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.

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.

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.

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.

Widespread regeneration failure in forests of Greater Yellowstone under scenarios of future climate and fire.

Changing climate and disturbance regimes are increasingly challenging the resilience of forest ecosystems around the globe. A powerful indicator for the loss of resilience is regeneration failure, that is, the inability of the prevailing tree species to regenerate after disturbance. Regeneration failure can result from the interplay among disturbance changes (e.g., larger and more frequent fires), altered climate conditions (e.g., increased drought), and functional traits (e.g., method of seed dispersal). This complexity makes projections of regeneration failure challenging. Here we applied a novel simulation approach assimilating data-driven fire projections with vegetation responses from process modeling by means of deep neural networks. We (i) quantified the future probability of regeneration failure; (ii) identified spatial hotspots of regeneration failure; and (iii) assessed how current forest types differ in their ability to regenerate under future climate and fire. We focused on the Greater Yellowstone Ecosystem (2.9 × 106  ha of forest) in the Rocky Mountains of the USA, which has experienced large wildfires in the past and is expected to undergo drastic changes in climate and fire in the future. We simulated four climate scenarios until 2100 at a fine spatial grain (100 m). Both wildfire activity and unstocked forest area increased substantially throughout the 21st century in all simulated scenarios. By 2100, between 28% and 59% of the forested area failed to regenerate, indicating considerable loss of resilience. Areas disproportionally at risk occurred where fires are not constrained by topography and in valleys aligned with predominant winds. High-elevation forest types not adapted to fire (i.e., Picea engelmannii-Abies lasiocarpa as well as non-serotinous Pinus contorta var. latifolia forests) were especially vulnerable to regeneration failure. We conclude that changing climate and fire could exceed the resilience of forests in a substantial portion of Greater Yellowstone, with profound implications for carbon, biodiversity, and recreation.

Do bark beetle outbreaks amplify or dampen future bark beetle disturbances in Central Europe?

Bark beetle outbreaks have intensified in many forests around the globe in recent years. Yet, the legacy of these disturbances for future forest development remains unclear. Bark beetle disturbances are expected to increase further because of climate change. Consequently, feedbacks within the disturbance regime are of growing interest, for example, whether bark beetle outbreaks are amplifying future bark beetle activity (through the initiation of an even-aged cohort of trees) or dampening it (through increased structural and compositional diversity).We studied bark beetle-vegetation-climate interactions in the Bavarian Forest National Park (Germany), an area characterised by unprecedented bark beetle activity in the recent past. We simulated the effect of future bark beetle outbreaks on forest structure and composition and analysed how disturbance-mediated forest dynamics influence future bark beetle activity under different scenarios of climate change. We used process-based simulation modelling in combination with machine learning to disentangle the long-term interactions between vegetation, climate and bark beetles at the landscape scale.Disturbances by the European spruce bark beetle were strongly amplified by climate change, increasing between 59% and 221% compared to reference climate. Bark beetle outbreaks reduced the dominance of Norway spruce (Picea abies (L.) Karst.) on the landscape, increasing compositional diversity. Disturbances decreased structural diversity within stands (α diversity) and increased structural diversity between stands (ß diversity). Overall, disturbance-mediated changes in forest structure and composition dampened future disturbance activity (a reduction of up to -67%), but were not able to fully compensate for the amplifying effect of climate change. Synthesis. Our findings indicate that the recent disturbance episode at the Bavarian Forest National Park was caused by a convergence of highly susceptible forest structures with climatic conditions favourable for bark beetle outbreaks. While future climate is increasingly conducive to massive outbreaks, the emerging landscape structure is less and less likely to support them. This study improves our understanding of the long-term legacies of ongoing bark beetle disturbances in Central Europe. It indicates that increased diversity provides an important dampening feedback, and suggests that preventing disturbances or homogenizing post-disturbance forests could elevate the future susceptibility to large-scale bark beetle outbreaks.

Simulating forest resilience: A review.

