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.

Peat loss collocates with a threshold in plant-mycorrhizal associations in drained peatlands encroached by trees.

Drainage-induced encroachment by trees may have major effects on the carbon balance of northern peatlands, and responses of microbial communities are likely to play a central mechanistic role. We profiled the soil fungal community and estimated its genetic potential for the decay of lignin and phenolics (class II peroxidase potential) along peatland drainage gradients stretching from interior locations (undrained, open) to ditched locations (drained, forested). Mycorrhizal fungi dominated the community across the gradients. When moving towards ditches, the dominant type of mycorrhizal association abruptly shifted from ericoid mycorrhiza to ectomycorrhiza at c. 120 m from the ditches. This distance corresponded with increased peat loss, from which more than half may be attributed to oxidation. The ectomycorrhizal genus Cortinarius dominated at the drained end of the gradients and its relatively higher genetic potential to produce class II peroxidases (together with Mycena) was positively associated with peat humification and negatively with carbon-to-nitrogen ratio. Our study is consistent with a plant-soil feedback mechanism, driven by a shift in the mycorrhizal type of vegetation, that potentially mediates changes in aerobic decomposition during postdrainage succession. Such feedback may have long-term legacy effects upon postdrainage restoration efforts and implication for tree encroachment onto carbon-rich soils globally.

Evidence for older carbon loss with lowered water tables and changing plant functional groups in peatlands.

A small imbalance in plant productivity and decomposition accounts for the carbon (C) accumulation capacity of peatlands. As climate changes, the continuity of peatland net C storage relies on rising primary production to offset increasing ecosystem respiration (ER) along with the persistence of older C in waterlogged peat. A lowering in the water table position in peatlands often increases decomposition rates, but concurrent plant community shifts can interactively alter ER and plant productivity responses. The combined effects of water table variation and plant communities on older peat C loss are unknown. We used a full-factorial 1-m3 mesocosm array with vascular plant functional group manipulations (Unmanipulated Control, Sedge only, and Ericaceous only) and water table depth (natural and lowered) treatments to test the effects of plants and water depth on CO2 fluxes, decomposition, and older C loss. We used Δ14 C and δ13 C of ecosystem CO2 respiration, bulk peat, plants, and porewater dissolved inorganic C to construct mixing models partitioning ER among potential sources. We found that the lowered water table treatments were respiring C fixed before the bomb spike (1955) from deep waterlogged peat. Lowered water table Sedge treatments had the oldest dissolved inorganic 14 C signature and the highest proportional peat contribution to ER. Decomposition assays corroborated sustained high rates of decomposition with lowered water tables down to 40 cm below the peat surface. Heterotrophic respiration exceeded plant respiration at the height of the growing season in lowered water table treatments. Rates of gross primary production were only impacted by vegetation, whereas ER was affected by vegetation and water table depth treatments. The decoupling of respiration and primary production with lowered water tables combined with older C losses suggests that climate and land-use-induced changes in peatland hydrology can increase the vulnerability of peatland C stores.

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.

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.

Peatland microbial community responses to plant functional group and drought are depth-dependent.

Peatlands store one-third of Earth's soil carbon, the stability of which is uncertain due to climate change-driven shifts in hydrology and vegetation, and consequent impacts on microbial communities that mediate decomposition. Peatland carbon cycling varies over steep physicochemical gradients characterizing vertical peat profiles. However, it is unclear how drought-mediated changes in plant functional groups (PFGs) and water table (WT) levels affect microbial communities at different depths. We combined a multiyear mesocosm experiment with community sequencing across a 70-cm depth gradient, to test the hypotheses that vascular PFGs (Ericaceae vs. sedges) and WT (high vs. low) structure peatland microbial communities in depth-dependent ways. Several key results emerged. (i) Both fungal and prokaryote (bacteria and archaea) community structure shifted with WT and PFG manipulation, but fungi were much more sensitive to PFG whereas prokaryotes were much more sensitive to WT. (ii) PFG effects were largely driven by Ericaceae, although sedge effects were evident in specific cases (e.g., methanotrophs). (iii) Treatment effects varied with depth: the influence of PFG was strongest in shallow peat (0-10, 10-20 cm), whereas WT effects were strongest at the surface and middle depths (0-10, 30-40 cm), and all treatment effects waned in the deepest peat (60-70 cm). Our results underline the depth-dependent and taxon-specific ways that plant communities and hydrologic variability shape peatland microbial communities, pointing to the importance of understanding how these factors integrate across soil profiles when examining peatland responses to climate change.

Impacts of historical ditching on peat volume and carbon in northern Minnesota USA peatlands.

