<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD Journal Publishing with OASIS Tables v3.0 20080202//EN" "https://jats.nlm.nih.gov/nlm-dtd/publishing/3.0/journalpub-oasis3.dtd">
<article xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:oasis="http://docs.oasis-open.org/ns/oasis-exchange/table" xml:lang="en" dtd-version="3.0" article-type="research-article">
  <front>
    <journal-meta><journal-id journal-id-type="publisher">BG</journal-id><journal-title-group>
    <journal-title>Biogeosciences</journal-title>
    <abbrev-journal-title abbrev-type="publisher">BG</abbrev-journal-title><abbrev-journal-title abbrev-type="nlm-ta">Biogeosciences</abbrev-journal-title>
  </journal-title-group><issn pub-type="epub">1726-4189</issn><publisher>
    <publisher-name>Copernicus Publications</publisher-name>
    <publisher-loc>Göttingen, Germany</publisher-loc>
  </publisher></journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.5194/bg-23-3777-2026</article-id><title-group><article-title>Addition of brackish water to tundra soils does not inhibit methane production: implications for Arctic coastal methane production</article-title><alt-title>Addition of brackish water to tundra soils does not inhibit methane production</alt-title>
      </title-group>
      <contrib-group>
        <contrib contrib-type="author" corresp="yes" rid="aff1 aff5 aff6">
          <name><surname>Roy-Lafontaine</surname><given-names>Alexie</given-names></name>
          <email>alexieroylafontaine@gmail.com</email>
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff2">
          <name><surname>Lee</surname><given-names>Rebecca</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff3 aff4 aff6">
          <name><surname>Douglas</surname><given-names>Peter M. J.</given-names></name>
          
        <ext-link>https://orcid.org/0000-0002-4282-6615</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff2">
          <name><surname>Whalen</surname><given-names>Dustin</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1 aff5 aff6">
          <name><surname>Pellerin</surname><given-names>André</given-names></name>
          
        <ext-link>https://orcid.org/0000-0003-3588-8372</ext-link></contrib>
        <aff id="aff1"><label>1</label><institution>Institut des Sciences de la Mer de Rimouski, Université du Québec à Rimouski, Rimouski, Québec, Canada</institution>
        </aff>
        <aff id="aff2"><label>2</label><institution>Geological Survey of Canada, Natural Resources Canada, Halifax, Nova Scotia, Canada</institution>
        </aff>
        <aff id="aff3"><label>3</label><institution>Department of Earth and Planetary Sciences and Geotop Research Centre, McGill University, Montréal, Quebec, Canada</institution>
        </aff>
        <aff id="aff4"><label>4</label><institution>Centre d'Études Nordiques, Université Laval, Québec, Quebec, Canada</institution>
        </aff>
        <aff id="aff5"><label>5</label><institution>Québec Océan, Université Laval, Québec, Quebec, Canada</institution>
        </aff>
        <aff id="aff6"><label>6</label><institution>Research Centre in Earth System Dynamics, Université du Québec à Montréal, Quebec, Canada</institution>
        </aff>
      </contrib-group>
      <author-notes><corresp id="corr1">Alexie Roy-Lafontaine (alexieroylafontaine@gmail.com)</corresp></author-notes><pub-date><day>10</day><month>June</month><year>2026</year></pub-date>
      
      <volume>23</volume>
      <issue>11</issue>
      <fpage>3777</fpage><lpage>3792</lpage>
      <history>
        <date date-type="received"><day>10</day><month>June</month><year>2025</year></date>
           <date date-type="rev-request"><day>26</day><month>June</month><year>2025</year></date>
           <date date-type="rev-recd"><day>19</day><month>March</month><year>2026</year></date>
           <date date-type="accepted"><day>23</day><month>March</month><year>2026</year></date>
      </history>
      <permissions>
        <copyright-statement>Copyright: © 2026 Alexie Roy-Lafontaine et al.</copyright-statement>
        <copyright-year>2026</copyright-year>
      <license license-type="open-access"><license-p>This work is licensed under the Creative Commons Attribution 4.0 International License. To view a copy of this licence, visit <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">https://creativecommons.org/licenses/by/4.0/</ext-link></license-p></license></permissions><self-uri xlink:href="https://bg.copernicus.org/articles/23/3777/2026/bg-23-3777-2026.html">This article is available from https://bg.copernicus.org/articles/23/3777/2026/bg-23-3777-2026.html</self-uri><self-uri xlink:href="https://bg.copernicus.org/articles/23/3777/2026/bg-23-3777-2026.pdf">The full text article is available as a PDF file from https://bg.copernicus.org/articles/23/3777/2026/bg-23-3777-2026.pdf</self-uri>
      <abstract><title>Abstract</title>

      <p id="d2e156">In Arctic regions where coastal sediments contain permafrost, global climate change drives processes such as erosion and subsidence. The contribution of these processes to carbon emissions, especially from ground subsidence, are still uncertain. Relative sea level rise can lead to more waterlogged environments, promoting anoxic degradation of organic matter but it can also lead to a greater exposure of coastal sediments to seawater. This could alter methane (CH<sub>4</sub>) production dynamics, although the controls remain poorly understood. For instance, sulfates contained in seawater may have a tampering effect on methanogenesis through competitive inhibition but the increase in microbial abundance could enhance methanogenesis. In this study, we present CH<sub>4</sub> production rates alongside geochemical analyses in a rapidly evolving coastal landscape near the community of Tuktoyaktuk, NWT, Canada, which is located in the continuous permafrost zone. To better constrain CH<sub>4</sub> production dynamics along the land to ocean continuum, sediment cores were collected from nearshore marine sediments and soil profiles were collected from the active layer of the coastal (intertidal) zone and inland soils. Anoxic incubations were performed, amended with brackish water to simulate the effect of seawater on the breakdown of organic matter and the production of CH<sub>4</sub>. We found marine sediments expectedly led to negligible CH<sub>4</sub> production rates, while the inland sites showed variable rates between null and 35 nmol cm<sup>−3</sup> d<sup>−1</sup>. The coastal (intertidal) zone had the highest rates reaching 415 nmol cm<sup>−3</sup> d<sup>−1</sup>. Interestingly, sulfate present in brackish water and sediments did not suppress methanogenesis in the incubations of the coastal and inland zones. Analyses of stable carbon isotopes from CH<sub>4</sub> produced in the incubation experiment indicated greater acetotrophy and higher organic matter lability in the coastal zone, possibly contributing to higher CH<sub>4</sub> production rates. This study highlights the potential for significant CH<sub>4</sub> emissions even with high sulfate concentrations which are classically thought to inhibit methanogenesis. This suggests that Arctic coastal microbial CH<sub>4</sub> production might be an understudied source to the atmosphere.</p>
  </abstract>
    
<funding-group>
<award-group id="gs1">
<funding-source>Natural Sciences and Engineering Research Council of Canada</funding-source>
<award-id>Discovery Grant</award-id>
</award-group>
<award-group id="gs2">
<funding-source>Natural Resources Canada</funding-source>
<award-id>Geological Survey of Canada - Nuna Project</award-id>
</award-group>
</funding-group>
</article-meta>
  </front>
<body>
      

<sec id="Ch1.S1" sec-type="intro">
  <label>1</label><title>Introduction</title>
      <p id="d2e299">Arctic coastal ecosystems are impacted by sea level rise, coastal erosion, land submersion, higher frequency in storm events and permafrost degradation (AMAP, 2017; Guimond et al., 2021; Irrgang et al., 2022; Lantuit et al., 2012; Lim et al., 2020). The amplification of coastal environmental changes has impacts on biogeochemical cycles (AMAP, 2017) and on organic matter (OM) degradation processes and fluxes at the land-ocean continuum (Tanski et al., 2021). Furthermore, the progressive thawing of permafrost exposes long frozen organic matter to microbial decomposition (Lapham et al., 2020; Pellerin et al., 2022; Schuur et al., 2015), leading to the release of greenhouse gases like carbon dioxide (CO<sub>2</sub>) and methane (CH<sub>4</sub>). Inputs and outputs of the Arctic carbon biogeochemical cycle are known to be reshaped by rapid environmental changes (Couture et al., 2018), but processes in coastal settings are still poorly understood.</p>
      <p id="d2e320">Rates of coastal change vary according to the morphology of coastal landscapes (Manson et al., 2019). The average rate of land retreat measured in the Tuktoyaktuk Coastlands (North-West Territories, Canada), our study site, between 1985 and 2020 was <inline-formula><mml:math id="M16" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>1.0 m yr<sup>−1</sup>, while processes of ground subsidence and submersion induced retreat rates higher than <inline-formula><mml:math id="M18" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>4 m yr<sup>−1</sup> (Costa et al., 2022) which can inundate large swaths of land. Inundated tundra flats and polygons are widespread landforms in the landscape (Costa et al., 2022). Polygon tundra flats are characterized by ice-wedge polygons, which are formed by the repeated thermal contraction and expansion of the upper layers of the permafrost (Steedman et al., 2017). At the surface, the polygons are expressed as minor topographic features separated by lower-lying, often wet or inundated channels called troughs (Fig. 1). Polygons can be classified as low-centered (with a low, wet center and raised rims) or as high-centered (with well-drained centers and lower well-drained rims) (Fig. 1), exhibiting strong thermal, hydrological and geochemical gradients (Vaughn et al., 2016).</p>

      <fig id="F1" specific-use="star"><label>Figure 1</label><caption><p id="d2e363">Schematic representation of polygonal tundra with peat accumulation as seen in continuous permafrost environments and sampling design for this study. High-centered polygons are associated with drier conditions, while low-centered polygons, troughs and pondlets are associated with humid or water-saturated conditions. Vegetation cover and OM reflect the hydrology of sites. Not to scale.</p></caption>
        <graphic xlink:href="https://bg.copernicus.org/articles/23/3777/2026/bg-23-3777-2026-f01.png"/>

      </fig>

      <p id="d2e373">During growing season, where atmospheric temperatures allow for active layer to thaw and vegetation to grow, hydrological conditions in polygons play a pivotal role in shaping the pathways of OM decomposition and consequently influence the resulting CO<sub>2</sub> and CH<sub>4</sub> production. Well drained oxic conditions allow microbes to decompose OM rapidly, leading to the production of CO<sub>2</sub> (Jones et al., 2020). Conversely, water saturation restricts oxygen availability, promoting anaerobic respiration and fermentation, inducing both CO<sub>2</sub> and CH<sub>4</sub> production (Lipson et al., 2012; Turetsky et al., 2008). Thus, coastal changes and higher atmospheric temperatures during open-water season can swiftly alter water saturation conditions in polygons, in many cases significantly enhancing fermentation and CH<sub>4</sub> production (Elberling et al., 2013; Holm et al., 2020; Treat et al., 2014).</p>
      <p id="d2e431">Furthermore, coastal changes can also influence the chemistry of the water within soils, which can affect OM degradation. In anaerobic conditions, OM degradation processes follow a sequence of electron acceptors of decreasing energetic yields with nitrate, manganese oxides, iron oxides and sulfate as the most abundant electron acceptors (Froelich et al., 1979). It is when all alternative electron acceptors are depleted that fermentation takes place, leading to the production of CH<sub>4</sub>; methanogenesis. For example, it has long been established that in beach, estuarine, and marsh mudflats on the Brittany coast (France), organic matter (OM) degradation is dominated by sulfate reduction, as the high sulfate content of seawater inhibits methanogenesis through competitive inhibition (Winfrey and Ward, 1983). In contrast, sediments beneath thermokarst lakes are anoxic and largely devoid of alternative electron acceptors, so OM degradation is almost entirely driven by methanogenesis (Sepulveda-Jauregui et al., 2015). These examples highlight that the chemical composition of the aqueous environment plays a critical role in controlling the pathways of OM degradation. CH<sub>4</sub> produced in soils or sediments can also be oxidized by anaerobic methanotrophic archaea and sulfate-reducing bacteria (Boetius et al., 2000; La et al., 2022) present in the soils or sediment, contributing to lower CH<sub>4</sub> emissions in coastal environments. Thus, on or near the coast, the interaction with seawater, which contains electron acceptors such as sulfate, can shift the OM mineralization pathway and the resulting CO<sub>2</sub> and CH<sub>4</sub> productions. Consequently, a nuanced understanding of biogeochemical processes and their drivers is paramount in determining the magnitude of permafrost carbon emissions, especially from coastal environments.</p>
      <p id="d2e479">Numerous CH<sub>4</sub> emissions monitoring programs are in operation, but remote-sensing methods lack the ability to comprehensively capture the microbial, biogeochemical and environmental processes involved. In specific regions, estimates of methane production from the breakdown of OM is possible by carefully studying degradation pathways and production rates (Pellerin et al., 2022; Heslop et al., 2015; Knoblauch et al., 2018; Treat et al., 2014). To reduce the knowledge gap of CH<sub>4</sub> biogeochemistry in coastal permafrost settings, we collected material from the active layer and taliks of water bodies for incubation experiments, which were coupled to physical and chemical characterizations. The main objective of this study was to assess microbial CH<sub>4</sub> production dynamics in a coastal permafrost setting and apply it at the landscape level, since methane production is well documented in inland thermokarst but is not well understood in a land-ocean interaction context. We hypothesized that methanogenesis in coastal active layer incubations would be suppressed by the addition of sulfate. Consequently, we discuss the influence of environmental conditions on microbial CH<sub>4</sub> production with an emphasis on brackish water addition in coastal soils and sediments along with the microbial pathways involved. We then apply these results at the landscape level to provide an estimate of CH<sub>4</sub> production in the event that a natural process like a storm inputs brackish water over a large area of polygonal patterned ground. We use the region around Tuktoyaktuk as an example.</p>
</sec>
<sec id="Ch1.S2">
  <label>2</label><title>Methods</title>
<sec id="Ch1.S2.SS1">
  <label>2.1</label><title>Site description and sampling</title>
      <p id="d2e542">Tuktoyaktuk (69°26<sup>′</sup>24<sup>′′</sup> N, 133°01<sup>′</sup>52<sup>′′</sup> W) is located in the Inuvik region of the North-West Territories, adjacent to the Arctic Ocean in the Kugmallit Bay, east of the Mackenzie Delta. The region experiences prolonged cold winters, short cool summers, and year-round low precipitation, fostering low-arctic tundra vegetation. Lying in the continuous permafrost zone, its coastal areas feature thick Quaternary and glaciogenic unconsolidated deposits (Rampton, 1988), where permafrost thickness averages 400 m (Hu et al., 2013) and is characterized by prevalent ground ice structures (Mackay and Dallimore, 1992; Martin et al., 2018; Murton, 1996; Rampton, 1988). The area has been ice-free for the past 13 000 years, with evidence indicating that early Holocene summer temperatures were up to 6 °C warmer than today, fostering vegetation and peat accumulation (Dallimore et al., 1997; Vardy et al., 1997). During that same period, sea level was considerably lower than it is today and the Tuktoyaktuk area was located approximately 100 km inland (Vardy et al., 1997). Currently, ground subsidence and coastal erosion are major causes of rapid land retreat (Hynes et al., 2014; Lapham et al., 2020; Lim et al., 2020). Combined with sea level rise (Hill et al., 1993), it is projected that a substantial amount of terrestrial soil will become part of the ocean seafloor either by erosion and deposition or by subsidence of land and submersion. Over the past 15 years, extensive studies on Tuktoyaktuk's coastal environment, driven by the region's vulnerability to climate change, highlighted challenges for the Inuvialuit population relying on hunting, fishing, trapping and harvesting (Andrachuk and Smit, 2012).</p>
      <p id="d2e587">Active layer samples were collected from two sites: an inland site, Reindeer Point (RP) and a coastal site, Toker Point (TP). Talik sediments were also collected at both RP and TP sites from polygonal troughs and pondlets and sediments from a marine site, Harbor, completed the transect from terrestrial to marine settings (Fig. 2). RP was selected as the inland site because it features a polygonal patterned ground typical of the region, and is located in a stable region not directly affected by coastal processes such as storm surges, tides, seawater intrusion, erosion etc. The thermokarst lake margin, about 300 m south of RP has remain unchanged since aerial photos began recording the evolution of the landscape in 1947 (Fig. S1 in the Supplement). TP was selected as the coastal site because of the strong coastal processes such as tides and storm surge that regularly lead to seawater intrusion in this polygonal patterned ground, strongly influenced by ground subsidence. The Harbor site was selected about 400 m offshore in the Harbor of Tuktoyaktuk where total water depth was 20 m and cold marine bottom waters were overlain by a 10 m surface brackish water layer. 25 cm sediment cores were collected using a UWITEC gravity corer. The sediments consisted of recently deposited silty sands originating from the strong erosional processes occurring in the region (Whalen et al., 2022). The site was accessible by small watercraft. At RP and TP sites, soil profiles were extracted from the active layer by digging a soil pit with a shovel. To retain an intact stratigraphic relationship, samples were taken from the wall of the soil pit. Biogenic ebullition gases were collected from pondlets at RP and TP. Pondlets were located within sampled polygonal patterned ground and are defined as small (1 to 3 m<sup>2</sup>) and shallow standing bodies of water, potentially draining seasonally. Samples were trapped using a plastic funnel attached to a 20 mL glass vial. Surface soil lying at the bottom of the pondlets (Fig. 1) were poked until the vial was filled with gas. Once full, vials were crimped with 20 mm butyl rubber stoppers and aluminum caps. Samples were kept frozen until the time of analyses.</p>

      <fig id="F2"><label>Figure 2</label><caption><p id="d2e601">Map of study area indicating the sampled sites with yellow dots (Esri, 2022; powered by Esri). Harbor site is located in the marine waters of Tuktoyaktuk, Toker Point site is located in the coastal (intertidal) zone and Reindeer Point site is located inland. High resolution satellite imagery and pictures of soil profiles for RP and TP sites available in the Supplement (Fig. S3).</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/3777/2026/bg-23-3777-2026-f02.png"/>