AIM: Simulation models are important tools for quantifying the resilience (i.e., persistence under changed environmental conditions) of forest ecosystems to global change. We synthesized the modelling literature on forest resilience, summarizing common models and applications in resilience research, and scrutinizing the implementation of important resilience mechanisms in these models. Models applied to assess resilience are highly diverse, and our goal was to assess how well they account for important resilience mechanisms identified in experimental and empirical research. LOCATION: Global. TIME PERIOD: 1994 to 2019. MAJOR TAXA STUDIED: Trees. METHODS: We reviewed the forest resilience literature using online databases, selecting 119 simulation modelling studies for further analysis. We identified a set of resilience mechanisms from the general resilience literature and analysed models for their representation of these mechanisms. Analyses were grouped by investigated drivers (resilience to what) and responses (resilience of what), as well as by the type of model being used. RESULTS: Models used to study forest resilience varied widely, from analytical approaches to complex landscape simulators. The most commonly addressed questions were associated with resilience of forest cover to fire. Important resilience mechanisms pertaining to regeneration, soil processes, and disturbance legacies were explicitly simulated in only 34 to 46% of the model applications. MAIN CONCLUSIONS: We found a large gap between processes identified as underpinning forest resilience in the theoretical and empirical literature, and those represented in models used to assess forest resilience. Contemporary forest models developed for other goals may be poorly suited for studying forest resilience during an era of accelerating change. Our results highlight the need for a new wave of model development to enhance understanding of and management for resilient forests.

The influence of climate change and canopy disturbances on landslide susceptibility in headwater catchments.

Forests have an important regulating function on water runoff and the occurrence of shallow landslides. Their structure and composition directly influence the risk of hydrogeomorphic processes, like floods with high sediment transport or debris flows. Climate change is substantially altering forest ecosystems, and for Central Europe an increase in natural disturbances from wind and insect outbreaks is expected for the future. How such changes impact the regulating function of forest ecosystems remains unclear. By combining methods from forestry, hydrology and geotechnical engineering we investigated possible effects of changing climate and disturbance regimes on shallow landslides. We simulated forest landscapes in two headwater catchments in the Eastern Alps of Austria under four different future climate scenarios over 200 years. Our results indicate that climate-mediated changes in forest dynamics can substantially alter the protective function of forest ecosystems. Climate change generally increased landslide risk in our simulations. Only when future warming coincided with drying landslide risk decreased relative to historic conditions. In depth analyses showed that an important driver of future landslide risk was the simulated vegetation composition. Trajectories away from flat rooting Norway spruce (Picea abies (L.) Karst.) forests currently dominating the system towards an increasing proportion of tree species with heart and taproot systems, increased root cohesion and reduced the soil volume mobilized in landslides. Natural disturbances generally reduced landslide risk in our simulations, with the positive effect of accelerated tree species change and increasing root cohesion outweighing a potential negative effect of disturbances on the water cycle. We conclude that while the efficacy of green infrastructure such as protective forests could be substantially reduced by climate change, such systems also have a strong inherent ability to adapt to changing conditions. Forest management should foster this adaptive capacity to strengthen the protective function of forests also under changing environmental conditions.

Climate change causes critical transitions and irreversible alterations of mountain forests.

Mountain forests are at particular risk of climate change impacts due to their temperature limitation and high exposure to warming. At the same time, their complex topography may help to buffer the effects of climate change and create climate refugia. Whether climate change can lead to critical transitions of mountain forest ecosystems and whether such transitions are reversible remain incompletely understood. We investigated the resilience of forest composition and size structure to climate change, focusing on a mountain forest landscape in the Eastern Alps. Using the individual-based forest landscape model iLand, we simulated ecosystem responses to a wide range of climatic changes (up to a 6°C increase in mean annual temperature and a 30% reduction in mean annual precipitation), testing for tipping points in vegetation size structure and composition under different topography scenarios. We found that at warming levels above +2°C a threshold was crossed, with the system tipping into an alternative state. The system shifted from a conifer-dominated landscape characterized by large trees to a landscape dominated by smaller, predominantly broadleaved trees. Topographic complexity moderated climate change impacts, smoothing and delaying the transitions between alternative vegetation states. We subsequently reversed the simulated climate forcing to assess the ability of the landscape to recover from climate change impacts. The forest landscape showed hysteresis, particularly in scenarios with lower precipitation. At the same mean annual temperature, equilibrium vegetation size structure and species composition differed between warming and cooling trajectories. Here we show that even moderate warming corresponding to current policy targets could result in critical transitions of forest ecosystems and highlight the importance of topographic complexity as a buffering agent. Furthermore, our results show that overshooting ambitious climate mitigation targets could be dangerous, as ecological impacts can be irreversible at millennial time scales once a tipping point has been crossed.

Norway spruce at the trailing edge: the effect of landscape configuration and composition on climate resilience.