Peatlands play a critical role in terrestrial carbon (C) storage, containing an estimated 30% of global soil C, despite occupying only 3% of global land area. Historic management of peatlands has led to widespread degradation and loss of important ecosystem services, including C sequestration. Legacy drainage features in the peatlands of northern Minnesota, USA were studied to assess the volume of peat and the amount of C lost in the ~100 years since drainage. Using high-resolution Light Detection and Ranging (LiDAR) data, we measured elevation changes adjacent to legacy ditches to model pre-ditch surface elevations, which were used to calculate peat volume loss. We established relationships between volume loss and site characteristics from existing geographic information systems datasets and used those relationships to scale volume loss to all mapped peatland ditches in northern Minnesota (USA). We estimated that 0.165 ± 0.009 km3 of peat have been lost along almost 4000 km of peatland ditches. Peat loss upslope of ditches was significantly less than downslope (P < 0.001). Mean width of the entire ditch-effect zone was 333 ± 8.32 m. Using our volume loss estimates, literature estimates of oxidation, and mean bulk density and peat C% values from Minnesota peatlands, we calculate a total historic loss 3.847 ± 0.364 Tg C. Assuming a constant oxidation rate during the 100 years since drainage, euic and dysic peatlands within the ditch effect zone have lost 0.26 ± 0.08 and 0.40 ± 0.13 Mg C ha-1 yr-1, respectively, comparable to IPCC estimates. Our spatially-explicit peat loss estimates could be incorporated into decision support tools to inform management decisions regarding peatland C and other ecosystem services.

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.

Towards a global assessment of pyrogenic carbon from vegetation fires.

The production of pyrogenic carbon (PyC; a continuum of organic carbon (C) ranging from partially charred biomass and charcoal to soot) is a widely acknowledged C sink, with the latest estimates indicating that ~50% of the PyC produced by vegetation fires potentially sequesters C over centuries. Nevertheless, the quantitative importance of PyC in the global C balance remains contentious, and therefore, PyC is rarely considered in global C cycle and climate studies. Here we examine the robustness of existing evidence and identify the main research gaps in the production, fluxes and fate of PyC from vegetation fires. Much of the previous work on PyC production has focused on selected components of total PyC generated in vegetation fires, likely leading to underestimates. We suggest that global PyC production could be in the range of 116-385 Tg C yr(-1) , that is ~0.2-0.6% of the annual terrestrial net primary production. According to our estimations, atmospheric emissions of soot/black C might be a smaller fraction of total PyC (<2%) than previously reported. Research on the fate of PyC in the environment has mainly focused on its degradation pathways, and its accumulation and resilience either in situ (surface soils) or in ultimate sinks (marine sediments). Off-site transport, transformation and PyC storage in intermediate pools are often overlooked, which could explain the fate of a substantial fraction of the PyC mobilized annually. We propose new research directions addressing gaps in the global PyC cycle to fully understand the importance of the products of burning in global C cycle dynamics.

Contributions of algae to GPP and DOC production in an Alaskan fen: effects of historical water table manipulations on ecosystem responses to a natural flood.

The role of algae in the metabolism of northern peatlands is largely unknown, as is how algae will respond to the rapid climate change being experienced in this region. In this study, we examined patterns in algal productivity, nutrients, and dissolved organic carbon (DOC) during an uncharacteristically wet summer in an Alaskan rich fen. Our sampling was conducted in three large-scale experimental plots where water table position had been manipulated (including both drying and wetting plots and a control) for the previous 4 years. This study allowed us to explore how much ecosystem memory of the antecedent water table manipulations governed algal responses to natural flooding. Despite no differences in water table position between the manipulated plots at the time of sampling, algal primary productivity was consistently higher in the lowered water table plot compared to the control or raised water table plots. In all plots, algal productivity peaked immediately following seasonal maxima in nutrient concentrations. We found a positive relationship between algal productivity and water-column DOC concentrations (r (2) = 0.85, P < 0.001). Using these data, we estimate that algae released approximately 19% of fixed carbon into the water column. Algal exudates were extremely labile in biodegradability assays, decreasing by more than 55% within the first 24 h of incubation. We suggest that algae can be an important component of the photosynthetic community in boreal peatlands and may become increasingly important for energy flow in a more variable climate with more intense droughts and flooding.

Variation in carbon and nitrogen concentrations among peatland categories at the global scale.

Peatlands account for 15 to 30% of the world's soil carbon (C) stock and are important controls over global nitrogen (N) cycles. However, C and N concentrations are known to vary among peatlands contributing to the uncertainty of global C inventories, but there are few global studies that relate peatland classification to peat chemistry. We analyzed 436 peat cores sampled in 24 countries across six continents and measured C, N, and organic matter (OM) content at three depths down to 70 cm. Sites were distinguished between northern (387) and tropical (49) peatlands and assigned to one of six distinct broadly recognized peatland categories that vary primarily along a pH gradient. Peat C and N concentrations, OM content, and C:N ratios differed significantly among peatland categories, but few differences in chemistry with depth were found within each category. Across all peatlands C and N concentrations in the 10-20 cm layer, were 440 ± 85.1 g kg-1 and 13.9 ± 7.4 g kg-1, with an average C:N ratio of 30.1 ± 20.8. Among peatland categories, median C concentrations were highest in bogs, poor fens and tropical swamps (446-532 g kg-1) and lowest in intermediate and extremely rich fens (375-414 g kg-1). The C:OM ratio in peat was similar across most peatland categories, except in deeper samples from ombrotrophic tropical peat swamps that were higher than other peatlands categories. Peat N concentrations and C:N ratios varied approximately two-fold among peatland categories and N concentrations tended to be higher (and C:N lower) in intermediate fens compared with other peatland types. This study reports on a unique data set and demonstrates that differences in peat C and OM concentrations among broadly classified peatland categories are predictable, which can aid future studies that use land cover assessments to refine global peatland C and N stocks.

