Site fertility and soil water-table level affect fungal biomass production and community composition in boreal peatland forests.

A substantial amount of below-ground carbon (C) is suggested to be associated with fungi, which may significantly affect the soil C balance in forested ecosystems. Ergosterol from in-growth mesh bags and litterbags was used to estimate fungal biomass production and community composition in drained peatland forests with differing fertility. Extramatrical mycelia (EMM) biomass production was generally higher in the nutrient-poor site, increased with deeper water table level and decreased along the length of the recovery time. EMM biomass production was of the same magnitude as in mineral-soil forests. Saprotrophic fungal biomass production was higher in the nutrient-rich site. Both ectomycorrhizal (ECM) and saprotrophic fungal community composition changed according to site fertility and water table level. ECM fungal community composition with different exploration types may explain the differences in fungal biomass production between peatland forests. Melanin-rich Hyaloscypha may indicate decreased turnover of biomass in nutrient-rich young peatland forest. Genera Lactarius and Laccaria may be important in nutrient rich and Piloderma in the nutrient-poor conditions, respectively. Furthermore, Paxillus involutus and Cortinarius sp. may be important generalists in all sites and responsible for EMM biomass production during the first summer months. Saprotrophs showed a functionally more diverse fungal community in the nutrient-rich site.

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

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

Warming impacts on boreal fen CO2 exchange under wet and dry conditions.

Northern peatlands form a major soil carbon (C) stock. With climate change, peatland C mineralization is expected to increase, which in turn would accelerate climate change. A particularity of peatlands is the importance of soil aeration, which regulates peatland functioning and likely modulates the responses to warming climate. Our aim is to assess the impacts of warming on a southern boreal and a sub-arctic sedge fen carbon dioxide (CO2 ) exchange under two plausible water table regimes: wet and moderately dry. We focused this study on minerotrophic treeless sedge fens, as they are common peatland types at boreal and (sub)arctic areas, which are expected to face the highest rates of climate warming. In addition, fens are expected to respond to environmental changes faster than the nutrient poor bogs. Our study confirmed that CO2 exchange is more strongly affected by drying than warming. Experimental water level draw-down (WLD) significantly increased gross photosynthesis and ecosystem respiration. Warming alone had insignificant impacts on the CO2 exchange components, but when combined with WLD it further increased ecosystem respiration. In the southern fen, CO2 uptake decreased due to WLD, which was amplified by warming, while at northern fen it remained stable. As a conclusion, our results suggest that a very small difference in the WLD may be decisive, whether the C sink of a fen decreases, or whether the system is able to adapt within its regime and maintain its functions. Moreover, the water table has a role in determining how much the increased temperature impacts the CO2 exchange.

Responses of phenology and biomass production of boreal fens to climate warming under different water-table level regimes.

Climate change affects peatlands directly through increased air temperatures and indirectly through changes in water-table level (WL). The interactions of these two still remain poorly known. We determined experimentally the separate and interactive effects of temperature and WL regime on factors of relevance for the inputs to the carbon cycle: plant community composition, phenology, biomass production, and shoot:root allocation in two wet boreal sedge-dominated fens, "southern" at 62°N and "northern" at 68°Ν. Warming (1.5°C higher average daily air temperature) was induced with open-top chambers and WL drawdown (WLD; 3-7 cm on average) by shallow ditches. Total biomass production varied from 250 to 520 g/m2 , with belowground production comprising 25%-63%. Warming was associated with minor effects on phenology and negligible effects on community composition, biomass production, and allocation. WLD clearly affected the contribution of different plant functional types (PFTs) in the community and the biomass they produced: shrubs benefited while forbs and mosses suffered. These responses did not depend on the warming treatment. Following WLD, aboveground biomass production decreased mainly due to reduced growth of mosses in the southern fen. Aboveground vascular plant biomass production remained unchanged but the contribution of different PFTs changed. The observed changes were also reflected in plant phenology, with different PFTs showing different responses. Belowground production increased following WLD in the northern fen only, but an increase in the contributions of shrubs and forbs was observed in both sites, while sedge contribution decreased. Moderate warming alone seems not able to drive significant changes in plant productivity or community composition in these wet ecosystems. However, if warming is accompanied by even modest WL drawdown, changes should be expected in the relative contribution of PFTs, which could lead to profound changes in the function of fens. Consequently, hydrological scenarios are of utmost importance when estimating their future function.

A synthesis of methane emissions from 71 northern, temperate, and subtropical wetlands.

