Increasing wildfires threaten historic carbon sink of boreal forest soils.

Boreal forest fires emit large amounts of carbon into the atmosphere primarily through the combustion of soil organic matter1-3. During each fire, a portion of this soil beneath the burned layer can escape combustion, leading to a net accumulation of carbon in forests over multiple fire events4. Climate warming and drying has led to more severe and frequent forest fires5-7, which threaten to shift the carbon balance of the boreal ecosystem from net accumulation to net loss1, resulting in a positive climate feedback8. This feedback will occur if organic-soil carbon that escaped burning in previous fires, termed 'legacy carbon', combusts. Here we use soil radiocarbon dating to quantitatively assess legacy carbon loss in the 2014 wildfires in the Northwest Territories of Canada2. We found no evidence for the combustion of legacy carbon in forests that were older than the historic fire-return interval of northwestern boreal forests9. In forests that were in dry landscapes and less than 60 years old at the time of the fire, legacy carbon that had escaped burning in the previous fire cycle was combusted. We estimate that 0.34 million hectares of young forests (<60 years) that burned in the 2014 fires could have experienced legacy carbon combustion. This implies a shift to a domain of carbon cycling in which these forests become a net source-instead of a sink-of carbon to the atmosphere over consecutive fires. As boreal wildfires continue to increase in size, frequency and intensity7, the area of young forests that experience legacy carbon combustion will probably increase and have a key role in shifting the boreal carbon balance.

Persistent net release of carbon dioxide and methane from an Alaskan lowland boreal peatland complex.

Permafrost degradation in peatlands is altering vegetation and soil properties and impacting net carbon storage. We studied four adjacent sites in Alaska with varied permafrost regimes, including a black spruce forest on a peat plateau with permafrost, two collapse scar bogs of different ages formed following thermokarst, and a rich fen without permafrost. Measurements included year-round eddy covariance estimates of net carbon dioxide (CO2 ), mid-April to October methane (CH4 ) emissions, and environmental variables. From 2011 to 2022, annual rainfall was above the historical average, snow water equivalent increased, and snow-season duration shortened due to later snow return. Seasonally thawed active layer depths also increased. During this period, all ecosystems acted as slight annual sources of CO2 (13-59 g C m-2 year-1 ) and stronger sources of CH4 (11-14 g CH4 m-2 from ~April to October). The interannual variability of net ecosystem exchange was high, approximately ±100 g C m-2 year-1 , or twice what has been previously reported across other boreal sites. Net CO2 release was positively related to increased summer rainfall and winter snow water equivalent and later snow return. Controls over CH4 emissions were related to increased soil moisture and inundation status. The dominant emitter of carbon was the rich fen, which, in addition to being a source of CO2 , was also the largest CH4 emitter. These results suggest that the future carbon-source strength of boreal lowlands in Interior Alaska may be determined by the area occupied by minerotrophic fens, which are expected to become more abundant as permafrost thaw increases hydrologic connectivity. Since our measurements occur within close proximity of each other (≤1 km2 ), this study also has implications for the spatial scale and data used in benchmarking carbon cycle models and emphasizes the necessity of long-term measurements to identify carbon cycle process changes in a warming climate.

Increasing fire and the decline of fire adapted black spruce in the boreal forest.

Intensifying wildfire activity and climate change can drive rapid forest compositional shifts. In boreal North America, black spruce shapes forest flammability and depends on fire for regeneration. This relationship has helped black spruce maintain its dominance through much of the Holocene. However, with climate change and more frequent and severe fires, shifts away from black spruce dominance to broadleaf or pine species are emerging, with implications for ecosystem functions including carbon sequestration, water and energy fluxes, and wildlife habitat. Here, we predict that such reductions in black spruce after fire may already be widespread given current trends in climate and fire. To test this, we synthesize data from 1,538 field sites across boreal North America to evaluate compositional changes in tree species following 58 recent fires (1989 to 2014). While black spruce was resilient following most fires (62%), loss of resilience was common, and spruce regeneration failed completely in 18% of 1,140 black spruce sites. In contrast, postfire regeneration never failed in forests dominated by jack pine, which also possesses an aerial seed bank, or broad-leaved trees. More complete combustion of the soil organic layer, which often occurs in better-drained landscape positions and in dryer duff, promoted compositional changes throughout boreal North America. Forests in western North America, however, were more vulnerable to change due to greater long-term climate moisture deficits. While we find considerable remaining resilience in black spruce forests, predicted increases in climate moisture deficits and fire activity will erode this resilience, pushing the system toward a tipping point that has not been crossed in several thousand years.

