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.

Modeling the response of Norway spruce tree-ring carbon and oxygen isotopes to selection harvest on a drained peatland forest.

Continuous cover forestry (CCF) has gained interest as an alternative to even-aged management particularly on drained peatland forests. However, relatively little is known about the physiological response of suppressed trees when larger trees are removed as a part of CCF practices. Consequently, studies concentrating on process-level modeling of the response of trees to selection harvesting are also rare. Here, we compared, modeled and measured harvest response of previously suppressed Norway spruce (Picea abies) trees to a selection harvest. We quantified the harvest response by collecting Norway spruce tree-ring samples in a drained peatland forest site and measuring the change in stable carbon and oxygen isotopic ratios of wood formed during 2010-20, including five post-harvest years. The measured isotopic ratios were compared with ecosystem-level process model predictions for ${\kern0em }^{13}$C discrimination and ${\kern0em }^{18}$O leaf water enrichment. We found that the model predicted similar but lower harvest response than the measurements. Furthermore, accounting for mesophyll conductance was important for capturing the variation in ${\kern0em }^{13}$C discrimination. In addition, we performed sensitivity analysis on the model, which suggests that the modeled ${\kern0em }^{13}$C discrimination is sensitive to parameters related to CO2 transport through stomata to the mesophyll.

Potential of continuous cover forestry on drained peatlands to increase the carbon sink in Finland.

Land-based mitigation measures are needed to achieve climate targets. One option is the mitigation of currently high greenhouse gas (GHG) emissions of nutrient-rich drained peatland forest soils. Continuous cover forestry (CCF) has been proposed as a measure to manage this GHG emission source; however, its emission reduction potential and impact on timber production at regional and national scales have not been quantified. To quantify the potential emission reduction, we simulated four management scenarios for Finnish forests: (i) The replacement of clear-cutting by selection harvesting on nutrient-rich drained peatlands (CCF) and (ii) the current forest management regime (BAU), and both at two harvest levels, namely (i) the mean annual harvesting (2016-2018) and (ii) the maximum sustainable yield. The simulations were conducted at the stand scale with a forest simulator (MELA) coupled with a hydrological model (SpaFHy), soil C model (Yasso07) and empirical GHG exchange models. Simulations showed that the management scenario that avoided clear-cutting on nutrient-rich drained peatlands (i.e. CCF) produced approximately 1 Tg CO2 eq. higher carbon sinks annually compared with BAU at equal harvest level for Finland. This emission reduction can be attributed to the maintenance of a higher biomass sink and to the mitigation of soil emissions from nutrient-rich drained peatland sites.

Role of land cover in Finland's greenhouse gas emissions.

We present regionally aggregated emissions of greenhouse gases (GHG) from five land cover categories in Finland: artificial surfaces, arable land, forest, waterbodies, and wetlands. Carbon (C) sequestration to managed forests and unmanaged wetlands was also assessed. Models FRES and ALas were applied for emissions (CO2, CH4, N2O) from artificial surfaces and agriculture, and PREBAS for forest growth and C balance. Empirical emission coefficients were used to estimate emissions from drained forested peatland (CH4, N2O), cropland (CO2), waterbodies (CH4, CO2), peat production sites and undrained mires (CH4, CO2, N2O). We calculated gross emissions of 147.2 ± 6.8 TgCO2eq yr-1 for 18 administrative units covering mainland Finland, using data representative of the period 2017-2025. Emissions from energy production, industrial processes, road traffic and other sources in artificial surfaces amounted to 45.7 ± 2.0 TgCO2eq yr-1. The loss of C in forest harvesting was the largest emission source in the LULUCF sector, in total 59.8 ± 3.3 TgCO2eq yr-1. Emissions from domestic livestock production, field cultivation and organic soils added up to 12.2 ± 3.5 TgCO2eq yr-1 from arable land. Rivers and lakes (13.4 ± 1.9 TgCO2eq yr-1) as well as undrained mires and peat production sites (14.7 ± 1.8 TgCO2eq yr-1) increased the total GHG fluxes. The C sequestration from the atmosphere was 93.2 ± 13.7 TgCO2eq yr-1. with the main sink in forest on mineral soil (79.9 ± 12.2 TgCO2eq yr-1). All sinks compensated 63% of total emissions and thus the net emissions were 53.9 ± 15.3 TgCO2eq yr-1, or a net GHG flux per capita of 9.8 MgCO2eq yr-1.