        </fig>

      <p id="d2e611">The inland site, (RP), was located 750 m from the coast and 2 km East of Tuktoyaktuk in a polygonal patterned ground. This patterned ground is located in a depression, surrounded by elevated plateaus with observable ground water flowing into the valley. In this area, low-centered polygons exhibited higher moisture levels compared to high-centered polygons. High-centered polygons were colonized by shrubs and small flowering plants like <italic>Ericaceae</italic>, while low-centered polygons were dominated by hydrophilic plants such as grasses and sedges. Wet troughs delimited the polygons, with vegetation reflecting waterlogged conditions. The mean active layer and talik thickness across RP was about 35 cm. Profile 10A was collected from a trough and presented water-saturated conditions with brown OM. Profiles 10B and 10D were collected from high-centered polygons and characterized by unsaturated conditions with dark brown OM and presence of roots until 20 cm depth. Profile 10C was collected from a low-centered polygon and consisted of reddish-brown peat throughout. Profiles 10A, 10B and 10D did not consist of peat.</p>
      <p id="d2e617">The coastal site (TP) is located 20 km NW of Tuktoyaktuk, featuring a polygonal patterned ground, largely colonized by <italic>Carex</italic> sp., a type of graminoid plant common near Arctic coastlines. The mean active layer and talik depths were 35 cm. The site's dynamics are influenced by the twice-daily ebb and flow of tides. Profile 07 was collected from a water-saturated low-centered polygon, located in the intertidal zone. The soil color was very dark greyish black. Profile 08 was collected from a water-saturated polygonal trough not immediately located in the intertidal zone, but which floods during storms The soil was characterized by dark greyish-brown OM mixed with sand. Finally, profile 09, was collected from the center of a higher-centered polygon situated in the middle intertidal zone. The active layer appeared water unsaturated. The soil from this site consisted of a mixture of black organic-rich material and sand. The sand found in samples from TP appeared to be wind-deposited from nearby dunes.</p>
</sec>
<sec id="Ch1.S2.SS2">
  <label>2.2</label><title>Sulfates and chloride concentrations in sediments</title>
      <p id="d2e631">The extraction of sulfate and chloride from sediments and soils pore-water was conducted through a leaching experiment following Lacelle et al. (2019). Frozen aliquots of sediments and soils were thawed at 4 °C overnight, then weighed, dried in the oven at 60 °C for 24 h and re-weighed to determine the densities. Aliquots of dried material were put in 50 mL falcon tubes with nanopure water following a <inline-formula><mml:math id="M41" display="inline"><mml:mrow><mml:mn mathvariant="normal">1</mml:mn><mml:mo>:</mml:mo><mml:mn mathvariant="normal">10</mml:mn></mml:mrow></mml:math></inline-formula> ratio. Tubes were then shaken for one hour to promote leaching of anions towards the aqueous phase of the solution. Once the leaching process was done, 2 mL of the aqueous solution was filtered using 0.2 <inline-formula><mml:math id="M42" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m pore size Whatman 25 mm GD/X syringe filters and transferred in disposable microtubes. Concentrations of sulfate and chloride were measured by ion chromatography using a Thermo Dionex Integrion at UQAR's Chemistry department facilities with a limit of detection of 0.01 <inline-formula><mml:math id="M43" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>g mL<sup>−1</sup>. The measured concentrations are expressed in mmol g<sup>−1</sup> per wet-weight of material (mmol g<sup>−1</sup> wweight<sup>−1</sup>). Only one measurement per sample was performed as stability tests revealed variability of less than 3 % between measured samples. The error on each value was calculated by the least squares method (Skoog et al., 2014).</p>
</sec>
<sec id="Ch1.S2.SS3">
  <label>2.3</label><title>Methane production rates in incubations</title>
      <p id="d2e719">Long-term sediment and soil incubations under anoxic conditions were used to assess CH<sub>4</sub> production rates over several months by measuring CH<sub>4</sub> accumulation in the vials' headspace. The objective was to simulate the increased connectivity between the land and the ocean in the coastal environment of the Canadian Arctic, which represents an important aspect of the ongoing regional environmental transition. Collected sediments and soil profiles were immediately sub-sampled based on depth, at 5 or 10 cm intervals, according to shifts in sedimentary units. To prepare incubations, about 4 mL of sediment and exactly 2 mL of brackish water (collected from the coast) were immediately transferred into 20 mL glass vials. Incubation vials were crimped with 20 mm blue chlorobutyl rubber stoppers and aluminum caps. The bottles were flushed with nitrogen gas (Alpha Gaz 1) at a rate of 300 mL min<sup>−1</sup> for 2 min in the field to replace the air with a nitrogen atmosphere. Four incubations were prepared for each sampled depth; 3 were kept for measurements of CH<sub>4</sub> production rates (triplicates) and one served for isotopic analyses. Incubations were kept at a constant temperature of 4 °C throughout the entire 339 d incubation period with no fluctuations. Substrate concentrations were not actively controlled or monitored, aside from repeated measurements of headspace methane. For logistical reasons, we were not able to measure CH<sub>4</sub> concentrations from the incubations in the first few weeks and the last measurement was conducted at day 339. The brackish water added to all incubations contained 5.7 <inline-formula><mml:math id="M53" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.0 mmol g<sup>−1</sup> wweight<sup>−1</sup> of sulfate and 28.7 <inline-formula><mml:math id="M56" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.5 mmol g<sup>−1</sup> wweight<sup>−1</sup> of chloride.</p>
      <p id="d2e833">Analyses of the CH<sub>4</sub> concentrations in the headspace of the vials were performed on a gas chromatograph (Agilent 8900) equipped with a flame ionization detector (GC-FID) at UQAR facilities. The GC-FID is equipped with a 100 <inline-formula><mml:math id="M60" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>L injection loop to ensure a consistent volume of sample is analyzed. To saturate the injection loop, 300 <inline-formula><mml:math id="M61" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>L are taken from the headspace of the vials and transferred to the injection loop with a gas-tight syringe. Prior to injection, samples were shaken for 30 s to equilibrate headspace and sediment gases. This procedure was done every 2 weeks for 16 weeks to measure CH<sub>4</sub> accumulation in the headspace. The resulting production rates were calculated from the linear accumulation measured during the incubation period, and values are expressed in nmol of CH<sub>4</sub> per cubic centimeters of wet material per day (nmol cm<sup>−3</sup> d<sup>−1</sup>). The density of the collected samples varied widely, with some being organic deposits and peat, while others contained higher mineral content. Consequently, the CH<sub>4</sub> production rates were expressed volumetrically to account for these discrepancies which are more representative of the volume they occupy in the soil, sedimentary columns and landscape. The limit of detection of the GC-FID is 0.3 ppm and all samples had higher concentrations. Each value represents the mean of triplicate measurements and the reported uncertainty on the measurement is the standard deviation on triplicates.</p>
      <p id="d2e913">To estimate the potential total active layer CH<sub>4</sub> production (<inline-formula><mml:math id="M68" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>), the active layer production rates were vertically integrated to obtain the total CH<sub>4</sub> production of each profile. Values are reported in mmol m<sup>−2</sup> d<sup>−1</sup> and were calculated using Eq. (1):

            <disp-formula id="Ch1.E1" content-type="numbered"><label>1</label><mml:math id="M72" display="block"><mml:mrow><mml:mi>T</mml:mi><mml:mo>=</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mn mathvariant="normal">1</mml:mn><mml:mn mathvariant="normal">100</mml:mn></mml:mfrac></mml:mstyle><mml:msubsup><mml:mo>∑</mml:mo><mml:mrow><mml:mi>i</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow><mml:mi>n</mml:mi></mml:msubsup><mml:msub><mml:mi>P</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>⋅</mml:mo><mml:msub><mml:mi>e</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mfenced close="]" open="["><mml:mrow><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">d</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:mfenced></mml:mrow></mml:math></disp-formula>

          where <inline-formula><mml:math id="M73" display="inline"><mml:mrow><mml:msub><mml:mi>P</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> represents CH<sub>4</sub> production rate in layer <inline-formula><mml:math id="M75" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula> (nmol cm<sup>−3</sup> d<sup>−1</sup>), <inline-formula><mml:math id="M78" display="inline"><mml:mrow><mml:msub><mml:mi>e</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> represents the thickness of layer <inline-formula><mml:math id="M79" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula> (cm) and <inline-formula><mml:math id="M80" display="inline"><mml:mi>n</mml:mi></mml:math></inline-formula> represents the numbers of layers in the profile.</p>
      <p id="d2e1105">Using aerial imagery from 2022, the polygonal tundra at RP was mapped in QGIS, allowing for the discrimination between high-centered polygons, low-centered polygons and throughs (Fig. S2). The total area of each geomorphological form was calculated based on the map data (Table S1). Landforms total areas were multiplied by the corresponding potential total active layer methane production (<inline-formula><mml:math id="M81" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>) to estimate the total CH<sub>4</sub> produced in the polygonal tundra of RP over a day (mol d<sup>−1</sup>).</p>
</sec>
<sec id="Ch1.S2.SS4">
  <label>2.4</label><title>Elemental and isotope composition of the sediment</title>
      <p id="d2e1144">The total organic carbon (TOC) content of the sediments was measured by combustion using an elemental analyzer (ECS 8020, NC Technologies) combined with a gas chromatograph equipped with a thermal conductivity detector at ULaval facilities (The International Research Laboratory Takuvik). A 100 mg aliquot of sediment was thawed and weighed for each sample. They were then dried in an oven at 60 °C for 48 h and re-weighed to determine their water content. Sediments were then ground using a granite mortar pestle and homogenized using a 1.18 mm pore size sieve to remove roots and rootlets. Instruments were cleaned with ethanol between manipulations. Inorganic carbon was removed from sediments by adding 2.2 mL of 12 M HCl in every sample. After reacting for 24 h, around 8 mg was encapsulated in tin foil capsules. Samples were kept in a desiccator until analyses. Values are expressed as % of carbon contained in the weighed sample (wt %).</p>
      <p id="d2e1147">The organic carbon (<inline-formula><mml:math id="M84" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C-TOC) isotopic compositions were measured at UOttawa facilities (Jàn Veizer Stable Isotope Laboratory) using EA-IRMS (Delta Advantage, Thermo Germany). The sample preparation method was the same used for elemental analyses. <inline-formula><mml:math id="M85" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C-TOC values are denoted as <inline-formula><mml:math id="M86" display="inline"><mml:mi mathvariant="italic">δ</mml:mi></mml:math></inline-formula> ‰ <inline-formula><mml:math id="M87" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula>10<sup>3</sup> ((<inline-formula><mml:math id="M89" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mi mathvariant="normal">sample</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>/<inline-formula><mml:math id="M90" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mi mathvariant="normal">standard</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) <inline-formula><mml:math id="M91" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula> 1), where <inline-formula><mml:math id="M92" display="inline"><mml:mi>R</mml:mi></mml:math></inline-formula> is <sup>13</sup>C <inline-formula><mml:math id="M94" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> <sup>12</sup>C and standards refer to the Vienna Pee Dee Belmnite (VPDP).</p>

      <fig id="F3"><label>Figure 3</label><caption><p id="d2e1260">Total organic carbon and sulfate (SO<inline-formula><mml:math id="M96" display="inline"><mml:mrow><mml:msubsup><mml:mi/><mml:mn mathvariant="normal">4</mml:mn><mml:mrow><mml:mn mathvariant="normal">2</mml:mn><mml:mo>-</mml:mo></mml:mrow></mml:msubsup></mml:mrow></mml:math></inline-formula>) concentrations in sediment or soil of different sites in this study. The datasets are separated into two for clarity. The upper part of the figure <bold>(a, b) </bold>displays the data of the marine site Harbor (profile 01 and 02), and the coastal site, Toker Point (profile 07, 08 and 09). The lower part of the figure <bold>(c, d)</bold> displays the data from the inland site Reindeer Point (profile 10A, 10B, 10C and 10D). The black horizontal dotted line in each graph represents the permafrost-active layer or talik interface except for the Harbor site, where the talik is much deeper but not measured. TOC data from Harbor site is not available. A uniform color pattern is used throughout this manuscript.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/3777/2026/bg-23-3777-2026-f03.png"/>

        </fig>

      <fig id="F4"><label>Figure 4</label><caption><p id="d2e1293">CH<sub>4</sub> production in incubations of soil and sediment with brackish water from <bold>(a)</bold> TP and <bold>(b)</bold> RP. Each datapoint represent the mean value of three incubations. The error bars in grey lines are equal to the standard deviation of the three separate incubations. Each profile corresponds to a specific landform. At Toker Point <bold>(a)</bold>, profile 07 is from a low-centered polygon, profile 08 is from a trough and profile 09 is from a high-centered polygon. At Reindeer point <bold>(b)</bold>, profile 10A is from a high-centered polygon, profile 10B is from a trough, profile 10C is from a low-centered polygon and profile 10D is from a high-centered polygon.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/3777/2026/bg-23-3777-2026-f04.png"/>

        </fig>

</sec>
<sec id="Ch1.S2.SS5">
  <label>2.5</label><title>Stable carbon isotopic composition of methane</title>
      <p id="d2e1332">One incubation vial was analyzed for stable carbon isotopic composition of headspace methane (<inline-formula><mml:math id="M98" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C–CH<sub>4</sub>). Stable carbon from methane ebullition samples collected from pondlets were also analyzed. Both types of samples were analyzed with a cavity ring-down spectrometer (PICARRO G2201-i isotopic CO<sub>2</sub> <inline-formula><mml:math id="M101" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> CH<sub>4</sub>) equipped with a 16-port distribution manifold and small sample introduction module (SSIM) at McGill (McGill Isotope Biogeochemistry Laboratory). Incubations were kept at 4 °C in the dark for 8 months to let the microbial community stabilize and produce sufficient CH<sub>4</sub> for analysis. To stay in the detection range of the analyzer (1.8–1000 ppm CH<sub>4</sub>), a small volume of the headspace, proportional to CH<sub>4</sub> concentration in sample, was drawn from the incubation vial (0.2–6 mL). The sample was introduced to the 16-port manifold with a 21 G needle connected to a disposable luer lock plastic syringe. Samples were diluted with zero air by the SSIM to reach a volume of 20 mL. Two or three measurements per sample were conducted depending on headspace concentration. Ebullition gases samples were analyzed following the same method. Measured values were corrected with internal certified methane standards (<inline-formula><mml:math id="M106" display="inline"><mml:mo lspace="0mm">-</mml:mo></mml:math></inline-formula>59 ‰ and <inline-formula><mml:math id="M107" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>42 ‰ ) from AirLiquide and stability of the analyzer was tested with injections of ambient air. Measured values were more precise than <inline-formula><mml:math id="M108" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula>1.2 ‰. All <inline-formula><mml:math id="M109" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C–CH<sub>4</sub> values are expressed relatively to VPDB. While those isotopic analyses results provide valuable insight into methane cycling processes, they should be interpreted with caution in the absence of biological replication.</p>

      <fig id="F5" specific-use="star"><label>Figure 5</label><caption><p id="d2e1452">Total CH<sub>4</sub> production rates (T) at Toker Point (Coastal) and Reindeer Point (Inland) organized by geomorphological forms of high-centered polygons, low-centered polygons and throughs. High-centered polygons at RP is the mean of two profiles. All other landforms at RP and TP are one profile. The uncertainty on T is propagated from the uncertainty of individual CH<sub>4</sub> production rates, not averages from replicate sites.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/3777/2026/bg-23-3777-2026-f05.png"/>

        </fig>

      <fig id="F6"><label>Figure 6</label><caption><p id="d2e1481">Isotopic composition of CH<sub>4</sub> produced in brackish water incubations from <bold>(a)</bold> TP and <bold>(b)</bold> RP. Each datapoint corresponds to the mean value of two or three measurements done on one incubation, depending on the headspace concentration. The dashed vertical lines correspond to in situ ebullition CH<sub>4</sub> collected in pondlets at each sampling site (<inline-formula><mml:math id="M115" display="inline"><mml:mi>n</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M116" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 1). These values give information on the pathways used by the soil microbes to produce CH<sub>4</sub>. <inline-formula><mml:math id="M118" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C between <inline-formula><mml:math id="M119" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>65 ‰ and <inline-formula><mml:math id="M120" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>50 ‰ is typically associated with acetoclastic methanogenesis, while <inline-formula><mml:math id="M121" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C between <inline-formula><mml:math id="M122" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>110 ‰ and <inline-formula><mml:math id="M123" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>60 ‰ is associated with hydrogenotrophic methanogenesis (Hornibrook et al., 1997, 2000). The grey error bars on each point represents the analytical uncertainty on the measured value. If not visible, the uncertainty is smaller than the point.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/3777/2026/bg-23-3777-2026-f06.png"/>