Wetlands are the largest natural source of atmospheric methane. Here, we assess controls on methane flux using a database of approximately 19 000 instantaneous measurements from 71 wetland sites located across subtropical, temperate, and northern high latitude regions. Our analyses confirm general controls on wetland methane emissions from soil temperature, water table, and vegetation, but also show that these relationships are modified depending on wetland type (bog, fen, or swamp), region (subarctic to temperate), and disturbance. Fen methane flux was more sensitive to vegetation and less sensitive to temperature than bog or swamp fluxes. The optimal water table for methane flux was consistently below the peat surface in bogs, close to the peat surface in poor fens, and above the peat surface in rich fens. However, the largest flux in bogs occurred when dry 30-day averaged antecedent conditions were followed by wet conditions, while in fens and swamps, the largest flux occurred when both 30-day averaged antecedent and current conditions were wet. Drained wetlands exhibited distinct characteristics, e.g. the absence of large flux following wet and warm conditions, suggesting that the same functional relationships between methane flux and environmental conditions cannot be used across pristine and disturbed wetlands. Together, our results suggest that water table and temperature are dominant controls on methane flux in pristine bogs and swamps, while other processes, such as vascular transport in pristine fens, have the potential to partially override the effect of these controls in other wetland types. Because wetland types vary in methane emissions and have distinct controls, these ecosystems need to be considered separately to yield reliable estimates of global wetland methane release.

Active afforestation of drained peatlands is not a viable option under the EU Nature Restoration Law.

The EU Nature Restoration Law (NRL) is critical for the restoration of degraded ecosystems and active afforestation of degraded peatlands has been suggested as a restoration measure under the NRL. Here, we discuss the current state of scientific evidence on the climate mitigation effects of peatlands under forestry. Afforestation of drained peatlands without restoring their hydrology does not fully restore ecosystem functions. Evidence on long-term climate benefits is lacking and it is unclear whether CO2 sequestration of forest on drained peatland can offset the carbon loss from the peat over the long-term. While afforestation may offer short-term gains in certain cases, it compromises the sustainability of peatland carbon storage. Thus, active afforestation of drained peatlands is not a viable option for climate mitigation under the EU Nature Restoration Law and might even impede future rewetting/restoration efforts. Instead, restoring hydrological conditions through rewetting is crucial for effective peatland restoration.

Fine-root biomass production and its contribution to organic matter accumulation in sedge fens under changing climate.

Climate change may affect the carbon sink function of peatlands through warming and drying. Fine-root biomass production (FRBP) of sedge fens, a widespread peatland habitat, is important in this context, since most of the biomass is below ground in these ecosystems. We examined the response of fine-root biomass production, depth distribution (10 cm intervals down to 60 cm), chemical characteristics, and decomposition along with other main litter types (sedge leaves, Sphagnum moss shoots) to an average May-to-October warming of 1.7 °C above ambient daily mean temperature and drying of 2-8 cm below ambient soil water-table level (WL) in two sedge fens situated in Northern and Southern Boreal zones. Warming was induced with open top chambers and drying with shallow ditching. Finally, we simulated short-term organic matter (OM) accumulation using net primary production and mass loss data. Total FRBP, and FRBP in deeper layers, was clearly higher in southern than northern fen. Drying significantly increased, and warming marginally increased, total FRBP, while warming significantly increased, and drying marginally increased, the proportional share of FRBP in deeper layers. Drying, especially, modified root chemistry as the relative proportions of fats, wax, lipids, lignin and other aromatics increased while the proportion of polysaccharides decreased. Warming did not affect the decomposition of any litter types, while drying reduced the decomposition of sedge leaf litter. Although drying increased OM accumulation from root litter at both fens, total OM accumulation decreased at the southern fen, while the northern fen with overall lower values showed no such pattern. Our results suggest that in warmer and/or modestly drier conditions, sedge fen FRBP will increase and/or be allocated to deeper soil layers. These changes along with the altered litter inputs may sustain the soil carbon sink function through OM accumulation, unless the WL falls below a tipping point.

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

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

Quantification of Plant Root Species Composition in Peatlands Using FTIR Spectroscopy.