Impacts of experimental alteration of water table regime and vascular plant community composition on peat mercury profiles and methylmercury production.

Climate change is expected to alter the hydrology and vascular plant communities in peatland ecosystems. These changes may have as yet unexplored impacts on peat mercury (Hg) concentrations and net methylmercury (MeHg) production. In this study, peat was collected from PEATcosm, an outdoor, controlled mesocosm experiment where peatland water table regimes and vascular plant functional groups were manipulated over several years to simulate potential climate change effects. Potential Hg(II) methylation and MeHg demethylation rate constants were assessed using enriched stable isotope incubations at the end of the study in 2015, and ambient peat total Hg (THg) and MeHg concentration depth profiles were tracked annually from 2011 to 2014. Peat THg and MeHg concentrations and the proportion of THg methylated (%MeHg) increased significantly within the zone of water table fluctuation when water tables were lowered, but potential Hg(II) methylation rate constants were similar regardless of water table treatment. When sedges dominate over ericaceous shrubs, MeHg concentrations and %MeHg became significantly elevated within the sedge rooting zone. Increased desorption of Hg(II) and MeHg from the solid phase peat into pore water occurred with a lowered water table and predominant sedge cover, likely due to greater aerobic peat decomposition. Deeper, more variable water tables and a transition to sedge-dominated communities coincided with increased MeHg accumulation within the zone of water table fluctuation. Sustained high water tables promoted the net downward migration of Hg(II) and MeHg. The simultaneous decrease in Hg(II) and MeHg concentrations in the near-surface peat and accumulation deeper in the peat profile, combined with the trends in Hg(II) and MeHg partitioning to mobile pore waters, suggest that changes to peatland hydrology and vascular plant functional groups redistribute peat Hg(II) and MeHg via vertical hydrochemical transport mechanisms.

Patterns and drivers of fungal community depth stratification in Sphagnum peat.

Peatlands store an immense pool of soil carbon vulnerable to microbial oxidation due to drought and intentional draining. We used amplicon sequencing and quantitative PCR to (i) examine how fungi are influenced by depth in the peat profile, water table and plant functional group at the onset of a multiyear mesocosm experiment, and (ii) test if fungi are correlated with abiotic variables of peat and pore water. We hypothesized that each factor influenced fungi, but that depth would have the strongest effect early in the experiment. We found that (i) communities were strongly depth stratified; fungi were four times more abundant in the upper (10-20 cm) than the lower (30-40 cm) depth, and dominance shifted from ericoid mycorrhizal fungi to saprotrophs and endophytes with increasing depth; (ii) the influence of plant functional group was depth dependent, with Ericaceae structuring the community in the upper peat only; (iii) water table had minor influences; and (iv) communities strongly covaried with abiotic variables, including indices of peat and pore water carbon quality. Our results highlight the importance of vertical stratification to peatland fungi, and the depth dependency of plant functional group effects, which must be considered when elucidating the role of fungi in peatland carbon dynamics.

The response of soil organic carbon of a rich fen peatland in interior Alaska to projected climate change.

It is important to understand the fate of carbon in boreal peatland soils in response to climate change because a substantial change in release of this carbon as CO2 and CH4 could influence the climate system. The goal of this research was to synthesize the results of a field water table manipulation experiment conducted in a boreal rich fen into a process-based model to understand how soil organic carbon (SOC) of the rich fen might respond to projected climate change. This model, the peatland version of the dynamic organic soil Terrestrial Ecosystem Model (peatland DOS-TEM), was calibrated with data collected during 2005-2011 from the control treatment of a boreal rich fen in the Alaska Peatland Experiment (APEX). The performance of the model was validated with the experimental data measured from the raised and lowered water-table treatments of APEX during the same period. The model was then applied to simulate future SOC dynamics of the rich fen control site under various CO2 emission scenarios. The results across these emissions scenarios suggest that the rate of SOC sequestration in the rich fen will increase between year 2012 and 2061 because the effects of warming increase heterotrophic respiration less than they increase carbon inputs via production. However, after 2061, the rate of SOC sequestration will be weakened and, as a result, the rich fen will likely become a carbon source to the atmosphere between 2062 and 2099. During this period, the effects of projected warming increase respiration so that it is greater than carbon inputs via production. Although changes in precipitation alone had relatively little effect on the dynamics of SOC, changes in precipitation did interact with warming to influence SOC dynamics for some climate scenarios.