        </fig>

</sec>
</sec>
<sec id="Ch1.S3">
  <label>3</label><title>Results</title>
<sec id="Ch1.S3.SS1">
  <label>3.1</label><title>Soil description and composition</title>
      <p id="d2e1605">TOC content in the sampled soils ranged from 2 wt % to 47 wt %, with no clear trend in relation to depth (Fig. 3a, c). The RP polygonal patterned ground featured organic soils with TOC content ranging from 14 wt % to 47 wt % (Fig. 3c). The TP coastal polygonal patterned ground also featured organic soils with TOC content ranging from 2 wt % to 37 wt % (Fig. 3a).</p>
      <p id="d2e1608">RP, the inland site, had low sulfate and chloride concentrations relative to TP, the coastal site (Figs. 3b, d and S2). Sulfates at RP ranged from null concentrations to 0.68 <inline-formula><mml:math id="M124" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.03 mmol g<sup>−1</sup> wweight<sup>−1</sup>, while at TP, profiles exhibited varying concentrations and patterns in relation to depth. Sulfate concentrations, ranged from 0.07 <inline-formula><mml:math id="M127" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.03 to 12.72 <inline-formula><mml:math id="M128" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.03 mmol g<sup>−1</sup> wweight<sup>−1</sup>. Profile 07, the low-centered polygon, exhibited the highest sulfates concentrations of all TP site at its surface (12.72 <inline-formula><mml:math id="M131" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.03 mmol g<sup>−1</sup> wweight<sup>−1</sup>), with concentrations decreasing drastically with depth, reaching 0.29 <inline-formula><mml:math id="M134" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.03 mmol g<sup>−1</sup> wweight<sup>−1</sup> at 25 cm (Fig. 3a). In profile 09, the high-centered polygon, sulfate concentrations increased with depth ranging from 0.09 <inline-formula><mml:math id="M137" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.03 mmol g<sup>−1</sup> wweight<sup>−1</sup> at 5 cm to 3.2 <inline-formula><mml:math id="M140" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.03 mmol g<sup>−1</sup> wweight<sup>−1</sup> at 25 cm. Finally, profile 08, characterized as a polygonal trough, had sulfate concentrations ranging from 0.07 <inline-formula><mml:math id="M143" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.03 to 0.75 <inline-formula><mml:math id="M144" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.03 mmol g<sup>−1</sup> wweight<sup>−1</sup>. The highest sulfate concentrations measured in this study were found in the sediments of the Harbor site, with a mean value of 16.6 mmol g<sup>−1</sup> wweight<sup>−1</sup> (Fig. 3a).</p>
</sec>
<sec id="Ch1.S3.SS2">
  <label>3.2</label><title>Methane production</title>
      <p id="d2e1878">Rates of CH<sub>4</sub> production in incubations of sediment and soil with brackish water were undertaken at the three studied sites: RP, TP and Harbor. Production rates ranged from null to 415.4 <inline-formula><mml:math id="M150" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 69.2 nmol cm<sup>−3</sup> d<sup>−1</sup> (Fig. 4) throughout all samples in this study. At RP, the maximum CH<sub>4</sub> production rate of 35.2 <inline-formula><mml:math id="M154" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 15.7 nmol cm<sup>−3</sup> d<sup>−1</sup> was measured in the trough profile (10B) at a depth of 20 cm. Lower values were obtained for the surface and at the talik-permafrost interface. The low-centered polygon (10C) had its maximum CH<sub>4</sub> production rate in the surface, decreasing with depth. High-centered polygons (10A and 10D) had very low production rates along their depth profiles ranging between null to 1.2 <inline-formula><mml:math id="M158" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.2 nmol cm<sup>−3</sup> d<sup>−1</sup>. Both water-saturated trough and low-centered polygon (10B, 10C) had relatively high CH<sub>4</sub> production rate compared with the high-centered polygon profiles (10A, 10D), which were water-unsaturated.</p>
      <p id="d2e2012">At TP, a maximum CH<sub>4</sub> production rate was recorded in profile 09, the high-centered polygon at 415.4 <inline-formula><mml:math id="M163" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 69.2 nmol cm<sup>−3</sup> d<sup>−1</sup> at the uppermost depth but it quickly decreased in the subsurface. Profile 08, the trough, and profile 07, the low-centered polygon, had lower sub-surface CH<sub>4</sub> production rates, but rates decreased less drastically with depth with values being relatively high at the permafrost-talik and permafrost-active layer interface, respectively. Profile 07 had values ranging from 27.9 <inline-formula><mml:math id="M167" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 1.5 to 92.8 <inline-formula><mml:math id="M168" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 21.2 nmol cm<sup>−3</sup> d<sup>−1</sup> and profile 08 had values ranging from 50.4 <inline-formula><mml:math id="M171" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 7.2 and 153.7 <inline-formula><mml:math id="M172" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 33.9 nmol cm<sup>−3</sup> d<sup>−1</sup> (Fig. 4). In general, at TP, the coastal site, much higher CH<sub>4</sub> production rates were measured than at RP, the inland site (Fig. 4). The mean CH<sub>4</sub> production rate measured at RP was 5.7 nmol cm<sup>−3</sup> d<sup>−1</sup>, while at TP it was 96.2 nmol cm<sup>−3</sup> d<sup>−1</sup>. The incubations with silty-clay Harbor sediments did not have measurable CH<sub>4</sub> production rates (Fig. S5).</p>
      <p id="d2e2218">Estimated total CH<sub>4</sub> production rates were calculated for each geomorphological landforms of RP and TP sites. At RP, the total CH<sub>4</sub> production estimated for the high-centered polygons (profile 10A and 10D), low-centered polygon (profile 10C) and trough (profile 10B) were 0.3 <inline-formula><mml:math id="M184" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.1, 2.4 <inline-formula><mml:math id="M185" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 1.0 and 5 <inline-formula><mml:math id="M186" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 2 mmol m<sup>−2</sup> d<sup>−1</sup>, respectively (Fig. 5). At TP, the total CH<sub>4</sub> production estimated for the high-centered polygon (profile 09), the low-centered polygon (profile 07) and the through (profile 08) were 41.5 <inline-formula><mml:math id="M190" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 6.9, 16.8 <inline-formula><mml:math id="M191" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.0 and 28.3 <inline-formula><mml:math id="M192" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 7.4 mmol m<sup>−2</sup> d<sup>−1</sup>, respectively (Fig. 5). In all landforms, the total CH<sub>4</sub> production rates were higher in the coastal site, TP, than the inland site, RP.</p>
</sec>
<sec id="Ch1.S3.SS3">
  <label>3.3</label><title>Isotopic composition of <sup>13</sup>C–CH<sub>4</sub></title>
      <p id="d2e2375">In parallel with CH<sub>4</sub> production rates, one incubation vial per depth was used to measure the stable carbon isotopic composition of the CH<sub>4</sub> produced. At RP, the <inline-formula><mml:math id="M200" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C–CH<sub>4</sub> of the first sampled depth (10 cm) ranged from <inline-formula><mml:math id="M202" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>81.3 ‰ to <inline-formula><mml:math id="M203" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>89.4 ‰ . At TP, the coastal site, the <inline-formula><mml:math id="M204" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C–CH<sub>4</sub> signature of the first sampled depth (5 cm) ranged from <inline-formula><mml:math id="M206" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>47.1 ‰ and <inline-formula><mml:math id="M207" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>51.6 ‰ . The values cluster together based on site, suggesting surface OM degradation processes are most similar within sites than between sites (Fig. 6). Profiles at RP became progressively enriched in <sup>13</sup>C with depth, except for profile 10C where a more depleted value was observed at 35 cm. Conversely, at TP, profiles became depleted in <sup>13</sup>C with depth, except for profile 08 where an enrichment was measured between 15 and 30 cm.</p>
      <p id="d2e2484">Ebullition samples from pondlets were also measured for stable isotopes. The ebullition samples represent the net <inline-formula><mml:math id="M210" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C signature of methane produced in the sediments of pondlets at RP and TP. At RP, CH<sub>4</sub> ebullition from a sampled thaw pond had a <inline-formula><mml:math id="M212" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C of <inline-formula><mml:math id="M213" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>66.1 ‰ (Fig. 6). At TP, CH<sub>4</sub> ebullition from a sampled pondlet had a <inline-formula><mml:math id="M215" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C–CH<sub>4</sub> of <inline-formula><mml:math id="M217" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>65.0 ‰ (Fig. 6).</p>
</sec>
</sec>
<sec id="Ch1.S4">
  <label>4</label><title>Discussion</title>
<sec id="Ch1.S4.SS1">
  <label>4.1</label><title>Addition of brackish water to anoxic incubations did not strongly suppress methanogenesis</title>
      <p id="d2e2579">Before discussing the effects of brackish water addition in incubation experiments, it is important to clarify the role of sulfate measured in situ within soil and sediment profiles. Across the studied sites, sulfate concentrations varied with depth and between landforms; however, this spatial variability did not show a consistent relationship with methane production rates measured in the incubations (Fig. S4). A few layers clearly contained higher sulfate amounts. However, layers characterized by higher or lower sulfate concentrations did not systematically correspond to lower or higher CH<sub>4</sub> production, indicating that in situ sulfate availability alone does not explain the observed patterns in methane production across profiles. This interpretation is subject to important limitations. Sulfate and chloride concentrations were measured at single points within each profile and were not replicated across multiple locations within the same landform, preventing resolution of fine-scale spatial heterogeneity in electron-acceptor availability. As a result, sulfate concentrations are interpreted here as first-order indicators of geochemical context rather than as spatially representative or mechanistic controls on methane production. Given these constraints, we focus the following discussion on the experimental addition of sulfate via brackish water during anoxic incubations, which evaluate how episodic marine influence may affect methane production potential in coastal permafrost environments.</p>
      <p id="d2e2591">Despite the addition of brackish water containing sulfate to incubations, the range of CH<sub>4</sub> production rates measured in this study is consistent with reports for anaerobic incubations of recently thawed permafrost soils, suggesting that the input of brackish water to some costal systems may not inhibit CH<sub>4</sub> production. For example, in the talik of Big Trail Lake, a young thermokarst lake in the interior of Alaska, CH<sub>4</sub> production rates based on incubations ranged between 4.7 and 16.1 nmol cm<sup>−3</sup> d<sup>−1</sup> (Pellerin et al., 2022), while in incubations from Vault Lake, another thermokarst lake in the interior of Alaska, CH<sub>4</sub> production rates varied between 11.1 and 275 nmol cm<sup>−3</sup> d<sup>−1</sup> (Heslop et al., 2015). In active layer incubations from the Yamal Peninsula in NW Siberia (Russia), CH<sub>4</sub> production rates of incubations varied between 0.1 and 33.8 nmol cm<sup>−3</sup> d<sup>−1</sup> (Heyer et al., 2002). This indicates that overall, the CH<sub>4</sub> production rates measured at both TP and RP are within the range observed in typical ice-rich permafrost settings and reasonable for the environment studied (Fig. 4). We note that our experimental design did not include parallel incubations without brackish water or with sulfate concentration gradients; therefore, our interpretation relies in part on comparison with previous incubations of Tuktoyaktuk soils conducted without brackish water addition (Lapham et al., 2020) and should be regarded as exploratory rather than definitive. Lapham et al. (2020) conducted sediment incubation experiments using a core collected from the coast of the Tuktoyaktuk Peninsula. In their study, CH<sub>4</sub> production was measured under anaerobic conditions at 15 °C, without the addition of water, over a 35 d period. The reported CH<sub>4</sub> production rate was 0.07 nmol cm<sup>−3</sup> d<sup>−1</sup>. Although their incubations were performed over a shorter duration and at a significantly higher temperature than those in the present study, the measured rate reflects CH<sub>4</sub> production under relatively natural, unamended conditions. This value is comparable to the lowest CH<sub>4</sub> production rates measured in our incubations at both the coastal (TP) and inland (RP) sites and provides a useful reference for CH<sub>4</sub> production under unamended conditions.</p>
      <p id="d2e2792">The novel aspect of this study is that it attempts to understand marine influence on OM degradation by addition of brackish water to sediment and soil incubations of a fully marine site (Harbor), one that is periodically submerged (TP) and never submerged (RP). This simulates the input of seawater to the active layer and taliks of tundra soils (RP) as well as providing reference sites with a high marine influence (Habor and TP). We hypothesized that the addition of locally obtained brackish water, which contained sulfate (5.7 mmol L<sup>−1</sup>), to the incubations, would suppress CH<sub>4</sub> production in RP, the inland site and potentially also at TP, the coastal site. This reasoning is because supplying sulfate to low sulfate organic-rich sediment would promote sulfate reduction, which is thermodynamically more favorable than methanogenesis, thereby competitively inhibiting it (Lovley and Klug, 1983; Oremland and Polcin, 1982). This hypothesis is also consistent with field observations; organic matter mineralization in brackish wetlands is consistently dominated by bacterial sulfate reduction (Bridgham et al., 2012; Torres-Alvarado et al., 2005) where little to no CH<sub>4</sub> emissions are observed (Pönisch et al., 2023; Petersen et al., 2023; Kroeger et al., 2017). However, recent field studies show that in coastal permafrost soils, inundation and low sulfate concentrations do not necessarily suppress methanogenesis (Jenrich et al., 2025; Jenrich et al., 2024; Yang et al., 2023). These contrasting observations reveal a key knowledge gap in how marine influence controls carbon mineralization pathways in permafrost systems. By experimentally testing brackish water additions across sites with contrasting marine exposure, our study provides new mechanistic insight into the regulation of OM degradation and CH<sub>4</sub> production under ongoing Arctic coastal change.</p>
      <p id="d2e2834">RP had low sulfate concentrations before addition of the brackish water but so did many of the profiles from TP (Fig. 3). In the Harbor sediments, no methane production was observed (Fig. S5). This is consistent with the competitive inhibition of methanogenesis by energetically favorable redox reactions with electron acceptors like oxygen, nitrate, iron oxides or sulfate that is typical of marine systems e.g. (Martens and Berner, 1974) as well as the potential for anaerobic oxidation of methane (AOM). Given that the Harbor sediments already had high sulfate concentrations, the lack of methane production with addition of brackish water was expected. However, strong CH<sub>4</sub> production was observed in the incubations of both the coastal site TP and the inland site RP, indicating that CH<sub>4</sub> production was not halted by the addition of sulfate via the brackish water addition at those sites. While sulfate reduction rates were not measured and therefore not demonstrated directly in our incubations, a strong sulfide smell was recorded when opening most of the incubations at the end of the experiment. This observation may indicate the coexistence of sulfate reduction and methanogenesis during the incubations. However, to rigorously assess this observation, future studies should include tracer-based sulfate reduction assays and microbial functional gene analysis.</p>
      <p id="d2e2856">Coexistence of sulfate and methanogenesis within complex sediment systems such as estuarine, coastal and salt marsh sediments, as well as thermokarst lasens has been widely reported (Lovley and Klug, 1983; Oremland and Polcin, 1982; Sela-Adler et al., 2017; Yang et al., 2023). Two main mechanisms are invoked to explain this co-existence in our incubation experiment: (1) noncompetitive methanogenesis (i.e. methylotrophic methanogenesis) and (2) syntrophic methanogenesis. (1) Noncompetitive substrates are substrates like methanol and methylamines that are used by methanogens alone and cannot be used with electron acceptors like sulfate (Lovley and Klug, 1983; Oremland and Polcin, 1982). Noncompetitive substrates are thus microbially converted to CH<sub>4</sub>, even in sediments with high sulfate concentrations (Maltby et al., 2018; Yuan et al., 2019). For example, in salt marshes, where high sulfate concentrations are often found, elevated CH<sub>4</sub> emissions are suggested to mainly stem from noncompetitive methanogenesis (Comer-Warner et al., 2022; Poffenbarger et al., 2011; Yuan et al., 2019). In the sulfate reducing zone of sediments from the Baltic Sea, where ample sulfate is found in the porewaters, seasonal methanogenesis rates were measured up to 1.3 nmol cm<sup>−3</sup> d<sup>−1</sup> due to noncompetitive substrates (Maltby et al., 2018). In permafrost soils, methanol, methylamines and the microorganisms capable of degrading them have been observed but their concentrations are typically low (Coolen and Orsi, 2015; Kramshøj et al., 2018). However, our study sites are on a coast undergoing a rapid transgression which may be driving imbalances between substrate supply and microbial abundances. The rates of methane production observed at RP and TP of up to 154 nmol cm<sup>−3</sup> d<sup>−1</sup> contrast with reported values for methylotrophic methanogenesis (Maltby et al., 2018). Based on these numbers, noncompetitive substrates likely play a small role in the total methane production at our study sites but further investigation into methylotrophic methane production in coastal environments will allow to document the overall role of methylotrophic methane production in coastal permafrost settings.</p>
      <p id="d2e2926">(2) Syntrophic methanogenesis occurs when molecular hydrogen produced by acetoclastic sulfate-reducing bacteria is used by hydrogenotrophic methanogens. In this syntrophy, the chemical energy is shared via interspecies hydrogen transfer (Ozuolmez et al., 2015). For instance, in permafrost soils of Sweden, it was demonstrated that syntrophic methanogenesis was favored in anoxic and water-saturated soils by an elevated abundance in methanogens and their syntrophic partners (Keuschnig et al., 2022). As the incubation experiment in our study at RP and TP featured water-saturated and anoxic environments, syntrophic methanogenesis could participate in the co-occurrence of sulfate-reduction and methanogenesis. This mechanism is consistent with most incubations producing methane with a <inline-formula><mml:math id="M250" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C value in the range of hydrogenotrophic methanogenesis (see below).</p>
      <p id="d2e2940">Measuring methane production through incubations inherently has limitations as they prevent continuous inputs of microorganisms, fresh OM and nutrients that would occur in the natural environment. This can create a “bottle effect”, which leads to restrictions in microbial community composition, limits the input of nutrients and leads to the accumulation of metabolites which would normally be degraded (Ionescu et al., 2015). Typically, overestimation of microbial processes rates is observed compared to in situ data (Sherr et al., 1999). The overestimation of CH<sub>4</sub> production rates by incubations relative to the in situ rates are difficult to assess because of a lack of data in permafrost environments (Heslop et al., 2020). Furthermore, a lag time between the start of anaerobic incubations and maximum CH<sub>4</sub> production rate is widely documented, which appears to be the case for both active layer and thawed permafrost incubations (Holm et al., 2020; Knoblauch et al., 2018; Knoblauch et al., 2013; Roy Chowdhury et al., 2015). Drier or water-unsaturated conditions lead to a longer lag time before the onset of maximum CH<sub>4</sub> production (Treat et al., 2014). Microbial community composition in the soil or sediment also exerts a strong control on the organic carbon degradation and has been shown to change throughout the incubations (Holm et al., 2020). Low initial methanogen population in soils can contribute to this lag time, but other factors such as disturbance of sediment during sampling, substrate availability and redox state can also contribute to the observed lag time in some incubations (Treat et al., 2014; Roy Chowdhury et al., 2015).</p>
      <p id="d2e2970">Furthermore, it is also possible that a “priming effect” from the addition of brackish water in incubations could have supercharged OM degradation with marine organic carbon, nutrients and microorganisms (Bianchi, 2011), which may have enhanced CH<sub>4</sub> production. However, this priming effect was not observed in the Harbor sediments which were amended with the same brackish water. Furthermore, CH<sub>4</sub> ebullition samples collected from pondlets adjacent to RP and TP exhibited broadly similar <inline-formula><mml:math id="M256" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C values to methane produced in incubations (Fig. 6), suggesting a similitude in microbial degradation pathways to methane in situ and in the incubations. Despite these uncertainties, our dataset shows clear depth trends and landscape-level variations, indicating that even under brackish water addition, local conditions will strongly influence CH<sub>4</sub> production.</p>
</sec>
<sec id="Ch1.S4.SS2">
  <label>4.2</label><title>CH<sub>4</sub> production pathways depend on hydrology and organic matter lability</title>
      <p id="d2e3029">The addition of brackish water resulted in incubation conditions being water-saturated in all cases, but it appears that biological and hydrological conditions of the polygonal patterned grounds influenced the magnitude of CH<sub>4</sub> production, nonetheless.</p>
      <p id="d2e3041">In all landforms, CH<sub>4</sub> production rates were lower at the inland site, RP than at TP, the coastal site (Fig. 4). Inland, low-centered polygons and troughs have typically higher CH<sub>4</sub> fluxes than unsaturated landforms like high-centered polygons (Roy Chowdhury et al., 2015; Martin et al., 2018; Zheng et al., 2018) which indicates they may also have higher CH<sub>4</sub> production rates. Within sites in our study, brackish water amended incubations of high-centered polygon soils had lower CH<sub>4</sub> production rates, while brackish water amended incubations of troughs and low-centered polygons had higher CH<sub>4</sub> production rates (Fig. 4). This indicates that for the degradation of organic matter into CH<sub>4</sub> in tundra soils, increasing seawater interactions through coastal processes, such as submersion due to subsidence or increased storm severity, resulting in the input of seawater in terrestrial soils, does not halt CH<sub>4</sub> production. It also shows that landforms and local hydrology remain important in controlling the microbial communities which affects the resulting CH<sub>4</sub> production. Differences among landforms and sites are generally large and clearly exceed the range of variability as shown by the uncertainty, supporting the use of means with standard deviations to convey contrasts without formal statistical tests. This approach allows us to highlight pronounced differences in methane production potential and geochemical context across coastal and inland sites.</p>
      <p id="d2e3117">Marine OM and nutrient inputs from tides and storm surges may contribute to the higher lability of OM and could fuel greater fermentation (Valdemarsen and Kristensen, 2010). It was reported that 8.7 % of the organic carbon in nearshore sediments of Herschel Island, Beaufort Sea, came from marine sources (Couture et al., 2018). This is relevant for the TP site because while <inline-formula><mml:math id="M268" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C signature of soils showed that terrestrial OM is dominant (Fig. S6), marine OM may get transported and deposited in coastal soils during high tides and storm surges. Although our analyses could not detect the presence of marine OM in TP soils, the higher CH<sub>4</sub> production rates recorded in the incubations of TP, relatively to those of RP could in part be explained by marine OM and nutrient inputs. Interestingly, the high-centered polygon at TP, profile 09 (Fig. 4), did not behave in a predictable manner, since it had very high CH<sub>4</sub> production rates on the surface. This elevated methane production rate coincided with the presence of substantial goose fecal deposits at TP, profile 09. While this observation suggests a potential local input of labile organic matter and nutrients (e.g., N and P) and possibly a distinct surface microbial community, no direct measurements were conducted to establish a mechanistic link. This site-specific observation is therefore reported as contextual field information rather than evidence of causation. Lower in the profile, CH<sub>4</sub> production rates were very low, characteristic of the CH<sub>4</sub> production rates observed in water-unsaturated high-centered polygons (Fig.  4). Therefore, in this instance, proximity with the coast may have influenced CH<sub>4</sub> production through the presence of fauna.</p>
      <p id="d2e3178">Stable carbon isotopic signature of CH<sub>4</sub> provides insights on the microbial processes involved in methanogenesis and on substrates used. <inline-formula><mml:math id="M275" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C–CH<sub>4</sub> between <inline-formula><mml:math id="M277" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>65 ‰ and <inline-formula><mml:math id="M278" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>50 ‰ is typically associated with acetoclastic methanogenesis, while <inline-formula><mml:math id="M279" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C–CH<sub>4</sub> between <inline-formula><mml:math id="M281" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>110 ‰ and <inline-formula><mml:math id="M282" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>60 ‰ is associated with hydrogenotrophic methanogenesis (Hornibrook et al., 1997, 2000). The stable isotopic signature of methylotrophic methanogenesis is between <inline-formula><mml:math id="M283" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>83 ‰ and <inline-formula><mml:math id="M284" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>72 ‰ (Penger et al., 2012), which overlaps with the hydrogenotrophic interval, precluding us from separating these two metabolic pathways. At RP, except for profile 10B, <inline-formula><mml:math id="M285" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C–CH<sub>4</sub> had more negative values, consistent with the processing of recalcitrant organic matter through the hydrogenotrophic production pathway (Heffernan et al., 2022; Hodgkins et al., 2014). Profile 10B, a polygonal trough, had less negative <sup>13</sup>C–CH<sub>4</sub> values more consistent with acetoclastic methanogenesis (Hornibrook et al., 1997). At TP, the coastal polygonal tundra, <inline-formula><mml:math id="M289" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C–CH<sub>4</sub> at 5 cm depth is less negative, consistent with methanogenesis with more labile organic carbon and the acetoclastic production pathway (Hodgkins et al., 2014), transitioning to more negative values, associated to hydrogenotrophic production with depth. This shift suggests an input of labile OM in TP surface and sub-surface soils. This may be due to the labile OM from abundant geese fecal matter that was observed in the surface. It is also possible that <italic>Carex</italic> sp., the dominant plant species of the site, may be a source of labile fermentation precursors (Galand et al., 2010; Liebner et al., 2015). To evaluate whether <inline-formula><mml:math id="M291" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C–CH<sub>4</sub> covaried with other geochemical properties measured in this study, <inline-formula><mml:math id="M293" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C–CH<sub>4</sub> values were examined alongside TOC content and sulfate concentrations; however, no consistent relationships were observed across landforms or depths (Fig. S4), indicating that methanogenic pathway signatures are not straightforwardly predicted by bulk TOC or sulfate availability at the scale investigated. However, it is clear that future work should integrate measurements of organic matter degradation, microbial community composition, and pore water chemistry to better resolve the mechanisms driving spatial variability in methane production.</p>
</sec>
<sec id="Ch1.S4.SS3">
  <label>4.3</label><title>Total CH<sub>4</sub> production rates are comparable to the net CH<sub>4</sub> fluxes measured in similar environments</title>
      <p id="d2e3403">In a polygonal terrain of the Tuktoyaktuk Coastlands, net CH<sub>4</sub> fluxes from the center of high-centered polygons and troughs derived from flux chambers were measured to be 1.9 <inline-formula><mml:math id="M298" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 20.4 and 13.0 <inline-formula><mml:math id="M299" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 20.4 mmol m<sup>−2</sup> d<sup>−1</sup> respectively (Martin et al., 2018). These overlap with values of estimated total CH<sub>4</sub> production derived from the brackish water amended incubation experiments (Fig. 5). It is clear from the large variations in measured CH<sub>4</sub> emissions from the study of Martin et al. (2018) that incubations to estimate total active layer CH<sub>4</sub> production rates can discern small differences due to local variations that stem mostly from the polygonal features. For example, at RP a comparable polygonal terrain located in the same study area of Martin et al. (2018), the total CH<sub>4</sub> production of high-centered polygons and trough were 0.3 <inline-formula><mml:math id="M306" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.1 and 5.0 <inline-formula><mml:math id="M307" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 2.0 mmol m<sup>−2</sup> d<sup>−1</sup> (Fig. 5), respectively which are significantly different. This indicates the role of polygonal forms in controlling the activity of microbial communities which controls CH<sub>4</sub> production and the potential to scale more accurately CH<sub>4</sub> production at the landscape level based on landform distributions.</p>
      <p id="d2e3547">Interestingly, TP, the coastal site, had an estimated total CH<sub>4</sub> production rate comparable to emissions of a St. Lawrence estuary salt marsh which had a CH<sub>4</sub> flux of 24 <inline-formula><mml:math id="M314" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 14.4 mmol m<sup>−2</sup> d<sup>−1</sup> (Comer-Warner et al., 2022). The St. Lawrence estuary salt marshes are affected by freeze-thaw cycles associated with seasons comparable to the freeze-thaw cycles observed in the active layer of Tuktoyaktuk coastlands despite lacking some characteristics features of our site like the presence of permafrost and rapid coastal erosion rates. CH<sub>4</sub> emissions and production within areas of coastal influence thus appear of similar magnitude. By comparison, mangrove forests, which are a major global source of CH<sub>4</sub> but a very different environment from coastal Arctic polygon terrain, had average CH<sub>4</sub> fluxes to the atmosphere of 0.3 <inline-formula><mml:math id="M320" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.1 mmol m<sup>−2</sup> d<sup>−1</sup> (Rosentreter et al., 2018). In another study, the average measured CH<sub>4</sub> flux from a Yangtze Estuary (China) tidal salt marsh, with a subtropical monsoon climate, was 2.4 mmol m<sup>−2</sup> d<sup>−1</sup> (Li et al., 2021). These reported values are similar to our study as well as other studies in the region. When considered alongside the global distribution of coastal wetlands, this similarity in flux magnitude becomes particularly relevant. Tropical coastal wetlands are dominated by mangroves (<inline-formula><mml:math id="M326" display="inline"><mml:mo lspace="0mm">∼</mml:mo></mml:math></inline-formula> 147 000 km<sup>2</sup>), whereas Arctic wetlands cover approximately 3.5<inline-formula><mml:math id="M328" display="inline"><mml:mrow><mml:mo>×</mml:mo><mml:msup><mml:mn mathvariant="normal">10</mml:mn><mml:mn mathvariant="normal">6</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> km<sup>2</sup> (Worthington et al., 2024). Even if only a small fraction of Arctic wetlands is located within coastal zones, their total extent is comparable to the global mangrove area (Worthington et al., 2024), suggesting that permafrost Arctic coastal wetlands could represent a non-negligible component of the global CH<sub>4</sub> budget and warrant further investigation.</p>

