Relative decline in density of Northern Hemisphere tree species in warm and arid regions of their climate niches.

Although climate change is expected to drive tree species toward colder and wetter regions of their distribution, broadscale empirical evidence is lacking. One possibility is that past and present human activities in forests obscure or alter the effects of climate. Here, using data from more than two million monitored trees from 73 widely distributed species, we quantify changes in tree species density within their climatic niches across Northern Hemisphere forests. We observe a reduction in mean density across species, coupled with a tendency toward increasing tree size. However, the direction and magnitude of changes in density exhibit considerable variability between species, influenced by stand development that results from previous stand-level disturbances. Remarkably, when accounting for stand development, our findings show a significant change in density toward cold and wet climatic conditions for 43% of the species, compared to only 14% of species significantly changing their density toward warm and arid conditions in both early- and late-development stands. The observed changes in climate-driven density showed no clear association with species traits related to drought tolerance, recruitment and dispersal capacity, or resource use, nor with the temperature or aridity affiliation of the species, leaving the underlying mechanism uncertain. Forest conservation policies and associated management strategies might want to consider anticipated long-term species range shifts alongside the integration of contemporary within-distribution density changes.

Phosphorus limitation promotes soil carbon storage in a boreal forest exposed to long-term nitrogen fertilization.

Forests play a crucial role in global carbon cycling by absorbing and storing significant amounts of atmospheric carbon dioxide. Although boreal forests contribute to approximately 45% of the total forest carbon sink, tree growth and soil carbon sequestration are constrained by nutrient availability. Here, we examine if long-term nutrient input enhances tree productivity and whether this leads to carbon storage or whether stimulated microbial decomposition of organic matter limits soil carbon accumulation. Over six decades, nitrogen, phosphorus, and calcium were supplied to a Pinus sylvestris-dominated boreal forest. We found that nitrogen fertilization alone or together with calcium and/or phosphorus increased tree biomass production by 50% and soil carbon sequestration by 65% compared to unfertilized plots. However, the nonlinear relationship observed between tree productivity and soil carbon stock across treatments suggests microbial regulation. When phosphorus was co-applied with nitrogen, it acidified the soil, increased fungal biomass, altered microbial community composition, and enhanced biopolymer degradation capabilities. While no evidence of competition between ectomycorrhizal and saprotrophic fungi has been observed, key functional groups with the potential to reduce carbon stocks were identified. In contrast, when nitrogen was added without phosphorus, it increased soil carbon sequestration because microbial activity was likely limited by phosphorus availability. In conclusion, the addition of nitrogen to boreal forests may contribute to global warming mitigation, but this effect is context dependent.

Statistical upscaling of ecosystem CO2 fluxes across the terrestrial tundra and boreal domain: Regional patterns and uncertainties.

The regional variability in tundra and boreal carbon dioxide (CO2 ) fluxes can be high, complicating efforts to quantify sink-source patterns across the entire region. Statistical models are increasingly used to predict (i.e., upscale) CO2 fluxes across large spatial domains, but the reliability of different modeling techniques, each with different specifications and assumptions, has not been assessed in detail. Here, we compile eddy covariance and chamber measurements of annual and growing season CO2 fluxes of gross primary productivity (GPP), ecosystem respiration (ER), and net ecosystem exchange (NEE) during 1990-2015 from 148 terrestrial high-latitude (i.e., tundra and boreal) sites to analyze the spatial patterns and drivers of CO2 fluxes and test the accuracy and uncertainty of different statistical models. CO2 fluxes were upscaled at relatively high spatial resolution (1 km2 ) across the high-latitude region using five commonly used statistical models and their ensemble, that is, the median of all five models, using climatic, vegetation, and soil predictors. We found the performance of machine learning and ensemble predictions to outperform traditional regression methods. We also found the predictive performance of NEE-focused models to be low, relative to models predicting GPP and ER. Our data compilation and ensemble predictions showed that CO2 sink strength was larger in the boreal biome (observed and predicted average annual NEE -46 and -29 g C m-2  yr-1 , respectively) compared to tundra (average annual NEE +10 and -2 g C m-2  yr-1 ). This pattern was associated with large spatial variability, reflecting local heterogeneity in soil organic carbon stocks, climate, and vegetation productivity. The terrestrial ecosystem CO2 budget, estimated using the annual NEE ensemble prediction, suggests the high-latitude region was on average an annual CO2 sink during 1990-2015, although uncertainty remains high.

