Leaf isotopes reveal tree diversity effects on the functional responses to the pan-European 2018 summer drought.

Recent droughts have strongly impacted forest ecosystems and are projected to increase in frequency, intensity, and duration in the future together with continued warming. While evidence suggests that tree diversity can regulate drought impacts in natural forests, few studies examine whether mixed tree plantations are more resistant to the impacts of severe droughts. Using natural variations in leaf carbon (C) and nitrogen (N) isotopic ratios, that is δ13C and δ15N, as proxies for drought response, we analyzed the effects of tree species richness on the functional responses of tree plantations to the pan-European 2018 summer drought in seven European tree diversity experiments. We found that leaf δ13C decreased with increasing tree species richness, indicating less drought stress. This effect was not related to drought intensity, nor desiccation tolerance of the tree species. Leaf δ15N increased with drought intensity, indicating a shift toward more open N cycling as water availability diminishes. Additionally, drought intensity was observed to alter the influence of tree species richness on leaf δ15N from weakly negative under low drought intensity to weakly positive under high drought intensity. Overall, our findings suggest that dual leaf isotope analysis helps understand the interaction between drought, nutrients, and species richness.

Green deserts, but not always: A global synthesis of native woody species regeneration under tropical tree monocultures.

Tree monocultures constitute an increasing fraction of the global tree cover and are the dominant tree-growing strategy of forest landscape restoration commitments. Their advantages to produce timber are well known, but their value for biodiversity is highly controversial and context dependent. Therefore, understanding whether, and in which conditions, they can harbor native species regeneration is crucial. Here, we conducted meta-analyses based on a global survey of the literature and on a database created with local, unpublished studies throughout Brazil to evaluate the regeneration potential of native species under tree monocultures and the way management influences this regeneration. Native woody species regeneration under tree monocultures harbors a substantial fraction of the diversity (on average 40% and 68% in the global and Brazilian surveys, respectively) and abundance (on average 25% and 60% in the global and Brazilian surveys, respectively) of regeneration observed in natural forests. Plantations with longer rotation lengths, composed of native tree species, and located adjacent to forest remnants harbor more species. Pine plantations harbor more native individuals than eucalypt plantations, and the abundance of regenerating trees is higher in sites with higher mean temperatures. Species-area curves revealed that the number of woody species under pine and eucalypt plantations in Brazil is 606 and 598 species, respectively, over an aggregated sampled area of ca. 12 ha. We highlight that the understory of tree monocultures can harbor a considerable diversity of regenerating native species at the landscape and regional scales, but this diversity strongly depends on management. Long-rotation length and favorable location are key factors for woody regeneration success under tropical tree monocultures. Therefore, tree monocultures can play a role in forest landscape restoration and conservation, but only if they are planned and managed for achieving this purpose.

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

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

Forest structure and composition alleviate human thermal stress.

Current climate change aggravates human health hazards posed by heat stress. Forests can locally mitigate this by acting as strong thermal buffers, yet potential mediation by forest ecological characteristics remains underexplored. We report over 14 months of hourly microclimate data from 131 forest plots across four European countries and compare these to open-field controls using physiologically equivalent temperature (PET) to reflect human thermal perception. Forests slightly tempered cold extremes, but the strongest buffering occurred under very hot conditions (PET >35°C), where forests reduced strong to extreme heat stress day occurrence by 84.1%. Mature forests cooled the microclimate by 12.1 to 14.5°C PET under, respectively, strong and extreme heat stress conditions. Even young plantations reduced those conditions by 10°C PET. Forest structure strongly modulated the buffering capacity, which was enhanced by increasing stand density, canopy height and canopy closure. Tree species composition had a more modest yet significant influence: that is, strongly shade-casting, small-leaved evergreen species amplified cooling. Tree diversity had little direct influences, though indirect effects through stand structure remain possible. Forests in general, both young and mature, are thus strong thermal stress reducers, but their cooling potential can be even further amplified, given targeted (urban) forest management that considers these new insights.

Blazing the trail: Social innovation supporting wildfire-resilient territories in Catalonia (Spain).