Permafrost thaw induces short-term increase in vegetation productivity in northwestern Canada.

Over the past several decades, various trends in vegetation productivity, from increases to decreases, have been observed throughout Arctic-Boreal ecosystems. While some of this variation can be explained by recent climate warming and increased disturbance, very little is known about the impacts of permafrost thaw on productivity across diverse vegetation communities. Active layer thickness data from 135 permafrost monitoring sites along a 10° latitudinal transect of the Northwest Territories, Canada, paired with a Landsat time series of normalized difference vegetation index from 1984 to 2019, were used to quantify the impacts of changing permafrost conditions on vegetation productivity. We found that active layer thickness contributed to the observed variation in vegetation productivity in recent decades in the northwestern Arctic-Boreal, with the highest rates of greening occurring at sites where the near-surface permafrost recently had thawed. However, the greening associated with permafrost thaw was not sustained after prolonged periods of thaw and appeared to diminish after the thaw front extended outside the plants' rooting zone. Highest rates of greening were found at the mid-transect sites, between 62.4° N and 65.2° N, suggesting that more southernly sites may have already surpassed the period of beneficial permafrost thaw, while more northern sites may have yet to reach a level of thaw that supports enhanced vegetation productivity. These results indicate that the response of vegetation productivity to permafrost thaw is highly dependent on the extent of active layer thickening and that increases in productivity may not continue in the coming decades.

Lowering water table reduces carbon sink strength and carbon stocks in northern peatlands.

Peatlands at high latitudes have accumulated >400 Pg carbon (C) because saturated soil and cold temperatures suppress C decomposition. This substantial amount of C in Arctic and Boreal peatlands is potentially subject to increased decomposition if the water table (WT) decreases due to climate change, including permafrost thaw-related drying. Here, we optimize a version of the Organizing Carbon and Hydrology In Dynamic Ecosystems model (ORCHIDEE-PCH4) using site-specific observations to investigate changes in CO2 and CH4 fluxes as well as C stock responses to an experimentally manipulated decrease of WT at six northern peatlands. The unmanipulated control peatlands, with the WT <20 cm on average (seasonal max up to 45 cm) below the surface, currently act as C sinks in most years (58 ± 34 g C m-2  year-1 ; including 6 ± 7 g C-CH4 m-2  year-1 emission). We found, however, that lowering the WT by 10 cm reduced the CO2 sink by 13 ± 15 g C m-2  year-1 and decreased CH4 emission by 4 ± 4 g CH4 m-2  year-1 , thus accumulating less C over 100 years (0.2 ± 0.2 kg C m-2 ). Yet, the reduced emission of CH4 , which has a larger greenhouse warming potential, resulted in a net decrease in greenhouse gas balance by 310 ± 360 g CO2-eq  m-2  year-1 . Peatlands with the initial WT close to the soil surface were more vulnerable to C loss: Non-permafrost peatlands lost >2 kg C m-2 over 100 years when WT is lowered by 50 cm, while permafrost peatlands temporally switched from C sinks to sources. These results highlight that reductions in C storage capacity in response to drying of northern peatlands are offset in part by reduced CH4 emissions, thus slightly reducing the positive carbon climate feedbacks of peatlands under a warmer and drier future climate scenario.

Cascading effects of predators on algal size structure.