A new method to estimate branch biomass from terrestrial laser scanning data by bridging tree structure models.

BACKGROUND AND AIMS: Branch biomass and other attributes are important for estimating the carbon budget of forest stands and characterizing crown structure. As destructive measuring is time-consuming and labour-intensive, terrestrial laser scanning (TLS) as a solution has been used to estimate branch biomass quickly and non-destructively. However, branch information extraction from TLS data alone is challenging due to occlusion and other defects, especially for estimating individual branch attributes in coniferous trees. METHODS: This study presents a method, entitled TSMtls, to estimate individual branch biomass non-destructively and accurately by combining tree structure models and TLS data. The TSMtls method constructs the stem-taper curve from TLS data, then uses tree structure models to determine the number, basal area and biomass of individual branches at whorl level. We estimated the tree structural model parameters from 122 destructively measured Scots pine (Pinus sylvestris) trees and tested the method on six Scots pine trees that were first TLS-scanned and later destructively measured. Additionally, we estimated the branch biomass using other TLS-based approaches for comparison. KEY RESULTS: Tree-level branch biomass estimates derived from TSMtls showed the best agreement with the destructive measurements [coefficient of variation of root mean square error (CV-RMSE) = 9.66 % and concordance correlation coefficient (CCC) = 0.99], outperforming the other TLS-based approaches (CV-RMSE 12.97-57.45 % and CCC 0.43-0.98 ). Whorl-level individual branch attributes estimates produced from TSMtls showed more accurate results than those produced from TLS data directly. CONCLUSIONS: The results showed that the TSMtls method proposed in this study holds promise for extension to more species and larger areas.

Treatment profile and long-term outcome of intensive care unit-admitted patients with liver cirrhosis or other liver disease in relation to alcohol consumption.

OBJECTIVE: To examine the impact of alcohol consumption on the treatment profile, mortality and causes of death in intensive care unit (ICU)-admitted patients with liver cirrhosis and other liver disease. METHODS: Data on liver disease and ICU treatment of patients with previously diagnosed liver disease between 2015 and 2017 were retrospectively collected from medical records at Oulu University Hospital, Finland. The median follow-up was 367 days. The causes of death were obtained from Statistics Finland. RESULTS: From 250 patients, high-risk alcohol consumption was present in 74.7% (71 of 95) cirrhotic patients and 43.2% (67 of 155) patients in the other liver disease group. Gastrointestinal causes were the most common admission causes. Despite the higher SOFA scores in the alcoholic liver cirrhosis patients compared with the non-alcoholic cirrhosis, there were no differences in the need for organ support, length of ICU stay or outcome between the groups or the subgroups. There were no differences in 1-year mortality between the cirrhosis groups (alcoholic cirrhosis 43.7% versus non-alcoholic cirrhosis 45.8%, p = 1.0) or between the other liver disease groups (patients with alcohol consumption 37.3% versus patients without alcohol consumption 36.4%, p = 1.0). The patients with high-risk alcohol consumption died more often due to liver disease, whereas the patients without high-risk alcohol consumption died often due to malignancies. CONCLUSIONS: We report no significant impact of alcohol consumption on the ICU treatment profile or mortality of patients with cirrhosis or other liver disease. The high mortality underlines the importance of preventive measures after ICU admission.

Continental mapping of forest ecosystem functions reveals a high but unrealised potential for forest multifunctionality.

Humans require multiple services from ecosystems, but it is largely unknown whether trade-offs between ecosystem functions prevent the realisation of high ecosystem multifunctionality across spatial scales. Here, we combined a comprehensive dataset (28 ecosystem functions measured on 209 forest plots) with a forest inventory dataset (105,316 plots) to extrapolate and map relationships between various ecosystem multifunctionality measures across Europe. These multifunctionality measures reflected different management objectives, related to timber production, climate regulation and biodiversity conservation/recreation. We found that trade-offs among them were rare across Europe, at both local and continental scales. This suggests a high potential for 'win-win' forest management strategies, where overall multifunctionality is maximised. However, across sites, multifunctionality was on average 45.8-49.8% below maximum levels and not necessarily highest in protected areas. Therefore, using one of the most comprehensive assessments so far, our study suggests a high but largely unrealised potential for management to promote multifunctional forests.

Climate- and successional-related changes in functional composition of European forests are strongly driven by tree mortality.