<table-wrap id="T1" specific-use="star"><label>Table 1</label><caption><p id="d2e3743">Total methane production in a context of brackish water addition in high-centered polygons, low-centered polygons and throughs during growing season applied to the spatial scale of the polygonal landscape of RP. Two samples were taken for the high-centered polygon. The mean active layer and talik depth of the region is 35 cm. The error represents the propagation of the analytical uncertainty from the incubations results.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="3">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="right"/>
     <oasis:colspec colnum="3" colname="col3" align="right"/>
     <oasis:thead>
       <oasis:row>
         <oasis:entry colname="col1">Geomorphological form</oasis:entry>
         <oasis:entry colname="col2">Relative area of each</oasis:entry>
         <oasis:entry colname="col3">Estimated Total CH<sub>4</sub> production</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2">landform (km<sup>2</sup>)</oasis:entry>
         <oasis:entry colname="col3">(mol d<sup>−1</sup>)</oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>
         <oasis:entry colname="col1">High-centered polygons</oasis:entry>
         <oasis:entry colname="col2">0.0803</oasis:entry>
         <oasis:entry colname="col3">20.7 <inline-formula><mml:math id="M334" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 10.3</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Low-centered polygons</oasis:entry>
         <oasis:entry colname="col2">0.119</oasis:entry>
         <oasis:entry colname="col3">284 <inline-formula><mml:math id="M335" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 123</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Troughs</oasis:entry>
         <oasis:entry colname="col2">0.0362</oasis:entry>
         <oasis:entry colname="col3">182 <inline-formula><mml:math id="M336" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 73.4</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>

      <p id="d2e3874">The calculated total methane production rates (<inline-formula><mml:math id="M337" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>) from TP and RP do not take into account aerobic and anaerobic oxidation of CH<sub>4</sub>, which will most likely reduce fluxes of CH<sub>4</sub> from these sites. Studies and models of Arctic soils emissions have highlighted that aerobic methanotrophy could consume more than half of the CH<sub>4</sub> produced in soils, greatly limiting surface emissions (Oh et al., 2020; Zheng et al., 2018). Furthermore, AOM has been shown to play an important role in attenuating CH<sub>4</sub> production in soils and sediments (Segarra et al., 2013; Winkel et al., 2019) but did not appear to influence significantly CH<sub>4</sub> production in incubations with thermokarst lake sediments (Lotem et al., 2023). While AOM represents a major sink for CH<sub>4</sub> in marine sediments (Knittel and Boetius, 2009; Reeburgh, 2009), the very different biogeochemical and hydrological characteristics of our coastal sites suggest that the role of AOM in these environments may diverge from that observed in fully marine systems. Recent work in coastal thermokarst lagoons, which can present key similarities to our coastal study sites due to episodic or persistent brackish water intrusion, have been shown to exhibit strong AOM control on CH<sub>4</sub> dynamics, particularly in sulfate-rich settings where AOM may constitute a major CH<sub>4</sub> sink (Yang et al., 2023). For the discussion of this study, we compared results of brackish water incubations to CH<sub>4</sub> emissions measured in other landscapes. Such comparisons provide valuable context by comparing long-term microbial production processes with net atmospheric fluxes. However, we emphasize that CH<sub>4</sub> production rates cannot be directly equated to CH<sub>4</sub> emissions.</p>
      <p id="d2e3985">To better frame the potential impact of brackish water addition at scale, we extrapolated its consequence to a 25 ha area of polygonal tundra surrounding RP (Fig. S2). This estimate simulates CH<sub>4</sub> production following the infiltration of brackish water into the terrestrial polygonal landscape around Tuktoyaktuk. This event could result from coastal flooding during storm surges, which are frequent in the Mackenzie River delta (Kokelj et al., 2012; Solomon, 2005). Taking into consideration the distribution of the polygonal features within RP and the relative areas of each landform, the CH<sub>4</sub> production rates in the active layer, excluding pondlets, for an area of 25 ha was calculated to be 487 mol d<sup>−1</sup> (Table 1) or 22 nmol m<sup>−2</sup> s<sup>−1</sup> and is consistent with the CH<sub>4</sub> emissions measured from various wetland types (Cui et al., 2024). The increasing sensitivity of wetlands to climate change and the preponderant role of carbon substrate availability in controlling global methane emissions (Hu et al., 2024) warrants further investigating CH<sub>4</sub> dynamics in thawing continuous permafrost landscapes and the role of coastal processes influencing these emissions. More polygonal tundra in various settings should be investigated as a comparison to the studied region. Further research on aerobic and anaerobic CH<sub>4</sub> oxidation is necessary to provide a more precise estimate of the CH<sub>4</sub> cycle inputs and outputs in a scope of the evaluation of its impacts on the greenhouse gas feedback loop.</p>
</sec>
</sec>
<sec id="Ch1.S5" sec-type="conclusions">
  <label>5</label><title>Conclusions</title>
      <p id="d2e4089">The primary hypothesis for this study was that an increase in waterlogged environments due to coastal flooding and inundation processes would not enhance CH<sub>4</sub> production because of sulfate present in coastal waters. However, our incubation experiments revealed high CH<sub>4</sub> production rates in the presence of sulfates. Additionally, waterlogged conditions attributed to the ebb and flow of tides, seems to favor anoxic OM degradation and may potentially provide inputs of fresh OM and nutrients from marine sources, contributing to the elevated CH<sub>4</sub> production rates measured in the coastal setting of TP. Moreover, no conclusive explanation for the co-occurrence of sulfate-reduction and methanogenesis in our brackish water incubations was identified, but based on evidence, we suggest syntrophic methanogenesis could support this co-occurrence. More investigation on methylotrophic methanogenesis in coastal soils are needed as it can be an important process in saline environments (Conrad, 2020). Future studies should investigate CH<sub>4</sub> oxidation processes in greater detail, as they could provide crucial insights into Arctic coastal carbon cycling in sediments and soils affected by changing sea level.</p>
</sec>

      
      </body>
    <back><notes notes-type="dataavailability"><title>Data availability</title>

      <p id="d2e4132">All raw data of incubation experiment and other analyses performed and generated by study are available in the Supplement.</p>
  </notes><app-group>
        <supplementary-material position="anchor"><p id="d2e4135">The supplement related to this article is available online at <inline-supplementary-material xlink:href="https://doi.org/10.5194/bg-23-3777-2026-supplement" xlink:title="zip">https://doi.org/10.5194/bg-23-3777-2026-supplement</inline-supplementary-material>.</p></supplementary-material>
        </app-group><notes notes-type="authorcontribution"><title>Author contributions</title>

      <p id="d2e4144">AP designed the experiment; ARL executed the experiments and analyses. AP, ARL, DW, RL participated in the fieldwork. PMJD provided lab space, equipment and insights for the stable carbon isotopes analyses on incubation CH<sub>4</sub>. RL performed all GIS analyses and maps. ARL performed the data interpretation and generated all figures. AP provided expertise on the writing and interpretation of figures. All authors reviewed and edited the manuscript.</p>
  </notes><notes notes-type="competinginterests"><title>Competing interests</title>

      <p id="d2e4159">The contact author has declared that none of the authors has any competing interests.</p>
  </notes><notes notes-type="disclaimer"><title>Disclaimer</title>

      <p id="d2e4165">Publisher's note: Copernicus Publications remains neutral with regard to jurisdictional claims made in the text, published maps, institutional affiliations, or any other geographical representation in this paper. The authors bear the ultimate responsibility for providing appropriate place names. Views expressed in the text are those of the authors and do not necessarily reflect the views of the publisher.</p>
  </notes><ack><title>Acknowledgements</title><p id="d2e4171">We thank Santiago Mareque for assistance during field sampling. Mathieu Babin and Thi Hao Bui are acknowledged for assisting with the laboratory work performed at Université du Québec à Rimouski and at McGill University, respectively. Takuvik Laboratory is acknowledged for providing analyses and results on <inline-formula><mml:math id="M363" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">δ</mml:mi><mml:mn mathvariant="normal">13</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>C and TOC content of sediments. We also thank the community of Tuktoyaktuk for providing wildlife monitors with insightful information on the territory during field sampling. This research was funded by NSERC Discovery Grant and Northern Supplement to AP. ARL acknowledges financial support from the NSERC Northern Scientific Training Program. PMJD acknowledges support from the NSERC Discovery Grant and the Canadian Foundation for Innovation.</p></ack><notes notes-type="financialsupport"><title>Financial support</title>

      <p id="d2e4187">This research has been supported by the Natural Sciences and Engineering Research Council of Canada (Discovery Grant) and the Natural Resources Canada (Geological Survey of Canada – Nuna Project grant).</p>
  </notes><notes notes-type="reviewstatement"><title>Review statement</title>

      <p id="d2e4193">This paper was edited by Susanne Liebner and reviewed by Maren Jenrich and one anonymous referee.</p>
  </notes><ref-list>
    <title>References</title>

