Changes in planned and unplanned canopy openings are linked in Europe's forests.

Canopy openings are increasing in Europe's forests, yet the contributions of anthropogenic and ecological agents of disturbance to this increase remain debated. Here we attribute the root cause of all stand-replacing canopy disturbances identified for Europe in the period 1986-2020 from Landsat data (417,000 km²), distinguishing between planned and unplanned canopy openings (i.e., disturbance by human land use versus by wind, bark beetles, and wildfire). We show that canopy openings by humans dominate the European forest disturbance regime, accounting for 82% of the area disturbed. Both planned and unplanned canopy openings increased in the early 21st century (+24% and +30% relative to the late 20th century). Their changes are linked, with simultaneous increases in planned and unplanned canopy openings on 68% of Europe's forest area. We conclude that an important direction for tackling disturbance change in policy and management is to break the link between planned and unplanned canopy openings in Europe's forests.

A harmonized database of European forest simulations under climate change.

Process-based forest models combine biological, physical, and chemical process understanding to simulate forest dynamics as an emergent property of the system. As such, they are valuable tools to investigate the effects of climate change on forest ecosystems. Specifically, they allow testing of hypotheses regarding long-term ecosystem dynamics and provide means to assess the impacts of climate scenarios on future forest development. As a consequence, numerous local-scale simulation studies have been conducted over the past decades to assess the impacts of climate change on forests. These studies apply the best available models tailored to local conditions, parameterized and evaluated by local experts. However, this treasure trove of knowledge on climate change responses remains underexplored to date, as a consistent and harmonized dataset of local model simulations is missing. Here, our objectives were (i) to compile existing local simulations on forest development under climate change in Europe in a common database, (ii) to harmonize them to a common suite of output variables, and (iii) to provide a standardized vector of auxiliary environmental variables for each simulated location to aid subsequent investigations. Our dataset of European stand- and landscape-level forest simulations contains over 1.1 million simulation runs representing 135 million simulation years for more than 13,000 unique locations spread across Europe. The data were harmonized to consistently describe forest development in terms of stand structure (dominant height), composition (dominant species, admixed species), and functioning (leaf area index). Auxiliary variables provided include consistent daily climate information (temperature, precipitation, radiation, vapor pressure deficit) as well as information on local site conditions (soil depth, soil physical properties, soil water holding capacity, plant-available nitrogen). The present dataset facilitates analyses across models and locations, with the aim to better harness the valuable information contained in local simulations for large-scale policy support, and for fostering a deeper understanding of the effects of climate change on forest ecosystems in Europe.

A climate-induced tree species bottleneck for forest management in Europe.

Large pulses of tree mortality have ushered in a major reorganization of Europe's forest ecosystems. To initiate a robust next generation of trees, the species that are planted today need to be climatically suitable throughout the entire twenty-first century. Here we developed species distribution models for 69 European tree species based on occurrence data from 238,080 plot locations to investigate the option space for current forest management in Europe. We show that the average pool of tree species continuously suitable throughout the century is smaller than that under current and end-of-century climate conditions, creating a tree species bottleneck for current management. If the need for continuous climate suitability throughout the lifespan of a tree planted today is considered, climate change shrinks the tree species pool available to management by between 33% and 49% of its current values (40% and 54% of potential end-of-century values), under moderate (Representative Concentration Pathway 2.6) and severe (Representative Concentration Pathway 8.5) climate change, respectively. This bottleneck could have strong negative impacts on timber production, carbon storage and biodiversity conservation, as only 3.18, 3.53 and 2.56 species of high potential for providing these functions remain suitable throughout the century on average per square kilometre in Europe. Our results indicate that the option space for silviculture is narrowing substantially because of climate change and that an important adaptation strategy in forestry-creating mixed forests-might be curtailed by widespread losses of climatically suitable tree species.

Ecosystem services at risk from disturbance in Europe's forests.

