Advanced research tools for fungal diversity and its impact on forest ecosystem

Fungi are dominant ecological participants in the forest ecosystems, which play a major role in recycling organic matter and channeling nutrients across trophic levels. Fungal populations are shaped by plant communities and environmental parameters, and in turn, fungal communities also impact the forest ecosystem through intrinsic participation of different fungal guilds. Mycorrhizal fungi result in conservation and stability of forest ecosystem, while pathogenic fungi can bring change in forest ecosystem, by replacing the dominant plant species with new or exotic plant species. Saprotrophic fungi, being ecological regulators in the forest ecosystem, convert dead tree logs into reusable constituents and complete the ecological cycles of nitrogen and carbon. However, fungal communities have not been studied in-depth with respect to functional, spatiotemporal, or environmental parameters. Previously, fungal diversity and its role in shaping the forest ecosystem were studied by traditional and laborious cultural methods, which were unable to achieve real-time results and draw a conclusive picture of fungal communities. This review highlights the latest advances in biological methods such as next-generation sequencing and meta’omics for observing fungal diversity in the forest ecosystem, the role of different fungal groups in shaping forest ecosystem, forest productivity, and nutrient cycling at global scales.

     

Global change and the boreal forest: thresholds, shifting states or gradual change?

Changes in boreal climate of the magnitude projected for the 21st century have always caused vegetation changes large enough to be societally important. However, the rates and patterns of vegetation change are difficult to predict. We review evidence suggesting that these vegetation changes may be gradual at the northern forest limit or where seed dispersal limits species distribution. However, forest composition may be quite resilient to climate change in the central portions of a species range until some threshold is surpassed. At this point, changes can be rapid and unexpected, often causing a switch to very different ecosystem types. Many of these triggers for change are amenable to management, suggesting that our choice of policies in the coming decades will substantially influence the ecological and societal consequences of current climatic change.     

Micro-evolution due to pollution: possible consequences for ecosystem responses to toxic stress

Polluting events can change community structure and ecosystem functioning. Selection of genetically inherited tolerance on exposed populations, here referred as micro-evolution due to pollution, has been recognized as one of the causes of these changes. However, there is a gap between studies addressing this process and those assessing effects at higher levels of biological organization. In this review we attempt to address these evolutionary considerations into the ecological risk assessment (ERA) of polluting events and to trigger the discussion about the consequences of this process for the ecosystem response to toxic stress. We provide clear evidence that pollution drives micro-evolutionary processes in several species. When this process occurs, populations inhabiting environments that become polluted may persist. However, due to the existence of ecological costs derived from the loss of genetic variability, negative pleiotropy with fitness traits and/or from physiological alterations, micro-evolution due to pollution may alter different properties of the affected populations. Despite the existence of empirical evidence showing that safety margins currently applied in the ERA process may account for pollution-induced genetic changes in tolerance, information regarding long-term ecological consequences at higher levels of biological organization due to ecological costs is not explicitly considered in these procedures. In relation to this, we present four testable hypotheses considering that micro-evolution due to pollution acts upon the variability of functional response traits of the exposed populations and generates changes on their functional effect traits, therefore, modifying the way species exploit their ecological niches and participate in the overall ecosystem functioning.     

Urban forests and pollution mitigation: analyzing ecosystem services and disservices

The purpose of this paper is to integrate the concepts of ecosystem services and disservices when assessing the efficacy of using urban forests for mitigating pollution. A brief review of the literature identifies some pollution mitigation ecosystem services provided by urban forests. Existing ecosystem services definitions and typologies from the economics and ecological literature are adapted and applied to urban forest management and the concepts of ecosystem disservices from natural and semi-natural systems are discussed. Examples of the urban forest ecosystem services of air quality and carbon dioxide sequestration are used to illustrate issues associated with assessing their efficacy in mitigating urban pollution. Development of urban forest management alternatives that mitigate pollution should consider scale, contexts, heterogeneity, management intensities and other social and economic co-benefits, tradeoffs, and costs affecting stakeholders and urban sustainability goals.     