      <ref id="bib1.bib1"><label>1</label><mixed-citation> AMAP: Snow, Water, Ice and Permafrost in the Arctic (SWIPA) 2017, Arctic Monitoring and Assessment Programme (AMAP), Oslo, Norway, 269 pp., ISBN 978-82-7971-101-8, 2017.</mixed-citation></ref>
      <ref id="bib1.bib2"><label>2</label><mixed-citation>Andrachuk, M. and Smit, B.: Community-based vulnerability assessment of Tuktoyaktuk, NWT, Canada to environmental and socio-economic changes, Reg. Environ. Change, 12, 867–885, <ext-link xlink:href="https://doi.org/10.1007/s10113-012-0299-0" ext-link-type="DOI">10.1007/s10113-012-0299-0</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bib3"><label>3</label><mixed-citation>Bianchi, T. S.: The role of terrestrially derived organic carbon in the coastal ocean: A changing paradigm and the priming effect, Proc. Natl. Acad. Sci. USA, 108, 19473–19481, <ext-link xlink:href="https://doi.org/10.1073/pnas.1017982108" ext-link-type="DOI">10.1073/pnas.1017982108</ext-link>, 2011.</mixed-citation></ref>
      <ref id="bib1.bib4"><label>4</label><mixed-citation>Boetius, A., Ravenschlag, K., Schubert, C. J., Rickert, D., Widdel, F., Gieseke, A., Amann, R., Jørgensen, B. B., Witte, U., and Pfannkuche, O.: A marine microbial consortium apparently mediating anaerobic oxidation of methane, Nature, 407, 623–626, <ext-link xlink:href="https://doi.org/10.1038/35036572" ext-link-type="DOI">10.1038/35036572</ext-link>, 2000.</mixed-citation></ref>
      <ref id="bib1.bib5"><label>5</label><mixed-citation>Bridgham, S. D., Cadillo-Quiroz, H., Keller, J. K., and Zhuang, Q.: Methane emissions from wetlands: biogeochemical, microbial, and modeling perspectives from local to global scales, Glob. Chang. Biol., 19, 1325–1346, <ext-link xlink:href="https://doi.org/10.1111/gcb.12131" ext-link-type="DOI">10.1111/gcb.12131</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bib6"><label>6</label><mixed-citation>Comer-Warner, S. A., Ullah, S., Ampuero Reyes, W., Krause, S., and Chmura, G. L.: Spartina alterniflora has the highest methane emissions in a St. Lawrence estuary salt marsh, Environ. Res. Ecol., 1, 011003, <ext-link xlink:href="https://doi.org/10.1088/2752-664X/ac706a" ext-link-type="DOI">10.1088/2752-664X/ac706a</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bib7"><label>7</label><mixed-citation>Conrad, R.: Importance of hydrogenotrophic, aceticlastic and methylotrophic methanogenesis for methane production in terrestrial, aquatic and other anoxic environments: A mini review, Pedosphere, 30, 25–39, <ext-link xlink:href="https://doi.org/10.1016/S1002-0160(18)60052-9" ext-link-type="DOI">10.1016/S1002-0160(18)60052-9</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bib8"><label>8</label><mixed-citation>Coolen, M. J. L. and Orsi, W. D.: The transcriptional response of microbial communities in thawing Alaskan permafrost soils, Front. Microbiol., 6, 197, <ext-link xlink:href="https://doi.org/10.3389/fmicb.2015.00197" ext-link-type="DOI">10.3389/fmicb.2015.00197</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bib9"><label>9</label><mixed-citation>Costa, B., Vieira, G., and Whalen, D.: The fast-changing coast of Tuktoyaktuk Peninsula (Beaufort Sea, Canada): geomorphological controls on changes between 1985 and 2020, EGU General Assembly 2022, Vienna, Austria, 23–27 May 2022, EGU22-2426, <ext-link xlink:href="https://doi.org/10.5194/egusphere-egu22-2426" ext-link-type="DOI">10.5194/egusphere-egu22-2426</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bib10"><label>10</label><mixed-citation>Couture, N. J., Irrgang, A., Pollard, W., Lantuit, H., and Fritz, M.: Coastal erosion of permafrost soils along the Yukon Coastal Plain and fluxes of organic carbon to the Canadian Beaufort Sea, J. Geophys. Res. Biogeosci., 123, 406–422, <ext-link xlink:href="https://doi.org/10.1002/2017JG004166" ext-link-type="DOI">10.1002/2017JG004166</ext-link>, 2018.</mixed-citation></ref>
      <ref id="bib1.bib11"><label>11</label><mixed-citation>Cui, S., Liu, P., Guo, H., Nielsen, C. K., Pullens, J. W. M., Chen, Q., Pugliese, L., and Wu, S.: Wetland hydrological dynamics and methane emissions, Commun. Earth Environ., 5, 1635, <ext-link xlink:href="https://doi.org/10.1038/s43247-024-01635-w" ext-link-type="DOI">10.1038/s43247-024-01635-w</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bib12"><label>12</label><mixed-citation>Dallimore, S. R., Wolfe, S. A., Matthews Jr., J. V., and Vincent, J.-S.: Mid-Wisconsinan eolian deposits of the Kittigazuit Formation, Tuktoyaktuk Coastlands, Northwest Territories, Canada, Can. J. Earth Sci., 34, 1421–1441, <ext-link xlink:href="https://doi.org/10.1139/e17-116" ext-link-type="DOI">10.1139/e17-116</ext-link>, 1997.</mixed-citation></ref>
      <ref id="bib1.bib13"><label>13</label><mixed-citation>Elberling, B., Michelsen, A., Schädel, C., Schuur, E. A. G., Christiansen, H. H., Berg, L., Tamstorf, M. P., and Sigsgaard, C.: Long-term CO<sub>2</sub> production following permafrost thaw, Nat. Clim. Chang., 3, 890–894, <ext-link xlink:href="https://doi.org/10.1038/nclimate1955" ext-link-type="DOI">10.1038/nclimate1955</ext-link>, 2013.</mixed-citation></ref>
      <ref id="bib1.bib14"><label>14</label><mixed-citation>Froelich, P., Klinkhammer, G., Bender, M., Luedtke, N., Heath, G., Cullen, D., Dauphin, P., Hammond, D., Hartman, B., and Maynard, V.: Early oxidation of organic matter in pelagic sediments of the eastern equatorial Atlantic: suboxic diagenesis, Geochim. Cosmochim. Acta, 43, 1075–1090, <ext-link xlink:href="https://doi.org/10.1016/0016-7037(79)90095-4" ext-link-type="DOI">10.1016/0016-7037(79)90095-4</ext-link>, 1979.</mixed-citation></ref>
      <ref id="bib1.bib15"><label>15</label><mixed-citation>Galand, P. E., Yrjälä, K., and Conrad, R.: Stable carbon isotope fractionation during methanogenesis in three boreal peatland ecosystems, Biogeosciences, 7, 3893–3900, <ext-link xlink:href="https://doi.org/10.5194/bg-7-3893-2010" ext-link-type="DOI">10.5194/bg-7-3893-2010</ext-link>, 2010.</mixed-citation></ref>
      <ref id="bib1.bib16"><label>16</label><mixed-citation>Guimond, J. A., Mohammed, A. A., Walvoord, M. A., Bense, V. F., and Kurylyk, B. L.: Saltwater intrusion intensifies coastal permafrost thaw, Geophys. Res. Lett., 48, e2021GL094776, <ext-link xlink:href="https://doi.org/10.1029/2021GL094776" ext-link-type="DOI">10.1029/2021GL094776</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bib17"><label>17</label><mixed-citation>Heffernan, L., Cavaco, M. A., Bhatia, M. P., Estop-Aragonés, C., Knorr, K.-H., and Olefeldt, D.: High peatland methane emissions following permafrost thaw: enhanced acetoclastic methanogenesis during early successional stages, Biogeosciences, 19, 3051–3071, <ext-link xlink:href="https://doi.org/10.5194/bg-19-3051-2022" ext-link-type="DOI">10.5194/bg-19-3051-2022</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bib18"><label>18</label><mixed-citation>Heslop, J. K., Walter Anthony, K. M., Sepulveda-Jauregui, A., Martinez-Cruz, K., Bondurant, A., Grosse, G., and Jones, M. C.: Thermokarst lake methanogenesis along a complete talik profile, Biogeosciences, 12, 4317–4331, <ext-link xlink:href="https://doi.org/10.5194/bg-12-4317-2015" ext-link-type="DOI">10.5194/bg-12-4317-2015</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bib19"><label>19</label><mixed-citation>Heslop, J. K., Walter Anthony, K. M., Winkel, M., Sepulveda-Jauregui, A., Martinez-Cruz, K., Bondurant, A., Grosse, G., and Liebner, S.: A synthesis of methane dynamics in thermokarst lake environments, Earth Sci. Rev., 210, 103365, <ext-link xlink:href="https://doi.org/10.1016/j.earscirev.2020.103365" ext-link-type="DOI">10.1016/j.earscirev.2020.103365</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bib20"><label>20</label><mixed-citation>Heyer, J., Berger, U., Kuzin, I. L., and Yakovlev, O. N.: Methane emissions from different ecosystem structures of the subarctic tundra in Western Siberia during midsummer and during the thawing period, Tellus B, 54, 231–249, <ext-link xlink:href="https://doi.org/10.1034/j.1600-0889.2002.01280.x" ext-link-type="DOI">10.1034/j.1600-0889.2002.01280.x</ext-link>, 2002.</mixed-citation></ref>
      <ref id="bib1.bib21"><label>21</label><mixed-citation>Hill, P. R., Héquette, A., and Ruz, M.-H.: Holocene sea-level history of the Canadian Beaufort shelf, Can. J. Earth Sci., 30, 103–108, <ext-link xlink:href="https://doi.org/10.1139/e93-009" ext-link-type="DOI">10.1139/e93-009</ext-link>, 1993.</mixed-citation></ref>
      <ref id="bib1.bib22"><label>22</label><mixed-citation>Hodgkins, S. B., Tfaily, M. M., McCalley, C. K., Logan, T. A., Crill, P. M., Saleska, S. R., Rich, V. I., and Chanton, J. P.: Changes in peat chemistry associated with permafrost thaw increase greenhouse gas production, Proc. Natl. Acad. Sci. USA, 111, 5819–5824, <ext-link xlink:href="https://doi.org/10.1073/pnas.1314641111" ext-link-type="DOI">10.1073/pnas.1314641111</ext-link>, 2014.</mixed-citation></ref>
      <ref id="bib1.bib23"><label>23</label><mixed-citation>Holm, S., Walz, J., Horn, F., Yang, S., Grigoriev, M. N., Wagner, D., Knoblauch, C., and Liebner, S.: Methanogenic response to long-term permafrost thaw is determined by paleoenvironment, FEMS Microbiol. Ecol., 96, fiaa021, <ext-link xlink:href="https://doi.org/10.1093/femsec/fiaa021" ext-link-type="DOI">10.1093/femsec/fiaa021</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bib24"><label>24</label><mixed-citation>Hornibrook, E. R., Longstaffe, F. J., and Fyfe, W. S.: Spatial distribution of microbial methane production pathways in temperate zone wetland soils: stable carbon and hydrogen isotope evidence, Geochim. Cosmochim. Acta, 61, 745–753, <ext-link xlink:href="https://doi.org/10.1016/S0016-7037(96)00368-7" ext-link-type="DOI">10.1016/S0016-7037(96)00368-7</ext-link>, 1997.</mixed-citation></ref>
      <ref id="bib1.bib25"><label>25</label><mixed-citation>Hornibrook, E. R. C., Longstaffe, F. J., and Fyfe, W. S.: Evolution of stable carbon isotope compositions for methane and carbon dioxide in freshwater wetlands and other anaerobic environments, Geochim. Cosmochim. Acta, 64, 1013–1027, <ext-link xlink:href="https://doi.org/10.1016/S0016-7037(99)00321-X" ext-link-type="DOI">10.1016/S0016-7037(99)00321-X</ext-link>, 2000.</mixed-citation></ref>
      <ref id="bib1.bib26"><label>26</label><mixed-citation>Hu, H., Chen, J., Zhou, F., Nie, M., Hou, D., Liu, H., Delgado-Baquerizo, M., Ni, H., Huang, W., Zhou, J., Song, X., Cao, X., Sun, B., Zhang, J., Crowther, T. W., and Liang, Y.: Relative increases in CH<sub>4</sub> and CO2 emissions from wetlands under global warming dependent on soil carbon substrates, Nat. Geosci., 17, 26–31, <ext-link xlink:href="https://doi.org/10.1038/s41561-023-01345-6" ext-link-type="DOI">10.1038/s41561-023-01345-6</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bib27"><label>27</label><mixed-citation>Hu, K., Issler, D., Chen, Z., and Brent, T.: Permafrost investigation by well logs, and seismic velocity and repeated shallow temperature surveys, Beaufort-Mackenzie Basin, Geological Survey of Canada, <ext-link xlink:href="https://doi.org/10.4095/293120" ext-link-type="DOI">10.4095/293120</ext-link>, 2013.</mixed-citation></ref>
      <ref id="bib1.bib28"><label>28</label><mixed-citation>Hynes, S., Solomon, S. M., and Whalen, D.: GIS compilation of coastline variability spanning 60 years in the Mackenzie Delta and Tuktoyaktuk in the Beaufort Sea, Geological Survey of Canada Open File 7685, <ext-link xlink:href="https://doi.org/10.4095/295579" ext-link-type="DOI">10.4095/295579</ext-link>, 2014.</mixed-citation></ref>
      <ref id="bib1.bib29"><label>29</label><mixed-citation>Ionescu, D., Bizic-Ionescu, M., Khalili, A., Malekmohammadi, R., Morad, M. R., de Beer, D., and Grossart, H.-P.: A new tool for long-term studies of POM-bacteria interactions: overcoming the century-old Bottle Effect, Sci. Rep., 5, 14706, <ext-link xlink:href="https://doi.org/10.1038/srep14706" ext-link-type="DOI">10.1038/srep14706</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bib30"><label>30</label><mixed-citation>Irrgang, A. M., Bendixen, M., Farquharson, L. M., Baranskaya, A. V., Erikson, L. H., Gibbs, A. E., Ogorodov, S. A., Overduin, P. P., Lantuit, H., Grigoriev, M. N., and Jones, B. M.: Drivers, dynamics and impacts of changing Arctic coasts, Nat. Rev. Earth Environ., 3, 39–54, <ext-link xlink:href="https://doi.org/10.1038/s43017-021-00232-1" ext-link-type="DOI">10.1038/s43017-021-00232-1</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bib31"><label>31</label><mixed-citation>Jenrich, M., Angelopoulos, M., Liebner, S., Treat, C. C., Knoblauch, C., Yang, S., Grosse, G., Giebeler, F., Jongejans, L. L., Grigoriev, M., and Strauss, J.: Greenhouse gas production and microbial response during the transition from terrestrial permafrost to a marine environment, Permafr. Periglac. Process., <ext-link xlink:href="https://doi.org/10.1002/ppp.2251" ext-link-type="DOI">10.1002/ppp.2251</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bib32"><label>32</label><mixed-citation>Jenrich, M., Wolter, J., Liebner, S., Knoblauch, C., Grosse, G., Giebeler, F., Whalen, D., and Strauss, J.: Rising Arctic seas and thawing permafrost: uncovering the carbon cycle impact in a thermokarst lagoon system in the outer Mackenzie Delta, Canada, Biogeosciences, 22, 2069–2086, <ext-link xlink:href="https://doi.org/10.5194/bg-22-2069-2025" ext-link-type="DOI">10.5194/bg-22-2069-2025</ext-link>, 2025.</mixed-citation></ref>
      <ref id="bib1.bib33"><label>33</label><mixed-citation>Jones, E. L., Hodson, A. J., Thornton, S. F., Redeker, K. R., Rogers, J., Wynn, P. M., Dixon, T. J., Bottrell, S. H., and O'Neill, H. B.: Biogeochemical processes in the active layer and permafrost of a high Arctic fjord valley, Front. Earth Sci., 8, 342, <ext-link xlink:href="https://doi.org/10.3389/feart.2020.00342" ext-link-type="DOI">10.3389/feart.2020.00342</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bib34"><label>34</label><mixed-citation>Keuschnig, C., Larose, C., Rudner, M., Pesqueda, A., Doleac, S., Elberling, B., Björk, R. G., Klemedtsson, L., and Björkman, M. P.: Reduced methane emissions in former permafrost soils driven by vegetation and microbial changes following drainage, Glob. Chang. Biol., 28, 3411–3425, <ext-link xlink:href="https://doi.org/10.1111/gcb.16137" ext-link-type="DOI">10.1111/gcb.16137</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bib35"><label>35</label><mixed-citation>Knittel, K. and Boetius, A.: Anaerobic oxidation of methane: progress with an unknown process, Annu. Rev. Microbiol., 63, 311–334, <ext-link xlink:href="https://doi.org/10.1146/annurev.micro.61.080706.093130" ext-link-type="DOI">10.1146/annurev.micro.61.080706.093130</ext-link>, 2009.</mixed-citation></ref>
      <ref id="bib1.bib36"><label>36</label><mixed-citation>Knoblauch, C., Beer, C., Sosnin, A., Wagner, D., and Pfeiffer, E.-M.: Predicting long-term carbon mineralization and trace gas production from thawing permafrost of Northeast Siberia, Glob. Chang. Biol., 19, 1160–1172, <ext-link xlink:href="https://doi.org/10.1111/gcb.12116" ext-link-type="DOI">10.1111/gcb.12116</ext-link>, 2013.</mixed-citation></ref>
      <ref id="bib1.bib37"><label>37</label><mixed-citation>Knoblauch, C., Beer, C., Liebner, S., Grigoriev, M. N., and Pfeiffer, E.-M.: Methane production as key to the greenhouse gas budget of thawing permafrost, Nat. Clim. Chang., 8, 309–312, <ext-link xlink:href="https://doi.org/10.1038/s41558-018-0095-z" ext-link-type="DOI">10.1038/s41558-018-0095-z</ext-link>, 2018.</mixed-citation></ref>
      <ref id="bib1.bib38"><label>38</label><mixed-citation>Kokelj, S. V., Lantz, T. C., Solomon, S., Pisaric, M. F., Keith, D., Morse, P., Thienpont, J. R., Smol, J. P., and Esagok, D.: Using multiple sources of knowledge to investigate northern environmental change: regional ecological impacts of a storm surge in the Outer Mackenzie Delta, N.W.T., Arctic, 65, 3, <ext-link xlink:href="https://doi.org/10.14430/arctic4214" ext-link-type="DOI">10.14430/arctic4214</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bib39"><label>39</label><mixed-citation>Kramshøj, M., Albers, C. N., Holst, T., Holzinger, R., Elberling, B., and Rinnan, R.: Biogenic volatile release from permafrost thaw is determined by the soil microbial sink, Nat. Commun., 9, 3412, <ext-link xlink:href="https://doi.org/10.1038/s41467-018-05824-y" ext-link-type="DOI">10.1038/s41467-018-05824-y</ext-link>, 2018.</mixed-citation></ref>
      <ref id="bib1.bib40"><label>40</label><mixed-citation>Kroeger, K. D., Crooks, S., Moseman-Valtierra, S., and Tang, J.: Restoring tides to reduce methane emissions in impounded wetlands: a new and potent blue carbon climate change intervention, Sci. Rep., 7, 12138, <ext-link xlink:href="https://doi.org/10.1038/s41598-017-12138-4" ext-link-type="DOI">10.1038/s41598-017-12138-4</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bib41"><label>41</label><mixed-citation>La, W., Han, X., Liu, C.-Q., Ding, H., Liu, M., Sun, F., Li, S., and Lang, Y.: Sulfate concentrations affect sulfate reduction pathways and methane consumption in coastal wetlands, Water Res., 217, 118441, <ext-link xlink:href="https://doi.org/10.1016/j.watres.2022.118441" ext-link-type="DOI">10.1016/j.watres.2022.118441</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bib42"><label>42</label><mixed-citation>Lacelle, D., Fontaine, M., Pellerin, A., Kokelj, S. V., and Clark, I. D.: Legacy of Holocene landscape changes on soil biogeochemistry: a perspective from paleo-active layers in northwestern Canada, J. Geophys. Res. Biogeosciences, 124, 2662–2679, <ext-link xlink:href="https://doi.org/10.1029/2018JG004916" ext-link-type="DOI">10.1029/2018JG004916</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bib43"><label>43</label><mixed-citation>Lantuit, H., Overduin, P. P., Couture, N., Wetterich, S., Aré, F., Atkinson, D., Brown, J., Cherkashov, G., Drozdov, D., Forbes, D. L., Graves-Gaylord, A., Grigoriev, M., Hubberten, H.-W., Jordan, J., Jorgenson, T., Ødegård, R. S., Ogorodov, S., Pollard, W. H., Rachold, V., and Vasiliev, A.: The Arctic coastal dynamics database: a new classification scheme and statistics on Arctic permafrost coastlines, Estuar. Coasts, 35, 383–400, <ext-link xlink:href="https://doi.org/10.1007/s12237-010-9362-6" ext-link-type="DOI">10.1007/s12237-010-9362-6</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bib44"><label>44</label><mixed-citation>Lapham, L. L., Dallimore, S. R., Magen, C., Henderson, L. C., Powers, L. C., Gonsior, M., Clark, B., Côté, M., Fraser, P., and Orcutt, B. N.: Microbial greenhouse gas dynamics associated with warming coastal permafrost, western Canadian Arctic, Front. Earth Sci., 8, 582103, <ext-link xlink:href="https://doi.org/10.3389/feart.2020.582103" ext-link-type="DOI">10.3389/feart.2020.582103</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bib45"><label>45</label><mixed-citation>Li, Y., Wang, D., Chen, Z., Chen, J., Hu, H., and Wang, R.: Methane emissions during the tide cycle of a Yangtze Estuary salt marsh, Atmosphere, 12, 245, <ext-link xlink:href="https://doi.org/10.3390/atmos12020245" ext-link-type="DOI">10.3390/atmos12020245</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bib46"><label>46</label><mixed-citation>Liebner, S., Ganzert, L., Kiss, A., Yang, S., Wagner, D., and Svenning, M. M.: Shifts in methanogenic community composition and methane fluxes along the degradation of discontinuous permafrost, Front. Microbiol., 6, 356, <ext-link xlink:href="https://doi.org/10.3389/fmicb.2015.00356" ext-link-type="DOI">10.3389/fmicb.2015.00356</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bib47"><label>47</label><mixed-citation>Lim, M., Whalen, D., Martin, J., Mann, P. J., Hayes, S., Fraser, P., Berry, H. B., and Ouellette, D.: Massive ice control on permafrost coast erosion and sensitivity, Geophys. Res. Lett., 47, e2020GL087917, <ext-link xlink:href="https://doi.org/10.1029/2020GL087917" ext-link-type="DOI">10.1029/2020GL087917</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bib48"><label>48</label><mixed-citation>Lipson, D. A., Zona, D., Raab, T. K., Bozzolo, F., Mauritz, M., and Oechel, W. C.: Water-table height and microtopography control biogeochemical cycling in an Arctic coastal tundra ecosystem, Biogeosciences, 9, 577–591, <ext-link xlink:href="https://doi.org/10.5194/bg-9-577-2012" ext-link-type="DOI">10.5194/bg-9-577-2012</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bib49"><label>49</label><mixed-citation>Lotem, N., Pellerin, A., Anthony, K. W., Gafni, A., Boyko, V., and Sivan, O.: Anaerobic oxidation of methane does not attenuate methane emissions from thermokarst lakes, Limnol. Oceanogr., 68, 1316–1330, <ext-link xlink:href="https://doi.org/10.1002/lno.12349" ext-link-type="DOI">10.1002/lno.12349</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bib50"><label>50</label><mixed-citation>Lovley, D. R. and Klug, M. J.: Sulfate reducers can outcompete methanogens at freshwater sulfate concentrations, Appl. Environ. Microbiol., 45, 187–192, <ext-link xlink:href="https://doi.org/10.1128/aem.45.1.187-192.1983" ext-link-type="DOI">10.1128/aem.45.1.187-192.1983</ext-link>, 1983.</mixed-citation></ref>
      <ref id="bib1.bib51"><label>51</label><mixed-citation>Mackay, J. R. and Dallimore, S. R.: Massive ice of the Tuktoyaktuk area, western Arctic coast, Canada, Can. J. Earth Sci., 29, 1235–1249, <ext-link xlink:href="https://doi.org/10.1139/e92-099" ext-link-type="DOI">10.1139/e92-099</ext-link>, 1992.</mixed-citation></ref>
      <ref id="bib1.bib52"><label>52</label><mixed-citation>Maltby, J., Steinle, L., Löscher, C. R., Bange, H. W., Fischer, M. A., Schmidt, M., and Treude, T.: Microbial methanogenesis in the sulfate-reducing zone of sediments in the Eckernförde Bay, SW Baltic Sea, Biogeosciences, 15, 137–157, <ext-link xlink:href="https://doi.org/10.5194/bg-15-137-2018" ext-link-type="DOI">10.5194/bg-15-137-2018</ext-link>, 2018.</mixed-citation></ref>
      <ref id="bib1.bib53"><label>53</label><mixed-citation>Manson, G. K., Couture, N. J., and James, T. S.: CanCoast 2.0: data and indices to describe the sensitivity of Canada's marine coasts to changing climate, Geological Survey of Canada Open File 8551, <ext-link xlink:href="https://doi.org/10.4095/314669" ext-link-type="DOI">10.4095/314669</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bib54"><label>54</label><mixed-citation>Martens, C. S. and Berner, R. A.: Methane production in the interstitial waters of sulfate-depleted marine sediments, Science, 185, 1167–1169, <ext-link xlink:href="https://doi.org/10.1126/science.185.4157.1167" ext-link-type="DOI">10.1126/science.185.4157.1167</ext-link>, 1974.</mixed-citation></ref>
      <ref id="bib1.bib55"><label>55</label><mixed-citation>Martin, A. F., Lantz, T. C., and Humphreys, E. R.: Ice wedge degradation and CO2 and CH<sub>4</sub> emissions in the Tuktoyaktuk Coastlands, Northwest Territories, Arct. Sci., 4, 130–145, <ext-link xlink:href="https://doi.org/10.1139/as-2016-0011" ext-link-type="DOI">10.1139/as-2016-0011</ext-link>, 2018.</mixed-citation></ref>
      <ref id="bib1.bib56"><label>56</label><mixed-citation>Murton, J. B.: Thermokarst-lake-basin sediments, Tuktoyaktuk Coastlands, western Arctic Canada, Sedimentology, 43, 737–760, <ext-link xlink:href="https://doi.org/10.1111/j.1365-3091.1996.tb02023.x" ext-link-type="DOI">10.1111/j.1365-3091.1996.tb02023.x</ext-link>, 1996.</mixed-citation></ref>
      <ref id="bib1.bib57"><label>57</label><mixed-citation>Oh, Y., Zhuang, Q., Liu, L., Welp, L. R., Lau, M. C. Y., Onstott, T. C., Medvigy, D., Bruhwiler, L., Dlugokencky, E. J., Hugelius, G., D'Imperio, L., and Elberling, B.: Reduced net methane emissions due to microbial methane oxidation in a warmer Arctic, Nat. Clim. Chang., 10, 317–321, <ext-link xlink:href="https://doi.org/10.1038/s41558-020-0734-z" ext-link-type="DOI">10.1038/s41558-020-0734-z</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bib58"><label>58</label><mixed-citation>Oremland, R. S. and Polcin, S.: Methanogenesis and sulfate reduction: competitive and noncompetitive substrates in estuarine sediments, Appl. Environ. Microbiol., 44, 1270–1276, <ext-link xlink:href="https://doi.org/10.1128/aem.44.6.1270-1276.1982" ext-link-type="DOI">10.1128/aem.44.6.1270-1276.1982</ext-link>, 1982.</mixed-citation></ref>
      <ref id="bib1.bib59"><label>59</label><mixed-citation>Ozuolmez, D., Na, H., Lever, M. A., Kjeldsen, K. U., Jørgensen, B. B., and Plugge, C. M.: Methanogenic archaea and sulfate reducing bacteria co-cultured on acetate: teamwork or coexistence?, Front. Microbiol., 6, 492, <ext-link xlink:href="https://doi.org/10.3389/fmicb.2015.00492" ext-link-type="DOI">10.3389/fmicb.2015.00492</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bib60"><label>60</label><mixed-citation>Pellerin, A., Lotem, N., Walter Anthony, K., Eliani Russak, E., Hasson, N., Røy, H., Chanton, J. P., and Sivan, O.: Methane production controls in a young thermokarst lake formed by abrupt permafrost thaw, Glob. Chang. Biol., 28, 3206–3221, <ext-link xlink:href="https://doi.org/10.1111/gcb.16151" ext-link-type="DOI">10.1111/gcb.16151</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bib61"><label>61</label><mixed-citation>Penger, J., Conrad, R., and Blaser, M.: Stable carbon isotope fractionation by methylotrophic methanogenic archaea, Appl. Environ. Microbiol., 78, 7596–7602, <ext-link xlink:href="https://doi.org/10.1128/AEM.01773-12" ext-link-type="DOI">10.1128/AEM.01773-12</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bib62"><label>62</label><mixed-citation>Petersen, S. G. G., Kristensen, E., and Quintana, C. O.: Greenhouse gas emissions from agricultural land before and after permanent flooding with seawater or freshwater, Estuar. Coasts, 46, 1459–1474, <ext-link xlink:href="https://doi.org/10.1007/s12237-023-01218-6" ext-link-type="DOI">10.1007/s12237-023-01218-6</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bib63"><label>63</label><mixed-citation>Poffenbarger, H. J., Needelman, B. A., and Megonigal, J. P.: Salinity influence on methane emissions from tidal marshes, Wetlands, 31, 831–842, <ext-link xlink:href="https://doi.org/10.1007/s13157-011-0197-0" ext-link-type="DOI">10.1007/s13157-011-0197-0</ext-link>, 2011.</mixed-citation></ref>
      <ref id="bib1.bib64"><label>64</label><mixed-citation>Pönisch, D. L., Breznikar, A., Gutekunst, C. N., Jurasinski, G., Voss, M., and Rehder, G.: Nutrient release and flux dynamics of CO<sub>2</sub>, CH<sub>4</sub>, and N<sub>2</sub>O in a coastal peatland driven by actively induced rewetting with brackish water from the Baltic Sea, Biogeosciences, 20, 295–323, <ext-link xlink:href="https://doi.org/10.5194/bg-20-295-2023" ext-link-type="DOI">10.5194/bg-20-295-2023</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bib65"><label>65</label><mixed-citation> Rampton, V. N.: Quaternary geology of the Tuktoyaktuk coastlands, Northwest Territories, Geological Survey of Canada, Memoir 423, 98 pp., Report No. EMR-M46-423E, MICROLOG-88-02809, 1988.</mixed-citation></ref>
      <ref id="bib1.bib66"><label>66</label><mixed-citation>Reeburgh, W. S.: Oceanic methane biogeochemistry, Chem. Rev., 107, 486–513, <ext-link xlink:href="https://doi.org/10.1021/cr050362v" ext-link-type="DOI">10.1021/cr050362v</ext-link>, 2009.</mixed-citation></ref>
      <ref id="bib1.bib67"><label>67</label><mixed-citation>Rosentreter, J. A., Maher, D. T., Erler, D. V., Murray, R. H., and Eyre, B. D.: Methane emissions partially offset blue carbon burial in mangroves, Sci. Adv., 4, eao4985, <ext-link xlink:href="https://doi.org/10.1126/sciadv.aao4985" ext-link-type="DOI">10.1126/sciadv.aao4985</ext-link>, 2018.</mixed-citation></ref>
      <ref id="bib1.bib68"><label>68</label><mixed-citation>Roy Chowdhury, T., Herndon, E. M., Phelps, T. J., Elias, D. A., Gu, B., Liang, L., Wullschleger, S. D., and Graham, D. E.: Stoichiometry and temperature sensitivity of methanogenesis and CO2 production from saturated polygonal tundra in Barrow, Alaska, Glob. Chang. Biol., 21, 722–737, <ext-link xlink:href="https://doi.org/10.1111/gcb.12762" ext-link-type="DOI">10.1111/gcb.12762</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bib69"><label>69</label><mixed-citation>Schuur, E. A. G., McGuire, A. D., Schädel, C., Grosse, G., Harden, J. W., Hayes, D. J., Hugelius, G., Koven, C. D., Kuhry, P., Lawrence, D. M., Natali, S. M., Olefeldt, D., Romanovsky, V. E., Schaefer, K., Turetsky, M. R., Treat, C. C., and Vonk, J. E.: Climate change and the permafrost carbon feedback, Nature, 520, 171–179, <ext-link xlink:href="https://doi.org/10.1038/nature14338" ext-link-type="DOI">10.1038/nature14338</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bib70"><label>70</label><mixed-citation>Segarra, K. E., Comerford, C., Slaughter, J., and Joye, S. B.: Impact of electron acceptor availability on the anaerobic oxidation of methane in coastal freshwater and brackish wetland sediments, Geochim. Cosmochim. Acta, 115, 15–30, <ext-link xlink:href="https://doi.org/10.1016/j.gca.2013.03.029" ext-link-type="DOI">10.1016/j.gca.2013.03.029</ext-link>, 2013.</mixed-citation></ref>
      <ref id="bib1.bib71"><label>71</label><mixed-citation>Sela-Adler, M., Ronen, Z., Herut, B., Antler, G., Vigderovich, H., Eckert, W., and Sivan, O.: Co-existence of methanogenesis and sulfate reduction with common substrates in sulfate-rich estuarine sediments, Front. Microbiol., 8, 766, <ext-link xlink:href="https://doi.org/10.3389/fmicb.2017.00766" ext-link-type="DOI">10.3389/fmicb.2017.00766</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bib72"><label>72</label><mixed-citation>Sepulveda-Jauregui, A., Walter Anthony, K. M., Martinez-Cruz, K., Greene, S., and Thalasso, F.: Methane and carbon dioxide emissions from 40 lakes along a north–south latitudinal transect in Alaska, Biogeosciences, 12, 3197–3223, <ext-link xlink:href="https://doi.org/10.5194/bg-12-3197-2015" ext-link-type="DOI">10.5194/bg-12-3197-2015</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bib73"><label>73</label><mixed-citation>Sherr, E., Sherr, B., and Sigmon, C.: Activity of marine bacteria under incubated and in situ conditions, Aquat. Microb. Ecol., 20, 213–223, <ext-link xlink:href="https://doi.org/10.3354/ame020213" ext-link-type="DOI">10.3354/ame020213</ext-link>, 1999.</mixed-citation></ref>
      <ref id="bib1.bib74"><label>74</label><mixed-citation> Skoog, D. A., West, D. M., Holler, F. J., and Crouch, S. R.: Fundamentals of analytical chemistry, 9th ed., Cengage Learning, Singapore, ISBN 978-0-495-55828-6, 2014.</mixed-citation></ref>
      <ref id="bib1.bib75"><label>75</label><mixed-citation>Solomon, S. M.: Spatial and temporal variability of shoreline change in the Beaufort-Mackenzie region, Northwest Territories, Canada, Geo-Mar. Lett., 25, 127–137, <ext-link xlink:href="https://doi.org/10.1007/s00367-004-0194-x" ext-link-type="DOI">10.1007/s00367-004-0194-x</ext-link>, 2005.</mixed-citation></ref>
      <ref id="bib1.bib76"><label>76</label><mixed-citation>Steedman, A. E., Lantz, T. C., and Kokelj, S. V.: Spatio-temporal variation in high-centre polygons and ice-wedge melt ponds, Tuktoyaktuk Coastlands, Northwest Territories, Permafr. Periglac. Process., 28, 66–78, <ext-link xlink:href="https://doi.org/10.1002/ppp.1880" ext-link-type="DOI">10.1002/ppp.1880</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bib77"><label>77</label><mixed-citation>Tanski, G., Bröder, L., Wagner, D., Knoblauch, C., Lantuit, H., Beer, C., Sachs, T., Fritz, M., Tesi, T., Koch, B. P., Haghipour, N., Eglinton, T. I., Strauss, J., and Vonk, J. E.: Permafrost carbon and CO<sub>2</sub> pathways differ at contrasting coastal erosion sites in the Canadian Arctic, Front. Earth Sci., 9, 630493, <ext-link xlink:href="https://doi.org/10.3389/feart.2021.630493" ext-link-type="DOI">10.3389/feart.2021.630493</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bib78"><label>78</label><mixed-citation> Torres-Alvarado, R., Ramírez-Vives, F., Fernández, F. J., and Barriga-Sosa, I.: Methanogenesis and methane oxidation in wetlands: Implications in the global carbon cycle, Hidrobiológica, 15, 327–349, 2005.</mixed-citation></ref>
      <ref id="bib1.bib79"><label>79</label><mixed-citation>Treat, C. C., Wollheim, W. M., Varner, R. K., Grandy, A. S., Talbot, J., and Frolking, S.: Temperature and peat type control CO<sub>2</sub> and CH<sub>4</sub> production in Alaskan permafrost peats, Glob. Chang. Biol., 20, 2674–2686, <ext-link xlink:href="https://doi.org/10.1111/gcb.12572" ext-link-type="DOI">10.1111/gcb.12572</ext-link>, 2014.</mixed-citation></ref>
      <ref id="bib1.bib80"><label>80</label><mixed-citation>Turetsky, M. R., Treat, C. C., Waldrop, M. P., Waddington, J. M., Harden, J. W., and McGuire, A. D.: Short-term response of methane fluxes and methanogen activity to water table and soil warming manipulations in an Alaskan peatland, J. Geophys. Res. Biogeosci., 113, G03S05, <ext-link xlink:href="https://doi.org/10.1029/2007JG000496" ext-link-type="DOI">10.1029/2007JG000496</ext-link>, 2008. </mixed-citation></ref>
      <ref id="bib1.bib81"><label>81</label><mixed-citation>Valdemarsen, T. B. and Kristensen, E.: Degradation of dissolved organic monomers and short-chain fatty acids in sandy marine sediment by fermentation and sulfate reduction, Geochim. Cosmochim. Acta, 74, 1593–1605, <ext-link xlink:href="https://doi.org/10.1016/j.gca.2009.12.009" ext-link-type="DOI">10.1016/j.gca.2009.12.009</ext-link>, 2010.</mixed-citation></ref>
      <ref id="bib1.bib82"><label>82</label><mixed-citation>Vardy, S. R., Warner, B. G., and Aravena, R.: Holocene climate effects on the development of a peatland on the Tuktoyaktuk Peninsula, Northwest Territories, Quat. Res., 47, 90–104, <ext-link xlink:href="https://doi.org/10.1006/qres.1996.1869" ext-link-type="DOI">10.1006/qres.1996.1869</ext-link>, 1997.</mixed-citation></ref>
      <ref id="bib1.bib83"><label>83</label><mixed-citation>Vaughn, L. J. S., Conrad, M. E., Bill, M., and Torn, M. S.: Isotopic insights into methane production, oxidation, and emissions in Arctic polygon tundra, Glob. Chang. Biol., 22, 3487–3502, <ext-link xlink:href="https://doi.org/10.1111/gcb.13281" ext-link-type="DOI">10.1111/gcb.13281</ext-link>, 2016.</mixed-citation></ref>
      <ref id="bib1.bib84"><label>84</label><mixed-citation>Whalen, D., Forbes, D. L., Kostylev, V., Lim, M., Fraser, P., Nedimović, M. R., and Stuckey, S.: Mechanisms, volumetric assessment, and prognosis for rapid coastal erosion of Tuktoyaktuk Island, an important natural barrier for the harbour and community, Can. J. Earth Sci., 59, 945–960, <ext-link xlink:href="https://doi.org/10.1139/cjes-2021-0101" ext-link-type="DOI">10.1139/cjes-2021-0101</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bib85"><label>85</label><mixed-citation>Winfrey, M. R. and Ward, D. M.: Substrates for sulfate reduction and methane production in intertidal sediments, Appl. Environ. Microbiol., 45, 193–199, <ext-link xlink:href="https://doi.org/10.1128/aem.45.1.193-199.1983" ext-link-type="DOI">10.1128/aem.45.1.193-199.1983</ext-link>, 1983.</mixed-citation></ref>
      <ref id="bib1.bib86"><label>86</label><mixed-citation>Winkel, M., Sepulveda-Jauregui, A., Martinez-Cruz, K., Heslop, J. K., Rijkers, R., Horn, F., Liebner, S., and Walter Anthony, K. M.: First evidence for cold-adapted anaerobic oxidation of methane in deep sediments of thermokarst lakes, Environ. Res. Commun., 1, 021002, <ext-link xlink:href="https://doi.org/10.1088/2515-7620/ab1042" ext-link-type="DOI">10.1088/2515-7620/ab1042</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bib87"><label>87</label><mixed-citation>Worthington, T. A., Spalding, M., Landis, E., Maxwell, T. L., Navarro, A., Smart, L. S., Murray, N. J.: The distribution of global tidal marshes from Earth observation data, Global Ecol. Biogeogr., 33, <ext-link xlink:href="https://doi.org/10.1111/geb.13852" ext-link-type="DOI">10.1111/geb.13852</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bib88"><label>88</label><mixed-citation>Yang, S., Anthony, S. E., Jenrich, M., in 't Zandt, M. H., Strauss, J., Overduin, P. P., Grosse, G., Angelopoulos, M., Biskaborn, B. K., Grigoriev, M. N., Wagner, D., Knoblauch, C., Jaeschke, A., Rethemeyer, J., and Liebner, S.: Microbial methane cycling in sediments of Arctic thermokarst lagoons, Glob. Chang. Biol., 29, 2714–2731, <ext-link xlink:href="https://doi.org/10.1111/gcb.16649" ext-link-type="DOI">10.1111/gcb.16649</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bib89"><label>89</label><mixed-citation>Yuan, J., Liu, D., Ji, Y., Xiang, J., Lin, Y., Wu, M., and Ding, W.: Spartina alterniflora invasion drastically increases methane production potential by shifting methanogenesis from hydrogenotrophic to methylotrophic pathway in a coastal marsh, J. Ecol., 107, 2436–2450, <ext-link xlink:href="https://doi.org/10.1111/1365-2745.13164" ext-link-type="DOI">10.1111/1365-2745.13164</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bib90"><label>90</label><mixed-citation>Zheng, J., RoyChowdhury, T., Yang, Z., Gu, B., Wullschleger, S. D., and Graham, D. E.: Impacts of temperature and soil characteristics on methane production and oxidation in Arctic tundra, Biogeosciences, 15, 6621–6635, <ext-link xlink:href="https://doi.org/10.5194/bg-15-6621-2018" ext-link-type="DOI">10.5194/bg-15-6621-2018</ext-link>, 2018.</mixed-citation></ref>

