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Forest models are instrumental for understanding and projecting the impact of climate change on forests. A considerable number of forest models have been developed in the last decades. However, few systematic and comprehensive model comparisons have been performed in Europe that combine an evaluation of modelled carbon and water fluxes and forest structure. We evaluate 13 widely used, state-of-the-art, stand-scale forest models against field measurements of forest structure and eddy-covariance data of carbon and water fluxes over multiple decades across an environmental gradient at nine typical European forest stands. We test the models' performance in three dimensions: accuracy of local predictions (agreement of modelled and observed annual data), realism of environmental responses (agreement of modelled and observed responses of daily gross primary productivity to temperature, radiation and vapour pressure deficit) and general applicability (proportion of European tree species covered). We find that multiple models are available that excel according to our three dimensions of model performance. For the accuracy of local predictions, variables related to forest structure have lower random and systematic errors than annual carbon and water flux variables. Moreover, the multi-model ensemble mean provided overall more realistic daily productivity responses to environmental drivers across all sites than any single individual model. The general applicability of the models is high, as almost all models are currently able to cover Europe's common tree species. We show that forest models complement each other in their response to environmental drivers and that there are several cases in which individual models outperform the model ensemble. Our framework provides a first step to capturing essential differences between forest models that go beyond the most commonly used accuracy of predictions. Overall, this study provides a point of reference for future model work aimed at predicting climate impacts and supporting climate mitigation and adaptation measures in forests., Global Change Biology, 28 (23), ISSN:1354-1013, ISSN:1365-2486

To understand forest dynamics under today’s changing environmental conditions, it is important to analyze the state of forests at large scales. Forest inventories are not available for all regions, so it is important to use other additional methods, e.g., remote sensing observations. Increasingly, remotely sensed data based on optical instruments and airborne LIDAR are becoming widely available for forests. There is great potential in analyzing these measurements and gaining an understanding of forest states. In this work, we combine the new-generation radiative transfer model mScope with the individual-based forest model FORMIND to generate reflectance spectra for forests. Combining the two models allows us to account for species diversity at different height layers in the forest. We compare the generated reflectances for forest stands in Finland, in the region of North Karelia, with Sentinel-2 measurements. We investigate which level of forest representation gives the best results and explore the influence of different calculation methods of mean leaf parameters. For the majority of the forest stands, we generated good reflectances with all levels of forest representation compared to the measured reflectance. Good correlations were also found for the vegetation indices (especially NDVI with R2=0.62). This work provides a forward modeling approach for relating forest reflectance to forest characteristics. With this tool, it is possible to analyze a large set of forest stands with corresponding reflectances. This opens up the possibility to understand how reflectance is related to succession and different forest conditions.

Ecosystem functions of temperate forests are expected to be severely impacted by future climate change - particularly hydro-meteorological extremes (heavy precipitation events, droughts, and heat waves) that will increase in frequency, duration, magnitude, and extent. Previous studies have shown that both structural and tree species diversity may act as buffers against the impacts of climate extremes.To better understand the influence of structural and tree species diversity, we use two models to analyze the influence of species and structural diversity on hydrologic dynamics during recent drought events. The well-equipped test sites are located in central Germany and represent typical forests in this area. One model is the individual forest gap model FORMIND. Using a newly developed technique, it allows us to analyze local heterogeneous patterns on a 2 meter scale of carbon and water cycling, flow, and water stress, and their relationship to structural and species diversity. The second model is mHM, a mesoscale hydrological model. We parameterize mHM for two catchments in central Germany (Nägelstedt and Upper Saale). We further analyze the relevance of local heterogeneity in structural and species diversity and the resulting local heterogeneity in water fluxes on the fluxes at the mesoscale.This two-model and interdisciplinary workflow allows us to consider various soil-plant-atmosphere interactions in drought disturbed ecohydrological systems

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., Forest models can help understanding the processes that shape forest functioning, structure and diversity, since they can can simulate forest dynamics over spatio‐temporal scales unreachable by most empirical investigations. Here we describe the development of three widely applied but contrasting forest mo−delling approaches — species distribution models, individual‐based models and dynamic global vegetation models. We provide an overview of recent model applications and pinpoint ten key questions that could, and should, be tackled with forest models in the next decade.

