cycle of carbon dioxide (CO2) since the 1960s, but the underlying mechanisms are not yet fully understood. The much stronger increase in high latitudes relative to low ones suggests that northern ecosystems are experiencing large changes in vegetation and carbon cycle dynamics. We found that the latitudinal gradient of the increasing CO2 amplitude is mainly driven by positive trends in photosynthetic carbon uptake caused by recent climate change and mediated by changing vegetation cover in northern ecosystems. Our results underscore the importance of climate-vegetation-carbon cycle feedbacks at high latitudes; moreover, they indicate that in recent decades, photosynthetic carbon uptake has reacted much more strongly to warming than have carbon release processes.

Model-data synthesis for the next generation of forest free-air CO2 enrichment (FACE) experiments.

The first generation of forest free-air CO2 enrichment (FACE) experiments has successfully provided deeper understanding about how forests respond to an increasing CO2 concentration in the atmosphere. Located in aggrading stands in the temperate zone, they have provided a strong foundation for testing critical assumptions in terrestrial biosphere models that are being used to project future interactions between forest productivity and the atmosphere, despite the limited inference space of these experiments with regards to the range of global ecosystems. Now, a new generation of FACE experiments in mature forests in different biomes and over a wide range of climate space and biodiversity will significantly expand the inference space. These new experiments are: EucFACE in a mature Eucalyptus stand on highly weathered soil in subtropical Australia; AmazonFACE in a highly diverse, primary rainforest in Brazil; BIFoR-FACE in a 150-yr-old deciduous woodland stand in central England; and SwedFACE proposed in a hemiboreal, Pinus sylvestris stand in Sweden. We now have a unique opportunity to initiate a model-data interaction as an integral part of experimental design and to address a set of cross-site science questions on topics including responses of mature forests; interactions with temperature, water stress, and phosphorus limitation; and the influence of biodiversity.

Codominant water control on global interannual variability and trends in land surface phenology and greenness.

Identifying the relative importance of climatic and other environmental controls on the interannual variability and trends in global land surface phenology and greenness is challenging. Firstly, quantifications of land surface phenology and greenness dynamics are impaired by differences between satellite data sets and phenology detection methods. Secondly, dynamic global vegetation models (DGVMs) that can be used to diagnose controls still reveal structural limitations and contrasting sensitivities to environmental drivers. Thus, we assessed the performance of a new developed phenology module within the LPJmL (Lund-Potsdam-Jena managed Lands) DGVM with a comprehensive ensemble of three satellite data sets of vegetation greenness and ten phenology detection methods, thereby thoroughly accounting for observational uncertainties. The improved and tested model allows us quantifying the relative importance of environmental controls on interannual variability and trends of land surface phenology and greenness at regional and global scales. We found that start of growing season interannual variability and trends are in addition to cold temperature mainly controlled by incoming radiation and water availability in temperate and boreal forests. Warming-induced prolongations of the growing season in high latitudes are dampened by a limited availability of light. For peak greenness, interannual variability and trends are dominantly controlled by water availability and land-use and land-cover change (LULCC) in all regions. Stronger greening trends in boreal forests of Siberia than in North America are associated with a stronger increase in water availability from melting permafrost soils. Our findings emphasize that in addition to cold temperatures, water availability is a codominant control for start of growing season and peak greenness trends at the global scale.

Effects of climate extremes on the terrestrial carbon cycle: concepts, processes and potential future impacts.

Extreme droughts, heat waves, frosts, precipitation, wind storms and other climate extremes may impact the structure, composition and functioning of terrestrial ecosystems, and thus carbon cycling and its feedbacks to the climate system. Yet, the interconnected avenues through which climate extremes drive ecological and physiological processes and alter the carbon balance are poorly understood. Here, we review the literature on carbon cycle relevant responses of ecosystems to extreme climatic events. Given that impacts of climate extremes are considered disturbances, we assume the respective general disturbance-induced mechanisms and processes to also operate in an extreme context. The paucity of well-defined studies currently renders a quantitative meta-analysis impossible, but permits us to develop a deductive framework for identifying the main mechanisms (and coupling thereof) through which climate extremes may act on the carbon cycle. We find that ecosystem responses can exceed the duration of the climate impacts via lagged effects on the carbon cycle. The expected regional impacts of future climate extremes will depend on changes in the probability and severity of their occurrence, on the compound effects and timing of different climate extremes, and on the vulnerability of each land-cover type modulated by management. Although processes and sensitivities differ among biomes, based on expert opinion, we expect forests to exhibit the largest net effect of extremes due to their large carbon pools and fluxes, potentially large indirect and lagged impacts, and long recovery time to regain previous stocks. At the global scale, we presume that droughts have the strongest and most widespread effects on terrestrial carbon cycling. Comparing impacts of climate extremes identified via remote sensing vs. ground-based observational case studies reveals that many regions in the (sub-)tropics are understudied. Hence, regional investigations are needed to allow a global upscaling of the impacts of climate extremes on global carbon-climate feedbacks.