  </ref-list></back>
    <!--<article-title-html>Addition of brackish water to tundra soils does not inhibit methane production: implications for Arctic coastal methane production</article-title-html>
<abstract-html/>
<ref-html id="bib1.bib1"><label>1</label><mixed-citation>
      
AMAP: Snow, Water, Ice and Permafrost in the Arctic (SWIPA) 2017, Arctic Monitoring and Assessment Programme (AMAP), Oslo, Norway, 269 pp., ISBN 978-82-7971-101-8, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib2"><label>2</label><mixed-citation>
      
Andrachuk, M. and Smit, B.: Community-based vulnerability assessment of
Tuktoyaktuk, NWT, Canada to environmental and socio-economic
changes, Reg. Environ. Change, 12,
867–885, <a href="https://doi.org/10.1007/s10113-012-0299-0" target="_blank">https://doi.org/10.1007/s10113-012-0299-0</a>, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib3"><label>3</label><mixed-citation>
      
Bianchi, T. S.: The role of terrestrially derived organic carbon in the
coastal ocean: A changing paradigm and the priming effect, Proc. Natl. Acad. Sci. USA, 108,
19473–19481, <a href="https://doi.org/10.1073/pnas.1017982108" target="_blank">https://doi.org/10.1073/pnas.1017982108</a>, 2011.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib4"><label>4</label><mixed-citation>
      
Boetius, A., Ravenschlag, K., Schubert, C. J., Rickert, D., Widdel, F.,
Gieseke, A., Amann, R., Jørgensen, B. B., Witte, U., and Pfannkuche, O.:
A marine microbial consortium apparently mediating anaerobic oxidation of
methane, Nature, 407, 623–626, <a href="https://doi.org/10.1038/35036572" target="_blank">https://doi.org/10.1038/35036572</a>, 2000.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib5"><label>5</label><mixed-citation>
      