Mediterranean territories have co-evolved and been shaped by fire throughout history. However, global environmental change conditions are increasing the size, intensity and severity of wildfires, which have gone from a regular natural disturbance to a serious threat for civil protection, surpassing firefighting capacities. Therefore, building resilience in fire-prone territories is an increasingly relevant policy and management objective. However, the notion of resilience has been criticized for paying insufficient attention to key social issues such as socio-political dynamics, power imbalances and societal change. At the same time, social science contributions to wildfire research are still rather limited. In this paper, we bridge social innovation theory to resilience theory in order to create a territorially embedded and socially sensitive framework for assessing socio-ecological resilience. From this perspective, we then examine how Forest Defence Groups (ADFs, by their Catalan acronym) have evolved from grassroots, bottom-up initiatives to well-established bottom-linked institutions and we evaluate their contributions to socio-ecological resilience in the territories where they operate. Our results show that ADFs contribute in several aspects to socio-ecological resilience and that the pave the way for opening up spaces of dialogue and collaboration through which local communities can engage with the issues that directly affect them, such as wildfires.

Relative importance of tree species richness, tree functional type, and microenvironment for soil macrofauna communities in European forests.

Soil fauna communities are major drivers of many forest ecosystem processes. Tree species diversity and composition shape soil fauna communities, but their relationships are poorly understood, notably whether or not soil fauna diversity depends on tree species diversity. Here, we characterized soil macrofauna communities from forests composed of either one or three tree species, located in four different climate zones and growing on different soil types. Using multivariate analysis and model averaging we investigated the relative importance of tree species richness, tree functional type (deciduous vs. evergreen), litter quality, microhabitat and microclimatic characteristics as drivers of soil macrofauna community composition and structure. We found that macrofauna communities in mixed forest stands were represented by a higher number of broad taxonomic groups that were more diverse and more evenly represented. We also observed a switch from earthworm-dominated to predator-dominated communities with increasing evergreen proportion in forest stands, which we interpreted as a result of a lower litter quality and a higher forest floor mass. Finally, canopy openness was positively related to detritivore abundance and biomass, leading to higher predator species richness and diversity probably through trophic cascade effects. Interestingly, considering different levels of taxonomic resolution in the analyses highlighted different facets of macrofauna response to tree species richness, likely a result of both different ecological niche range and methodological constraints. Overall, our study supports the positive effects of tree species richness on macrofauna diversity and abundance through multiple changes in resource quality and availability, microhabitat, and microclimate modifications.

Mapping tree species vulnerability to multiple threats as a guide to restoration and conservation of tropical dry forests.

Understanding the vulnerability of tree species to anthropogenic threats is important for the efficient planning of restoration and conservation efforts. We quantified and compared the effects of future climate change and four current threats (fire, habitat conversion, overgrazing and overexploitation) on the 50 most common tree species of the tropical dry forests of northwestern Peru and southern Ecuador. We used an ensemble modelling approach to predict species distribution ranges, employed freely accessible spatial datasets to map threat exposures, and developed a trait-based scoring approach to estimate species-specific sensitivities, using differentiated trait weights in accordance with their expected importance in determining species sensitivities to specific threats. Species-specific vulnerability maps were constructed from the product of the exposure maps and the sensitivity estimates. We found that all 50 species face considerable threats, with an average of 46% of species' distribution ranges displaying high or very high vulnerability to at least one of the five threats. Our results suggest that current levels of habitat conversion, overexploitation and overgrazing pose larger threats to most of the studied species than climate change. We present a spatially explicit planning strategy for species-specific restoration and conservation actions, proposing management interventions to focus on (a) in situ conservation of tree populations and seed collection for tree planting activities in areas with low vulnerability to climate change and current threats; (b) ex situ conservation or translocation of populations in areas with high climate change vulnerability; and (c) active planting or assisted regeneration in areas under high current threat vulnerability but low climate change vulnerability, provided that interventions are in place to lower threat pressure. We provide an online, user-friendly tool to visualize both the vulnerability maps and the maps indicating priority restoration and conservation actions.

Should we Leave Nature Unattended or Assist through Enrichment to Foster Climate Change Mitigation? Exclosure Management in the Highlands of Ethiopia.