Modelling changes in forest soil chemistry at 16 Swedish coniferous forest sites following deposition reduction

The dynamic forest ecosystem model ForSAFE was applied at 16 coniferous forest sites in Sweden to investigate past and future changes in soil chemistry following changes in atmospheric deposition. The simulation shows a considerable historical soil acidification. Acidification in the southwest, where deposition has been greatest, was more expressed in the deepest soil layers, while it was more evenly distributed through the soil profile in central Sweden, and was greater in the upper soil layers in the north. The simulation also shows that a slight recovery took place after the reduction in emissions, but was counteracted by the effect of harvesting. The simulation predicts an increase in the number of acidified sites in the future. The results also suggest that future acidification will be mainly due to the enhanced tree growth resulting from the chronic high deposition of nitrogen and the removal of soil base cations through harvesting.     

Forest responses to tropospheric ozone and global climate change: an analysis

In this paper an analysis is provided on: what we know, what we need to know, and what we need to do, to further our understanding of the relationships between tropospheric ozone (O(3)), global climate change and forest responses. The relationships between global geographic distributions of forest ecosystems and potential geographic regions of high photochemical smog by the year 2025 AD are described. While the emphasis is on the effects of tropospheric O(3) on forest ecosystems, discussion is presented to understand such effects in the context of global climate change. One particular strong point of this paper is the audit of published surface O(3) data by photochemical smog region that reveals important forest/woodland geographic regions where little or no O(3) data exist even though the potential threat to forests in those regions appears to be large. The concepts and considerations relevant to the examination of ecosystem responses as a whole, rather than simply tree stands alone are reviewed. A brief argument is provided to stimulate the modification of the concept of simple cause and effect relationships in viewing total ecosystems. Our knowledge of O(3) exposure and its effects on the energy, nutrient and hydrological flow within the ecosystem are described. Modeling strategies for such systems are reviewed. A discussion of responses of forests to potential multiple climatic changes is provided. An important concept in this paper is that changes in water exchange processes throughout the hydrological cycle can be used as early warning indicators of forest responses to O(3). Another strength of this paper is the integration of information on structural and functional processes of ecosystems and their responses to O(3). An admitted weakness of this analysis is that the information on integrated ecosystem responses is based overwhelmingly on the San Bernardino Forest ecosystem research program of the 1970s because of a lack of similar studies. In the final analysis, it is recommended that systems ecology be applied in examining the joint effects of O(3), carbon dioxide and ultraviolet-B radiation on forest ecosystems.     

Nitrogen dynamics of a mountain forest on dolomitic limestone--a scenario-based risk assessment

The dominant nitrogen (N) fluxes were simulated in a mountain forest ecosystem on dolomitic bedrock in the Austrian Alps. Based on an existing small-scale climate model the simulation encompassed the present situation and a 50-yr projection. The investigated scenarios were current climate, current N deposition (SC1) and future climate (+2.5 degrees C and +10% annual precipitation) with three levels of N deposition (SC2, 3, 4). The microbially mediated N transformation, including the emission of nitrogen oxides, was calculated with PnET-N-DNDC. Soil hydrology was calculated with HYDRUS and was used to estimate the leaching of nitrate. The expected change of the forest ecosystem due to changes of the climate and the N availability was simulated with PICUS. The incentive for the project was the fact that forests on dolomitic limestone stock on shallow Rendzic Leptosols that are rich in soil organic matter are considered highly sensitive to the expected environmental changes. The simulation results showed a strong effect due to increased temperatures and to elevated levels of N deposition. The outflux of N, both as nitrate (6-25kg Nha(-1)yr(-1)) and nitrogen oxides (1-2kg Nha(-1)yr(-1)), from the forest ecosystem are expected to increase. Temperature exerts a stronger effect on the N(2)O emission than the increased rate of N deposition. The main part of the N emission will occur as N(2) (15kg Nha(-1)yr(-1)). The total N loss is partially offset by increased rates of N uptake in the biomass due to an increase in forest productivity.     