Effect of forest management choices on carbon sequestration and biodiversity at national scale.

Forest management methods and harvest intensities influence wood production, carbon sequestration and biodiversity. We devised different management scenarios by means of stakeholder analysis and incorporated them in the forest growth simulator PREBAS. To analyse impacts of harvest intensity, we used constraints on total harvest: business as usual, low harvest, intensive harvest and no harvest. We carried out simulations on a wall-to-wall grid in Finland until 2050. Our objectives were to (1) test how the management scenarios differed in their projections, (2) analyse the potential wood production, carbon sequestration and biodiversity under the different harvest levels, and (3) compare different options of allocating the scenarios and protected areas. Harvest level was key to carbon stocks and fluxes regardless of management actions and moderate changes in proportion of strictly protected forest. In contrast, biodiversity was more dependent on other management variables than harvesting levels, and relatively independent of carbon stocks and fluxes.

Quantification of forest carbon flux and stock uncertainties under climate change and their use in regionally explicit decision making: Case study in Finland.

Uncertainties are essential, yet often neglected, information for evaluating the reliability in forest carbon balance projections used in national and regional policy planning. We analysed uncertainties in the forest net biome exchange (NBE) and carbon stocks under multiple management and climate scenarios with a process-based ecosystem model. Sampled forest initial state values, model parameters, harvest levels and global climate models (GCMs) served as inputs in Monte Carlo simulations, which covered forests of the 18 regions of mainland Finland over the period 2015-2050. Under individual scenarios, the results revealed time- and region-dependent variability in the magnitude of uncertainty and mean values of the NBE projections. The main sources of uncertainty varied with time, by region and by the amount of harvested wood. Combinations of uncertainties in the representative concentration pathways scenarios, GCMs, forest initial values and model parameters were the main sources of uncertainty at the beginning, while the harvest scenarios dominated by the end of the simulation period, combined with GCMs and climate scenarios especially in the north. Our regionally explicit uncertainty analysis was found a useful approach to reveal the variability in the regional potentials to reach a policy related, future target level of NBE, which is important information when planning realistic and regionally fair national policy actions.

Soil GHG dynamics after water level rise - Impacts of selection harvesting in peatland forests.

Managed boreal peatlands are widespread and economically important, but they are a large source of greenhouse gases (GHGs). Peatland GHG emissions are related to soil water-table level (WT), which controls the vertical distribution of aerobic and anaerobic processes and, consequently, sinks and sources of GHGs in soils. On forested peatlands, selection harvesting reduces stand evapotranspiration and it has been suggested that the resulting WT rise decreases soil net emissions, while the tree growth is maintained. We monitored soil concentrations of CO2, CH4, N2O and O2 by depth down to 80 cm, and CO2 and CH4 fluxes from soil in two nutrient-rich Norway spruce dominated peatlands in Southern Finland to examine the responses of soil GHG dynamics to WT rise. Selection harvesting raised WT by 14 cm on both sites, on average, mean WTs of the monitoring period being 73 cm for unharvested control and 59 cm for selection harvest. All soil gas concentrations were associated with proximity to WT. Both CH4 and CO2 showed remarkable vertical concentration gradients, with high values in the deepest layer, likely due to slow gas transfer in wet peat. CH4 was efficiently consumed in peat layers near and above WT where it reached sub-atmospheric concentrations, indicating sustained oxidation of CH4 from both atmospheric and deeper soil origins also after harvesting. Based on soil gas concentration data, surface peat (top 25/30 cm layer) contributed most to the soil-atmosphere CO2 fluxes and harvesting slightly increased the CO2 source in deeper soil (below 45/50 cm), which could explain the small CO2 flux differences between treatments. N2O production occurred above WT, and it was unaffected by harvesting. Overall, the WT rise obtained with selection harvesting was not sufficient to reduce soil GHG emissions, but additional hydrological regulation would have been needed.

Functional traits and climate drive interspecific differences in disturbance-induced tree mortality.