In order to foster the potential of exclosures to sequester carbon, it is understood that they are increasingly assisted through enrichment planting. To study the impact of the enrichment planting on carbon sequestration process, five exclosures with enrichment planting and five pure naturally regenerated exclosures were selected. Along parallel transects, 20 × 20 m plots were laid at 100 m intervals where all woody vegetations were counted and measured for their diameter and total height. For soil sampling, five subplots at the center and four at each corner of the plots were established. The samples were collected at a depth of 0-0.2 m, and this procedure was repeated for each plot. In this case, when good management practices were implemented (such as Wukro exclosures), significant differences in organic soil carbon above the ground and the total carbon between naturally regenerated and enriched exclosures (P < 0.05) were found. The mean estimates of the above ground carbon, soil carbon, and total carbon were respectively 8.08, 31.04, and 39.12 ton/ha for natural regeneration vs. 7.94, 31.00, and 38.93 ton/ha for enriched regeneration. Lower altitudes had significantly higher soil organic carbon (P < 0.05) than the higher altitudes. However, the slope had an insignificant effect on carbon distribution. Enriched exclosures performed more poorly in carbon sequestration. This was possibly due to the disturbances caused by mass plantation and poor post plantation follow up, since improved performance (P < 0.05) was seen in one enriched exclosure with better management practices.

Biotic predictors complement models of bat and bird responses to climate and tree diversity in European forests.

Bats and birds are key providers of ecosystem services in forests. How climate and habitat jointly shape their communities is well studied, but whether biotic predictors from other trophic levels may improve bird and bat diversity models is less known, especially across large bioclimatic gradients. Here, we achieved multi-taxa surveys in 209 mature forests replicated in six European countries from Spain to Finland, to investigate the importance of biotic predictors (i.e. the abundance or activity of defoliating insects, spiders, earthworms and wild ungulates) for bat and bird taxonomic and functional diversity. We found that nine out of 12 bird and bat diversity metrics were best explained when biotic factors were added to models including climate and habitat variables, with a mean gain in explained variance of 38% for birds and 15% for bats. Tree functional diversity was the most important habitat predictor for birds, while bats responded more to understorey structure. The best biotic predictors for birds were spider abundance and defoliating insect activity, while only bat functional evenness responded positively to insect herbivory. Accounting for potential biotic interactions between bats, birds and other taxa of lower trophic levels will help to understand how environmental changes along large biogeographical gradients affect higher-level predator diversity in forest ecosystems.

Biotic homogenization can decrease landscape-scale forest multifunctionality.

Many experiments have shown that local biodiversity loss impairs the ability of ecosystems to maintain multiple ecosystem functions at high levels (multifunctionality). In contrast, the role of biodiversity in driving ecosystem multifunctionality at landscape scales remains unresolved. We used a comprehensive pan-European dataset, including 16 ecosystem functions measured in 209 forest plots across six European countries, and performed simulations to investigate how local plot-scale richness of tree species (α-diversity) and their turnover between plots (ß-diversity) are related to landscape-scale multifunctionality. After accounting for variation in environmental conditions, we found that relationships between α-diversity and landscape-scale multifunctionality varied from positive to negative depending on the multifunctionality metric used. In contrast, when significant, relationships between ß-diversity and landscape-scale multifunctionality were always positive, because a high spatial turnover in species composition was closely related to a high spatial turnover in functions that were supported at high levels. Our findings have major implications for forest management and indicate that biotic homogenization can have previously unrecognized and negative consequences for large-scale ecosystem multifunctionality.

Tree diversity mitigates defoliation after a drought-induced tipping point.