The sensitivity of current and future forest managers to climate-induced changes in ecological processes

Climate vulnerability of managed forest ecosystems is not only determined by ecological processes but also influenced by the adaptive capacity of forest managers. To better understand adaptive behaviour, we conducted a questionnaire study among current and future forest managers (i.e. active managers and forestry students) in Austria. We found widespread belief in climate change (94.7 % of respondents), and no significant difference between current and future managers. Based on intended responses to climate-induced ecosystem changes, we distinguished four groups: highly sensitive managers (27.7 %), those mainly sensitive to changes in growth and regeneration processes (46.7 %), managers primarily sensitive to regeneration changes (11.2 %), and insensitive managers (14.4 %). Experiences and beliefs with regard to disturbance-related tree mortality were found to particularly influence a manager’s sensitivity to climate change. Our findings underline the importance of the social dimension of climate change adaptation, and suggest potentially strong adaptive feedbacks between ecosystems and their managers.

Electronic supplementary material

The online version of this article (doi:10.1007/s13280-015-0737-6) contains supplementary material, which is available to authorized users.     

Modeling plant response to tropospheric ozone: a critical review

The advantages and disadvantages, benefits and limitations, of a number of published mathematical models representing the effects of ozone on crops and native vegetation are described. Several levels of modeling are addressed: word models, graphic models, mathematical models, and computer simulation implementation. Special attention is given to evaluating: (1) how the interaction between ozone exposure and vegetation effects is quantified, (2) the status of field testing of the model, and (3) the adequacy of information for enabling other investigators to replicate the model for further testing. Original contributions, not previously published, are made in this evaluation in the form of: (1) graphic model flow charts for published models, (2) clarification of mathematical equations for existing models, (3) graphic forms of functional relations comprising portions of models, and (4) graphic displays of model output performance versus observed data. The models that are evaluated cover acute exposure-response models, statistical and mechanistic-process models, including a partial model of ambient exposure versus ozone flux, and uptake. They also cover chronic exposure statistical approaches, including time-series modeling, mechanistic-process models, 'disintegrated' models of forest system simulations, chronic flux density-uptake-response, and models for regional effects assessment in forests and agricultural lands.     

Sensitivity of a forest ecosystem model to climate parametrization schemes

An analysis of the climate parametrization scheme adopted by conventional forest gap models revealed that most models assume a constant climate and are difficult to calibrate consistently. Tree growth showed unrealistically sensitive threshold effects along ecological gradients of temperature and precipitation. A new parametrization was compared with its predecessors in terms of the model's capability to predict realistic steady state species compositions at three test sites in the Alps. Applying the new model variant ForClim to some climate-change scenarios suggests that forest gap models are highly sensitive to climate pametrizations, regardless of the realism with which they simulate forests for the current climate. Moreover, the precision of climate scenarios based on General Circulation Models (GCM), for example, falls short of ForClim's sensitivity. Climate-dependent processes in forest gap models should be rehearsed before these models are used in impact studies of climatic change.     

A conceptual framework: Redefining forest soil's critical acid loads under a changing climate