With climate change, natural disturbances such as storm or fire are reshuffled, inducing pervasive shifts in forest dynamics. To predict how it will impact forest structure and composition, it is crucial to understand how tree species differ in their sensitivity to disturbances. In this study, we investigated how functional traits and species mean climate affect their sensitivity to disturbances while controlling for tree size and stand structure. With data on 130,594 trees located on 7617 plots that were disturbed by storm, fire, snow, biotic or other disturbances from the French, Spanish, and Finnish National Forest Inventory, we modeled annual mortality probability for 40 European tree species as a function of tree size, dominance status, disturbance type, and intensity. We tested the correlation of our estimated species probability of disturbance mortality with their traits and their mean climate niches. We found that different trait combinations controlled species sensitivity to disturbances. Storm-sensitive species had a high height-dbh ratio, low wood density and high maximum growth, while fire-sensitive species had low bark thickness and high P50. Species from warmer and drier climates, where fires are more frequent, were more resistant to fire. The ranking in disturbance sensitivity between species was overall consistent across disturbance types. Productive conifer species were the most disturbance sensitive, while Mediterranean oaks were the least disturbance sensitive. Our study identified key relations between species functional traits and disturbance sensitivity, that allows more reliable predictions of how changing climate and disturbance regimes will impact future forest structure and species composition at large spatial scales.

Accuracy, realism and general applicability of European forest models.

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

A model bridging waterlogging, stomatal behavior and water use in trees in drained peatland.

Waterlogging causes hypoxic or anoxic conditions in soils, which lead to decreases in root and stomatal hydraulic conductance. Although these effects have been observed in a variety of plant species, they have not been quantified continuously over a range of water table depths (WTD) or soil water contents (SWC). To provide a quantitative theoretical framework for tackling this issue, we hypothesized similar mathematical descriptions of waterlogging and drought effects on whole-tree hydraulics and constructed a hierarchical model by connecting optimal stomata and soil-to-leaf hydraulic conductance models. In the model, the soil-to-root conductance is non-monotonic with WTD to reflect both the limitations by water under low SWC and by hypoxic effects associated with inhibited oxygen diffusion under high SWC. The model was parameterized using priors from literature and data collected over four growing seasons from Scots pine (Pinus sylvestris L.) trees grown in a drained peatland in Finland. Two reference models (RMs) were compared with the new model, RM1 with no belowground hydraulics and RM2 with no waterlogging effects. The new model was more accurate than the RMs in predicting transpiration rate (fitted slope of measured against modeled transpiration rate = 0.991 vs 0.979 (RM1) and 0.984 (RM2), R2 = 0.801 vs 0.665 (RM1) and 0.776 (RM2)). Particularly, RM2's overestimation of transpiration rate under shallow water table conditions (fitted slope = 0.908, R2 = 0.697) was considerably reduced by the new model (fitted slope = 0.956, R2 = 0.711). The limits and potential improvements of the model are discussed.

Correction: Mapping the probability of forest snow disturbances in Finland.

[This corrects the article DOI: 10.1371/journal.pone.0254876.].

Mapping the probability of forest snow disturbances in Finland.

The changing forest disturbance regimes emphasize the need for improved damage risk information. Here, our aim was to (1) improve the current understanding of snow damage risks by assessing the importance of abiotic factors, particularly the modelled snow load on trees, versus forest properties in predicting the probability of snow damage, (2) produce a snow damage probability map for Finland. We also compared the results for winters with typical snow load conditions and a winter with exceptionally heavy snow loads. To do this, we used damage observations from the Finnish national forest inventory (NFI) to create a statistical snow damage occurrence model, spatial data layers from different sources to use the model to predict the damage probability for the whole country in 16 x 16 m resolution. Snow damage reports from forest owners were used for testing the final map. Our results showed that best results were obtained when both abiotic and forest variables were included in the model. However, in the case of the high snow load winter, the model with only abiotic predictors performed nearly as well as the full model and the ability of the models to identify the snow damaged stands was higher than in other years. The results showed patterns of forest adaptation to high snow loads, as spruce stands in the north were less susceptible to damage than in southern areas and long-term snow load reduced the damage probability. The model and the derived wall-to-wall map were able to discriminate damage from no-damage cases on a good level (AUC > 0.7). The damage probability mapping approach identifies the drivers of snow disturbances across forest landscapes and can be used to spatially estimate the current and future disturbance probabilities in forests, informing practical forestry and decision-making and supporting the adaptation to the changing disturbance regimes.

Sources and sinks of greenhouse gases in the landscape: Approach for spatially explicit estimates.