CONTEXT: Norway spruce (Picea abies) is one of the most widespread tree species in Europe's forests. Due to its high economic value it has been strongly favored by management, especially at the trailing edge of its natural distribution. However, disturbances from wind and bark beetles are increasingly impacting these forests, and their resilience under climate change has been called into question recently. OBJECTIVES: We quantified the effects of landscape configuration and composition on (1) the risk from natural disturbances, and (2) on the overall resilience of Norway spruce to changing climate at the trailing edge. METHODS: We simulated the dynamics of a 9183 ha forest landscape in Eastern Austria over 190 years. We used the simulation model iLand to experimentally study a wide range of landscape compositions and configurations under five different climate scenarios. RESULTS: Natural disturbances increased considerably under all future climate scenarios. Dispersing Norway spruce throughout the landscape in mixed stands resulted in the highest levels of climate resilience. Reducing the percentage of Norway spruce on the landscape increased the resilience of the remaining Norway spruce trees, yet landscape configuration generally had a stronger effect on resilience than composition. CONCLUSIONS: The resilience of Norway spruce at the trailing edge of its distribution is challenged by climate change, and considerable efforts are needed to sustain these ecosystems. While currently discussed adaptation measures focus largely on the stand level, we show that modifying landscape composition and configuration can be used to foster Norway spruce resilience while maintaining socio-economically relevant proportions of Norway spruce.

Reducing rotation age to address increasing disturbances in Central Europe: Potential and limitations.

Forest disturbance regimes are intensifying in many parts of the globe. In order to mitigate disturbance impacts a number of management responses have been proposed, yet their effectiveness in addressing changing disturbance regimes remains largely unknown. The strong positive relationship between forest age and the vulnerability to disturbances such as windthrows and bark beetle infestations suggests that a reduced rotation length can be a potent means for mitigating the impacts of natural disturbances. However, disturbance mitigation measures such as shortened rotation lengths (SRL) can also have undesired consequences on ecosystem services and biodiversity, which need to be considered in their application. Here, we used the process-based landscape and disturbance model iLand to investigate the effects of SRL on the vulnerability of a 16,000 ha forest landscape in Central Europe to wind and bark beetle disturbances. We experimentally reduced the current rotation length (between 100 and 115 years) by up to -40% in 10% increments, and studied effects on disturbance dynamics under current and future climate conditions over a 200-year simulation period. Simultaneously, we quantified the collateral effects of SRL on forest carbon stocks and indicators of biodiversity. Shortening the rotation length by 40% decreased disturbances by 14%. This effect was strongly diminished under future climate change, reducing the mitigating effect of shortened rotation to < 6%. Collateral effects were severe in the initial decades after implementation: Reducing the rotation length by 40% caused a spike in harvested timber volume (+ 92%), decreased total forest carbon storage by 6% and reduced the number of large trees on the landscape by 20%. The long-term effects of SRL were less pronounced. At the same time, SRL caused an increase in tree species diversity. Shortening rotation length can reduce the impact of wind and bark beetle disturbances, but the overall efficiency of the measure is limited and decreases under climate change. Given the potential for undesired collateral effects we conclude that a reduction of the rotation length is no panacea for managing increasing disturbances, and should be applied in combination with other management measures reducing risks and fostering resilience.

Can wildland fire management alter 21st-century subalpine fire and forests in Grand Teton National Park, Wyoming, USA?

In subalpine forests of the western United States that historically experienced infrequent, high-severity fire, whether fire management can shape 21st-century fire regimes and forest dynamics to meet natural resource objectives is not known. Managed wildfire use (i.e., allowing lightning-ignited fires to burn when risk is low instead of suppressing them) is one approach for maintaining natural fire regimes and fostering mosaics of forest structure, stand age, and tree-species composition, while protecting people and property. However, little guidance exists for where and when this strategy may be effective with climate change. We simulated most of the contiguous forest in Grand Teton National Park, Wyoming, USA to ask: (1) how would subalpine fires and forest structure be different if fires had not been suppressed during the last three decades? And (2) what is the relative influence of climate change vs. fire management strategy on future fire and forests? We contrasted fire and forests from 1989 to 2098 under two fire management scenarios (managed wildfire use and fire suppression), two general circulation models (CNRM-CM5 and GFDL-ESM2M), and two representative concentration pathways (8.5 and 4.5). We found little difference between management scenarios in the number, size, or severity of fires during the last three decades. With 21st-century warming, fire activity increased rapidly, particularly after 2050, and followed nearly identical trajectories in both management scenarios. Area burned per year between 2018 and 2099 was 1,700% greater than in the last three decades (1989-2017). Large areas of forest were abruptly lost; only 65% of the original 40,178 ha of forest remained by 2098. However, forests stayed connected and fuels were abundant enough to support profound increases in burning through this century. Our results indicate that strategies emphasizing managed wildfire use, rather than suppression, will not alter climate-induced changes to fire and forests in subalpine landscapes of western North America. This suggests that managers may continue to have flexibility to strategically suppress subalpine fires without concern for long-term consequences, in distinct contrast with dry conifer forests of the Rocky Mountains and mixed conifer forest of California where maintaining low fuel loads is essential for sustaining frequent, low-severity surface fire regimes.