Federal agencies of several nations have or are currently developing guidelines for critical forest soil acid loads. These guidelines are used to establish regulations designed to maintain atmospheric acid inputs below levels shown to damage forests and streams. Traditionally, when the critical soil acid load exceeds the amount of acid that the ecosystem can absorb, it is believed to potentially impair forest health. The excess over the critical soil acid load is termed the exceedance, and the larger the exceedance, the greater the risk of ecosystem damage. This definition of critical soil acid load applies to exposure of the soil to a single, long-term pollutant (i.e., acidic deposition). However, ecosystems can be simultaneously under multiple ecosystem stresses and a single critical soil acid load level may not accurately reflect ecosystem health risk when subjected to multiple, episodic environmental stress. For example, the Appalachian Mountains of western North Carolina receive some of the highest rates of acidic deposition in the eastern United States, but these levels are considered to be below the critical acid load (CAL) that would cause forest damage. However, the area experienced a moderate three-year drought from 1999 to 2002, and in 2001 red spruce (Picea rubens Sarg.) trees in the area began to die in large numbers. The initial survey indicated that the affected trees were killed by the southern pine beetle (Dendroctonus frontalis Zimm.). This insect is not normally successful at colonizing these tree species because the trees produce large amounts of oleoresin that exclude the boring beetles. Subsequent investigations revealed that long-term acid deposition may have altered red spruce forest structure and function. There is some evidence that elevated acid deposition (particularly nitrogen) reduced tree water uptake potential, oleoresin production, and caused the trees to become more susceptible to insect colonization during the drought period. While the ecosystem was not in exceedance of the CAL, long-term nitrogen deposition pre-disposed the forest to other ecological stress. In combination, insects, drought, and nitrogen ultimately combined to cause the observed forest mortality. If any one of these factors were not present, the trees would likely not have died. This paper presents a conceptual framework of the ecosystem consequences of these interactions as well as limited plot level data to support this concept. Future assessments of the use of CAL studies need to account for multiple stress impacts to better understand ecosystem response.     

High Arctic ecosystem states: Conceptual models of vegetation change to guide long-term monitoring and research

Vegetation change has consequences for terrestrial ecosystem structure and functioning and may involve climate feedbacks. Hence, when monitoring ecosystem states and changes thereof, the vegetation is often a primary monitoring target. Here, we summarize current understanding of vegetation change in the High Arctic—the World’s most rapidly warming region—in the context of ecosystem monitoring. To foster development of deployable monitoring strategies, we categorize different kinds of drivers (disturbances or stresses) of vegetation change either as pulse (i.e. drivers that occur as sudden and short events, though their effects may be long lasting) or press (i.e. drivers where change in conditions remains in place for a prolonged period, or slowly increases in pressure). To account for the great heterogeneity in vegetation responses to climate change and other drivers, we stress the need for increased use of ecosystem-specific conceptual models to guide monitoring and ecological studies in the Arctic. We discuss a conceptual model with three hypothesized alternative vegetation states characterized by mosses, herbaceous plants, and bare ground patches, respectively. We use moss-graminoid tundra of Svalbard as a case study to discuss the documented and potential impacts of different drivers on the possible transitions between those states. Our current understanding points to likely additive effects of herbivores and a warming climate, driving this ecosystem from a moss-dominated state with cool soils, shallow active layer and slow nutrient cycling to an ecosystem with warmer soil, deeper permafrost thaw, and faster nutrient cycling. Herbaceous-dominated vegetation and (patchy) bare ground would present two states in response to those drivers. Conceptual models are an operational tool to focus monitoring efforts towards management needs and identify the most pressing scientific questions. We promote greater use of conceptual models in conjunction with a state-and-transition framework in monitoring to ensure fit for purpose approaches. Defined expectations of the focal systems’ responses to different drivers also facilitate linking local and regional monitoring efforts to international initiatives, such as the Circumpolar Biodiversity Monitoring Program.     

Representative boreal forest habitats in northern Europe,and a revised model for ecosystem management and biodiversity conservation

The natural range of variation of ecosystems provides reference conditions for sustainable management and biodiversity conservation. We review how the understanding of natural reference conditions of boreal forests in northern Europe has changed from earlier perceptions of even-aged dynamics driven by stand-replacing disturbances towards current understanding highlighting the role of non-stand-replacing disturbances and the resultant complex forest dynamics and structures. We show how earlier views and conceptual models of forest disturbance dynamics, including the influential ASIO model, provide estimates of reference conditions that are outside the natural range of variation. Based on a research synthesis, we present a revised forest reference model incorporating the observed complexity of ecosystem dynamics and the prevalence of old forests. Finally, we outline a management model and demonstrate its use in forest ecosystem management and show how regional conservation area needs can be estimated. We conclude that attaining favourable conservation status in northern Europe’s boreal forests requires increasing emphasis on ecosystem management and conservation for old forest characteristics.Electronic supplementary materialThe online version of this article (10.1007/s13280-020-01444-3) contains supplementary material, which is available to authorized users.     