Climate change mitigation is a global response that requires actions at the local level. Quantifying local sources and sinks of greenhouse gases (GHG) facilitate evaluating mitigation options. We present an approach to collate spatially explicit estimated fluxes of GHGs (carbon dioxide, methane and nitrous oxide) for main land use sectors in the landscape, to aggregate, and to calculate the net emissions of an entire region. Our procedure was developed and tested in a large river basin in Finland, providing information from intensively studied eLTER research sites. To evaluate the full GHG balance, fluxes from natural ecosystems (lakes, rivers, and undrained mires) were included together with fluxes from anthropogenic activities, agriculture and forestry. We quantified the fluxes based on calculations with an anthropogenic emissions model (FRES) and a forest growth and carbon balance model (PREBAS), as well as on emission coefficients from the literature regarding emissions from lakes, rivers, undrained mires, peat extraction sites and cropland. Spatial data sources included CORINE land use data, soil map, lake and river shorelines, national forest inventory data, and statistical data on anthropogenic activities. Emission uncertainties were evaluated with Monte Carlo simulations. Artificial surfaces were the most emission intensive land-cover class. Lakes and rivers were about as emission intensive as arable land. Forests were the dominant land cover in the region (66%), and the C sink of the forests decreased the total emissions of the region by 72%. The region's net emissions amounted to 4.37 ± 1.43 Tg CO2-eq yr-1, corresponding to a net emission intensity 0.16 Gg CO2-eq km-2 yr-1, and estimated per capita net emissions of 5.6 Mg CO2-eq yr-1. Our landscape approach opens opportunities to examine the sensitivities of important GHG fluxes to changes in land use and climate, management actions, and mitigation of anthropogenic emissions.

Extending the range of applicability of the semi-empirical ecosystem flux model PRELES for varying forest types and climate.

Applications of ecosystem flux models on large geographical scales are often limited by model complexity and data availability. Here we calibrated and evaluated a semi-empirical ecosystem flux model, PREdict Light-use efficiency, Evapotranspiration and Soil water (PRELES), for various forest types and climate conditions, based on eddy covariance data from 55 sites. A Bayesian approach was adopted for model calibration and uncertainty quantification. We applied the site-specific calibrations and multisite calibrations to nine plant functional types (PFTs) to obtain the site-specific and PFT-specific parameter vectors for PRELES. A systematically designed cross-validation was implemented to evaluate calibration strategies and the risks in extrapolation. The combination of plant physiological traits and climate patterns generated significant variation in vegetation responses and model parameters across but not within PFTs, implying that applying the model without PFT-specific parameters is risky. But within PFT, the multisite calibrations performed as accurately as the site-specific calibrations in predicting gross primary production (GPP) and evapotranspiration (ET). Moreover, the variations among sites within one PFT could be effectively simulated by simply adjusting the parameter of potential light-use efficiency (LUE), implying significant convergence of simulated vegetation processes within PFT. The hierarchical modelling of PRELES provides a compromise between satellite-driven LUE and physiologically oriented approaches for extrapolating the geographical variation of ecosystem productivity. Although measurement errors of eddy covariance and remotely sensed data propagated a substantial proportion of uncertainty or potential biases, the results illustrated that PRELES could reliably capture daily variations of GPP and ET for contrasting forest types on large geographical scales if PFT-specific parameterizations were applied.

Refining the role of phenology in regulating gross ecosystem productivity across European peatlands.

The role of plant phenology as a regulator for gross ecosystem productivity (GEP) in peatlands is empirically not well constrained. This is because proxies to track vegetation development with daily coverage at the ecosystem scale have only recently become available and the lack of such data has hampered the disentangling of biotic and abiotic effects. This study aimed at unraveling the mechanisms that regulate the seasonal variation in GEP across a network of eight European peatlands. Therefore, we described phenology with canopy greenness derived from digital repeat photography and disentangled the effects of radiation, temperature and phenology on GEP with commonality analysis and structural equation modeling. The resulting relational network could not only delineate direct effects but also accounted for possible effect combinations such as interdependencies (mediation) and interactions (moderation). We found that peatland GEP was controlled by the same mechanisms across all sites: phenology constituted a key predictor for the seasonal variation in GEP and further acted as a distinct mediator for temperature and radiation effects on GEP. In particular, the effect of air temperature on GEP was fully mediated through phenology, implying that direct temperature effects representing the thermoregulation of photosynthesis were negligible. The tight coupling between temperature, phenology and GEP applied especially to high latitude and high altitude peatlands and during phenological transition phases. Our study highlights the importance of phenological effects when evaluating the future response of peatland GEP to climate change. Climate change will affect peatland GEP especially through changing temperature patterns during plant phenologically sensitive phases in high latitude and high altitude regions.