Leaf and stem economics spectra drive diversity of functional plant traits in a dynamic global vegetation model.

Functional diversity is critical for ecosystem dynamics, stability and productivity. However, dynamic global vegetation models (DGVMs) which are increasingly used to simulate ecosystem functions under global change, condense functional diversity to plant functional types (PFTs) with constant parameters. Here, we develop an individual- and trait-based version of the DGVM LPJmL (Lund-Potsdam-Jena managed Land) called LPJmL- flexible individual traits (LPJmL-FIT) with flexible individual traits) which we apply to generate plant trait maps for the Amazon basin. LPJmL-FIT incorporates empirical ranges of five traits of tropical trees extracted from the TRY global plant trait database, namely specific leaf area (SLA), leaf longevity (LL), leaf nitrogen content (Narea ), the maximum carboxylation rate of Rubisco per leaf area (vcmaxarea), and wood density (WD). To scale the individual growth performance of trees, the leaf traits are linked by trade-offs based on the leaf economics spectrum, whereas wood density is linked to tree mortality. No preselection of growth strategies is taking place, because individuals with unique trait combinations are uniformly distributed at tree establishment. We validate the modeled trait distributions by empirical trait data and the modeled biomass by a remote sensing product along a climatic gradient. Including trait variability and trade-offs successfully predicts natural trait distributions and achieves a more realistic representation of functional diversity at the local to regional scale. As sites of high climatic variability, the fringes of the Amazon promote trait divergence and the coexistence of multiple tree growth strategies, while lower plant trait diversity is found in the species-rich center of the region with relatively low climatic variability. LPJmL-FIT enables to test hypotheses on the effects of functional biodiversity on ecosystem functioning and to apply the DGVM to current challenges in ecosystem management from local to global scales, that is, deforestation and climate change effects.

Climate extremes and the carbon cycle.

The terrestrial biosphere is a key component of the global carbon cycle and its carbon balance is strongly influenced by climate. Continuing environmental changes are thought to increase global terrestrial carbon uptake. But evidence is mounting that climate extremes such as droughts or storms can lead to a decrease in regional ecosystem carbon stocks and therefore have the potential to negate an expected increase in terrestrial carbon uptake. Here we explore the mechanisms and impacts of climate extremes on the terrestrial carbon cycle, and propose a pathway to improve our understanding of present and future impacts of climate extremes on the terrestrial carbon budget.

Development of probability density functions for future South American rainfall.

*We estimate probability density functions (PDFs) for future rainfall in five regions of South America, by weighting the predictions of the 24 Coupled Model Intercomparison Archive Project 3 (CMIP3) General Circulation Models (GCMs). The models are rated according to their relative abilities to reproduce the inter-annual variability in seasonal rainfall. *The relative weighting of the climate models is updated sequentially according to Bayes' theorem, based on the biases in the mean of the predicted time-series and the distributional fit of the bias-corrected time-series. *Depending on the season and the region, we find very different rankings of the GCMs, with no single model doing well in all cases. However, in some regions and seasons, differential weighting of the models leads to significant shifts in the derived rainfall PDFs. *Using a combination of the relative model weightings for each season we have also derived a set of overall model weightings for each region that can be used to produce PDFs of forest biomass from the simulations of the Lund-Potsdam-Jena Dynamic Global Vegetation Model for managed land (LPJmL).

Estimating the risk of Amazonian forest dieback.

*Climate change will very likely affect most forests in Amazonia during the course of the 21st century, but the direction and intensity of the change are uncertain, in part because of differences in rainfall projections. In order to constrain this uncertainty, we estimate the probability for biomass change in Amazonia on the basis of rainfall projections that are weighted by climate model performance for current conditions. *We estimate the risk of forest dieback by using weighted rainfall projections from 24 general circulation models (GCMs) to create probability density functions (PDFs) for future forest biomass changes simulated by a dynamic vegetation model (LPJmL). *Our probabilistic assessment of biomass change suggests a likely shift towards increasing biomass compared with nonweighted results. Biomass estimates range between a gain of 6.2 and a loss of 2.7 kg carbon m(-2) for the Amazon region, depending on the strength of CO(2) fertilization. *The uncertainty associated with the long-term effect of CO(2) is much larger than that associated with precipitation change. This underlines the importance of reducing uncertainties in the direct effects of CO(2) on tropical ecosystems.

Ecosystem service supply and vulnerability to global change in Europe.

Global change will alter the supply of ecosystem services that are vital for human well-being. To investigate ecosystem service supply during the 21st century, we used a range of ecosystem models and scenarios of climate and land-use change to conduct a Europe-wide assessment. Large changes in climate and land use typically resulted in large changes in ecosystem service supply. Some of these trends may be positive (for example, increases in forest area and productivity) or offer opportunities (for example, "surplus land" for agricultural extensification and bioenergy production). However, many changes increase vulnerability as a result of a decreasing supply of ecosystem services (for example, declining soil fertility, declining water availability, increasing risk of forest fires), especially in the Mediterranean and mountain regions.