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.

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.

Past and future impacts of land-use changes on ecosystem services in Austria.

Environmental and socio-economic developments induce land-use changes with potentially negative impacts on human well-being. To counteract undesired developments, a profound understanding of the complex relationships between drivers, land use, and ecosystem services is needed. Yet, national studies examining extended time periods are still rare. Based on the Special Report on land use, land management and climate change by the Austrian Panel on Climate Change (APCC), we use the Driver-Pressure-State-Impact-Response (DPSIR) framework to (1) identify the main drivers of land-use change, (2) describe past and future land-use changes in Austria between 1950 and 2100, (3) report related impacts on ecosystem services, and (4) discuss management responses. Our findings indicate that socio-economic drivers (e.g., economic growth, political systems, and technological developments) have influenced past land-use changes the most. The intensification of agricultural land use and urban sprawl have primarily led to declining ecosystem services in the lowlands. In mountain regions, the abandonment of mountain grassland has prompted a shift from provisioning to regulating services. However, simulations indicate that accelerating climate change will surpass socio-economic drivers in significance towards the end of this century, particularly in intensively used agricultural areas. Although climate change-induced impacts on ecosystem services remain uncertain, it can be expected that the range of land-use management options will be restricted in the future. Consequently, policymaking should prioritize the development of integrated land-use planning to safeguard ecosystem services, accounting for future environmental and socio-economic uncertainties.

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.

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.

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.

Natural disturbance regimes as a guide for sustainable forest management in Europe.

In Europe, forest management has controlled forest dynamics to sustain commodity production over multiple centuries. Yet over-regulation for growth and yield diminishes resilience to environmental stress as well as threatens biodiversity, leading to increasing forest susceptibility to an array of disturbances. These trends have stimulated interest in alternative management systems, including natural dynamics silviculture (NDS). NDS aims to emulate natural disturbance dynamics at stand and landscape scales through silvicultural manipulations of forest structure and landscape patterns. We adapted a "Comparability Index" (CI) to assess convergence/divergence between natural disturbances and forest management effects. We extended the original CI concept based on disturbance size and frequency by adding the residual structure of canopy trees after a disturbance as a third dimension. We populated the model by compiling data on natural disturbance dynamics and management from 13 countries in Europe, covering four major forest types (i.e., spruce, beech, oak, and pine-dominated forests). We found that natural disturbances are highly variable in size, frequency, and residual structure, but European forest management fails to encompass this complexity. Silviculture in Europe is skewed toward even-aged systems, used predominately (72.9% of management) across the countries assessed. The residual structure proved crucial in the comparison of natural disturbances and silvicultural systems. CI indicated the highest congruence between uneven-aged silvicultural systems and key natural disturbance attributes. Even so, uneven-aged practices emulated only a portion of the complexity associated with natural disturbance effects. The remaining silvicultural systems perform poorly in terms of retention compared to tree survivorship after natural disturbances. We suggest that NDS can enrich Europe's portfolio of management systems, for example where wood production is not the primary objective. NDS is especially relevant to forests managed for habitat quality, risk reduction, and a variety of ecosystem services. We suggest a holistic approach integrating NDS with more conventional practices.

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.

Forest structure, not climate, is the primary driver of functional diversity in northeastern North America.

Functional diversity (FD), represented by plant traits, is fundamentally linked to an ecosystem's capacity to respond to environmental change. Yet, little is known about the spatial distribution of FD and its drivers. These knowledge gaps prevent the development of FD-based forest management approaches to increase the trait diversity insurance (i.e., the response diversity) against future environmental fluctuations and disturbances. Our study helps fill these knowledge gaps by (i) mapping the current FD distribution, (ii) and analyzing FD drivers across northeastern North America. Following the stress-dominance hypothesis, we expected a strong environmental filtering effect on FD. Moreover, we expected abundant species to determine the bulk of FD distributions as suggested by the mass-ratio hypothesis. We combined a literature and database review of 44 traits for 43 tree species with terrestrial inventory data of 48,426 plots spanning an environmental gradient from northern boreal to temperate biomes. We evaluated the statistical influence of 25 covariates related to forest structure, climate, topography, soils, and stewardship on FD by employing an ensemble approach consisting of 90 non-parametric models. Temperate forests and the boreal-temperate ecotone east and northeast of the Great Lakes were identified as FD hotspots. Environmental filtering by climate was of secondary importance, with forest structure explaining most of the FD distribution of tree species in northeastern North America. Thus, our study provides only partial support for the stress-dominance hypothesis. Species abundance weightings altered trait diversity distributions and drivers only marginally, supporting the mass-ratio hypothesis. Our results suggest that forest management could increase FD without requiring knowledge of functional ecology by fostering stand structural complexity instead. Further, mixing species from different functional groups identified in this study can enhance the trait diversity insurance of forests to an uncertain future.