The presence of edible and inedible prey species in a food web can influence the strength that nutrients (bottom-up) or herbivores (top-down) have on primary production. In boreal peatlands, wetter more nutrient-rich conditions associated with ongoing climate change are expanding consumer access to aquatic habitat and promoting sources of primary production (i.e., algae) that are susceptible to trophic regulation. Here, we used an in situ mesocosm experiment to evaluate the consequences of enhanced nutrient availability and food-web manipulation (herbivore and predator exclusion) on algal assemblage structure in an Alaskan fen. Owing to the potential for herbivores to selectively consume edible algae (small cells) in favor of more resistant forms, we predicted that the proportion of less-edible algae (large cells) would determine the strength of top-down or bottom-up effects. Consistent with these expectations, we observed an increase in algal-cell size in the presence of herbivores (2-tiered food web) that was absent in the presence of a trophic cascade (3-tiered food web), suggesting that predators indirectly prevented morphological changes in the algal assemblage by limiting herbivory. Increases in algal-cell size with herbivory were driven by a greater proportion of filamentous green algae and nitrogen-fixing cyanobacteria, whose size and morphological characteristics mechanically minimize consumption. While consumer-driven shifts in algal assemblage structure were significant, they did not prevent top-down regulation of biofilm development by herbivores. Our findings show that increasing wet periods in northern peatlands will provide new avenues for trophic regulation of algal production, including directly through consumption and indirectly via a trophic cascade.

Letter: Trophic interactions regulate peatland carbon cycling.

Peatlands are the most efficient natural ecosystems for long-term storage of atmospheric carbon. Our understanding of peatland carbon cycling is based entirely on bottom-up controls regulated by low nutrient availability. Recent studies have shown that top-down controls through predator-prey dynamics can influence ecosystem function, yet this has not been evaluated in peatlands to date. Here, we used a combination of nutrient enrichment and trophic-level manipulation to test the hypothesis that interactions between nutrient availability (bottom-up) and predation (top-down) influence peatland carbon fluxes. Elevated nutrients stimulated bacterial biomass and organic matter decomposition. In the absence of top-down regulation, carbon dioxide (CO2 ) respiration driven by greater decomposition was offset by elevated algal productivity. Herbivores accelerated CO2 emissions by removing algal biomass, while predators indirectly reduced CO2 emissions by muting herbivory in a trophic cascade. This study demonstrates that trophic interactions can mitigate CO2 emissions associated with elevated nutrient levels in northern peatlands.

Natural selection on a carbon cycling trait drives ecosystem engineering by Sphagnum (peat moss).

Sphagnum peat mosses have an extraordinary impact on the global carbon cycle as they control long-term carbon sequestration in boreal peatland ecosystems. Sphagnum species engineer peatlands, which harbour roughly a quarter of all terrestrial carbon, through peat accumulation by constructing their own niche that allows them to outcompete other plants. Interspecific variation in peat production, largely resulting from differences in tissue decomposability, is hypothesized to drive niche differentiation along microhabitat gradients thereby alleviating competitive pressure. However, little empirical evidence exists for the role of selection in the creation and maintenance of such gradients. In order to document how niche construction and differentiation evolved in Sphagnum, we quantified decomposability for 54 species under natural conditions and used phylogenetic comparative methods to model the evolution of this carbon cycling trait. We show that decomposability tracks the phylogenetic diversification of peat mosses, that natural selection favours different levels of decomposability corresponding to optimum niche and that divergence in this trait occurred early in the evolution of the genus prior to the divergence of most extant species. Our results demonstrate the evolution of ecosystem engineering via natural selection on an extended phenotype, of a fundamental ecosystem process, and one of the Earth's largest soil carbon pools.

The Rhizosphere Responds: Rich Fen Peat and Root Microbial Ecology after Long-Term Water Table Manipulation.