Global change impacts on disturbances can strongly compromise the capacity of forests to provide ecosystem services to society. In addition, many ecosystem services in Europe are simultaneously provided by forests, emphasizing the importance of multifunctionality in forest ecosystem assessments. To address disturbances in forest ecosystem policies and management, spatially explicit risk analyses that consider multiple disturbances and ecosystem services are needed. However, we do not yet know which ecosystem services are most at risk from disturbances in Europe, where the respective risk hotspots are, nor which of the main disturbance agents are most detrimental to the provisioning of multiple ecosystem services from Europe's forests. Here, we quantify the risk of losing important ecosystem services (timber supply, carbon storage, soil erosion control and outdoor recreation) to forest disturbances (windthrows, bark beetle outbreaks and wildfires) in Europe on a continental scale. We find that up to 12% of Europe's ecosystem service supply is at risk from current disturbances. Soil erosion control is the ecosystem service at the highest risk, and windthrow is the disturbance agent posing the highest risk. Disturbances challenge forest multifunctionality by threatening multiple ecosystem services simultaneously on 19.8 Mha (9.7%) of Europe's forests. Our results highlight priority areas for risk management aiming to safeguard the sustainable provisioning of forest ecosystem services.

Resilience and vulnerability: distinct concepts to address global change in forests.

Resilience and vulnerability are important concepts to understand, anticipate, and manage global change impacts on forest ecosystems. However, they are often used confusingly and inconsistently, hampering a synthetic understanding of global change, and impeding communication with managers and policy-makers. Both concepts are powerful and have complementary strengths, reflecting their different history, methodological approach, components, and spatiotemporal focus. Resilience assessments address the temporal response to disturbance and the mechanisms driving it. Vulnerability assessments focus on spatial patterns of exposure and susceptibility, and explicitly address adaptive capacity and stakeholder preferences. We suggest applying the distinct concepts of resilience and vulnerability where they provide particular leverage, and deduce a number of lessons learned to facilitate the next generation of global change assessments.

The anthropogenic imprint on temperate and boreal forest demography and carbon turnover.

Aim: The sweeping transformation of the biosphere by humans over the last millennia leaves only limited windows into its natural state. Much of the forests that dominated temperate and southern boreal regions have been lost and those that remain typically bear a strong imprint of forestry activities and past land-use change, which have changed forest age structure and composition. Here, we ask how would the dynamics, structure and function of temperate and boreal forests differ in the absence of forestry and the legacies of land-use change? Location: Global. Time Period: 2001-2014, integrating over the legacy of disturbance events from 1875 to 2014. Major Taxa Studied: Trees. Methods: We constructed an empirical model of natural disturbance probability as a function of community traits and climate, based on observed disturbance rate and form across 77 protected forest landscapes distributed across three continents. Coupling this within a dynamic vegetation model simulating forest composition and structure, we generated estimates of stand-replacing disturbance return intervals in the absence of forestry for northern hemisphere temperate and boreal forests. We then applied this model to calculate forest stand age structure and carbon turnover rates. Results: Comparison with observed disturbance rates revealed human activities to have almost halved the median return interval of stand-replacing disturbances across temperate forest, with more moderate changes in the boreal region. The resulting forests are typically much younger, especially in northern Europe and south-eastern North America, resulting in a 32% reduction in vegetation carbon turnover time across temperate forests and a 7% reduction for boreal forests. Conclusions: The current northern hemisphere temperate forest age structure is dramatically out of equilibrium with its natural disturbance regimes. Shifts towards more nature-based approaches to forest policy and management should more explicitly consider the current disturbance surplus, as it substantially impacts carbon dynamics and litter (including deadwood) stocks.

Projected climate and canopy change lead to thermophilization and homogenization of forest floor vegetation in a hotspot of plant species richness.