Effects of disturbance patterns and deadwood on the microclimate in European beech forests.

More frequent and severe disturbances increasingly open the forest canopy and initiate tree regeneration. Simultaneously, increasing weather extremes, such as drought and heat, are threatening species adapted to cool and moist climate. The magnitude of the microclimatic buffering capacity of forest canopies to mitigate hot and dry weather conditions and its disturbance-induced reduction remains poorly quantified. Also, the influence of disturbance legacies (e.g., deadwood) on forest microclimate is unresolved. In a unique manipulation experiment we investigated (i) the microclimatic buffering capacity of forest canopies in years with different climatic conditions; (ii) the impacts of spatial disturbance patterns on surface light and microclimate; and (iii) the effect of deadwood presence and type on microclimate. Treatments included two disturbance patterns (i.e., aggregated and distributed), four deadwood types (i.e., standing, downed, standing and downed, removed), and one untreated control (i.e., nine treatments in total), replicated at five sites dominated by European beech (Fagus sylvatica L.) in southeastern Germany. We measured forest floor light conditions and derived diurnal extremes and variation in temperature (T) and vapor pressure deficit (VPD) during four consecutive summer seasons (2016 - 2019). The buffering capacity of intact forest canopies was higher in warm and dry years. Surface light was significantly higher in spatially aggregated disturbance gaps compared to distributed disturbances of similar severity. An increase in surface light by 10 % relative to closed canopies elevated Tmax and VPDmax by 0.42°C and 0.04 kPa, respectively. Deadwood presence and type did not affect the forest microclimate significantly. Microclimatic buffering under forest canopies can dampen the effects of climate change. However, increasing canopy disturbances result in more light penetrating the canopy, reducing the microclimatic buffering capacity of forests. We conclude that forest management should foster microclimatic buffering in forests as one element of a multi-pronged strategy to counter climate change.

The climate sensitivity of carbon, timber, and species richness covaries with forest age in boreal-temperate North America.

Climate change threatens the provisioning of forest ecosystem services and biodiversity (ESB). The climate sensitivity of ESB may vary with forest development from young to old-growth conditions as structure and composition shift over time and space. This study addresses knowledge gaps hindering implementation of adaptive forest management strategies to sustain ESB. We focused on a number of ESB indicators to (a) analyze associations among carbon storage, timber growth rate, and species richness along a forest development gradient; (b) test the sensitivity of these associations to climatic changes; and (c) identify hotspots of climate sensitivity across the boreal-temperate forests of eastern North America. From pre-existing databases and literature, we compiled a unique dataset of 18,507 forest plots. We used a full Bayesian framework to quantify responses of nine ESB indicators. The Bayesian models were used to assess the sensitivity of these indicators and their associations to projected increases in temperature and precipitation. We found the strongest association among the investigated ESB indicators in old forests (>170 years). These forests simultaneously support high levels of carbon storage, timber growth, and species richness. Older forests also exhibit low climate sensitivity of associations among ESB indicators as compared to younger forests. While regions with a currently low combined ESB performance benefitted from climate change, regions with a high ESB performance were particularly vulnerable to climate change. In particular, climate sensitivity was highest east and southeast of the Great Lakes, signaling potential priority areas for adaptive management. Our findings suggest that strategies aimed at enhancing the representation of older forest conditions at landscape scales will help sustain ESB in a changing world.

Forest stand productivity derived from site conditions: an assessment of old Douglas-fir stands (Pseudotsuga menziesii (Mirb.) Franco var. menziesii) in Central Europe.