Bridgham, S. D., Cadillo-Quiroz, H., Keller, J. K., and Zhuang, Q.: Methane
emissions from wetlands: biogeochemical, microbial, and modeling
perspectives from local to global scales, Glob. Chang. Biol., 19,
1325–1346, <a href="https://doi.org/10.1111/gcb.12131" target="_blank">https://doi.org/10.1111/gcb.12131</a>, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib6"><label>6</label><mixed-citation>
      
Comer-Warner, S. A., Ullah, S., Ampuero Reyes, W., Krause, S., and Chmura,
G. L.: Spartina alterniflora has the highest methane emissions in a St.
Lawrence estuary salt marsh, Environ. Res. Ecol., 1,
011003, <a href="https://doi.org/10.1088/2752-664X/ac706a" target="_blank">https://doi.org/10.1088/2752-664X/ac706a</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib7"><label>7</label><mixed-citation>
      
Conrad, R.: Importance of hydrogenotrophic, aceticlastic and methylotrophic
methanogenesis for methane production in terrestrial, aquatic and other
anoxic environments: A mini review, Pedosphere, 30,
25–39, <a href="https://doi.org/10.1016/S1002-0160(18)60052-9" target="_blank">https://doi.org/10.1016/S1002-0160(18)60052-9</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib8"><label>8</label><mixed-citation>
      
Coolen, M. J. L. and Orsi, W. D.: The transcriptional response of microbial
communities in thawing Alaskan permafrost soils, Front. Microbiol.,
6, 197, <a href="https://doi.org/10.3389/fmicb.2015.00197" target="_blank">https://doi.org/10.3389/fmicb.2015.00197</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib9"><label>9</label><mixed-citation>
      
Costa, B., Vieira, G., and Whalen, D.: The fast-changing coast of Tuktoyaktuk Peninsula (Beaufort Sea, Canada): geomorphological controls on changes between 1985 and 2020, EGU General Assembly 2022, Vienna, Austria, 23–27 May 2022, EGU22-2426, <a href="https://doi.org/10.5194/egusphere-egu22-2426" target="_blank">https://doi.org/10.5194/egusphere-egu22-2426</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib10"><label>10</label><mixed-citation>
      
Couture, N. J., Irrgang, A., Pollard, W., Lantuit, H., and Fritz, M.:
Coastal erosion of permafrost soils along the Yukon Coastal Plain and fluxes
of organic carbon to the Canadian Beaufort Sea, J. Geophys. Res. Biogeosci., 123,
406–422, <a href="https://doi.org/10.1002/2017JG004166" target="_blank">https://doi.org/10.1002/2017JG004166</a>, 2018.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib11"><label>11</label><mixed-citation>
      
Cui, S., Liu, P., Guo, H., Nielsen, C. K., Pullens, J. W. M., Chen, Q.,
Pugliese, L., and Wu, S.: Wetland hydrological dynamics and methane
emissions, Commun. Earth Environ., 5,
1635, <a href="https://doi.org/10.1038/s43247-024-01635-w" target="_blank">https://doi.org/10.1038/s43247-024-01635-w</a>, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib12"><label>12</label><mixed-citation>
      
Dallimore, S. R., Wolfe, S. A., Matthews Jr., J. V., and Vincent, J.-S.:
Mid-Wisconsinan eolian deposits of the Kittigazuit Formation, Tuktoyaktuk
Coastlands, Northwest Territories, Canada, Can. J. Earth Sci., 34, 1421–1441, <a href="https://doi.org/10.1139/e17-116" target="_blank">https://doi.org/10.1139/e17-116</a>, 1997.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib13"><label>13</label><mixed-citation>
      
Elberling, B., Michelsen, A., Schädel, C., Schuur, E. A. G.,
Christiansen, H. H., Berg, L., Tamstorf, M. P., and Sigsgaard, C.: Long-term
CO<sub>2</sub> production following permafrost thaw, Nat. Clim. Chang., 3,
890–894, <a href="https://doi.org/10.1038/nclimate1955" target="_blank">https://doi.org/10.1038/nclimate1955</a>, 2013.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib14"><label>14</label><mixed-citation>
      
Froelich, P., Klinkhammer, G., Bender, M., Luedtke, N., Heath, G., Cullen,
D., Dauphin, P., Hammond, D., Hartman, B., and Maynard, V.: Early oxidation
of organic matter in pelagic sediments of the eastern equatorial Atlantic:
suboxic diagenesis, Geochim. Cosmochim. Acta, 43,
1075–1090, <a href="https://doi.org/10.1016/0016-7037(79)90095-4" target="_blank">https://doi.org/10.1016/0016-7037(79)90095-4</a>, 1979.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib15"><label>15</label><mixed-citation>
      
Galand, P. E., Yrjälä, K., and Conrad, R.: Stable carbon isotope fractionation during methanogenesis in three boreal peatland ecosystems, Biogeosciences, 7, 3893–3900, <a href="https://doi.org/10.5194/bg-7-3893-2010" target="_blank">https://doi.org/10.5194/bg-7-3893-2010</a>, 2010.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib16"><label>16</label><mixed-citation>
      
Guimond, J. A., Mohammed, A. A., Walvoord, M. A., Bense, V. F., and Kurylyk,
B. L.: Saltwater intrusion intensifies coastal permafrost thaw, Geophys. Res. Lett., 48, e2021GL094776, <a href="https://doi.org/10.1029/2021GL094776" target="_blank">https://doi.org/10.1029/2021GL094776</a>,
2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib17"><label>17</label><mixed-citation>
      
Heffernan, L., Cavaco, M. A., Bhatia, M. P., Estop-Aragonés, C., Knorr, K.-H., and Olefeldt, D.: High peatland methane emissions following permafrost thaw: enhanced acetoclastic methanogenesis during early successional stages, Biogeosciences, 19, 3051–3071, <a href="https://doi.org/10.5194/bg-19-3051-2022" target="_blank">https://doi.org/10.5194/bg-19-3051-2022</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib18"><label>18</label><mixed-citation>
      
Heslop, J. K., Walter Anthony, K. M., Sepulveda-Jauregui, A., Martinez-Cruz, K., Bondurant, A., Grosse, G., and Jones, M. C.: Thermokarst lake methanogenesis along a complete talik profile, Biogeosciences, 12, 4317–4331, <a href="https://doi.org/10.5194/bg-12-4317-2015" target="_blank">https://doi.org/10.5194/bg-12-4317-2015</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib19"><label>19</label><mixed-citation>
      
Heslop, J. K., Walter Anthony, K. M., Winkel, M., Sepulveda-Jauregui, A.,
Martinez-Cruz, K., Bondurant, A., Grosse, G., and Liebner, S.: A synthesis
of methane dynamics in thermokarst lake environments, Earth Sci. Rev.,
210, 103365, <a href="https://doi.org/10.1016/j.earscirev.2020.103365" target="_blank">https://doi.org/10.1016/j.earscirev.2020.103365</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib20"><label>20</label><mixed-citation>
      
Heyer, J., Berger, U., Kuzin, I. L., and Yakovlev, O. N.: Methane emissions
from different ecosystem structures of the subarctic tundra in Western
Siberia during midsummer and during the thawing period, Tellus B, 54,
231–249, <a href="https://doi.org/10.1034/j.1600-0889.2002.01280.x" target="_blank">https://doi.org/10.1034/j.1600-0889.2002.01280.x</a>, 2002.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib21"><label>21</label><mixed-citation>
      
Hill, P. R., Héquette, A., and Ruz, M.-H.: Holocene sea-level history of
the Canadian Beaufort shelf, Can. J. Earth Sci., 30,
103–108, <a href="https://doi.org/10.1139/e93-009" target="_blank">https://doi.org/10.1139/e93-009</a>, 1993.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib22"><label>22</label><mixed-citation>
      
Hodgkins, S. B., Tfaily, M. M., McCalley, C. K., Logan, T. A., Crill, P. M.,
Saleska, S. R., Rich, V. I., and Chanton, J. P.: Changes in peat chemistry
associated with permafrost thaw increase greenhouse gas production,
Proc. Natl. Acad. Sci. USA, 111,
5819–5824, <a href="https://doi.org/10.1073/pnas.1314641111" target="_blank">https://doi.org/10.1073/pnas.1314641111</a>, 2014.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib23"><label>23</label><mixed-citation>
      
Holm, S., Walz, J., Horn, F., Yang, S., Grigoriev, M. N., Wagner, D.,
Knoblauch, C., and Liebner, S.: Methanogenic response to long-term
permafrost thaw is determined by paleoenvironment, FEMS Microbiol. Ecol., 96, fiaa021, <a href="https://doi.org/10.1093/femsec/fiaa021" target="_blank">https://doi.org/10.1093/femsec/fiaa021</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib24"><label>24</label><mixed-citation>
      
Hornibrook, E. R., Longstaffe, F. J., and Fyfe, W. S.: Spatial distribution
of microbial methane production pathways in temperate zone wetland soils:
stable carbon and hydrogen isotope evidence, Geochim. Cosmochim. Acta, 61, 745–753, <a href="https://doi.org/10.1016/S0016-7037(96)00368-7" target="_blank">https://doi.org/10.1016/S0016-7037(96)00368-7</a>, 1997.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib25"><label>25</label><mixed-citation>
      
Hornibrook, E. R. C., Longstaffe, F. J., and Fyfe, W. S.: Evolution of
stable carbon isotope compositions for methane and carbon dioxide in
freshwater wetlands and other anaerobic environments, Geochim. Cosmochim. Acta, 64,
1013–1027, <a href="https://doi.org/10.1016/S0016-7037(99)00321-X" target="_blank">https://doi.org/10.1016/S0016-7037(99)00321-X</a>, 2000.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib26"><label>26</label><mixed-citation>
      
Hu, H., Chen, J., Zhou, F., Nie, M., Hou, D., Liu, H., Delgado-Baquerizo,
M., Ni, H., Huang, W., Zhou, J., Song, X., Cao, X., Sun, B., Zhang, J.,
Crowther, T. W., and Liang, Y.: Relative increases in CH<sub>4</sub> and CO2 emissions
from wetlands under global warming dependent on soil carbon substrates,
Nat. Geosci., 17, 26–31, <a href="https://doi.org/10.1038/s41561-023-01345-6" target="_blank">https://doi.org/10.1038/s41561-023-01345-6</a>,
2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib27"><label>27</label><mixed-citation>
      
Hu, K., Issler, D., Chen, Z., and Brent, T.: Permafrost investigation by
well logs, and seismic velocity and repeated shallow temperature surveys,
Beaufort-Mackenzie Basin, Geological Survey of
Canada, <a href="https://doi.org/10.4095/293120" target="_blank">https://doi.org/10.4095/293120</a>, 2013.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib28"><label>28</label><mixed-citation>
      
Hynes, S., Solomon, S. M., and Whalen, D.: GIS compilation of coastline
variability spanning 60 years in the Mackenzie Delta and Tuktoyaktuk in the
Beaufort Sea, Geological Survey of Canada Open File
7685, <a href="https://doi.org/10.4095/295579" target="_blank">https://doi.org/10.4095/295579</a>, 2014.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib29"><label>29</label><mixed-citation>
      
Ionescu, D., Bizic-Ionescu, M., Khalili, A., Malekmohammadi, R., Morad, M.
R., de Beer, D., and Grossart, H.-P.: A new tool for long-term studies of
POM-bacteria interactions: overcoming the century-old Bottle Effect,
Sci. Rep., 5, 14706, <a href="https://doi.org/10.1038/srep14706" target="_blank">https://doi.org/10.1038/srep14706</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib30"><label>30</label><mixed-citation>
      
Irrgang, A. M., Bendixen, M., Farquharson, L. M., Baranskaya, A. V.,
Erikson, L. H., Gibbs, A. E., Ogorodov, S. A., Overduin, P. P., Lantuit, H.,
Grigoriev, M. N., and Jones, B. M.: Drivers, dynamics and impacts of
changing Arctic coasts, Nat. Rev. Earth Environ., 3,
39–54, <a href="https://doi.org/10.1038/s43017-021-00232-1" target="_blank">https://doi.org/10.1038/s43017-021-00232-1</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib31"><label>31</label><mixed-citation>
      
Jenrich, M., Angelopoulos, M., Liebner, S., Treat, C. C., Knoblauch, C.,
Yang, S., Grosse, G., Giebeler, F., Jongejans, L. L., Grigoriev, M., and
Strauss, J.: Greenhouse gas production and microbial response during the
transition from terrestrial permafrost to a marine environment, Permafr. Periglac. Process.,
<a href="https://doi.org/10.1002/ppp.2251" target="_blank">https://doi.org/10.1002/ppp.2251</a>, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib32"><label>32</label><mixed-citation>
      
Jenrich, M., Wolter, J., Liebner, S., Knoblauch, C., Grosse, G., Giebeler, F., Whalen, D., and Strauss, J.: Rising Arctic seas and thawing permafrost: uncovering the carbon cycle impact in a thermokarst lagoon system in the outer Mackenzie Delta, Canada, Biogeosciences, 22, 2069–2086, <a href="https://doi.org/10.5194/bg-22-2069-2025" target="_blank">https://doi.org/10.5194/bg-22-2069-2025</a>, 2025.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib33"><label>33</label><mixed-citation>
      
Jones, E. L., Hodson, A. J., Thornton, S. F., Redeker, K. R., Rogers, J.,
Wynn, P. M., Dixon, T. J., Bottrell, S. H., and O'Neill, H. B.:
Biogeochemical processes in the active layer and permafrost of a high Arctic
fjord valley, Front. Earth Sci., 8,
342, <a href="https://doi.org/10.3389/feart.2020.00342" target="_blank">https://doi.org/10.3389/feart.2020.00342</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib34"><label>34</label><mixed-citation>
      
Keuschnig, C., Larose, C., Rudner, M., Pesqueda, A., Doleac, S., Elberling,
B., Björk, R. G., Klemedtsson, L., and Björkman, M. P.: Reduced
methane emissions in former permafrost soils driven by vegetation and
microbial changes following drainage, Glob. Chang. Biol., 28,
3411–3425, <a href="https://doi.org/10.1111/gcb.16137" target="_blank">https://doi.org/10.1111/gcb.16137</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib35"><label>35</label><mixed-citation>
      
Knittel, K. and Boetius, A.: Anaerobic oxidation of methane: progress with
an unknown process, Annu. Rev. Microbiol., 63,
311–334, <a href="https://doi.org/10.1146/annurev.micro.61.080706.093130" target="_blank">https://doi.org/10.1146/annurev.micro.61.080706.093130</a>, 2009.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib36"><label>36</label><mixed-citation>
      
Knoblauch, C., Beer, C., Sosnin, A., Wagner, D., and Pfeiffer, E.-M.:
Predicting long-term carbon mineralization and trace gas production from
thawing permafrost of Northeast Siberia, Glob. Chang. Biol., 19,
1160–1172, <a href="https://doi.org/10.1111/gcb.12116" target="_blank">https://doi.org/10.1111/gcb.12116</a>, 2013.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib37"><label>37</label><mixed-citation>
      
Knoblauch, C., Beer, C., Liebner, S., Grigoriev, M. N., and Pfeiffer, E.-M.:
Methane production as key to the greenhouse gas budget of thawing
permafrost, Nat. Clim. Chang., 8,
309–312, <a href="https://doi.org/10.1038/s41558-018-0095-z" target="_blank">https://doi.org/10.1038/s41558-018-0095-z</a>, 2018.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib38"><label>38</label><mixed-citation>
      
Kokelj, S. V., Lantz, T. C., Solomon, S., Pisaric, M. F., Keith, D., Morse,
P., Thienpont, J. R., Smol, J. P., and Esagok, D.: Using multiple sources of
knowledge to investigate northern environmental change: regional ecological
impacts of a storm surge in the Outer Mackenzie Delta, N.W.T., Arctic, 65,
3, <a href="https://doi.org/10.14430/arctic4214" target="_blank">https://doi.org/10.14430/arctic4214</a>, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib39"><label>39</label><mixed-citation>
      
Kramshøj, M., Albers, C. N., Holst, T., Holzinger, R., Elberling, B., and
Rinnan, R.: Biogenic volatile release from permafrost thaw is determined by
the soil microbial sink, Nat. Commun., 9,
3412, <a href="https://doi.org/10.1038/s41467-018-05824-y" target="_blank">https://doi.org/10.1038/s41467-018-05824-y</a>, 2018.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib40"><label>40</label><mixed-citation>
      
Kroeger, K. D., Crooks, S., Moseman-Valtierra, S., and Tang, J.: Restoring
tides to reduce methane emissions in impounded wetlands: a new and potent
blue carbon climate change intervention, Sci. Rep., 7,
12138, <a href="https://doi.org/10.1038/s41598-017-12138-4" target="_blank">https://doi.org/10.1038/s41598-017-12138-4</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib41"><label>41</label><mixed-citation>
      
La, W., Han, X., Liu, C.-Q., Ding, H., Liu, M., Sun, F., Li, S., and Lang,
Y.: Sulfate concentrations affect sulfate reduction pathways and methane
consumption in coastal wetlands, Water Res., 217,
118441, <a href="https://doi.org/10.1016/j.watres.2022.118441" target="_blank">https://doi.org/10.1016/j.watres.2022.118441</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib42"><label>42</label><mixed-citation>
      
Lacelle, D., Fontaine, M., Pellerin, A., Kokelj, S. V., and Clark, I. D.:
Legacy of Holocene landscape changes on soil biogeochemistry: a perspective
from paleo-active layers in northwestern Canada, J. Geophys. Res. Biogeosciences, 124,
2662–2679, <a href="https://doi.org/10.1029/2018JG004916" target="_blank">https://doi.org/10.1029/2018JG004916</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib43"><label>43</label><mixed-citation>
      
Lantuit, H., Overduin, P. P., Couture, N., Wetterich, S., Aré, F.,
Atkinson, D., Brown, J., Cherkashov, G., Drozdov, D., Forbes, D. L.,
Graves-Gaylord, A., Grigoriev, M., Hubberten, H.-W., Jordan, J., Jorgenson,
T., Ødegård, R. S., Ogorodov, S., Pollard, W. H., Rachold, V., and
Vasiliev, A.: The Arctic coastal dynamics database: a new classification
scheme and statistics on Arctic permafrost coastlines, Estuar. Coasts,
35, 383–400, <a href="https://doi.org/10.1007/s12237-010-9362-6" target="_blank">https://doi.org/10.1007/s12237-010-9362-6</a>, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib44"><label>44</label><mixed-citation>
      
Lapham, L. L., Dallimore, S. R., Magen, C., Henderson, L. C., Powers, L. C.,
Gonsior, M., Clark, B., Côté, M., Fraser, P., and Orcutt, B. N.:
Microbial greenhouse gas dynamics associated with warming coastal
permafrost, western Canadian Arctic, Front. Earth Sci., 8,
582103, <a href="https://doi.org/10.3389/feart.2020.582103" target="_blank">https://doi.org/10.3389/feart.2020.582103</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib45"><label>45</label><mixed-citation>
      
Li, Y., Wang, D., Chen, Z., Chen, J., Hu, H., and Wang, R.: Methane
emissions during the tide cycle of a Yangtze Estuary salt marsh, Atmosphere,
12, 245, <a href="https://doi.org/10.3390/atmos12020245" target="_blank">https://doi.org/10.3390/atmos12020245</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib46"><label>46</label><mixed-citation>
      