Intense droughts combined with increased temperatures are one of the major threats to forest persistence in the 21st century. Despite the direct impact of climate change on forest growth and shifts in species abundance, the effect of altered demography on changes in the composition of functional traits is not well known. We sought to (1) quantify the recent changes in functional composition of European forests; (2) identify the relative importance of climate change, mean climate and forest development for changes in functional composition; and (3) analyse the roles of tree mortality and growth underlying any functional changes in different forest types. We quantified changes in functional composition from the 1980s to the 2000s across Europe by two dimensions of functional trait variation: the first dimension was mainly related to changes in leaf mass per area and wood density (partially related to the trait differences between angiosperms and gymnosperms), and the second dimension was related to changes in maximum tree height. Our results indicate that climate change and mean climatic effects strongly interacted with forest development and it was not possible to completely disentangle their effects. Where recent climate change was not too extreme, the patterns of functional change generally followed the expected patterns under secondary succession (e.g. towards late-successional short-statured hardwoods in Mediterranean forests and taller gymnosperms in boreal forests) and latitudinal gradients (e.g. larger proportion of gymnosperm-like strategies at low water availability in forests formerly dominated by broad-leaved deciduous species). Recent climate change generally favoured the dominance of angiosperm-like related traits under increased temperature and intense droughts. Our results show functional composition changes over relatively short time scales in European forests. These changes are largely determined by tree mortality, which should be further investigated and modelled to adequately predict the impacts of climate change on forest function.

Ecosystem services of boreal forests - Carbon budget mapping at high resolution.

The carbon (C) cycle of forests produces ecosystem services (ES) such as climate regulation and timber production. Mapping these ES using simple land cover -based proxies might add remarkable inaccuracy to the estimates. A framework to map the current status of the C budget of boreal forested landscapes was developed. The C stocks of biomass and soil and the annual change in these stocks were quantified in a 20 × 20 m resolution at the regional level on mineral soils in southern Finland. The fine-scale variation of the estimates was analyzed geo-statistically. The reliability of the estimates was evaluated by comparing them to measurements from the national multi-source forest inventory. The C stocks of forests increased slightly from the south coast to inland whereas the changes in these stocks were more uniform. The spatial patches of C stocks were larger than those of C stock changes. The patch size of the C stocks reflected the spatial variation in the environmental conditions, and that of the C stock changes the typical area of forest management compartments. The simulated estimates agreed well with the measurements indicating a good mapping framework performance. The mapping framework is the basis for evaluating the effects of forest management alternatives on C budget at high resolution across large spatial scales. It will be coupled with the assessment of other ES and biodiversity to study their relationships. The framework integrated a wide suite of simulation models and extensive inventory data. It provided reliable estimates of the human influence on C cycle in forested landscapes.

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.

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.

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.

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

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

Estimating spatial variation in the effects of climate change on the net primary production of Japanese cedar plantations based on modeled carbon dynamics.

Spatiotemporal prediction of the response of planted forests to a changing climate is increasingly important for the sustainable management of forest ecosystems. In this study, we present a methodology for estimating spatially varying productivity in a planted forest and changes in productivity with a changing climate in Japan, with a focus on Japanese cedar (Cryptomeria japonica D. Don) as a representative tree species of this region. The process-based model Biome-BGC was parameterized using a plant trait database for Japanese cedar and a Bayesian optimization scheme. To compare productivity under historical (1996-2000) and future (2096-2100) climatic conditions, the climate scenarios of two representative concentration pathways (i.e., RCP2.6 and RCP8.5) were used in five global climate models (GCMs) with approximately 1-km resolution. The seasonality of modeled fluxes, namely gross primary production, ecosystem respiration, net ecosystem exchange, and soil respiration, improved after two steps of parameterization. The estimated net primary production (NPP) of stands aged 36-40 years under the historical climatic conditions of the five GCMs was 0.77 ± 0.10 kgC m-2 year-1 (mean ± standard deviation), in accordance with the geographical distribution of forest NPP estimated in previous studies. Under the RCP2.6 and RCP8.5 scenarios, the mean NPP of the five GCMs increased by 0.04 ± 0.07 and 0.14 ± 0.11 kgC m-2 year-1, respectively. The increases in annual NPP were small in the southwestern region because of the decreases in summer NPP and the small increases in winter NPP under the RCP2.6 and RCP8.5 scenarios, respectively. Under the RCP2.6 scenario, Japanese cedar was at risk in the southwestern region, in accordance with previous studies, and monitoring and silvicultural practices should be modified accordingly.

Forest carbon allocation modelling under climate change.