KEY MESSAGE: Douglas-fir growth correlates with the climate, the soil moisture regime, and the soil nutrient status, reflecting a broad physiological amplitude. Even though planting this non-native tree species is suggested as a viable strategy to improve adaptiveness of European forests to a more extreme climate and to assure future productivity, the expected temperature increase may induce a decline in forest stand productivity for Douglas-fir in already warm and dry regions. CONTEXT: Tree species selection is one of the most important forest management decisions to enhance forest productivity and stand stability on a given site. Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. menziesii), a non-native species from north-western America, is seen as an important additional species option for adapting Central European forests to a changing climate. AIMS: This study assesses Douglas-fir forest productivity derived from site conditions. We investigate climatic and physico-chemical soil characteristics and productivity of 28 mature Douglas-fir stands growing on siliceous, as well as carbonate bedrock material in southern Germany and north-eastern Austria. METHODS: The importance of climatic and physico-chemical soil characteristics was analyzed with the machine learning method Random Forests. RESULTS: The results show that Douglas-fir growth correlates with climate, soil moisture, and soil nutrient availability derived from ten climatic and physico-chemical soil parameters. CONCLUSION: The broad pH optimum between 4.5 and 7.2 reflects the broad physiological amplitude of Douglas-fir, and no significant differences were detectable between carbonate and siliceous bedrock. We also conclude that climate change may induce a forest stand productivity decline, because lower productivity with the highest mean summer temperature across our study range was observed at the warmest sites in Eastern Austria.

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.

Harnessing landscape heterogeneity for managing future disturbance risks in forest ecosystems.

In order to prevent irreversible impacts of climate change on the biosphere it is imperative to phase out the use of fossil fuels. Consequently, the provisioning of renewable resources such as timber and biomass from forests is an ecosystem service of increasing importance. However, risk factors such as changing disturbance regimes are challenging the continuous provisioning of ecosystem services, and are thus a key concern in forest management. We here used simulation modeling to study different risk management strategies in the context of timber production under changing climate and disturbance regimes, focusing on a 8127 ha forest landscape in the Northern Front Range of the Alps in Austria. We show that under a continuation of historical management, disturbances from wind and bark beetles increase by +39.5% on average over 200 years in response to future climate change. Promoting mixed forests and climate-adapted tree species as well as increasing management intensity effectively reduced future disturbance risk. Analyzing the spatial patterns of disturbance on the landscape, we found a highly uneven distribution of risk among stands (Gini coefficients up to 0.466), but also a spatially variable effectiveness of silvicultural risk reduction measures. This spatial variability in the contribution to and control of risk can be used to inform disturbance management: Stands which have a high leverage on overall risk and for which risks can effectively be reduced (24.4% of the stands in our simulations) should be a priority for risk mitigation measures. In contrast, management should embrace natural disturbances for their beneficial effects on biodiversity in areas which neither contribute strongly to landscape-scale risk nor respond positively to risk mitigation measures (16.9% of stands). We here illustrate how spatial heterogeneity in forest landscapes can be harnessed to address both positive and negative effects of changing natural disturbance regimes in ecosystem management.

Impacts of salvage logging on biodiversity: a meta-analysis.

Logging to "salvage" economic returns from forests affected by natural disturbances has become increasingly prevalent globally. Despite potential negative effects on biodiversity, salvage logging is often conducted, even in areas otherwise excluded from logging and reserved for nature conservation, inter alia because strategic priorities for post-disturbance management are widely lacking.A review of the existing literature revealed that most studies investigating the effects of salvage logging on biodiversity have been conducted less than 5 years following natural disturbances, and focused on non-saproxylic organisms.A meta-analysis across 24 species groups revealed that salvage logging significantly decreases numbers of species of eight taxonomic groups. Richness of dead wood dependent taxa (i.e. saproxylic organisms) decreased more strongly than richness of non-saproxylic taxa. In contrast, taxonomic groups typically associated with open habitats increased in the number of species after salvage logging.By analysing 134 original species abundance matrices, we demonstrate that salvage logging significantly alters community composition in 7 of 17 species groups, particularly affecting saproxylic assemblages.Synthesis and applications. Our results suggest that salvage logging is not consistent with the management objectives of protected areas. Substantial changes, such as the retention of dead wood in naturally disturbed forests, are needed to support biodiversity. Future research should investigate the amount and spatio-temporal distribution of retained dead wood needed to maintain all components of biodiversity.

Forest disturbances under climate change.

Forest disturbances are sensitive to climate. However, our understanding of disturbance dynamics in response to climatic changes remains incomplete, particularly regarding large-scale patterns, interaction effects and dampening feedbacks. Here we provide a global synthesis of climate change effects on important abiotic (fire, drought, wind, snow and ice) and biotic (insects and pathogens) disturbance agents. Warmer and drier conditions particularly facilitate fire, drought and insect disturbances, while warmer and wetter conditions increase disturbances from wind and pathogens. Widespread interactions between agents are likely to amplify disturbances, while indirect climate effects such as vegetation changes can dampen long-term disturbance sensitivities to climate. Future changes in disturbance are likely to be most pronounced in coniferous forests and the boreal biome. We conclude that both ecosystems and society should be prepared for an increasingly disturbed future of forests.