Ensuring ecosystem resilience is an intuitive approach to safeguard the functioning of ecosystems and hence the future provisioning of ecosystem services (ES). However, resilience is a multi-faceted concept that is difficult to operationalize. Focusing on resilience mechanisms, such as diversity, network architectures or adaptive capacity, has recently been suggested as means to operationalize resilience. Still, the focus on mechanisms is not specific enough. We suggest a conceptual framework, resilience trinity, to facilitate management based on resilience mechanisms in three distinctive decision contexts and time-horizons: i) reactive, when there is an imminent threat to ES resilience and a high pressure to act, ii) adjustive, when the threat is known in general but there is still time to adapt management, and iii) provident, when time horizons are very long and the nature of the threats is uncertain, leading to a low willingness to act. Resilience has different interpretations and implications at these different time horizons, which also prevail in different disciplines. Social ecology, ecology, and engineering are often implicitly focussing on provident, adjustive, or reactive resilience, respectively, but these different notions and of resilience and their corresponding social, ecological, and economic trade-offs need to be reconciled. Otherwise, we keep risking unintended consequences of reactive actions, or shying away from provident action because of uncertainties that cannot be reduced. The suggested trinity of time horizons and their decision contexts could help ensuring that longer-term management actions are not missed while urgent threats to ES are given priority.

International audience; Plant trait variability, emerging from eco-evolutionary dynamics that range from alleles to macroecological scales, is one of the most elusive, but possibly most consequential, aspects of biodiversity. Plasticity, epigenetics, and genetic diversity are major determinants of how plants will respond to climate change, yet these processes are rarely represented in current vegetation models. Here, we provide an overview of the challenges associated with understanding the causes and consequences of plant trait variability, and review current developments to include plasticity and evolutionary mechanisms in vegetation models. We also present a roadmap of research priorities to develop a next generation of vegetation models with flexible traits. Including trait variability in vegetation models is necessary to better represent biosphere responses to global change.

Forests provide important ecosystem services such as carbon sequestration. Forest landscapes are intrinsically heterogeneous—a problem for biomass and productivity assessment using remote sensing. Forest structure constitutes valuable additional information for the improved estimation of these variables. However, survey of forest structure by remote sensing remains a challenge which results mainly from the differences in forest structure metrics derived by using remote sensing compared to classical structural metrics from field data. To understand these differences, remote sensing measurements were linked with an individual-based forest model. Forest structure was analyzed by lidar remote sensing using metrics for the horizontal and vertical structures. To investigate the role of forest structure for biomass and productivity estimations in temperate forests, 25 lidar metrics of 375,000 simulated forest stands were analyzed. For the lidar-based metrics, top-of-canopy height arose as the best predictor for describing horizontal forest structure. The standard deviation of the vertical foliage profile was the best predictor for the vertical heterogeneity of a forest. Forest structure was also an important factor for the determination of forest biomass and aboveground wood productivity. In particular, horizontal structure was essential for forest biomass estimation. Predicting aboveground wood productivity must take into account both horizontal and vertical structures. In a case study based on these findings, forest structure, biomass and aboveground wood productivity are mapped for whole of Germany. The dominant type of forest in Germany is dense but less vertically structured forest stands. The total biomass of all German forests is 2.3 Gt, and the total aboveground woody productivity is 43 Mt/year. Future remote sensing missions will have the capability to provide information on forest structure (e.g., from lidar or radar). This will lead to more accurate assessments of forest biomass and productivity. These estimations can be used to evaluate forest ecosystems related to climate regulation and biodiversity protection.