Understanding the processes that underlie drought-related tree vitality loss is essential for anticipating future forest dynamics, and for developing management plans aiming at increasing the resilience of forests to climate change. Forest vitality has been continuously monitored in Europe since the acid rain alert in the 1980s, and the intensive monitoring plots of ICP Forests offer the opportunity to investigate the effects of air pollution and climate change on forest condition. By making use of over 100 long-term monitoring plots, where crown defoliation has been assessed extensively since 1990, we discovered a progressive shift from a negative to a positive effect of species richness on forest health. The observed tipping point in the balance of net interactions, from competition to facilitation, has never been reported from real ecosystems outside experimental conditions; and the strong temporal consistency of our observations with increasing drought stress emphasizes its climate change relevance. Furthermore, we show that higher species diversity has reduced the severity of defoliation in the long term. Our results confirm the greater resilience of diverse forests to future climate change-induced stress. More generally, they add to an accumulating body of evidence on the large potential of tree species mixtures to face manifold disturbances in a changing world.

EFO-LCI: A New Life Cycle Inventory Database of Forestry Operations in Europe.

Life cycle assessment (LCA) has become a common methodology to analyze environmental impacts of forestry systems. Although LCA has been widely applied to forestry since the 90s, the LCAs are still often based on generic Life Cycle Inventory (LCI). With the purpose of improving LCA practices in the forestry sector, we developed a European Life Cycle Inventory of Forestry Operations (EFO-LCI) and analyzed the available information to check if within the European forestry sector national differences really exist. We classified the European forests on the basis of "Forest Units" (combinations of tree species and silvicultural practices). For each Forest Unit, we constructed the LCI of their forest management practices on the basis of a questionnaire filled out by national silvicultural experts. We analyzed the data reported to evaluate how they vary over Europe and how they affect LCA results and made freely available the inventory data collected for future use. The study shows important variability in rotation length, type of regeneration, amount and assortments of wood products harvested, and machinery used due to the differences in management practices. The existing variability on these activities sensibly affect LCA results of forestry practices and raw wood production. Although it is practically unfeasible to collect site-specific data for all the LCAs involving forest-based products, the use of less generic LCI data of forestry practice is desirable to improve the reliability of the studies. With the release of EFO-LCI we made a step toward the construction of regionalized LCI for the European forestry sector.

Biodiversity and ecosystem functioning relations in European forests depend on environmental context.

The importance of biodiversity in supporting ecosystem functioning is generally well accepted. However, most evidence comes from small-scale studies, and scaling-up patterns of biodiversity-ecosystem functioning (B-EF) remains challenging, in part because the importance of environmental factors in shaping B-EF relations is poorly understood. Using a forest research platform in which 26 ecosystem functions were measured along gradients of tree species richness in six regions across Europe, we investigated the extent and the potential drivers of context dependency of B-EF relations. Despite considerable variation in species richness effects across the continent, we found a tendency for stronger B-EF relations in drier climates as well as in areas with longer growing seasons and more functionally diverse tree species. The importance of water availability in driving context dependency suggests that as water limitation increases under climate change, biodiversity may become even more important to support high levels of functioning in European forests.

The effect of increasing lifespan and recycling rate on carbon storage in wood products from theoretical model to application for the European wood sector.

The use of wood products is often promoted as a climate change mitigation option to reduce atmospheric carbon dioxide concentrations. In previous literature, we identified longevity and recycling rate as two determining factors that influence the carbon stock in wood products, but no studies have predicted the effect of improved wood use on carbon storage over time. In this study, we aimed at evaluating changes in the lifespan and the recycling rate as two options for enhancing carbon stock in wood products for different time horizons. We first explored the behaviour over time of both factors in a theoretical simulation, and then calculated their effect for the European wood sector of the future. The theoretical simulation shows that the carbon stock in wood products increases linearly when increasing the average lifespan of wood products and exponentially when improving the recycling rate. The emissions savings under the current use of wood products in Europe in 2030 were estimated at 57.65 Mt carbon dioxide (CO2) per year. This amount could be increased 5 Mt CO2 if average lifespan increased 19.54 % or if recycling rate increased 20.92 % in 2017. However, the combination of both strategies could increase the emissions saving almost 5 Mt CO2 more by 2030. Incrementing recycling rate of paper and paperboard is the best short-term strategy (2030) to reduce emissions, but elongating average lifespan of wood-based panels is a better strategy for longer term periods (2046).

Modelling carbon stocks and fluxes in the wood product sector: a comparative review.