Mountain forests are plant diversity hotspots, but changing climate and increasing forest disturbances will likely lead to far-reaching plant community change. Projecting future change, however, is challenging for forest understory plants, which respond to forest structure and composition as well as climate. Here, we jointly assessed the effects of both climate and forest change, including wind and bark beetle disturbances, using the process-based simulation model iLand in a protected landscape in the northern Alps (Berchtesgaden National Park, Germany), asking: (1) How do understory plant communities respond to 21st-century change in a topographically complex mountain landscape, representing a hotspot of plant species richness? (2) How important are climatic changes (i.e., direct climate effects) versus forest structure and composition changes (i.e., indirect climate effects and recovery from past land use) in driving understory responses at landscape scales? Stacked individual species distribution models fit with climate, forest, and soil predictors (248 species currently present in the landscape, derived from 150 field plots stratified by elevation and forest development, overall area under the receiving operator characteristic curve = 0.86) were driven with projected climate (RCP4.5 and RCP8.5) and modeled forest variables to predict plant community change. Nearly all species persisted in the landscape in 2050, but on average 8% of the species pool was lost by the end of the century. By 2100, landscape mean species richness and understory cover declined (-13% and -8%, respectively), warm-adapted species increasingly dominated plant communities (i.e., thermophilization, +12%), and plot-level turnover was high (62%). Subalpine forests experienced the greatest richness declines (-16%), most thermophilization (+17%), and highest turnover (67%), resulting in plant community homogenization across elevation zones. Climate rather than forest change was the dominant driver of understory responses. The magnitude of unabated 21st-century change is likely to erode plant diversity in a species richness hotspot, calling for stronger conservation and climate mitigation efforts.

Regeneration in European beech forests after drought: the effects of microclimate, deadwood and browsing.

With progressing climate change, increasing weather extremes will endanger tree regeneration. Canopy openings provide light for tree establishment, but also reduce the microclimatic buffering effect of forests. Thus, disturbances can have both positive and negative impacts on tree regeneration. In 2015, three years before an extreme drought episode hit Central Europe, we established a manipulation experiment with a factorial block design in European beech (Fagus sylvatica L.)-dominated forests. At five sites located in southeastern Germany, we conducted three censuses of tree regeneration after implementing two different canopy disturbances (aggregated and distributed canopy openings), and four deadwood treatments (retaining downed, standing, downed + standing deadwood and removing all deadwood), as well as in one untreated control plot. In addition, we measured understory light levels and recorded local air temperature and humidity over five years. We (i) tested the effects of experimental disturbance and deadwood treatments on regeneration and (ii) identified the drivers of regeneration density as well as seedling species and structural diversity. Regeneration density increased over time. Aggregated canopy openings supported species and structural diversity, but reduced regeneration density. Tree regeneration was positively associated with understory light levels, while maximum vapor pressure deficit influenced tree regeneration negatively. Deadwood and browsing impacts on regeneration varied and were inconclusive. Our study indicates that despite the drought episode regeneration in beech-dominated forests persisted under moderately disturbed canopies. However, the positive effect of increased light availability on tree regeneration might have been offset by harsher microclimate after canopies have been disturbed. Supplementary Information: The online version contains supplementary material available at 10.1007/s10342-022-01520-1.

Assessing the Economic Resilience of Different Management Systems to Severe Forest Disturbance.

Given the drastic changes in the environment, resilience is a key focus of ecosystem management. Yet, the quantification of the different dimensions of resilience remains challenging, particularly for long-lived systems such as forests. Here we present an analytical framework to study the economic resilience of different forest management systems, focusing on the rate of economic recovery after severe disturbance. Our framework quantifies the post-disturbance gain in the present value of a forest relative to a benchmark system as an indicator of economic resilience. Forest values and silvicultural interventions were determined endogenously from an optimization model and account for risks affecting tree survival. We consider the effects of differences in forest structure and tree growth post disturbance on economic resilience. We demonstrate our approach by comparing the economic resilience of continuous cover forestry against a clear fell system for typical conditions in Central Europe. Continuous cover forestry had both higher economic return and higher economic resilience than the clear fell system. The economic recovery from disturbance in the continuous cover system was between 18.2 and 51.5% faster than in the clear fell system, resulting in present value gains of between 1733 and 4535 € ha-1. The advantage of the continuous cover system increased with discount rate and stand age, and was driven by differences in both stand structure and economic return. We conclude that continuous cover systems can help to address the economic impacts of increasing disturbances in forest management.