Evidence of plant root biomass and production in peatlands at the level of species or plant functional type (PFT) is needed for defining ecosystem functioning and predicting its future development. However, such data are limited due to methodological difficulties and the toilsomeness of separating roots from peat. We developed Fourier transform infrared (FTIR) spectroscopy based calibration models for quantifying the mass proportions of several common peatland species, and alternatively, the PFTs that these species represented, in composite root samples. We further tested whether woody roots could be classified into diameter classes, and whether dead and living roots could be separated. We aimed to solve whether general models applicable in different studies can be developed, and what would be the best way to build such models. FTIR spectra were measured from dried and powdered roots: both "pure roots", original samples of 25 species collected in the field, and "root mixtures", artificial composite samples prepared by mixing known amounts of pure roots of different species. Partial least squares regression was used to build the calibration models. The general applicability of the models was tested using roots collected in different sites or times. Our main finding is that pure roots can replace complex mixtures as calibration data. Using pure roots, we constructed generally applicable models for quantification of roots of the main PFTs of northern peatlands. The models provided accurate estimates even for far distant sites, with root mean square error (RMSE) 1.4-6.6% for graminoids, forbs and ferns. For shrubs and trees the estimates were less accurate due to higher within-species heterogeneity, partly related to variation in root diameter. Still, we obtained RMSE 3.9-10.8% for total woody roots, but up to 20.1% for different woody-root types. Species-level and dead-root models performed well within the calibration dataset but provided unacceptable estimates for independent samples, limiting their routine application in field conditions. Our PFT-level models can be applied on roots separated from soil for biomass determination or from ingrowth cores for estimating root production. We present possibilities for further development of species-level or dead-root models using the pure-root approach.

Responses of aerobic microbial communities and soil respiration to water-level drawdown in a northern boreal fen.

On a global basis, peatlands are a major reserve of carbon (C). Hydrological changes can affect the decomposition processes in peatlands and in turn can alter their C balance. Since 1959, a groundwater extraction plant has generated a water-level gradient at our study site that has gradually changed part of the wet fen into a dry peatland forest. The average water-level drawdown of the gradient (from a pristine 9 cm to 26 cm in the dry end) is close to an estimate predicted by an increase in mean global temperature of 3 degrees C. We studied the total microbial community of the aerobic surface peat in four locations along the gradient through phospholipid fatty acid and PCR-DGGE methods. Additionally, field measurements of soil respiration showed a threefold increase in the C-emission rate at the driest location compared with the wettest one, indicating enhanced decomposition. Also, both fungal and bacterial biomass increased in the drier locations. At the species level, the fungal community changed due to water-level drawdown whereas actinobacteria were less sensitive to drying. The majority of fungal sequences were similar to ectomycorrhizal (ECM) fungi, which dominated throughout the gradient. Our results indicate that ECM fungi might act as important facultative decomposers in organic-rich environments such as peatlands.

Microbial ecology in a future climate: effects of temperature and moisture on microbial communities of two boreal fens.

Impacts of warming with open-top chambers on microbial communities in wet conditions and in conditions resulting from moderate water-level drawdown (WLD) were studied across 0-50 cm depth in northern and southern boreal sedge fens. Warming alone decreased microbial biomass especially in the northern fen. Impact of warming on microbial PLFA and fungal ITS composition was more obvious in the northern fen and linked to moisture regime and sample depth. Fungal-specific PLFA increased in the surface peat in the drier regime and decreased in layers below 10 cm in the wet regime after warming. OTUs representing Tomentella and Lactarius were observed in drier regime and Mortierella in wet regime after warming in the northern fen. The ectomycorrhizal fungi responded only to WLD. Interestingly, warming together with WLD decreased archaeal 16S rRNA copy numbers in general, and fungal ITS copy numbers in the northern fen. Expectedly, many results indicated that microbial response on warming may be linked to the moisture regime. Results indicated that microbial community in the northern fen representing Arctic soils would be more sensitive to environmental changes. The response to future climate change clearly may vary even within a habitat type, exemplified here by boreal sedge fen.

Quantifying beetle-mediated effects on gas fluxes from dung pats.

Agriculture is one of the largest contributors of the anthropogenic greenhouse gases (GHGs) responsible for global warming. Measurements of gas fluxes from dung pats suggest that dung is a source of GHGs, but whether these emissions are modified by arthropods has not been studied. A closed chamber system was used to measure the fluxes of carbon dioxide (CO2), methane (CH4) and nitrous oxide (N2O) from dung pats with and without dung beetles on a grass sward. The presence of dung beetles significantly affected the fluxes of GHGs from dung pats. Most importantly, fresh dung pats emitted higher amounts of CO2 and lower amounts of CH4 per day in the presence than absence of beetles. Emissions of N2O showed a distinct peak three weeks after the start of the experiment--a pattern detected only in the presence of beetles. When summed over the main grazing season (June-July), total emissions of CH4 proved significantly lower, and total emissions of N2O significantly higher in the presence than absence of beetles. While clearly conditional on the experimental conditions, the patterns observed here reveal a potential impact of dung beetles on gas fluxes realized at a small spatial scale, and thereby suggest that arthropods may have an overall effect on gas fluxes from agriculture. Dissecting the exact mechanisms behind these effects, mapping out the range of conditions under which they occur, and quantifying effect sizes under variable environmental conditions emerge as key priorities for further research.