Plant functional diversity (FD) is an important component of biodiversity. Evidence shows that FD strongly determines ecosystem functioning and stability and also regulates various ecosystem services that underpin human well-being. Given the importance of FD, it is critical to monitor its variations in an explicit manner across space and time, a highly demanding task that cannot be resolved solely by field data. Today, high hopes are placed on satellite-based observations to complement field plot data. The promise is that multiscale monitoring of plant FD, ecosystem functioning, and their services is now possible at global scales in near real-time. However, non-trivial scale challenges remain to be overcome before plant ecology can capitalize on the latest advances in Earth Observation (EO). Here, we articulate the existing scale challenges in linking field and satellite data and further elaborated in detail how to address these challenges via the latest innovations in optical and radar sensor technologies and image analysis algorithms. Addressing these challenges not only requires novel remote sensing theories and algorithms but also urges more effective communication between remote sensing scientists and field ecologists to foster mutual understanding of the existing challenges. Only through a collaborative approach can we achieve the global plant functional diversity monitoring goal.

Accurate accounting of aboveground biomass density (AGBD) is crucial for carbon cycle, biodiversity, and climate change science. The Global Ecosystem Dynamics Investigation (GEDI), which maps global AGBD from waveform lidar, is the first of a new generation of Earth observation missions designed to improve carbon accounting. This paper explores the possibility that lidar waveforms may not be unique to AGBD—that forest stands with different AGBD may produce highly similar waveforms—and we hypothesize that non-uniqueness may contribute to the large uncertainties in AGBD predictions. Our analysis integrates simulated GEDI waveforms from 428 in situ stem maps with output from an individual-based forest gap model, which we use to generate a database of potential forest stands and simulate GEDI waveforms from those stands. We use this database to predict the AGBD of the 428 in situ stem maps via two different methods: a linear regression from waveform metrics, and a waveform-matching approach that accounts for waveform-AGBD non-uniqueness. We find that some in situ waveforms are more unique to AGBD than others, which notably impacts AGBD prediction uncertainty (7–411 Mg ha−1, average of 167 Mg ha−1). We also find that forest structure complexity may influence the non-uniqueness effect; stands with low structural complexity are more unique to AGBD than more mature stands with multiple cohorts and canopy layers. These findings suggest that the non-uniqueness phenomena may be introduced by the measuring characteristics of waveform lidar in combination with how forest structure manifests at small scales, and we discuss how this complexity may complicate uncertainty estimation in AGBD prediction. This analysis suggests a limit to the accuracy and precision of AGBD predictions from lidar waveforms seen in empirical studies, and underscores the need for further exploration of the relationships between lidar remote sensing measurements, forest structure, and AGBD.

Tropical forests play an important role in the global carbon cycle. High-resolution remote sensing techniques, e.g., spaceborne lidar, can measure complex tropical forest structures, but it remains a challenge how to interpret such information for the assessment of forest biomass and productivity. Here, we develop an approach to estimate basal area, aboveground biomass and productivity within Amazonia by matching 770,000 GLAS lidar (ICESat) profiles with forest simulations considering spatial heterogeneous environmental and ecological conditions. This allows for deriving frequency distributions of key forest attributes for the entire Amazon. This detailed interpretation of remote sensing data improves estimates of forest attributes by 20–43% as compared to (conventional) estimates using mean canopy height. The inclusion of forest modeling has a high potential to close a missing link between remote sensing measurements and the 3D structure of forests, and may thereby improve continent-wide estimates of biomass and productivity., Improving estimates of forest biomass based on remote sensing data is important to assess global carbon cycling. Here the authors develop an approach to use forest gap models to simulate lidar waveforms and compare the outputs with ICESAT-1 GLAS profiles, showing improved estimates across the Amazon basin.