A balancing act: Principles, criteria and indicator framework to operationalize social-ecological resilience of forests.

Against a background of intensifying climate-induced disturbances, the need to enhance the resilience of forests and forest management is gaining urgency. In forest management, multiple trade-offs exist between different demands as well as across and within temporal and spatial scales. However, methods to assess resilience that consider these trade-offs are presently lacking. Here we propose a hierarchical framework of principles, criteria, and indicators to assess the resilience of a social-ecological system by focusing on the mechanisms behind resilience. This hierarchical framework balances trade-offs between mechanisms, different parts of the social-ecological system, ecosystem services, and spatial as well as temporal scales. The framework was developed to be used in a participatory manner in forest management planning. It accounts for the major parts of the forest-related social-ecological system and considers the multiple trade-offs involved. We demonstrate the utility of the framework by applying it to a landscape dominated by Norway spruce (Picea abies (L.) Karst.) in Central Europe, managed for three different management goals. The framework highlights how forest resilience varies with the pursued management goals and related management strategies. The framework is flexible and can be applied to various forest management contexts as part of a participatory process with stakeholders. It thus is an important step towards operationalizing social-ecological resilience in forest management systems.

Significant increase in natural disturbance impacts on European forests since 1950.

Over the last decades, the natural disturbance is increasingly putting pressure on European forests. Shifts in disturbance regimes may compromise forest functioning and the continuous provisioning of ecosystem services to society, including their climate change mitigation potential. Although forests are central to many European policies, we lack the long-term empirical data needed for thoroughly understanding disturbance dynamics, modeling them, and developing adaptive management strategies. Here, we present a unique database of >170,000 records of ground-based natural disturbance observations in European forests from 1950 to 2019. Reported data confirm a significant increase in forest disturbance in 34 European countries, causing on an average of 43.8 million m3 of disturbed timber volume per year over the 70-year study period. This value is likely a conservative estimate due to under-reporting, especially of small-scale disturbances. We used machine learning techniques for assessing the magnitude of unreported disturbances, which are estimated to be between 8.6 and 18.3 million m3 /year. In the last 20 years, disturbances on average accounted for 16% of the mean annual harvest in Europe. Wind was the most important disturbance agent over the study period (46% of total damage), followed by fire (24%) and bark beetles (17%). Bark beetle disturbance doubled its share of the total damage in the last 20 years. Forest disturbances can profoundly impact ecosystem services (e.g., climate change mitigation), affect regional forest resource provisioning and consequently disrupt long-term management planning objectives and timber markets. We conclude that adaptation to changing disturbance regimes must be placed at the core of the European forest management and policy debate. Furthermore, a coherent and homogeneous monitoring system of natural disturbances is urgently needed in Europe, to better observe and respond to the ongoing changes in forest disturbance regimes.

Increasing aridity causes larger and more severe forest fires across Europe.

Area burned has decreased across Europe in recent decades. This trend may, however, reverse under ongoing climate change, particularly in areas not limited by fuel availability (i.e. temperate and boreal forests). Investigating a novel remote sensing dataset of 64,448 fire events that occurred across Europe between 1986 and 2020, we find a power-law relationship between maximum fire size and area burned, indicating that large fires contribute disproportionally to fire activity in Europe. We further show a robust positive correlation between summer vapor pressure deficit and both maximum fire size (R2  = .19) and maximum burn severity (R2  = .12). Europe's fire regimes are thus highly sensitive to changes in future climate, with the probability for extreme fires more than doubling by the end of the century. Our results suggest that climate change will challenge current fire management approaches and could undermine the ability of Europe's forests to provide ecosystem services to society.

A climate risk analysis of Earth's forests in the 21st century.

Earth's forests harbor extensive biodiversity and are currently a major carbon sink. Forest conservation and restoration can help mitigate climate change; however, climate change could fundamentally imperil forests in many regions and undermine their ability to provide such mitigation. The extent of climate risks facing forests has not been synthesized globally nor have different approaches to quantifying forest climate risks been systematically compared. We combine outputs from multiple mechanistic and empirical approaches to modeling carbon, biodiversity, and disturbance risks to conduct a synthetic climate risk analysis for Earth's forests in the 21st century. Despite large uncertainty in most regions we find that some forests are consistently at higher risk, including southern boreal forests and those in western North America and parts of the Amazon.