Hydrologic shifts due to climate change will affect the cycling of carbon (C) stored in boreal peatlands. Carbon cycling in these systems is carried out by microorganisms and plants in close association. This study investigated the effects of experimentally manipulated water tables (lowered and raised) and plant functional groups on the peat and root microbiomes in a boreal rich fen. All samples were sequenced and processed for bacterial, archaeal (16S DNA genes; V4), and fungal (internal transcribed spacer 2 [ITS2]) DNA. Depth had a strong effect on microbial and fungal communities across all water table treatments. Bacterial and archaeal communities were most sensitive to the water table treatments, particularly at the 10- to 20-cm depth; this area coincides with the rhizosphere or rooting zone. Iron cyclers, particularly members of the family Geobacteraceae, were enriched around the roots of sedges, horsetails, and grasses. The fungal community was affected largely by plant functional group, especially cinquefoils. Fungal endophytes (particularly Acephala spp.) were enriched in sedge and grass roots, which may have underappreciated implications for organic matter breakdown and cycling. Fungal lignocellulose degraders were enriched in the lowered water table treatment. Our results were indicative of two main methanogen communities, a rooting zone community dominated by the archaeal family Methanobacteriaceae and a deep peat community dominated by the family Methanomicrobiaceae. IMPORTANCE This study demonstrated that roots and the rooting zone in boreal fens support organisms likely capable of methanogenesis, iron cycling, and fungal endophytic association and are directly or indirectly affecting carbon cycling in these ecosystems. These taxa, which react to changes in the water table and associate with roots and, particularly, graminoids, may gain greater biogeochemical influence, as projected higher precipitation rates could lead to an increased abundance of sedges and grasses in boreal fens.

Multi-omics of permafrost, active layer and thermokarst bog soil microbiomes.

Over 20% of Earth's terrestrial surface is underlain by permafrost with vast stores of carbon that, once thawed, may represent the largest future transfer of carbon from the biosphere to the atmosphere. This process is largely dependent on microbial responses, but we know little about microbial activity in intact, let alone in thawing, permafrost. Molecular approaches have recently revealed the identities and functional gene composition of microorganisms in some permafrost soils and a rapid shift in functional gene composition during short-term thaw experiments. However, the fate of permafrost carbon depends on climatic, hydrological and microbial responses to thaw at decadal scales. Here we use the combination of several molecular 'omics' approaches to determine the phylogenetic composition of the microbial communities, including several draft genomes of novel species, their functional potential and activity in soils representing different states of thaw: intact permafrost, seasonally thawed active layer and thermokarst bog. The multi-omics strategy reveals a good correlation of process rates to omics data for dominant processes, such as methanogenesis in the bog, as well as novel survival strategies for potentially active microbes in permafrost.

Wildfire combustion and carbon stocks in the southern Canadian boreal forest: Implications for a warming world.

Boreal wildfires are increasing in intensity, extent, and frequency, potentially intensifying carbon emissions and transitioning the region from a globally significant carbon sink to a source. The productive southern boreal forests of central Canada already experience relatively high frequencies of fire, and as such may serve as an analog of future carbon dynamics for more northern forests. Fire-carbon dynamics in southern boreal systems are relatively understudied, with limited investigation into the drivers of pre-fire carbon stocks or subsequent combustion. As part of NASA's Arctic-Boreal Vulnerability Experiment, we sampled 79 stands (47 burned, 32 unburned) throughout central Saskatchewan to characterize above- and belowground carbon stocks and combustion rates in relation to historical land use, vegetation characteristics, and geophysical attributes. We found southern boreal forests emitted an average of 3.3 ± 1.1 kg C/m2 from field sites. The emissions from southern boreal stands varied as a function of stand age, fire weather conditions, ecozone, and soil moisture class. Sites affected by historical timber harvesting had greater combustion rates due to faster carbon stock recovery rates than sites recovering from wildfire events, indicating that different boreal forest land use practices can generate divergent carbon legacy effects. We estimate the 2015 fire season in Saskatchewan emitted a total of 36.3 ± 15.0 Tg C, emphasizing the importance of southern boreal fires for regional carbon budgets. Using the southern boreal as an analog, the northern boreal may undergo fundamental shifts in forest structure and carbon dynamics, becoming dominated by stands <70 years old that hold 2-7 kg C/m2 less than current mature northern boreal stands. Our latitudinal approach reinforces previous studies showing that northern boreal stands are at a high risk of holding less carbon under changing disturbance conditions.