Spatial configuration matters when removing windfelled trees to manage bark beetle disturbances in Central European forest landscapes.

Windfelled Norway spruce (Picea abies) trees play a crucial role in triggering large-scale outbreaks of the European spruce bark beetle Ips typographus. Outbreak management therefore strives to remove windfelled trees to reduce the risk of outbreaks, a measure referred to as sanitation logging (SL). Although this practice has been traditionally applied, its efficiency in preventing outbreaks remains poorly understood. We used the landscape simulation model iLand to investigate the effects of different spatial configurations and intensities of SL of windfelled trees on the subsequent disturbance by bark beetles. We studied differences between SL applied evenly across the landscape, focused on the vicinity of roads (scenario of limited logging resources) and concentrated in a contiguous block (scenario of spatially diversified management objectives). We focused on a 16 050 ha forest landscape in Central Europe. The removal of >80% of all windfelled trees is required to substantially reduce bark beetle disturbances. Focusing SL on the vicinity of roads created a "fire break effect" on bark beetle spread, and was moderately efficient in reducing landscape-scale bark beetle disturbance. Block treatments substantially reduced outbreaks in treated areas. Leaving parts of the landscape untreated (e.g., conservation areas) had no significant amplifying effect on outbreaks in managed areas. Climate change increased bark beetle disturbances and reduced the effect of SL. Our results suggest that past outbreak management methods will not be sufficient to counteract climate-mediated increases in bark beetle disturbance.

Harnessing Deep Learning in Ecology: An Example Predicting Bark Beetle Outbreaks.

Addressing current global challenges such as biodiversity loss, global change, and increasing demands for ecosystem services requires improved ecological prediction. Recent increases in data availability, process understanding, and computing power are fostering quantitative approaches in ecology. However, flexible methodological frameworks are needed to utilize these developments towards improved ecological prediction. Deep learning is a rapidly evolving branch of machine learning, yet has received only little attention in ecology to date. It refers to the training of deep neural networks (DNNs), i.e. artificial neural networks consisting of many layers and a large number of neurons. We here provide a reproducible example (including code and data) of designing, training, and applying DNNs for ecological prediction. Using bark beetle outbreaks in conifer-dominated forests as an example, we show that DNNs are well able to predict both short-term infestation risk at the local scale and long-term outbreak dynamics at the landscape level. We furthermore highlight that DNNs have better overall performance than more conventional approaches to predicting bark beetle outbreak dynamics. We conclude that DNNs have high potential to form the backbone of a comprehensive disturbance forecasting system. More broadly, we argue for an increased utilization of the predictive power of DNNs for a wide range of ecological problems.

A scalable model of vegetation transitions using deep neural networks.