Post-disturbance reorganization of forest ecosystems in a changing world.

Forest ecosystems are strongly impacted by continuing climate change and increasing disturbance activity, but how forest dynamics will respond remains highly uncertain. Here, we argue that a short time window after disturbance (i.e., a discrete event that disrupts prevailing ecosystem structure and composition and releases resources) is pivotal for future forest development. Trees that establish during this reorganization phase can shape forest structure and composition for centuries, providing operational early indications of forest change. While forest change has been fruitfully studied through a lens of resilience, profound ecological changes can be masked by a resilience versus regime shift dichotomy. We present a framework for characterizing the full spectrum of change after disturbance, analyzing forest reorganization along dimensions of forest structure (number, size, and spatial arrangement of trees) and composition (identity and diversity of tree species). We propose four major pathways through which forest cover can persist but reorganize following disturbance: resilience (no change in structure and composition), restructuring (structure changes but composition does not), reassembly (composition changes but structure does not), and replacement (structure and composition both change). Regime shifts occur when vegetation structure and composition are altered so profoundly that the emerging trajectory leads to nonforest. We identify fundamental processes underpinning forest reorganization which, if disrupted, deflect ecosystems away from resilience. To understand and predict forest reorganization, assessing these processes and the traits modulating them is crucial. A new wave of experiments, measurements, and models emphasizing the reorganization phase will further the capacity to anticipate future forest dynamics.

Accelerating Mountain Forest Dynamics in the Alps.

Climate change alters forest development pathways, with consequences for ecosystem services and biodiversity. As the rate of warming increases, ecosystem change is expected to accelerate. However, ecosystem dynamics can have many causes unrelated to climate (for example, disturbance and stand development legacies). The compound effects of multiple drivers remain largely unclear. Here, we assessed forest dynamics over 28 years at Berchtesgaden National Park (BGNP), Germany, quantifying the spatiotemporal patterns and unraveling the drivers of forest change. We analyzed high-density forest inventory data, consisting of three consecutive censuses of 3759 permanent sample plots (132,866 tree records in total). We used semi-variograms to analyze spatial patterns of change, and boosted regression trees to quantify the effect of 30 covariates on changes in nine indicators of forest structure and composition. Over the 28 years investigated, the forests of BGNP were becoming denser, structurally more complex, and more species rich. Changes in forest structure were more pronounced and spatially correlated on the landscape than changes in tree species composition. Change rates of all indicators increased over time, signifying an acceleration of forest dynamics since the 1980s. Legacies and climate were the most important drivers of change, but had diverging impacts. Although forest change accelerated with increasing temperature, high legacy levels typical for late development stages dampened it. We here provide evidence for accelerating forest dynamics in mountain forests of the Alps, with potentially far-reaching consequences for biodiversity and ecosystem processes. We highlight that unmanaged forest development toward old-growth conditions could counteract climate-mediated acceleration of forest change. Supplementary Information: The online version contains supplementary material available at 10.1007/s10021-021-00674-0.

Will forest dynamics continue to accelerate throughout the 21st century in the Northern Alps?