Climate change decreases the cooling effect from postfire albedo in boreal North America.

Fire is a primary disturbance in boreal forests and generates both positive and negative climate forcings. The influence of fire on surface albedo is a predominantly negative forcing in boreal forests, and one of the strongest overall, due to increased snow exposure in the winter and spring months. Albedo forcings are spatially and temporally heterogeneous and depend on a variety of factors related to soils, topography, climate, land cover/vegetation type, successional dynamics, time since fire, season, and fire severity. However, how these variables interact to influence albedo is not well understood, and quantifying these relationships and predicting postfire albedo becomes increasingly important as the climate changes and management frameworks evolve to consider climate impacts. Here we developed a MODIS-derived 'blue sky' albedo product and a novel machine learning modeling framework to predict fire-driven changes in albedo under historical and future climate scenarios across boreal North America. Converted to radiative forcing (RF), we estimated that fires generate an annual mean cooling of -1.77 ± 1.35 W/m2 from albedo under historical climate conditions (1971-2000) integrated over 70 years postfire. Increasing postfire albedo along a south-north climatic gradient was offset by a nearly opposite gradient in solar insolation, such that large-scale spatial patterns in RF were minimal. Our models suggest that climate change will lead to decreases in mean annual postfire albedo, and hence a decreasing strength of the negative RF, a trend dominated by decreased snow cover in spring months. Considering the range of future climate scenarios and model uncertainties, we estimate that for fires burning in the current era (2016) the cooling effect from long-term postfire albedo will be reduced by 15%-28% due to climate change.

Tradeoffs between phenotypic plasticity and local adaptation influence the ecophysiology of the moss, Sphagnum magellanicum.

Bryophytes are a diverse plant group and are functionally different from vascular plants. Yet, their peculiarities are rarely considered in the theoretical frameworks for plants. Currently, we lack information about the magnitude and the importance of intraspecific variability in the ecophysiology of bryophytes and how these might translate to local adaptation-a prerequisite for adaptive evolution. Capitalizing on two ecologically distinct (hummock and hollow) phenotypes of Sphagnum magellanicum, we explored the magnitude and pattern of intraspecific variability in this species and asked whether the environmental-mediated changes in shoot and physiological traits are due to phenotypic plasticity or local adaptation. Size, pigmentation, and habitat type that distinguished the phenotypes in the field did not influence the trait responses under a transplant and factorial experiment. In addition, the magnitude and pattern of trait variability (e.g., branch, stem and capitulum mass) changed with the treatments, which suggest that trait responses were due largely to phenotypic plasticity. The trait responses also suggest that the ecophysiological needs for mosses to grow in clumps, where they maintain a uniform growth may have an overriding effect over the potential for a fixed adaptive response to environmental heterogeneity, which would constrain local adaptation. We conclude that extending the trait-based framework to mosses or making comparisons between mosses and vascular plants under any theoretical framework would only be meaningful to the extent that growth form and dispersal strategies are considered.

The response of boreal peatland community composition and NDVI to hydrologic change, warming, and elevated carbon dioxide.