The impacts of climate change and disturbance on spatio-temporal trajectories of biodiversity in a temperate forest landscape.

1. The ongoing changes to climate challenge the conservation of forest biodiversity. Yet, in thermally limited systems, such as temperate forests, not all species groups might be affected negatively. Furthermore, simultaneous changes in the disturbance regime have the potential to mitigate climate-related impacts on forest species. Here, we (i) investigated the potential long-term effect of climate change on biodiversity in a mountain forest landscape, (ii) assessed the effects of different disturbance frequencies, severities and sizes and (iii) identified biodiversity hotspots at the landscape scale to facilitate conservation management. 2. We employed the model iLand to dynamically simulate the tree vegetation on 13 865 ha of the Kalkalpen National Park in Austria over 1000 years, and investigated 36 unique combinations of different disturbance and climate scenarios. We used simulated changes in tree cover and composition as well as projected temperature and precipitation to predict changes in the diversity of Araneae, Carabidae, ground vegetation, Hemiptera, Hymenoptera, Mollusca, saproxylic beetles, Symphyta and Syrphidae, using empirical response functions. 3. Our findings revealed widely varying responses of biodiversity indicators to climate change. Five indicators showed overall negative effects, with Carabidae, saproxylic beetles and tree species diversity projected to decrease by more than 33%. Six indicators responded positively to climate change, with Hymenoptera, Mollusca and Syrphidae diversity projected to increase more than twofold. 4. Disturbances were generally beneficial for the studied indicators of biodiversity. Our results indicated that increasing disturbance frequency and severity have a positive effect on biodiversity, while increasing disturbance size has a moderately negative effect. Spatial hotspots of biodiversity were currently found in low- to mid-elevation areas of the mountainous study landscape, but shifted to higher-elevation zones under changing climate conditions. 5.Synthesis and applications. Our results highlight that intensifying disturbance regimes may alleviate some of the impacts of climate change on forest biodiversity. However, the projected shift in biodiversity hotspots is a challenge for static conservation areas. In this regard, overlapping hotspots under current and expected future conditions highlight priority areas for robust conservation management.

Disturbances catalyze the adaptation of forest ecosystems to changing climate conditions.

The rates of anthropogenic climate change substantially exceed those at which forest ecosystems - dominated by immobile, long-lived organisms - are able to adapt. The resulting maladaptation of forests has potentially detrimental effects on ecosystem functioning. Furthermore, as many forest-dwelling species are highly dependent on the prevailing tree species, a delayed response of the latter to a changing climate can contribute to an extinction debt and mask climate-induced biodiversity loss. However, climate change will likely also intensify forest disturbances. Here, we tested the hypothesis that disturbances foster the reorganization of ecosystems and catalyze the adaptation of forest composition to climate change. Our specific objectives were (i) to quantify the rate of autonomous forest adaptation to climate change, (ii) examine the role of disturbance in the adaptation process, and (iii) investigate spatial differences in climate-induced species turnover in an unmanaged mountain forest landscape (Kalkalpen National Park, Austria). Simulations with a process-based forest landscape model were performed for 36 unique combinations of climate and disturbance scenarios over 1000 years. We found that climate change strongly favored European beech and oak species (currently prevailing in mid- to low-elevation areas), with novel species associations emerging on the landscape. Yet, it took between 357 and 706 years before the landscape attained a dynamic equilibrium with the climate system. Disturbances generally catalyzed adaptation and decreased the time needed to attain equilibrium by up to 211 years. However, while increasing disturbance frequency and severity accelerated adaptation, increasing disturbance size had the opposite effect. Spatial analyses suggest that particularly the lowest and highest elevation areas will be hotspots of future species change. We conclude that the growing maladaptation of forests to climate and the long lead times of autonomous adaptation need to be considered more explicitly in the ongoing efforts to safeguard biodiversity and ecosystem services provisioning.

The impact of future forest dynamics on climate: interactive effects of changing vegetation and disturbance regimes.