In times of rapid global change, anticipating vegetation changes and assessing their impacts is of key relevance to managers and policy makers. Yet, predicting vegetation dynamics often suffers from an inherent scale mismatch, with abundant data and process understanding being available at a fine spatial grain, but the relevance for decision-making is increasing with spatial extent.We present a novel approach for scaling vegetation dynamics (SVD), using deep learning to predict vegetation transitions. Vegetation is discretized into a large number (103-106) of potential states based on its structure, composition and functioning. Transition probabilities between states are estimated via a deep neural network (DNN) trained on observed or simulated vegetation transitions in combination with environmental variables. The impact of vegetation transitions on important ecological indicators is quantified by probabilistically linking attributes such as carbon storage and biodiversity to vegetation states.Here, we describe the SVD approach and present results of applying the framework in a meta-modelling context. We trained a DNN using simulations of a process-based forest landscape model for a complex mountain forest landscape under different climate scenarios. Subsequently, we evaluated the ability of SVD to project long-term vegetation dynamics and the resulting changes in forest carbon storage and biodiversity. SVD captured spatial (e.g. elevational gradients) and temporal (e.g. species succession) patterns of vegetation dynamics well, and responded realistically to changing environmental conditions. In addition, we tested the computational efficiency of the approach, highlighting the utility of SVD for country- to continental scale applications. SVD is the-to our knowledge-first vegetation model harnessing deep neural networks. The approach has high predictive accuracy and is able to generalize well beyond training data. SVD was designed to run on widely available input data (e.g. vegetation states defined from remote sensing, gridded global climate datasets) and exceeds the computational performance of currently available highly optimized landscape models by three to four orders of magnitude. We conclude that SVD is a promising approach for combining detailed process knowledge on fine-grained ecosystem processes with the increasingly available big ecological datasets for improved large-scale projections of vegetation dynamics.

Tree mortality submodels drive simulated long-term forest dynamics: assessing 15 models from the stand to global scale.

Models are pivotal for assessing future forest dynamics under the impacts of changing climate and management practices, incorporating representations of tree growth, mortality, and regeneration. Quantitative studies on the importance of mortality submodels are scarce. We evaluated 15 dynamic vegetation models (DVMs) regarding their sensitivity to different formulations of tree mortality under different degrees of climate change. The set of models comprised eight DVMs at the stand scale, three at the landscape scale, and four typically applied at the continental to global scale. Some incorporate empirically derived mortality models, and others are based on experimental data, whereas still others are based on theoretical reasoning. Each DVM was run with at least two alternative mortality submodels. Model behavior was evaluated against empirical time series data, and then, the models were subjected to different scenarios of climate change. Most DVMs matched empirical data quite well, irrespective of the mortality submodel that was used. However, mortality submodels that performed in a very similar manner against past data often led to sharply different trajectories of forest dynamics under future climate change. Most DVMs featured high sensitivity to the mortality submodel, with deviations of basal area and stem numbers on the order of 10-40% per century under current climate and 20-170% under climate change. The sensitivity of a given DVM to scenarios of climate change, however, was typically lower by a factor of two to three. We conclude that (1) mortality is one of the most uncertain processes when it comes to assessing forest response to climate change, and (2) more data and a better process understanding of tree mortality are needed to improve the robustness of simulated future forest dynamics. Our study highlights that comparing several alternative mortality formulations in DVMs provides valuable insights into the effects of process uncertainties on simulated future forest dynamics.

What drives the future supply of regulating ecosystem services in a mountain forest landscape?

Forest ecosystems provide a wide variety of ecosystem services to society. In harsh mountain environments, the regulating services of forests are of particular importance. Managing mountain forests for regulating services is a cost- and labor intensive endeavor. Yet, also unmanaged forests regulate the environment. In the context of evidence-based decision making it is thus important to scrutinize if current management recommendations improve the supply of regulating ecosystem services over unmanaged development trajectories. A further issue complicating decision making in the context of regulating ecosystem services is their high sensitivity to climate change. Climate-mediated increases in natural disturbances, for instance, could strongly reduce the supply of regulating services from forests in the future. Given the profound environmental changes expected for the coming decades it remains unclear whether forest management will still be able to significantly control the future trajectories of mountain forest development, or whether the management effect will be superseded by a much stronger climate and disturbance effect. Here, our objectives were (i) to quantify the future regulating service supply from a 6456 ha landscape in the Stubai valley in Tyrol, Austria, and (ii) to assess the relative importance of management, climate, and natural disturbances on the future supply of regulating ecosystem services. We focused our analysis on climate regulation, water regulation, and erosion regulation, and used the landscape simulation model iLand to quantify their development under different climate scenarios and management strategies. Our results show that unmanaged forests are efficient in providing regulating ecosystem services. Both climate regulation and erosion regulation were higher in unmanaged systems compared to managed systems, while water regulation was slightly enhanced by management. Overall, direct effects of climate change had a stronger influence on the future supply of regulating services than management and natural disturbances. The ability of management to control ecosystem service supply decreased sharply with the severity of future climate change. This finding highlights that forest management could be severely stymied in the future if climate change continues to proceed at its current rate. An improved quantitative understanding of the drivers of future ecosystem service supply is needed to more effectively combine targeted management efforts and natural ecosystem dynamics towards sustaining the benefits society derives from forests in a rapidly changing world.