Widespread changes in arctic and boreal Normalized Difference Vegetation Index (NDVI) values captured by satellite platforms indicate that northern ecosystems are experiencing rapid ecological change in response to climate warming. Increasing temperatures and altered hydrology are driving shifts in ecosystem biophysical properties that, observed by satellites, manifest as long-term changes in regional NDVI. In an effort to examine the underlying ecological drivers of these changes, we used field-scale remote sensing of NDVI to track peatland vegetation in experiments that manipulated hydrology, temperature, and carbon dioxide (CO2 ) levels. In addition to NDVI, we measured percent cover by species and leaf area index (LAI). We monitored two peatland types broadly representative of the boreal region. One site was a rich fen located near Fairbanks, Alaska, at the Alaska Peatland Experiment (APEX), and the second site was a nutrient-poor bog located in Northern Minnesota within the Spruce and Peatland Responses Under Changing Environments (SPRUCE) experiment. We found that NDVI decreased with long-term reductions in soil moisture at the APEX site, coincident with a decrease in photosynthetic leaf area and the relative abundance of sedges. We observed increasing NDVI with elevated temperature at the SPRUCE site, associated with an increase in the relative abundance of shrubs and a decrease in forb cover. Warming treatments at the SPRUCE site also led to increases in the LAI of the shrub layer. We found no strong effects of elevated CO2 on community composition. Our findings support recent studies suggesting that changes in NDVI observed from satellite platforms may be the result of changes in community composition and ecosystem structure in response to climate warming.

Wildfire severity reduces richness and alters composition of soil fungal communities in boreal forests of western Canada.

Wildfire is the dominant disturbance in boreal forests and fire activity is increasing in these regions. Soil fungal communities are important for plant growth and nutrient cycling postfire but there is little understanding of how fires impact fungal communities across landscapes, fire severity gradients, and stand types in boreal forests. Understanding relationships between fungal community composition, particularly mycorrhizas, and understory plant composition is therefore important in predicting how future fire regimes may affect vegetation. We used an extreme wildfire event in boreal forests of Canada's Northwest Territories to test drivers of fungal communities and assess relationships with plant communities. We sampled soils from 39 plots 1 year after fire and 8 unburned plots. High-throughput sequencing (MiSeq, ITS) revealed 2,034 fungal operational taxonomic units. We found soil pH and fire severity (proportion soil organic layer combusted), and interactions between these drivers were important for fungal community structure (composition, richness, diversity, functional groups). Where fire severity was low, samples with low pH had higher total fungal, mycorrhizal, and saprotroph richness compared to where severity was high. Increased fire severity caused declines in richness of total fungi, mycorrhizas, and saprotrophs, and declines in diversity of total fungi and mycorrhizas. The importance of stand age (a surrogate for fire return interval) for fungal composition suggests we could detect long-term successional patterns even after fire. Mycorrhizal and plant community composition, richness, and diversity were weakly but significantly correlated. These weak relationships and the distribution of fungi across plots suggest that the underlying driver of fungal community structure is pH, which is modified by fire severity. This study shows the importance of edaphic factors in determining fungal community structure at large scales, but suggests these patterns are mediated by interactions between fire and forest stand composition.

The Sphagnome Project: enabling ecological and evolutionary insights through a genus-level sequencing project.

Considerable progress has been made in ecological and evolutionary genetics with studies demonstrating how genes underlying plant and microbial traits can influence adaptation and even 'extend' to influence community structure and ecosystem level processes. Progress in this area is limited to model systems with deep genetic and genomic resources that often have negligible ecological impact or interest. Thus, important linkages between genetic adaptations and their consequences at organismal and ecological scales are often lacking. Here we introduce the Sphagnome Project, which incorporates genomics into a long-running history of Sphagnum research that has documented unparalleled contributions to peatland ecology, carbon sequestration, biogeochemistry, microbiome research, niche construction, and ecosystem engineering. The Sphagnome Project encompasses a genus-level sequencing effort that represents a new type of model system driven not only by genetic tractability, but by ecologically relevant questions and hypotheses.

Cross-scale controls on carbon emissions from boreal forest megafires.

