Peat formation potential of temperate fens increases with hydrological stability.

Peat formation is the key process responsible for carbon sequestration in peatlands. In rich fens, peat is formed by brown mosses and belowground biomass of vascular plants. However, the impact of ecohydrological settings on the contribution of mosses and belowground biomass to peat formation remains an open question. We established seven transects in well-preserved fens in NE Poland along an ecohydrological gradient from mesotrophic sedge-moss communities with stable water levels, to more eutrophic tall sedge communities with higher water level fluctuations. In each transect, we measured the production of brown mosses (using the plug method), aboveground vascular plant biomass (one year after cutting) and belowground biomass (using ingrowth cores). Decomposition rates of all biomass fractions were assessed using litter bags. The first-year surplus of potentially peat-forming fractions, i.e., mosses and belowground biomass, decreased with increasing water level fluctuations and along a vegetation gradient from sedge-moss to tall sedge communities. Moss production was highest in the sedge-moss fen with a stable water level at the ground surface. We did not detect any difference in belowground biomass production across the gradient but found it to be consistently higher in the upper 0-5 cm than in the deeper layers. The decomposition rate also showed no response to the gradient, but differed between biomass types, with aboveground biomass of vascular plants decomposing 2.5 times faster than belowground biomass and mosses. Pattern of peat formation potential along the ecohydrological gradient in rich fen was strongly driven by brown moss production. Sedge-moss fens with a stable water level at the ground surface have the highest peat formation capacity compared to other vegetation types. In the part of the gradient that is poorer in nutrients, vascular plants invest in belowground production, and mosses dominate the aboveground layer.

Stable isotopes (δ13C, δ15N) and biomarkers as indicators of the hydrological regime of fens in a European east-west transect.

Peatland degradation is tightly connected to hydrological changes and microbial metabolism. To better understand these metabolism processes, more information is needed on how microbial communities and substrate cycling are affected by changing hydrological regimes. These activities should be imprinted in stable isotope bulk values (δ 15N, δ 13C) due to specific isotopic fractionation by different microbial communities, their metabolic pathways and nutrient sources. We hypothesize that stable isotope values and microbial abundance are correlated and act as indicators of different hydrological regimes. We sampled an East-West transect across European fens in 14 areas and conducted a stable isotope (δ 13C, δ 15N) and membrane fatty acid (mFA) analysis. Within each area an undrained, drained and rewetted site was selected. Rewetted sites were separated based on when rewetting occurred. We found differences in the upper layers of all sites in microbial-derived mFAs and stable isotope values corresponding to hydrological regimes. The highest and lowest quantities of microbial-derived mFAs were measured in undrained and drained sites, respectively. Fungal-derived mFAs were especially lower in drained sites. Simultaneously, δ15N stable isotope values were highest in drained sites. In addition, stable isotope values and microbial-derived mFAs showed distinct depth trends. In undrained sites stable isotopes values slightly increased with depth. In drained sites, δ15N values decreased downwards, whereas δ13C values increased. Overall microbial-derived mFAs decreased with depth. These patterns presumably result from anoxic conditions and high peat recalcitrance in the deeper layers. In sites with short time of rewetting, the microbial-derived mFAs and stable isotope values were similar to values of drained sites, while with increasing rewetting time values shifted to those of undrained sites. We conclude that biomarkers indicate that stable isotope values reflect specific microbial metabolic processes, which differ with hydrological regimes, and thus could indicate both drainage and rewetting in fens.

Can nutrient uptake by Carex counteract eutrophication in fen peatlands?

Ground- and surface-water-fed peatlands (i.e., fens) of temperate Europe face high anthropogenic nutrient loads from atmospheric deposition, agricultural catchment areas, and from peat decomposition, if drained. As a result, nitrogen loads may exceed a fen's natural nutrient removal capacity, leading to increased eutrophication of adjacent water bodies. Therefore, it is important to address possible means to decrease a fen's nutrient load, including nutrient uptake by fen plants. To assess how much fen plants can contribute to nutrient removal by uptake, nutrient stocks of above- and below-ground biomass need to be quantified. Therefore, we investigated nitrogen, phosphorous, and potassium uptake capacities of sedges (Carex species), which are common dominants in fen plant communities. We grew specimens of five Carex species with varying preferences in nutrient availability under controlled, different nutrient levels. We show that Carex above-ground biomass harvest can remove up to one third of a system's total nitrogen even at high loads of about 40 g nitrogen m-2. Species-specific differences in biomass production, rather than preferences in nutrient availability under natural conditions, were drivers of standing nutrient stocks: Highly productive species, i.e., C. acutiformis and C. rostrata, had highest nutrient standing stocks across all nutrient levels. Amounts of nutrients stored in shoots increased almost linearly with increasing nutrient levels, whereas below-ground nutrient stocks species-specifically increased, saturated, or decreased, with increasing nutrient levels. As a rough estimate, depending on the species, 6-16 cycles of annual above-ground harvest would suffice to decrease nitrogen concentrations from the highest to the lowest level used in this study. Overall, our results indicate that Carex biomass harvest can be an efficient means to counteract anthropogenic nitrogen eutrophication in fens.