In addition to forest ecosystems, wood products are carbon pools that can be strategically managed to mitigate climate change. Wood product models (WPMs) simulating the carbon balance of wood production, use and end of life can complement forest growth models to evaluate the mitigation potential of the forest sector as a whole. WPMs can be used to compare scenarios of product use and explore mitigation strategies. A considerable number of WPMs have been developed in the last three decades, but there is no review available analysing their functionality and performance. This study analyses and compares 41 WPMs. One surprising initial result was that we discovered the erroneous implementation of a few concepts and assumptions in some of the models. We further described and compared the models using six model characteristics (bucking allocation, industrial processes, carbon pools, product removal, recycling and substitution effects) and three model-use characteristics (system boundaries, model initialization and evaluation of results). Using a set of indicators based on the model characteristics, we classified models using a hierarchical clustering technique and differentiated them according to their increasing degrees of complexity and varying levels of user support. For purposes of simulating carbon stock in wood products, models with a simple structure may be sufficient, but to compare climate change mitigation options, complex models are needed. The number of models has increased substantially over the last ten years, introducing more diversity and accuracy. Calculation of substitution effects and recycling has also become more prominent. However, the lack of data is still an important constraint for a more realistic estimation of carbon stocks and fluxes. Therefore, if the sector wants to demonstrate the environmental quality of its products, it should make it a priority to provide reliable life cycle inventory data, particularly regarding aspects of time and location.

Environmental impact assessment and monetary ecosystem service valuation of an ecosystem under different future environmental change and management scenarios; a case study of a Scots pine forest.

For a sustainable future, we must sustainably manage not only the human/industrial system but also ecosystems. To achieve the latter goal, we need to predict the responses of ecosystems and their provided services to management practices under changing environmental conditions via ecosystem models and use tools to compare the estimated provided services between the different scenarios. However, scientific articles have covered a limited amount of estimated ecosystem services and have used tools to aggregate services that contain a significant amount of subjective aspects and that represent the final result in a non-tangible unit such as 'points'. To resolve these matters, this study quantifies the environmental impact (on human health, natural systems and natural resources) in physical units and uses an ecosystem service valuation based on monetary values (including ecosystem disservices with associated negative monetary values). More specifically, the paper also focuses on the assessment of ecosystem services related to pollutant removal/generation flows, accounting for the inflow of eutrophying nitrogen (N) when assessing the effect of N leached to groundwater. Regarding water use/provisioning, evapotranspiration is alternatively considered a disservice because it implies a loss of (potential) groundwater. These approaches and improvements, relevant to all ecosystems, are demonstrated using a Scots pine stand from 2010 to 2089 for a combination of three environmental change and three management scenarios. The environmental change scenarios considered interannual climate variability trends and included alterations in temperature, precipitation, nitrogen deposition, wind speed, Particulate matter (PM) concentration and CO2 concentration. The addressed flows/ecosystem services, including disservices, are as follows: particulate matter removal, freshwater loss, CO2 sequestration, wood production, NOx emissions, NH3 uptake and nitrogen pollution/removal. The monetary ecosystem service valuation yields a total average estimate of 361-1242 euro ha(-1) yr(-1). PM2.5 (<2.5 µm) removal is the key service, with a projected value of 622-1172 euro ha(-1) yr(-1). Concerning environmental impact assessment, with net CO2 uptake being the most relevant contributing flow, a loss prevention of 0.014-0.029 healthy life years ha(-1) yr(-1) is calculated for the respective flows. Both assessment methods favor the use of the least intensive management scenario due to its resulting higher CO2 sequestration and PM removal, which are the most important services of the considered ones.

Fragmentation and management of Ethiopian moist evergreen forest drive compositional shifts of insect communities visiting wild Arabica coffee flowers.