Plant functional diversity (FD) is an important component of biodiversity that characterizes the variability of functional traits within a community, landscape, or even large spatial scales. It can influence ecosystem processes and stability. Hence, it is important to understand how and why FD varies within and between ecosystems, along resources availability gradients and climate gradients, and across vegetation successional stages. Usually, FD is assessed through labor-intensive field measurements, while assessing FD from space may provide a way to monitor global FD changes in a consistent, time and resource efficient way. The potential of operational satellites for inferring FD, however, remains to be demonstrated. Here we studied the relationships between FD and spectral reflectance measurements taken by ESA's Sentinel-2 satellite over 117 field plots located in 6 European countries, with 46 plots having in-situ sampled leaf traits and the other 71 using traits from the TRY database. These field plots represent major European forest types, from boreal forests in Finland to Mediterranean mixed forests in Spain. Based on in-situ data collected in 2013 we computed functional dispersion (FDis), a measure of FD, using foliar and whole-plant traits of known ecological significance. These included five foliar traits: leaf nitrogen concentration (N%), leaf carbon concentration (%C), specific leaf area (SLA), leaf dry matter content (LDMC), leaf area (LA). In addition they included three whole-plant traits: tree height (H), crown cross-sectional area (CCSA), and diameter-at-breast-height (DBH). We applied partial least squares regression using Sentinel-2 surface reflectance measured in 2015 as predictive variables to model in-situ FDis measurements. We predicted, in cross-validation, 55% of the variation in the observed FDis. We also showed that the red-edge, near infrared and shortwave infrared regions of Sentinel-2 are more important than the visible region for predicting FDis. An initial 30-m resolution mapping of FDis revealed large local FDis variation within each forest type. The novelty of this study is the effective integration of spaceborne and in-situ measurements at a continental scale, and hence represents a key step towards achieving rapid global biodiversity monitoring schemes., This work was primarily supported by an iDiv-Flexpool project “Observing effects of biodiversity on ecosystem functioning across time, space, and wavelength: oBEF-Across” and a European Union H2020 project “Detecting changes in essential ecosystem and biodiversity properties - towards a Biosphere Atmosphere Change Index: BACI” (grant agreement number: 640176). The data from FunDivEUROPE network was collected with the financial support from the European Union Seventh Framework Programme (FP7/2007-2013) (grant agreement number: 265171). Remeasurements across all FunDivEUROPE plots were financed by EU H2020 project Soil4Europe (Bioidversa 2017-2019). We thank Prof. Dr. Michael Scherer-Lorenzen and Prof. Dr. Kris Verheyen from FunDivEUROPE network for their generous help with field data during the paper revision. DEPM and MM and MDM also acknowledge the support of the Trustee European Commission project no H2020-MSCA-ITN-2016 – 721995. The free use of Sentinel-2 data is enabled by ESA's Copernicus Open Access Hub. The in-situ plant traits data collected over Finnish, Romanian, and Spanish sites were supported by a Marie-Curie Fellowship (DIVERFOR, FP7-PEOPLE-2011-IEF. No. 302445) to R. Benavides. The free access of plant trait data was supported by the TRY initiative (http://www.try-db.org)

Ensuring ecosystem resilience is an intuitive approach to safeguard future provisioning of ecosystem services (ES). However, resilience is an ambiguous concept and difficult to operationalize. Focusing on resilience mechanisms, such as diversity, network architectures or adaptive capacity, has recently been suggested as means to operationalize resilience. Still, the focus on mechanisms is not specific enough because the usefulness of a mechanism is context-dependent. We suggest a conceptual framework, resilience trinity, to facilitate management of resilience mechanisms in three distinctive decision contexts and time-horizons. i) reactive, when there is an imminent threat to ES resilience and a high pressure to act, ii) adjustive, when the threat is known in general but there is still time to adapt management, and iii) provident when time horizons are very long and the nature of the threats is uncertain, leading to a low willingness to act. This emphasizes that resilience has different interpretations and implications at different time horizons which however need to be reconciled. The inclusion of time into resilience thinking ensures that longer-term management actions are not missed while urgent threats to ES are given priority.

Forests represent an important pool in the global carbon cycle. However, biomass stocks and carbon fluxes are variable due to the fact that forest dynamics are driven by processes that act on different spatial and temporal scales. Estimating forest biomass and productivity for larger regions is therefore a major challenge. In this study, horizontal and vertical forest structure is used to improve forest ecosystem service mapping by remote sensing. By linking remote sensing techniques with vegetation modelling (here FORMIND) and forest inventories, forest structure maps were derived for Germany (resolution 4 km). Using these maps, the role of forest structure for selected ecosystem services of forests has been investigated. For forest state estimations (like biomass) horizontal forest structure plays a key role while for productivity estimations both horizontal and vertical structures are relevant. This concept of forest structure classification in combination with forest modelling and remote sensing has high potential for applications at continental scales as future remote sensing missions will provide information on forest structure.