Observational evidence suggests that forests in the Northern Alps are changing at an increasing rate as a consequence of climate change. Yet, it remains unclear whether the acceleration of forest change will continue in the future, or whether downregulating feedbacks will eventually decouple forest dynamics from climate change. Here we studied future forest dynamics at Berchtesgaden National Park, Germany by means of a process-based forest landscape model, simulating an ensemble of 22 climate projections until the end of the 21st century. Our objectives were (i) to assess whether the observed acceleration of forest dynamics will continue in the future, (ii) to analyze how uncertainty in future climate translates to variation in future forest disturbance, structure, and composition, and (iii) to determine the main drivers of future forest dynamics. We found that forest dynamics continue to accelerate in the coming decades, with a trend towards denser, structurally more complex and more species rich forests. However, changes in forest structure leveled off in the second half of the 21st century regardless of climate scenario. In contrast, climate scenarios caused trajectories of tree species change to diverge in the second half of the 21st century, with stabilization under RCP 2.6 and RCP 4.5 scenarios and accelerated loss of conifers under RCP 8.5. Disturbance projections were 3 to 20 times more variable than future climate, whereas projected future forest structure and composition varied considerably less than climate. Indirect effects of climate change via alterations of the disturbance regime had a stronger impact on future forest dynamics than direct effects. Our findings suggest that dampening feedbacks within forest dynamics will decelerate forest change in the second half of the 21st century. However, warming beyond the levels projected under RCP 4.5 might profoundly alter future forest disturbance and composition, challenging conservation efforts and ecosystem service supply.

Using historical spy satellite photographs and recent remote sensing data to identify high-conservation-value forests.

High-conservation-value forests (HCVFs) are critically important for biodiversity and ecosystem service provisioning, but they face many threats. Where systematic HCVF inventories are missing, such as in parts of Eastern Europe, these forests remain largely unacknowledged and therefore often unprotected. We devised a novel, transferable approach for detecting HCVFs based on integrating historical spy satellite images, contemporary remote sensing data (Landsat), and information on current potential anthropogenic pressures (e.g., road infrastructure, population density, demand for fire wood, terrain). We applied the method to the Romanian Carpathians, for which we mapped forest continuity (1955-2019), canopy structural complexity, and anthropogenic pressures. We identified 738,000 ha of HCVF. More than half of this area was identified as susceptible to current anthropogenic pressures and lacked formal protection. By providing a framework for broad-scale HCVF monitoring, our approach facilitates integration of HCVF into forest conservation and management. This is urgently needed to achieve the goals of the European Union's Biodiversity Strategy to maintain valuable forest ecosystems.

Uso de Fotografías Históricas de Satélites Espía y Datos Recientes de Telemetría para Identificar Bosques de Alto Valor para la Conservación Resumen Los bosques de alto valor para la conservación (BAVC) tienen una importancia crítica para el suministro de servicios ambientales y biodiversidad pero enfrentan muchas amenazas. En donde hacen falta inventarios sistemáticos de los BAVC, como en partes del este de Europa, estos bosques siguen siendo ignorados y por lo tanto carecen de protección. Diseñamos una estrategia novedosa y transferible para la detección de BAVC con base en la integración de imágenes de satélites espía, datos contemporáneos de telemetría (Landsat) e información sobre las presiones antropogénicas actuales (p. ej.: infraestructura vial, densidad poblacional, demanda de leña, terreno). Aplicamos el método en los Cárpatos rumanos, para los cuales mapeamos la continuidad forestal (1955 - 2019), la complejidad estructural del dosel y las presiones antropogénicas. Identificamos 738,000 ha de BAVC. Más de la mitad de esta área fue identificada como susceptible a las actuales presiones antropogénicas y además carecía de protección formal. Mediante la aportación de un marco de trabajo para el monitoreo a escala amplia de los BAVC, nuestra estrategia facilita la integración de los BAVC dentro de la gestión y conservación de los bosques. Lo último es una necesidad urgente para alcanzar las metas de la Estrategia de Biodiversidad de la Unión Europea para mantener los ecosistemas boscosos valiosos.

Arthropod dark taxa provide new insights into diversity responses to bark beetle infestations.