Ecosystem Services Related to Carbon Cycling - Modeling Present and Future Impacts in Boreal Forests.

Forests regulate climate, as carbon, water and nutrient fluxes are modified by physiological processes of vegetation and soil. Forests also provide renewable raw material, food, and recreational possibilities. Rapid climate warming projected for the boreal zone may change the provision of these ecosystem services. We demonstrate model based estimates of present and future ecosystem services related to carbon cycling of boreal forests. The services were derived from biophysical variables calculated by two dynamic models. Future changes in the biophysical variables were driven by climate change scenarios obtained as results of a sample of global climate models downscaled for Finland, assuming three future pathways of radiative forcing. We introduce continuous monitoring on phenology to be used in model parametrization through a webcam network with automated image processing features. In our analysis, climate change impacts on key boreal forest ecosystem services are both beneficial and detrimental. Our results indicate an increase in annual forest growth of about 60% and an increase in annual carbon sink of roughly 40% from the reference period (1981-2010) to the end of the century. The vegetation active period was projected to start about 3 weeks earlier and end ten days later by the end of the century compared to currently. We found a risk for increasing drought, and a decrease in the number of soil frost days. Our results show a considerable uncertainty in future provision of boreal forest ecosystem services.

Carbon flux from decomposing wood and its dependency on temperature, wood N2 fixation rate, moisture and fungal composition in a Norway spruce forest.

Globally 40-70 Pg of carbon (C) are stored in coarse woody debris on the forest floor. Climate change may reduce the function of this stock as a C sink in the future due to increasing temperature. However, current knowledge on the drivers of wood decomposition is inadequate for detailed predictions. To define the factors that control wood respiration rate of Norway spruce and to produce a model that adequately describes the decomposition process of this species as a function of time, we used an unprecedentedly diverse analytical approach, which included measurements of respiration, fungal community sequencing, N2 fixation rate, nifH copy number, 14 C-dating as well as N%, δ13 C and C% values of wood. Our results suggest that climate change will accelerate C flux from deadwood in boreal conditions, due to the observed strong temperature dependency of deadwood respiration. At the research site, the annual C flux from deadwood would increase by 27% from the current 117 g C/kg wood with the projected climate warming (RCP4.5). The second most important control on respiration rate was the stage of wood decomposition; at early stages of decomposition low nitrogen content and low wood moisture limited fungal activity while reduced wood resource quality decreased the respiration rate at the final stages of decomposition. Wood decomposition process was best described by a Sigmoidal model, where after 116 years of wood decomposition mass loss of 95% was reached. Our results on deadwood decomposition are important for C budget calculations in ecosystem and climate change models. We observed for the first time that the temperature dependency of N2 fixation, which has a major role at providing N for wood-inhabiting fungi, was not constant but varied between wood density classes due to source supply and wood quality. This has significant consequences on projecting N2 fixation rates for deadwood in changing climate.

Tree differences in primary and secondary growth drive convergent scaling in leaf area to sapwood area across Europe.

Trees scale leaf (AL ) and xylem (AX ) areas to couple leaf transpiration and carbon gain with xylem water transport. Some species are known to acclimate in AL  : AX balance in response to climate conditions, but whether trees of different species acclimate in AL  : AX in similar ways over their entire (continental) distributions is unknown. We analyzed the species and climate effects on the scaling of AL vs AX in branches of conifers (Pinus sylvestris, Picea abies) and broadleaved (Betula pendula, Populus tremula) sampled across a continental wide transect in Europe. Along the branch axis, AL and AX change in equal proportion (isometric scaling: b Ëœ 1) as for trees. Branches of similar length converged in the scaling of AL vs AX with an exponent of b = 0.58 across European climates irrespective of species. Branches of slow-growing trees from Northern and Southern regions preferentially allocated into new leaf rather than xylem area, with older xylem rings contributing to maintaining total xylem conductivity. In conclusion, trees in contrasting climates adjust their functional balance between water transport and leaf transpiration by maintaining biomass allocation to leaves, and adjusting their growth rate and xylem production to maintain xylem conductance.

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.