Comparison of carbon balances between continuous-cover and clear-cut forestry in Sweden

Continuous-cover forestry (CCF) has been recognized for the production of multiple ecosystem services, and is seen as an alternative to clear-cut forestry (CF). Despite the increasing interest, it is still not well described how CCF would affect the carbon balance and the resulting climate benefit from the forest in relation to CF. This study compares carbon balances of CF and CCF, applied as two alternative land-use strategies for a heterogeneous Norway spruce (Picea abies) stand. We use a set of models to analyze the long-term effects of different forest management and wood use strategies in Sweden on carbon dioxide emissions and carbon stock changes. The results show that biomass growth and yield is more important than the choice of silvicultural system per se. When comparing CF and CCF assuming similar growth, extraction and product use, only minor differences in long-term climate benefit were found between the two principally different silvicultural systems.     

Reconnecting Cities to the Biosphere: Stewardship of Green Infrastructure and Urban Ecosystem Services

Within-city green infrastructure can offer opportunities and new contexts for people to become stewards of ecosystem services. We analyze cities as social–ecological systems, synthesize the literature, and provide examples from more than 15 years of research in the Stockholm urban region, Sweden. The social–ecological approach spans from investigating ecosystem properties to the social frameworks and personal values that drive and shape human interactions with nature. Key findings demonstrate that urban ecosystem services are generated by social–ecological systems and that local stewards are critically important. However, land-use planning and management seldom account for their role in the generation of urban ecosystem services. While the small scale patchwork of land uses in cities stimulates intense interactions across borders much focus is still on individual patches. The results highlight the importance and complexity of stewardship of urban biodiversity and ecosystem services and of the planning and governance of urban green infrastructure.     

The carbon budget of Canadian forests: a sensitivity analysis of changes in disturbance regimes, growth rates, and decomposition rates

Ecosystem responses to climate changes will affect the exchange of carbon (C) with the atmosphere, thus providing feedback for future climate response. We have developed a C budget model of Canadian forests and forest sector activities and used sensitivity analysis runs with changes in productivity, decomposition, and disturbance regimes to assess the sensitivity of the Canadian forest sector C budget over the next century. The model operates on data derived from Canada's National Forest Biomass Inventory, from the Oak Ridge National Laboratory global soil C data base, and from Canadian data bases that document areas annually disturbed by fire, insects, and harvesting. It simulates the dynamics of biomass and soil C pools (including detritus and coarse woody debris) as they are affected by growth, decomposition, and disturbances. For the reference run of the model, we assumed unchanging climate and disturbance regimes. Under these conditions, total ecosystem C increased by 2 Gt C (2.3%) over the 100-year simulation period. In the sensitivity analysis, we explored the effects of changes in the area annually disturbed by fire and insect-induced stand mortality (-60 to +300%), growth rates (-10 to +20%), decomposition rates (-10 to +25%), and combined changes in growth and decomposition rates. In every model run, the change of total ecosystem C relative to the reference run was less than 10%. Combined changes to growth and decomposition rates yielded very small deviations from the results of the reference run (-0.8 to +1.2%). Because disturbance regime changes affect forest age-class structure as well as forest dynamics, they are expected to affect C budgets strongly. Total ecosystem C, however, is slightly more sensitive to changes in growth and decomposition parameters than to changes in disturbance regimes. Although the sensitivity analysis results suggest that C budgets are little affected by the range of parameter changes implemented here, we must emphasize that our sensitivity analyses do not account for potentially important processes, such as regeneration failure or the shifts in forest distribution.     