Climate warming and drying is associated with increased wildfire disturbance and the emergence of megafires in North American boreal forests. Changes to the fire regime are expected to strongly increase combustion emissions of carbon (C) which could alter regional C balance and positively feedback to climate warming. In order to accurately estimate C emissions and thereby better predict future climate feedbacks, there is a need to understand the major sources of heterogeneity that impact C emissions at different scales. Here, we examined 211 field plots in boreal forests dominated by black spruce (Picea mariana) or jack pine (Pinus banksiana) of the Northwest Territories (NWT), Canada after an unprecedentedly large area burned in 2014. We assessed both aboveground and soil organic layer (SOL) combustion, with the goal of determining the major drivers in total C emissions, as well as to develop a high spatial resolution model to scale emissions in a relatively understudied region of the boreal forest. On average, 3.35 kg C m-2 was combusted and almost 90% of this was from SOL combustion. Our results indicate that black spruce stands located at landscape positions with intermediate drainage contribute the most to C emissions. Indices associated with fire weather and date of burn did not impact emissions, which we attribute to the extreme fire weather over a short period of time. Using these results, we estimated a total of 94.3 Tg C emitted from 2.85 Mha of burned area across the entire 2014 NWT fire complex, which offsets almost 50% of mean annual net ecosystem production in terrestrial ecosystems of Canada. Our study also highlights the need for fine-scale estimates of burned area that represent small water bodies and regionally specific calibrations of combustion that account for spatial heterogeneity in order to accurately model emissions at the continental scale.

A decade of boreal rich fen greenhouse gas fluxes in response to natural and experimental water table variability.

Rich fens are common boreal ecosystems with distinct hydrology, biogeochemistry and ecology that influence their carbon (C) balance. We present growing season soil chamber methane emission (FCH4 ), ecosystem respiration (ER), net ecosystem exchange (NEE) and gross primary production (GPP) fluxes from a 9-years water table manipulation experiment in an Alaskan rich fen. The study included major flood and drought years, where wetting and drying treatments further modified the severity of droughts. Results support previous findings from peatlands that drought causes reduced magnitude of growing season FCH4 , GPP and NEE, thus reducing or reversing their C sink function. Experimentally exacerbated droughts further reduced the capacity for the fen to act as a C sink by causing shifts in vegetation and thus reducing magnitude of maximum growing season GPP in subsequent flood years by ~15% compared to control plots. Conversely, water table position had only a weak influence on ER, but dominant contribution to ER switched from autotrophic respiration in wet years to heterotrophic in dry years. Droughts did not cause inter-annual lag effects on ER in this rich fen, as has been observed in several nutrient-poor peatlands. While ER was dependent on soil temperatures at 2 cm depth, FCH4 was linked to soil temperatures at 25 cm. Inter-annual variability of deep soil temperatures was in turn dependent on wetness rather than air temperature, and higher FCH4 in flooded years was thus equally due to increased methane production at depth and decreased methane oxidation near the surface. Short-term fluctuations in wetness caused significant lag effects on FCH4 , but droughts caused no inter-annual lag effects on FCH4 . Our results show that frequency and severity of droughts and floods can have characteristic effects on the exchange of greenhouse gases, and emphasize the need to project future hydrological regimes in rich fens.

The Sphagnum microbiome: new insights from an ancient plant lineage.

57 I. 57 II. 58 III. 59 IV. 59 V. 61 VI. 62 63 References 63 SUMMARY: Peat mosses of the genus Sphagnum play a major role in global carbon storage and dominate many northern peatland ecosystems, which are currently being subjected to some of the most rapid climate changes on Earth. A rapidly expanding database indicates that a diverse community of microorganisms is intimately associated with Sphagnum, inhabiting the tissues and surface of the plant. Here we summarize the current state of knowledge regarding the Sphagnum microbiome and provide a perspective for future research directions. Although the majority of the microbiome remains uncultivated and its metabolic capabilities uncharacterized, prokaryotes and fungi have the potential to act as mutualists, symbionts, or antagonists of Sphagnum. For example, methanotrophic and nitrogen-fixing bacteria may benefit the plant host by providing up to 20-30% of Sphagnum carbon and nitrogen, respectively. Next-generation sequencing approaches have enabled the detailed characterization of microbiome community composition in peat mosses. However, as with other ecologically or economically important plants, our knowledge of Sphagnum-microbiome associations is in its infancy. In order to attain a predictive understanding of the role of the microbiome in Sphagnum productivity and ecosystem function, the mechanisms of plant-microbiome interactions and the metabolic potential of constituent microbial populations must be revealed.