Natural disturbances are increasing around the globe, also impacting protected areas. Although previous studies have indicated that natural disturbances result in mainly positive effects on biodiversity, these analyses mostly focused on a few well established taxonomic groups, and thus uncertainty remains regarding the comprehensive impact of natural disturbances on biodiversity. Using Malaise traps and meta-barcoding, we studied a broad range of arthropod taxa, including dark and cryptic taxa, along a gradient of bark beetle disturbance severities in five European national parks. We identified order-level community thresholds of disturbance severity and classified barcode index numbers (BINs; a cluster system for DNA sequences, where each cluster corresponds to a species) as negative or positive disturbance indicators. Negative indicator BINs decreased above thresholds of low to medium disturbance severity (20%-30% of trees killed), whereas positive indicator BINs benefited from high disturbance severity (76%-98%). BINs allocated to a species name contained nearly as many positive as negative disturbance indicators, but dark and cryptic taxa, particularly Diptera and Hymenoptera in our data, contained higher numbers of negative disturbance indicator BINs. Analyses of changes in the richness of BINs showed variable responses of arthropods to disturbance severity at lower taxonomic levels, whereas no significant signal was detected at the order level due to the compensatory responses of the underlying taxa. We conclude that the analyses of dark taxa can offer new insights into biodiversity responses to disturbances. Our results suggest considerable potential for forest management to foster arthropod diversity, for example by maintaining both closed-canopy forests (>70% cover) and open forests (<30% cover) on the landscape.

The evolution, complexity and diversity of models of long-term forest dynamics.

To assess the impacts of climate change on vegetation from stand to global scales, models of forest dynamics that include tree demography are needed. Such models are now available for 50 years, but the currently existing diversity of model formulations and its evolution over time are poorly documented. This hampers systematic assessments of structural uncertainties in model-based studies.We conducted a meta-analysis of 28 models, focusing on models that were used in the past five years for climate change studies. We defined 52 model attributes in five groups (basic assumptions, growth, regeneration, mortality and soil moisture) and characterized each model according to these attributes. Analyses of model complexity and diversity included hierarchical cluster analysis and redundancy analysis.Model complexity evolved considerably over the past 50 years. Increases in complexity were largest for growth processes, while complexity of modelled establishment processes increased only moderately. Model diversity was lowest at the global scale, and highest at the landscape scale. We identified five distinct clusters of models, ranging from very simple models to models where specific attribute groups are rendered in a complex manner and models that feature high complexity across all attributes.Most models in use today are not balanced in the level of complexity with which they represent different processes. This is the result of different model purposes, but also reflects legacies in model code, modelers' preferences, and the 'prevailing spirit of the epoch'. The lack of firm theories, laws and 'first principles' in ecology provides high degrees of freedom in model development, but also results in high responsibilities for model developers and the need for rigorous model evaluation. Synthesis. The currently available model diversity is beneficial: convergence in simulations of structurally different models indicates robust projections, while convergence of similar models may convey a false sense of certainty. The existing model diversity-with the exception of global models-can be exploited for improved projections based on multiple models. We strongly recommend balanced further developments of forest models that should particularly focus on establishment and mortality processes, in order to provide robust information for decisions in ecosystem management and policymaking.

The contribution of insects to global forest deadwood decomposition.

The amount of carbon stored in deadwood is equivalent to about 8 per cent of the global forest carbon stocks1. The decomposition of deadwood is largely governed by climate2-5 with decomposer groups-such as microorganisms and insects-contributing to variations in the decomposition rates2,6,7. At the global scale, the contribution of insects to the decomposition of deadwood and carbon release remains poorly understood7. Here we present a field experiment of wood decomposition across 55 forest sites and 6 continents. We find that the deadwood decomposition rates increase with temperature, and the strongest temperature effect is found at high precipitation levels. Precipitation affects the decomposition rates negatively at low temperatures and positively at high temperatures. As a net effect-including the direct consumption by insects and indirect effects through interactions with microorganisms-insects accelerate the decomposition in tropical forests (3.9% median mass loss per year). In temperate and boreal forests, we find weak positive and negative effects with a median mass loss of 0.9 per cent and -0.1 per cent per year, respectively. Furthermore, we apply the experimentally derived decomposition function to a global map of deadwood carbon synthesized from empirical and remote-sensing data, obtaining an estimate of 10.9 ± 3.2 petagram of carbon per year released from deadwood globally, with 93 per cent originating from tropical forests. Globally, the net effect of insects may account for 29 per cent of the carbon flux from deadwood, which suggests a functional importance of insects in the decomposition of deadwood and the carbon cycle.