Carbon allocation plays a key role in ecosystem dynamics and plant adaptation to changing environmental conditions. Hence, proper description of this process in vegetation models is crucial for the simulations of the impact of climate change on carbon cycling in forests. Here we review how carbon allocation modelling is currently implemented in 31 contrasting models to identify the main gaps compared with our theoretical and empirical understanding of carbon allocation. A hybrid approach based on combining several principles and/or types of carbon allocation modelling prevailed in the examined models, while physiologically more sophisticated approaches were used less often than empirical ones. The analysis revealed that, although the number of carbon allocation studies over the past 10 years has substantially increased, some background processes are still insufficiently understood and some issues in models are frequently poorly represented, oversimplified or even omitted. Hence, current challenges for carbon allocation modelling in forest ecosystems are (i) to overcome remaining limits in process understanding, particularly regarding the impact of disturbances on carbon allocation, accumulation and utilization of nonstructural carbohydrates, and carbon use by symbionts, and (ii) to implement existing knowledge of carbon allocation into defence, regeneration and improved resource uptake in order to better account for changing environmental conditions.

How do trees respond to species mixing in experimental compared to observational studies?

For decades, ecologists have investigated the effects of tree species diversity on tree productivity at different scales and with different approaches ranging from observational to experimental study designs. Using data from five European national forest inventories (16,773 plots), six tree species diversity experiments (584 plots), and six networks of comparative plots (169 plots), we tested whether tree species growth responses to species mixing are consistent and therefore transferrable between those different research approaches. Our results confirm the general positive effect of tree species mixing on species growth (16% on average) but we found no consistency in species-specific responses to mixing between any of the three approaches, even after restricting comparisons to only those plots that shared similar mixtures compositions and forest types. These findings highlight the necessity to consider results from different research approaches when selecting species mixtures that should maximize positive forest biodiversity and functioning relationships.

Testing dependence between growth and needle litterfall in Scots pine--a case study in northern Finland.

Both drought and fungal disease increase needle litterfall of Scots pine (Pinus sylvestris L.) trees, but most factors causing annual variation in needle litterfall are poorly understood. We hypothesized that radial growth and weather conditions favorable to growth correlate positively with needle litterfall with a lag equal to the number of needle cohorts (here being 5-6). We studied the time series of needle litterfall, stem increment, pollen cone litter and daily weather conditions in a Scots pine stand over 43 years (1961-2004). The cross-correlations of standardized time series were estimated with various lags. Model predictions of annual needle litterfall were tested against independent data. Changes in annual growth and needle litterfall correlated with lags of 0 and 4 years. The best predictors for needle litterfall were May to mid July temperature sum with a lag of 4 years, May rainfall with a lag of 2 years and September temperature with a lag of 6 years. Pollen cone litter correlated negatively with needle litterfall with a lag of 2 years. The study provided empirical evidence that needle litterfall of Scots pine in northern Finland is influenced by needle production and needle mass development that occurred 4 to 6 years earlier.

Erratum: Complementarity effects on tree growth are contingent on tree size and climatic conditions across Europe.

This corrects the article DOI: 10.1038/srep32233.

Actual European forest management by region, tree species and owner based on 714,000 re-measured trees in national forest inventories.

BACKGROUND: European forests have a long record of management. However, the diversity of the current forest management across nations, tree species and owners, is hardly understood. Often when trying to simulate future forest resources under alternative futures, simply the yield table style of harvesting is applied. It is now crucially important to come to grips with actual forest management, now that demand for wood is increasing and the EU Land Use, Land Use Change and Forestry Regulation has been adopted requiring 'continuation of current management practices' as a baseline to set the Forest Reference Level carbon sink. METHODS: Based on a large dataset of 714,000 re-measured trees in National Forest inventories from 13 regions, we are now able to analyse actual forest harvesting. CONCLUSIONS: From this large set of repeated tree measurements we can conclude that there is no such thing as yield table harvesting in Europe. We found general trends of increasing harvest probability with higher productivity of the region and the species, but with important deviations related to local conditions like site accessibility, state of the forest resource (like age), specific subsidies, importance of other forest services, and ownership of the forest. As a result, we find a huge diversity in harvest regimes. Over the time period covered in our inventories, the average harvest probability over all regions was 2.4% yr-1 (in number of trees) and the mortality probability was 0.4% yr-1. Our study provides underlying and most actual data that can serve as a basis for quantifying 'continuation of current forest management'. It can be used as a cornerstone for the base period as required for the Forest Reference Level for EU Member States.