Coffea arabica is an indigenous understorey shrub of the moist evergreen Afromontane forest of SW Ethiopia. Coffee cultivation here occurs under different forest management intensities, ranging from almost no intervention in the 'forest coffee' system to far-reaching interventions that include the removal of competing shrubs and selective thinning of the upper canopy in the 'semi-forest coffee' system. We investigated whether increasing forest management intensity and fragmentation result in impacts upon potential coffee pollination services through examining shifts in insect communities that visit coffee flowers. Overall, we netted 2,976 insect individuals on C. arabica flowers, belonging to sixteen taxonomic groups, comprising 10 insect orders. Taxonomic richness of the flower-visiting insects significantly decreased and pollinator community changed with increasing forest management intensity and fragmentation. The relative abundance of honey bees significantly increased with increasing forest management intensity and fragmentation, likely resulting from the introduction of bee hives in the most intensively managed forests. The impoverishment of the insect communities through increased forest management intensity and fragmentation potentially decreases the resilience of the coffee production system as pollination increasingly relies on honey bees alone. This may negatively affect coffee productivity in the long term as global pollination services by managed honey bees are expected to decline under current climate change scenarios. Coffee agroforestry management practices should urgently integrate pollinator conservation measures.

The effect of drought stress on heterozygosity-fitness correlations in pedunculate oak (Quercus robur).

BACKGROUND AND AIMS: The interaction between forest fragmentation and predicted climate change may pose a serious threat to tree populations. In small and spatially isolated forest fragments, increased homozygosity may directly affect individual tree fitness through the expression of deleterious alleles. Climate change-induced drought stress may exacerbate these detrimental genetic consequences of forest fragmentation, as the fitness response to low levels of individual heterozygosity is generally thought to be stronger under environmental stress than under optimal conditions. METHODS: To test this hypothesis, a greenhouse experiment was performed in which various transpiration and growth traits of 6-month-old seedlings of Quercus robur differing in multilocus heterozygosity (MLH) were recorded for 3 months under a well-watered and a drought stress treatment. Heterozygosity-fitness correlations (HFC) were examined by correlating the recorded traits of individual seedlings to their MLH and by studying their response to drought stress. KEY RESULTS: Weak, but significant, effects of MLH on several fitness traits were obtained, which were stronger for transpiration variables than for the recorded growth traits. High atmospheric stress (measured as vapour pressure deficit) influenced the strength of the HFCs of the transpiration variables, whereas only a limited effect of the irrigation treatment on the HFCs was observed. CONCLUSIONS: Under ongoing climate change, increased atmospheric stress in the future may strengthen the negative fitness responses of trees to low MLH. This indicates the necessity to maximize individual multilocus heterozygosity in forest tree breeding programmes.

Multilayered modeling of particulate matter removal by a growing forest over time, from plant surface deposition to washoff via rainfall.

Airborne fine particulate matter (PM) is responsible for the most severe health effects induced by air pollution in Europe. Vegetation, and forests in particular, can play a role in mitigating this pollution since they have a large surface area to filter PM out of the air. Many studies have solely focused on dry deposition of PM onto the tree surface, but deposited PM can be resuspended to the air or may be washed off by precipitation dripping from the plants to the soil. It is only the latter process that represents a net-removal from the atmosphere. To quantify this removal all these processes should be accounted for, which is the case in our modeling framework. Practically, a multilayered PM removal model for forest canopies is developed. In addition, the framework has been integrated into an existing forest growth model in order to account for changes in PM removal efficiency during forest growth. A case study was performed on a Scots pine stand in Belgium (Europe), resulting for 2010 in a dry deposition of 31 kg PM2.5 (PM < 2.5 µm) ha(-1) yr(-1) from which 76% was resuspended and 24% washed off. For different future emission reduction scenarios from 2010 to 2030, with altering PM2.5 air concentration, the avoided health costs due to PM2.5 removal was estimated to range from 915 to 1075 euro ha(-1) yr(-1). The presented model could even be used to predict nutrient input via particulate matter though further research is needed to improve and better validate the model.

Nitrogen saturation and net ecosystem production.

Magnani et al. found that net carbon (C) sequestration of temperate and boreal forests is clearly driven by nitrogen (N) deposition. From the positive relationship between average net ecosystem production (NEP) and wet N deposition, the authors further conclude that "no signs of N saturation were apparent" in the studied forests and that this is "casting doubts on the risk of widespread ecosystem nitrogen saturation". Nitrogen additions can clearly alter net ecosystem production, but net ecosystem production cannot be used as an indicator of N saturation.