Liebner, S., Ganzert, L., Kiss, A., Yang, S., Wagner, D., and Svenning, M.
M.: Shifts in methanogenic community composition and methane fluxes along
the degradation of discontinuous permafrost, Front. Microbiol., 6,
356, <a href="https://doi.org/10.3389/fmicb.2015.00356" target="_blank">https://doi.org/10.3389/fmicb.2015.00356</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib47"><label>47</label><mixed-citation>
      
Lim, M., Whalen, D., Martin, J., Mann, P. J., Hayes, S., Fraser, P., Berry,
H. B., and Ouellette, D.: Massive ice control on permafrost coast erosion
and sensitivity, Geophys. Res. Lett., 47,
e2020GL087917, <a href="https://doi.org/10.1029/2020GL087917" target="_blank">https://doi.org/10.1029/2020GL087917</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib48"><label>48</label><mixed-citation>
      
Lipson, D. A., Zona, D., Raab, T. K., Bozzolo, F., Mauritz, M., and Oechel, W. C.: Water-table height and microtopography control biogeochemical cycling in an Arctic coastal tundra ecosystem, Biogeosciences, 9, 577–591, <a href="https://doi.org/10.5194/bg-9-577-2012" target="_blank">https://doi.org/10.5194/bg-9-577-2012</a>, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib49"><label>49</label><mixed-citation>
      
Lotem, N., Pellerin, A., Anthony, K. W., Gafni, A., Boyko, V., and Sivan,
O.: Anaerobic oxidation of methane does not attenuate methane emissions from
thermokarst lakes, Limnol. Oceanogr., 68,
1316–1330, <a href="https://doi.org/10.1002/lno.12349" target="_blank">https://doi.org/10.1002/lno.12349</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib50"><label>50</label><mixed-citation>
      
Lovley, D. R. and Klug, M. J.: Sulfate reducers can outcompete methanogens
at freshwater sulfate concentrations, Appl. Environ. Microbiol., 45, 187–192, <a href="https://doi.org/10.1128/aem.45.1.187-192.1983" target="_blank">https://doi.org/10.1128/aem.45.1.187-192.1983</a>,
1983.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib51"><label>51</label><mixed-citation>
      
Mackay, J. R. and Dallimore, S. R.: Massive ice of the Tuktoyaktuk area,
western Arctic coast, Canada, Can. J. Earth Sci., 29,
1235–1249, <a href="https://doi.org/10.1139/e92-099" target="_blank">https://doi.org/10.1139/e92-099</a>, 1992.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib52"><label>52</label><mixed-citation>
      
Maltby, J., Steinle, L., Löscher, C. R., Bange, H. W., Fischer, M. A., Schmidt, M., and Treude, T.: Microbial methanogenesis in the sulfate-reducing zone of sediments in the Eckernförde Bay, SW Baltic Sea, Biogeosciences, 15, 137–157, <a href="https://doi.org/10.5194/bg-15-137-2018" target="_blank">https://doi.org/10.5194/bg-15-137-2018</a>, 2018.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib53"><label>53</label><mixed-citation>
      
Manson, G. K., Couture, N. J., and James, T. S.: CanCoast 2.0: data and
indices to describe the sensitivity of Canada's marine coasts to changing
climate, Geological Survey of Canada Open File
8551, <a href="https://doi.org/10.4095/314669" target="_blank">https://doi.org/10.4095/314669</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib54"><label>54</label><mixed-citation>
      
Martens, C. S. and Berner, R. A.: Methane production in the interstitial
waters of sulfate-depleted marine sediments, Science, 185,
1167–1169, <a href="https://doi.org/10.1126/science.185.4157.1167" target="_blank">https://doi.org/10.1126/science.185.4157.1167</a>, 1974.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib55"><label>55</label><mixed-citation>
      
Martin, A. F., Lantz, T. C., and Humphreys, E. R.: Ice wedge degradation and
CO2 and CH<sub>4</sub> emissions in the Tuktoyaktuk Coastlands, Northwest Territories,
Arct. Sci., 4, 130–145, <a href="https://doi.org/10.1139/as-2016-0011" target="_blank">https://doi.org/10.1139/as-2016-0011</a>, 2018.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib56"><label>56</label><mixed-citation>
      
Murton, J. B.: Thermokarst-lake-basin sediments, Tuktoyaktuk Coastlands,
western Arctic Canada, Sedimentology, 43,
737–760, <a href="https://doi.org/10.1111/j.1365-3091.1996.tb02023.x" target="_blank">https://doi.org/10.1111/j.1365-3091.1996.tb02023.x</a>, 1996.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib57"><label>57</label><mixed-citation>
      
Oh, Y., Zhuang, Q., Liu, L., Welp, L. R., Lau, M. C. Y., Onstott, T. C.,
Medvigy, D., Bruhwiler, L., Dlugokencky, E. J., Hugelius, G., D'Imperio, L.,
and Elberling, B.: Reduced net methane emissions due to microbial methane
oxidation in a warmer Arctic, Nat. Clim. Chang., 10,
317–321, <a href="https://doi.org/10.1038/s41558-020-0734-z" target="_blank">https://doi.org/10.1038/s41558-020-0734-z</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib58"><label>58</label><mixed-citation>
      
Oremland, R. S. and Polcin, S.: Methanogenesis and sulfate reduction:
competitive and noncompetitive substrates in estuarine sediments, Appl. Environ. Microbiol., 44,
1270–1276, <a href="https://doi.org/10.1128/aem.44.6.1270-1276.1982" target="_blank">https://doi.org/10.1128/aem.44.6.1270-1276.1982</a>, 1982.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib59"><label>59</label><mixed-citation>
      
Ozuolmez, D., Na, H., Lever, M. A., Kjeldsen, K. U., Jørgensen, B. B.,
and Plugge, C. M.: Methanogenic archaea and sulfate reducing bacteria
co-cultured on acetate: teamwork or coexistence?, Front. Microbiol.,
6, 492, <a href="https://doi.org/10.3389/fmicb.2015.00492" target="_blank">https://doi.org/10.3389/fmicb.2015.00492</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib60"><label>60</label><mixed-citation>
      
Pellerin, A., Lotem, N., Walter Anthony, K., Eliani Russak, E., Hasson, N.,
Røy, H., Chanton, J. P., and Sivan, O.: Methane production controls in a
young thermokarst lake formed by abrupt permafrost thaw, Glob. Chang. Biol., 28, 3206–3221, <a href="https://doi.org/10.1111/gcb.16151" target="_blank">https://doi.org/10.1111/gcb.16151</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib61"><label>61</label><mixed-citation>
      
Penger, J., Conrad, R., and Blaser, M.: Stable carbon isotope fractionation
by methylotrophic methanogenic archaea, Appl. Environ. Microbiol., 78, 7596–7602, <a href="https://doi.org/10.1128/AEM.01773-12" target="_blank">https://doi.org/10.1128/AEM.01773-12</a>, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib62"><label>62</label><mixed-citation>
      
Petersen, S. G. G., Kristensen, E., and Quintana, C. O.: Greenhouse gas
emissions from agricultural land before and after permanent flooding with
seawater or freshwater, Estuar. Coasts, 46,
1459–1474, <a href="https://doi.org/10.1007/s12237-023-01218-6" target="_blank">https://doi.org/10.1007/s12237-023-01218-6</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib63"><label>63</label><mixed-citation>
      
Poffenbarger, H. J., Needelman, B. A., and Megonigal, J. P.: Salinity
influence on methane emissions from tidal marshes, Wetlands, 31,
831–842, <a href="https://doi.org/10.1007/s13157-011-0197-0" target="_blank">https://doi.org/10.1007/s13157-011-0197-0</a>, 2011.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib64"><label>64</label><mixed-citation>
      
Pönisch, D. L., Breznikar, A., Gutekunst, C. N., Jurasinski, G., Voss, M., and Rehder, G.: Nutrient release and flux dynamics of CO<sub>2</sub>, CH<sub>4</sub>, and N<sub>2</sub>O in a coastal peatland driven by actively induced rewetting with brackish water from the Baltic Sea, Biogeosciences, 20, 295–323, <a href="https://doi.org/10.5194/bg-20-295-2023" target="_blank">https://doi.org/10.5194/bg-20-295-2023</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib65"><label>65</label><mixed-citation>
      
Rampton, V. N.: Quaternary geology of the Tuktoyaktuk coastlands, Northwest Territories, Geological Survey of Canada, Memoir 423, 98 pp., Report No. EMR-M46-423E, MICROLOG-88-02809, 1988.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib66"><label>66</label><mixed-citation>
      
Reeburgh, W. S.: Oceanic methane biogeochemistry, Chem. Rev., 107,
486–513, <a href="https://doi.org/10.1021/cr050362v" target="_blank">https://doi.org/10.1021/cr050362v</a>, 2009.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib67"><label>67</label><mixed-citation>
      
Rosentreter, J. A., Maher, D. T., Erler, D. V., Murray, R. H., and Eyre, B.
D.: Methane emissions partially offset blue carbon burial in mangroves,
Sci. Adv., 4, eao4985, <a href="https://doi.org/10.1126/sciadv.aao4985" target="_blank">https://doi.org/10.1126/sciadv.aao4985</a>, 2018.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib68"><label>68</label><mixed-citation>
      
Roy Chowdhury, T., Herndon, E. M., Phelps, T. J., Elias, D. A., Gu, B.,
Liang, L., Wullschleger, S. D., and Graham, D. E.: Stoichiometry and
temperature sensitivity of methanogenesis and CO2 production from saturated
polygonal tundra in Barrow, Alaska, Glob. Chang. Biol., 21,
722–737, <a href="https://doi.org/10.1111/gcb.12762" target="_blank">https://doi.org/10.1111/gcb.12762</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib69"><label>69</label><mixed-citation>
      
Schuur, E. A. G., McGuire, A. D., Schädel, C., Grosse, G., Harden, J.
W., Hayes, D. J., Hugelius, G., Koven, C. D., Kuhry, P., Lawrence, D. M.,
Natali, S. M., Olefeldt, D., Romanovsky, V. E., Schaefer, K., Turetsky, M.
R., Treat, C. C., and Vonk, J. E.: Climate change and the permafrost carbon
feedback, Nature, 520, 171–179, <a href="https://doi.org/10.1038/nature14338" target="_blank">https://doi.org/10.1038/nature14338</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib70"><label>70</label><mixed-citation>
      
Segarra, K. E., Comerford, C., Slaughter, J., and Joye, S. B.: Impact of
electron acceptor availability on the anaerobic oxidation of methane in
coastal freshwater and brackish wetland sediments, Geochim. Cosmochim. Acta, 115, 15–30, <a href="https://doi.org/10.1016/j.gca.2013.03.029" target="_blank">https://doi.org/10.1016/j.gca.2013.03.029</a>,
2013.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib71"><label>71</label><mixed-citation>
      
Sela-Adler, M., Ronen, Z., Herut, B., Antler, G., Vigderovich, H., Eckert,
W., and Sivan, O.: Co-existence of methanogenesis and sulfate reduction with
common substrates in sulfate-rich estuarine sediments, Front. Microbiol., 8, 766, <a href="https://doi.org/10.3389/fmicb.2017.00766" target="_blank">https://doi.org/10.3389/fmicb.2017.00766</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib72"><label>72</label><mixed-citation>
      
Sepulveda-Jauregui, A., Walter Anthony, K. M., Martinez-Cruz, K., Greene, S., and Thalasso, F.: Methane and carbon dioxide emissions from 40 lakes along a north–south latitudinal transect in Alaska, Biogeosciences, 12, 3197–3223, <a href="https://doi.org/10.5194/bg-12-3197-2015" target="_blank">https://doi.org/10.5194/bg-12-3197-2015</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib73"><label>73</label><mixed-citation>
      
Sherr, E., Sherr, B., and Sigmon, C.: Activity of marine bacteria under
incubated and in situ conditions, Aquat. Microb. Ecol., 20,
213–223, <a href="https://doi.org/10.3354/ame020213" target="_blank">https://doi.org/10.3354/ame020213</a>, 1999.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib74"><label>74</label><mixed-citation>
      
Skoog, D. A., West, D. M., Holler, F. J., and Crouch, S. R.: Fundamentals of analytical chemistry, 9th ed., Cengage Learning, Singapore, ISBN 978-0-495-55828-6, 2014.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib75"><label>75</label><mixed-citation>
      
Solomon, S. M.: Spatial and temporal variability of shoreline change in the Beaufort-Mackenzie region, Northwest Territories, Canada, Geo-Mar. Lett., 25, 127–137, <a href="https://doi.org/10.1007/s00367-004-0194-x" target="_blank">https://doi.org/10.1007/s00367-004-0194-x</a>, 2005.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib76"><label>76</label><mixed-citation>
      
Steedman, A. E., Lantz, T. C., and Kokelj, S. V.: Spatio-temporal variation
in high-centre polygons and ice-wedge melt ponds, Tuktoyaktuk Coastlands,
Northwest Territories, Permafr. Periglac. Process., 28,
66–78, <a href="https://doi.org/10.1002/ppp.1880" target="_blank">https://doi.org/10.1002/ppp.1880</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib77"><label>77</label><mixed-citation>
      
Tanski, G., Bröder, L., Wagner, D., Knoblauch, C., Lantuit, H., Beer,
C., Sachs, T., Fritz, M., Tesi, T., Koch, B. P., Haghipour, N., Eglinton, T.
I., Strauss, J., and Vonk, J. E.: Permafrost carbon and CO<sub>2</sub> pathways differ
at contrasting coastal erosion sites in the Canadian Arctic, Front. Earth Sci., 9, 630493, <a href="https://doi.org/10.3389/feart.2021.630493" target="_blank">https://doi.org/10.3389/feart.2021.630493</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib78"><label>78</label><mixed-citation>
      
Torres-Alvarado, R., Ramírez-Vives, F., Fernández, F. J., and Barriga-Sosa, I.: Methanogenesis and methane oxidation in wetlands: Implications in the global carbon cycle, Hidrobiológica, 15, 327–349, 2005.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib79"><label>79</label><mixed-citation>
      
Treat, C. C., Wollheim, W. M., Varner, R. K., Grandy, A. S., Talbot, J., and
Frolking, S.: Temperature and peat type control CO<sub>2</sub> and CH<sub>4</sub> production in
Alaskan permafrost peats, Glob. Chang. Biol., 20,
2674–2686, <a href="https://doi.org/10.1111/gcb.12572" target="_blank">https://doi.org/10.1111/gcb.12572</a>, 2014.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib80"><label>80</label><mixed-citation>
      
Turetsky, M. R., Treat, C. C., Waldrop, M. P., Waddington, J. M., Harden, J.
W., and McGuire, A. D.: Short-term response of methane fluxes and methanogen
activity to water table and soil warming manipulations in an Alaskan
peatland, J. Geophys. Res. Biogeosci., 113,
G03S05, <a href="https://doi.org/10.1029/2007JG000496" target="_blank">https://doi.org/10.1029/2007JG000496</a>, 2008.


    </mixed-citation></ref-html>
<ref-html id="bib1.bib81"><label>81</label><mixed-citation>
      
Valdemarsen, T. B. and Kristensen, E.: Degradation of dissolved organic
monomers and short-chain fatty acids in sandy marine sediment by
fermentation and sulfate reduction, Geochim. Cosmochim. Acta, 74,
1593–1605, <a href="https://doi.org/10.1016/j.gca.2009.12.009" target="_blank">https://doi.org/10.1016/j.gca.2009.12.009</a>, 2010.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib82"><label>82</label><mixed-citation>
      
Vardy, S. R., Warner, B. G., and Aravena, R.: Holocene climate effects on
the development of a peatland on the Tuktoyaktuk Peninsula, Northwest
Territories, Quat. Res., 47,
90–104, <a href="https://doi.org/10.1006/qres.1996.1869" target="_blank">https://doi.org/10.1006/qres.1996.1869</a>, 1997.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib83"><label>83</label><mixed-citation>
      
Vaughn, L. J. S., Conrad, M. E., Bill, M., and Torn, M. S.: Isotopic
insights into methane production, oxidation, and emissions in Arctic polygon
tundra, Glob. Chang. Biol., 22,
3487–3502, <a href="https://doi.org/10.1111/gcb.13281" target="_blank">https://doi.org/10.1111/gcb.13281</a>, 2016.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib84"><label>84</label><mixed-citation>
      
Whalen, D., Forbes, D. L., Kostylev, V., Lim, M., Fraser, P., Nedimović,
M. R., and Stuckey, S.: Mechanisms, volumetric assessment, and prognosis for
rapid coastal erosion of Tuktoyaktuk Island, an important natural barrier
for the harbour and community, Can. J. Earth Sci., 59,
945–960, <a href="https://doi.org/10.1139/cjes-2021-0101" target="_blank">https://doi.org/10.1139/cjes-2021-0101</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib85"><label>85</label><mixed-citation>
      
Winfrey, M. R. and Ward, D. M.: Substrates for sulfate reduction and methane
production in intertidal sediments, Appl. Environ. Microbiol.,
45, 193–199, <a href="https://doi.org/10.1128/aem.45.1.193-199.1983" target="_blank">https://doi.org/10.1128/aem.45.1.193-199.1983</a>, 1983.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib86"><label>86</label><mixed-citation>
      
Winkel, M., Sepulveda-Jauregui, A., Martinez-Cruz, K., Heslop, J. K.,
Rijkers, R., Horn, F., Liebner, S., and Walter Anthony, K. M.: First
evidence for cold-adapted anaerobic oxidation of methane in deep sediments
of thermokarst lakes, Environ. Res. Commun., 1,
021002, <a href="https://doi.org/10.1088/2515-7620/ab1042" target="_blank">https://doi.org/10.1088/2515-7620/ab1042</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib87"><label>87</label><mixed-citation>
      
Worthington, T. A., Spalding, M., Landis, E., Maxwell, T. L., Navarro, A., Smart, L. S., Murray, N. J.: The distribution of global tidal marshes from Earth observation data, Global Ecol. Biogeogr., 33, <a href="https://doi.org/10.1111/geb.13852" target="_blank">https://doi.org/10.1111/geb.13852</a>, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib88"><label>88</label><mixed-citation>
      
Yang, S., Anthony, S. E., Jenrich, M., in 't Zandt, M. H., Strauss, J.,
Overduin, P. P., Grosse, G., Angelopoulos, M., Biskaborn, B. K., Grigoriev,
M. N., Wagner, D., Knoblauch, C., Jaeschke, A., Rethemeyer, J., and Liebner,
S.: Microbial methane cycling in sediments of Arctic thermokarst lagoons,
Glob. Chang. Biol., 29, 2714–2731, <a href="https://doi.org/10.1111/gcb.16649" target="_blank">https://doi.org/10.1111/gcb.16649</a>,
2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib89"><label>89</label><mixed-citation>
      
Yuan, J., Liu, D., Ji, Y., Xiang, J., Lin, Y., Wu, M., and Ding, W.:
Spartina alterniflora invasion drastically increases methane production
potential by shifting methanogenesis from hydrogenotrophic to methylotrophic
pathway in a coastal marsh, J. Ecol., 107,
2436–2450, <a href="https://doi.org/10.1111/1365-2745.13164" target="_blank">https://doi.org/10.1111/1365-2745.13164</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib90"><label>90</label><mixed-citation>
      
Zheng, J., RoyChowdhury, T., Yang, Z., Gu, B., Wullschleger, S. D., and Graham, D. E.: Impacts of temperature and soil characteristics on methane production and oxidation in Arctic tundra, Biogeosciences, 15, 6621–6635, <a href="https://doi.org/10.5194/bg-15-6621-2018" target="_blank">https://doi.org/10.5194/bg-15-6621-2018</a>, 2018.

    </mixed-citation></ref-html>--></article>