Currently, the temperate forest biome cools the earth's climate and dampens anthropogenic climate change. However, climate change will substantially alter forest dynamics in the future, affecting the climate regulation function of forests. Increasing natural disturbances can reduce carbon uptake and evaporative cooling, but at the same time increase the albedo of a landscape. Simultaneous changes in vegetation composition can mitigate disturbance impacts, but also influence climate regulation directly (e.g., via albedo changes). As a result of a number of interactive drivers (changes in climate, vegetation, and disturbance) and their simultaneous effects on climate-relevant processes (carbon exchange, albedo, latent heat flux) the future climate regulation function of forests remains highly uncertain. Here we address these complex interactions to assess the effect of future forest dynamics on the climate system. Our specific objectives were (1) to investigate the long-term interactions between changing vegetation composition and disturbance regimes under climate change, (2) to quantify the response of climate regulation to changes in forest dynamics, and (3) to identify the main drivers of the future influence of forests on the climate system. We investigated these issues using the individual-based forest landscape and disturbance model (iLand). Simulations were run over 200 yr for Kalkalpen National Park (Austria), assuming different future climate projections, and incorporating dynamically responding wind and bark beetle disturbances. To consistently assess the net effect on climate the simulated responses of carbon exchange, albedo, and latent heat flux were expressed as contributions to radiative forcing. We found that climate change increased disturbances (+27.7% over 200 yr) and specifically bark beetle activity during the 21st century. However, negative feedbacks from a simultaneously changing tree species composition (+28.0% broadleaved species) decreased disturbance activity in the long run (-10.1%), mainly by reducing the host trees available for bark beetles. Climate change and the resulting future forest dynamics significantly reduced the climate regulation function of the landscape, increasing radiative forcing by up to +10.2% on average over 200 yr. Overall, radiative forcing was most strongly driven by carbon exchange. We conclude that future changes in forest dynamics can cause amplifying climate feedbacks from temperate forest ecosystems.

Natural disturbance impacts on ecosystem services and biodiversity in temperate and boreal forests.

In many parts of the world forest disturbance regimes have intensified recently, and future climatic changes are expected to amplify this development further in the coming decades. These changes are increasingly challenging the main objectives of forest ecosystem management, which are to provide ecosystem services sustainably to society and maintain the biological diversity of forests. Yet a comprehensive understanding of how disturbances affect these primary goals of ecosystem management is still lacking. We conducted a global literature review on the impact of three of the most important disturbance agents (fire, wind, and bark beetles) on 13 different ecosystem services and three indicators of biodiversity in forests of the boreal, cool- and warm-temperate biomes. Our objectives were to (i) synthesize the effect of natural disturbances on a wide range of possible objectives of forest management, and (ii) investigate standardized effect sizes of disturbance for selected indicators via a quantitative meta-analysis. We screened a total of 1958 disturbance studies published between 1981 and 2013, and reviewed 478 in detail. We first investigated the overall effect of disturbances on individual ecosystem services and indicators of biodiversity by means of independence tests, and subsequently examined the effect size of disturbances on indicators of carbon storage and biodiversity by means of regression analysis. Additionally, we investigated the effect of commonly used approaches of disturbance management, i.e. salvage logging and prescribed burning. We found that disturbance impacts on ecosystem services are generally negative, an effect that was supported for all categories of ecosystem services, i.e. supporting, provisioning, regulating, and cultural services (P < 0.001). Indicators of biodiversity, i.e. species richness, habitat quality and diversity indices, on the other hand were found to be influenced positively by disturbance (P < 0.001). Our analyses thus reveal a 'disturbance paradox', documenting that disturbances can put ecosystem services at risk while simultaneously facilitating biodiversity. A detailed investigation of disturbance effect sizes on carbon storage and biodiversity further underlined these divergent effects of disturbance. While a disturbance event on average causes a decrease in total ecosystem carbon by 38.5% (standardized coefficient for stand-replacing disturbance), it on average increases overall species richness by 35.6%. Disturbance-management approaches such as salvage logging and prescribed burning were neither found significantly to mitigate negative effects on ecosystem services nor to enhance positive effects on biodiversity, and thus were not found to alleviate the disturbance paradox. Considering that climate change is expected to intensify natural disturbance regimes, our results indicate that biodiversity will generally benefit from such changes while a sustainable provisioning of ecosystem services might come increasingly under pressure. This underlines that disturbance risk and resilience require increased attention in ecosystem management in the future, and that new approaches to addressing the disturbance paradox in management are needed.