Trade-offs between temporal stability and level of forest ecosystem services provisioning under climate change.

The ability of forests to continuously provide ecosystem services (ES) is threatened by rapid changes in climate and disturbance regimes. Consequently, these changes present a considerable challenge for forest managers. Management of forests often focuses on maximizing the level of ES provisioning over extended time frames (i.e., rotation periods of more than 100 yr). However, temporal stability is also crucial for many ES, for example, in the context of a steady provisioning of resources to the industry, or the protection of human infrastructure against natural hazards. How temporal stability and the level of ES provisioning are related is of increasing interest, particularly since changing climate and disturbance regimes amplify temporal variability in forest ecosystems. In this simulation study, we investigated whether forest management can simultaneously achieve high levels and temporal stability of ES provisioning. Specifically, we quantified (1) trade-offs between ES stability and level of ES provisioning, and (2) the effect of tree species diversity on ES stability. Simulating a wide range of future climate scenarios and management strategies, we found a negative relationship between temporal stability and level of ES provisioning for timber production, carbon cycling, and site protection in a landscape in the Austrian Alps. Tree species diversity had a predominantly positive effect on ES stability. We conclude that attempts to maximize the level of ES provisioning may increase its temporal variability, and thus threaten the continuity of ES supply. Consequently, considerations of stability need to be more explicitly included in forest management planning under increasingly variable future conditions.

Invasive alien pests threaten the carbon stored in Europe's forests.

Forests mitigate climate change by sequestering large amounts of carbon (C). However, forest C storage is not permanent, and large pulses of tree mortality can thwart climate mitigation efforts. Forest pests are increasingly redistributed around the globe. Yet, the potential future impact of invasive alien pests on the forest C cycle remains uncertain. Here we show that large parts of Europe could be invaded by five detrimental alien pests already under current climate. Climate change increases the potential range of alien pests particularly in Northern and Eastern Europe. We estimate the live C at risk from a potential future invasion as 1027 Tg C (10% of the European total), with a C recovery time of 34 years. We show that the impact of introduced pests could be as severe as the current natural disturbance regime in Europe, calling for increased efforts to halt the introduction and spread of invasive alien species.

It takes a few to tango: changing climate and fire regimes can cause regeneration failure of two subalpine conifers.

Environmental change is accelerating in the 21st century, but how multiple drivers may interact to alter forest resilience remains uncertain. In forests affected by large high-severity disturbances, tree regeneration is a resilience linchpin that shapes successional trajectories for decades. We modeled stands of two widespread western U.S. conifers, Douglas-fir (Pseudotsuga menziesii var. glauca), and lodgepole pine (Pinus contorta var. latifolia), in Yellowstone National Park (Wyoming, USA) to ask (1) What combinations of distance to seed source, fire return interval, and warming-drying conditions cause postfire tree-regeneration failure? (2) If postfire tree regeneration was successful, how does early tree density differ under future climate relative to historical climate? We conducted a stand-level (1 ha) factorial simulation experiment using the individual-based forest process model iLand to identify combinations of fire return interval (11-100 yr), distance to seed source (50-1,000 m), and climate (historical, mid-21st century, late-21st century) where trees failed to regenerate by 30-yr postfire. If regeneration was successful, we compared stand densities between climate periods. Simulated postfire regeneration were surprisingly resilient to changing climate and fire drivers. Douglas-fir regeneration failed more frequently (55%) than lodgepole pine (28% and 16% for non-serotinous and serotinous stands, respectively). Distance to seed source was an important driver of regeneration failure for Douglas-fir and non-serotinous lodgepole pine; regeneration never failed when stands were 50 m from a seed source and nearly always failed when stands were 1 km away. Regeneration of serotinous lodgepole pine only failed when fire return intervals were ≤20 yr and stands were far (1 km) from a seed source. Warming climate increased regeneration success for Douglas-fir but did not affect lodgepole pine. If regeneration was successful, postfire density varied with climate. Douglas-fir and serotinous lodgepole pine regeneration density both increased under 21st-century climate but in response to different climate variables (growing season length vs. cold limitation). Results suggest that, given a warmer future with larger and more frequent fires, a greater number of stands that fail to regenerate after fires combined with increasing density in stands where regeneration is successful could produce a more coarse-grained forest landscape.