International audience; 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-dimensionalassessments are still to be recommended.

Rising temperatures due to climate change influence the wood production of forests. Observations show that some temperate forests increase their productivity, whereas others reduce their productivity. This study focuses on how species composition and forest structure properties influence the temperature sensitivity of aboveground wood production (AWP). It further investigates which forests will increase their productivity the most with rising temperatures. We described forest structure by leaf area index, forest height and tree height heterogeneity. Species composition was described by a functional diversity index (Rao's Q) and a species distribution index (ΩAWP). ΩAWP quantified how well species are distributed over the different forest layers with regard to AWP. We analysed 370 170 forest stands generated with a forest gap model. These forest stands covered a wide range of possible forest types. For each stand, we estimated annual aboveground wood production and performed a climate sensitivity analysis based on 320 different climate time series (of 1-year length). The scenarios differed in mean annual temperature and annual temperature amplitude. Temperature sensitivity of wood production was quantified as the relative change in productivity resulting from a 1 ∘C rise in mean annual temperature or annual temperature amplitude. Increasing ΩAWP positively influenced both temperature sensitivity indices of forest, whereas forest height showed a bell-shaped relationship with both indices. Further, we found forests in each successional stage that are positively affected by temperature rise. For such forests, large ΩAWP values were important. In the case of young forests, low functional diversity and small tree height heterogeneity were associated with a positive effect of temperature on wood production. During later successional stages, higher species diversity and larger tree height heterogeneity were an advantage. To achieve such a development, one could plant below the closed canopy of even-aged, pioneer trees a climax-species-rich understorey that will build the canopy of the mature forest. This study highlights that forest structure and species composition are both relevant for understanding the temperature sensitivity of wood production.

Background: Capturing the response of forest ecosystems to inter-annual climate variability is a great challenge. In this study, we tested the capability of an individual-based forest gap model to display carbon fluxes at yearly and daily time scales. The forest model was applied to a spruce forest to simulate the gross primary production (GPP), respiration and net ecosystem exchange (NEE). We analyzed how the variability in climate affected simulated carbon fluxes at the scale of the forest model.[br/][br/] Results: Six years were simulated at a daily time scale and compared to the observed eddy covariance (EC) data. In general, the seasonal cycle of the individual carbon fluxes was correctly described by the forest model. However, the estimated GPP differed from the observed data on the days of extreme climatic conditions. Two new parameterizations were developed: one resulting from a numerical calibration, and the other resulting from a filtering method. We suggest new parameter values and even a new function for the temperature limitation of photosynthesis.[br/][br/] Conclusions: The forest model reproduced the observed carbon fluxes of a forest ecosystem quite well. Of the three parameterizations, the calibrated model version performed best. However, the filtering approach showed that calibrated parameter values do not necessarily correctly display the individual functional relations. The concept of simulating forest dynamics at the individual base is a valuable tool for simulating the NEE, GPP and respiration of forest ecosystems.

We parametrize the maintenance respiration of a single tree depending on reference climate parameters (light, temperature and precipitation) and the observed stem diameter increase resulting from that climate. The simulated biomass increment results from photosynthesis under the given climate scenario and is then reduced by maintenance and growth respiration. We incorporate this new carbon allocation algorithm into the established individual based gap model FORMIND to reproduce the biomass development of typical central European forest stands. Yield tables for northern Germany recorded over the last century are used for our parametrizations, along with the climate of the area at the time of recording. The model simulates eight tree species based on data on pine, spruce, beech, oak, ash, poplar, birch and robinia. The model dynamics emerge from tree competition, growth and mortality. These processes are calculated on an annual scale. The climate variables (global radiation, air temperature and precipitation) are entered into the model in daily resolution. This new version of FORMIND version reproduces the forest biomass development represented in the yield tables for northern Germany as well as those for western France. The modeled annual fluxes of gross primary production, woody net primary production and autotrophic respiration correspond with results from eddy flux measurements. Therefore, this version of FORMIND with the new carbon allocation is a suitable tool to investigate the carbon flux, biomass development and potential yield of forests at the individual tree level in the temperate climate zone.