Past, current and future fire frequency in the Canadian boreal forest: implications for sustainable forest management

Over the past decades, there has been an increasing interest in the development of forest management approaches that are based on an understanding of historical natural disturbance dynamics. The rationale for such an approach is that management to favor landscape compositions and stand structures similar to those of natural ecosystems should also maintain biological diversity and essential ecological functions. In fire-dominated landscapes, this approach is possible only if current and future fire frequencies are sufficiently low, comparing to pre-industrial fire frequency, that we can substitute fire by forest management. We address this question by comparing current and future fire frequency to historical reconstruction of fire frequency from studies realized in the Canadian boreal forest. Current and simulated future fire frequencies using 2 and 3 x CO2 scenarios are lower than the historical fire frequency for many sites, suggesting that forest management could potentially be used to recreate the forest age structure of fire-controlled pre-industrial landscapes. There are however, important limitations to the current even-age management.     

Advances of air pollution science: From forest decline to multiple-stress effects on forest ecosystem services

Over the past 20 years, the focus of forest science on air pollution has moved from forest decline to a holistic framework of forest health, and from the effects on forest production to the ecosystem services provided by forest ecosystems. Hence, future research should focus on the interacting factorial impacts and resulting antagonistic and synergistic responses of forest trees and ecosystems. The synergistic effects of air pollution and climatic changes, in particular elevated ozone, altered nitrogen, carbon and water availability, must be key issues for research. Present evidence suggests air pollution will become increasingly harmful to forests under climate change, which requires integration amongst various stressors (abiotic and biotic factors, including competition, parasites and fire), effects on forest services (production, biodiversity protection, soil protection, sustained water balance, socio-economical relevance) and assessment approaches (research, monitoring, modeling) to be fostered.     

Marine ecosystem modeling beyond the box: using GIS to study carbon fluxes in a coastal ecosystem

Studies of carbon fluxes in marine ecosystems are often done by using box model approaches with basin size boxes, or highly resolved 3D models, and an emphasis on the pelagic component of the ecosystem. Those approaches work well in the ocean proper, but can give rise to considerable problems when applied to coastal systems, because of the scale of certain ecological niches and the fact that benthic organisms are the dominant functional group of the ecosystem. In addition, 3D models require an extensive modeling effort. In this project, an intermediate approach based on a high resolution (20x20 m) GIS data-grid has been developed for the coastal ecosystem in the Laxemar area (Baltic Sea, Sweden) based on a number of different site investigations. The model has been developed in the context of a safety assessment project for a proposed nuclear waste repository, in which the fate of hypothetically released radionuclides from the planned repository is estimated. The assessment project requires not only a good understanding of the ecosystem dynamics at the site, but also quantification of stocks and flows of matter in the system. The data-grid was then used to set up a carbon budget describing the spatial distribution of biomass, primary production, net ecosystem production and thus where carbon sinks and sources are located in the area. From these results, it was clear that there was a large variation in ecosystem characteristics within the basins and, on a larger scale, that the inner areas are net producing and the outer areas net respiring, even in shallow phytobenthic communities. Benthic processes had a similar or larger influence on carbon fluxes as advective processes in inner areas, whereas the opposite appears to be true in the outer basins. As many radionuclides are expected to follow the pathways of organic matter in the environment, these findings enhance our abilities to realistically describe and predict their fate in the ecosystem.     

Mercury Concentrations in Lentic Fish Populations Related to Ecosystem and Watershed Characteristics

Predicting mercury (Hg) concentrations of fishes at large spatial scales is a fundamental environmental challenge with the potential to improve human health. In this study, mercury concentrations were examined for five species across 161 lakes and ecosystem, and watershed parameters were investigated as explanatory variables in statistical models. For all species, Hg concentrations were significantly, positively related to wetland coverage. For three species (largemouth bass, pike, and walleye), Hg concentrations were significantly, negatively related to lake trophic state index (TSI), suggestive of growth biodilution. There were no significant relationships between ecosystem size and mercury concentrations. However, Hg concentrations were strongly, positively related to ecosystem size across species. Scores of small or remote lakes that have never been tested could be prioritized for testing using models akin to those presented in this article. Such an approach could also be useful for exploring how Hg concentrations of fishes might respond to natural or anthropogenic changes to ecosystems over time.