Recovery of fen peatland microbiomes and predicted functional profiles after rewetting.

Many of the world's peatlands have been affected by water table drawdown and subsequent loss of organic matter. Rewetting has been proposed as a measure to restore peatland functioning and to halt carbon loss, but its effectiveness is subject to debate. An important prerequisite for peatland recovery is a return of typical microbial communities, which drive key processes. To evaluate the effect of rewetting, we investigated 13 fen peatland areas across a wide (>1500 km) longitudinal gradient in Europe, in which we compared microbial communities between drained, undrained, and rewetted sites. There was a clear difference in microbial communities between drained and undrained fens, regardless of location. Community recovery upon rewetting was substantial in the majority of sites, and predictive functional profiling suggested a concomitant recovery of biogeochemical peatland functioning. However, communities in rewetted sites were only similar to those of undrained sites when soil organic matter quality (as expressed by cellulose fractions) and quantity were still sufficiently high. We estimate that a minimum organic matter content of ca. 70% is required to enable microbial recovery. We conclude that peatland recovery after rewetting is conditional on the level of drainage-induced degradation: severely altered physicochemical peat properties may preclude complete recovery for decades.

Are we restoring functional fens? - The outcomes of restoration projects in fens re-analysed with plant functional traits.

In peatland restoration we often lack an information whether re-established ecosystems are functionally similar to non-degraded ones. We re-analysed the long-term outcomes of restoration on vegetation and plant functional traits in 38 European fens restored by rewetting (18 sites) and topsoil removal (20 sites). We used traits related to nutrient acquisition strategies, competitiveness, seed traits, and used single- and multi-trait metrics. A separate set of vegetation records from near-natural fens with diverse plant communities was used to generate reference values to aid the comparisons. We found that both restoration methods enhanced the similarity of species composition to non-degraded systems but trait analysis revealed differences between the two approaches. Traits linked to nutrient acquisition strategies indicated that topsoil removal was more effective than rewetting. After topsoil removal competitive species in plant communities had decreased, while stress-tolerant species had increased. A substantial reduction in nutrient availability ruled out the effect of initial disturbance. An ability to survive and grow in anoxic conditions was enhanced after restoration, but the reference values were not achieved. Rewetting was more effective than topsoil removal in restricting variation in traits values permitted in re-developing vegetation. We found no indication of a shift towards reference in seed traits, which suggested that dispersal constraint and colonization deficit can be a widespread phenomena. Two functional diversity indices: functional richness and functional dispersion showed response to restoration and shifted values towards reference mires and away from the degraded systems. We concluded that targeting only one type of environmental stressor does not lead to a recovery of fens, as it provides insufficient level of stress to restore a functional ecosystem. In general, restoration efforts do not ensure the re-establishment and long-term persistence of fens. Restoration efforts result in recovery of fen ecosystems, confirmed with our functional trait analysis, although more rigid actions are needed for restoring fully functional mires, by achieving high and constant levels of anoxia and nutrient stresses.

Soil Iron Content as a Predictor of Carbon and Nutrient Mobilization in Rewetted Fens.

Rewetted, previously drained fens often remain sources rather than sinks for carbon and nutrients. To date, it is poorly understood which soil characteristics stimulate carbon and nutrient mobilization upon rewetting. Here, we assess the hypothesis that a large pool of iron in the soil negatively affects fen restoration success, as flooding-induced iron reduction (Fe3+ to Fe2+) causes a disproportionate breakdown of organic matter that is coupled with a release of inorganic compounds. We collected intact soil cores in two iron-poor and two iron-rich drained fens, half of which were subjected to a rewetting treatment while the other half was kept drained. Prolonged drainage led to the mobilization of nitrate (NO3-, > 1 mmol L-1) in all cores, regardless of soil iron content. In the rewetted iron-rich cores, a sharp increase in pore water iron (Fe) concentrations correlated with concentrations of inorganic carbon (TIC, > 13 mmol L-1) and dissolved organic carbon (DOC, > 16 mmol L-1). Additionally, ammonium (NH4+) accumulated up to phytotoxic concentrations of 1 mmol L-1 in the pore water of the rewetted iron-rich cores. Disproportionate mobilization of Fe, TIC, DOC and NH4+ was absent in the rewetted iron-poor cores, indicating a strong interaction between waterlogging and iron-mediated breakdown of organic matter. Concentrations of dissolved phosphorus (P) rose slightly in all cores upon rewetting, but remained low throughout the experiment. Our results suggest that large pools of iron in the top soil of drained fens can hamper the restoration of the fen's sink-service for ammonium and carbon upon rewetting. We argue that negative effects of iron should be most apparent in fens with fluctuating water levels, as temporary oxygenation allows frequent regeneration of Fe3+. We conclude that rewetting of iron-poor fens may be more feasible for restoration.