Forest management requires prediction of forest growth, but there is no general agreement about which models best predict growth, how to quantify model parameters, and how to assess the uncertainty of model predictions. In this paper, we show how Bayesian calibration (BC), Bayesian model comparison (BMC) and Bayesian model averaging (BMA) can help address these issues. We used six models, ranging from simple parameter-sparse models to complex process-based models: 3PG, 4C, ANAFORE, BASFOR, BRIDGING and FORMIND. For each model, the initial degree of uncertainty about parameter values was expressed in a prior probability distribution. Inventory data for Scots pine on tree height and diameter, with estimates of measurement uncertainty, were assembled for twelve sites, from four countries: Austria, Belgium, Estonia and Finland. From each country, we used data from two sites of the National Forest Inventories (NFIs), and one Permanent Sample Plot (PSP). The models were calibrated using the NFI-data and tested against the PSP-data. Calibration was done both per country and for all countries simultaneously, thus yielding country-specific and generic parameter distributions. We assessed model performance by sampling from prior and posterior distributions and comparing the growth predictions of these samples to the observations at the PSPs. We found that BC reduced uncertainties strongly in all but the most complex model. Surprisingly, country-specific BC did not lead to clearly better within-country predictions than generic BC. BMC identified the BRIDGING model, which is of intermediate complexity, as the most plausible model before calibration, with 4C taking its place after calibration. In this BMC, model plausibility was quantified as the relative probability of a model being correct given the information in the PSP-data. We discuss how the method of model initialisation affects model performance. Finally, we show how BMA affords a robust way of predicting forest growth that accounts for both parametric and model structural uncertainty. (C) 2012 Elsevier B.V. All rights reserved.

While various relationships between productivity and biodiversity are found in forests, the processes underlying these relationships remain unclear and theory struggles to coherently explain them. In this work, we analyse diversity–productivity relationships through an examination of forest structure (described by basal area and tree height heterogeneity). We use a new modelling approach, called ‘forest factory’, which generates various forest stands and calculates their annual productivity (above-ground wood increment). Analysing approximately 300 000 forest stands, we find that mean forest productivity does not increase with species diversity. Instead forest structure emerges as the key variable. Similar patterns can be observed by analysing 5054 forest plots of the German National Forest Inventory. Furthermore, we group the forest stands into nine forest structure classes, in which we find increasing, decreasing, invariant and even bell-shaped relationships between productivity and diversity. In addition, we introduce a new index, called optimal species distribution, which describes the ratio of realized to the maximal possible productivity (by shuffling species identities). The optimal species distribution and forest structure indices explain the obtained productivity values quite well ( R 2 between 0.7 and 0.95), whereby the influence of these attributes varies within the nine forest structure classes.

Forests worldwide are threatened by various environmental and anthropogenic hazards, especially tropical forests. Knowledge on the impacts of these hazards on forest structure and dynamics has been compiled in empirical studies. However, the results of these studies are often not sufficient for long-term projections and extrapolations to large spatial scales especially for unprecedented environmental conditions, which require both the identification and understanding of key underlying processes. Forest models bridge this gap by incorporating multiple ecological processes in a dynamic framework (i.e. including a realistic model structure) and addressing the complexity of forest ecosystems. Here, we describe the evolution of the individual-based and process-based forest gap model FORMIND and its application to tropical forests. At its core, the model includes physiological processes on tree level (photosynthesis, respiration, tree growth, mortality, regeneration, competition). During the past two decades, FORMIND has been used to address various scientific questions arising from different forest types by continuously extending the model structure. The model applications thus provided understanding in three main aspects: (1) the grouping of single tree species into plant functional types is a successful approach to reduce complexity in vegetation models, (2) structural realism was necessary to analyze impacts of natural and anthropogenic disturbances such as logging, fragmentation, or drought, and (3) complex ecological processes such as carbon fluxes in tropical forests – starting from the individual tree level up to the entire forest ecosystem – can be explored as a function of forest structure, species composition and disturbance regime. Overall, this review shows how the evolution of long-term modelling projects not only provides scientific understanding of forest ecosystems, but also provides benefits for ecological theory and empirical study design. peerReviewed