More is not always better: peat moss mixtures slightly enhance peatland stability.

Terrestrial wetland ecosystems challenge biodiversity-ecosystem function theory, which generally links high species diversity to stable ecosystem functions. An open question in ecosystem ecology is whether assemblages of co-occurring peat mosses contribute to the stability of peatland ecosystem processes. We conducted a two-species (Sphagnum cuspidatum, Sphagnum medium) replacement series mesocosm experiment to evaluate the resistance, resilience, and recovery rates of net ecosystem CO2 exchange (NEE) under mild and deep water table drawdown. Our results show a positive effect of mild water table drawdown on NEE with no apparent role for peat moss mixture. Our study indicates that the carbon uptake capacity by peat moss mixtures is rather resilient to mild water table drawdown, but seriously affected by deeper drought conditions. Co-occurring peat moss species seem to enhance the resilience of the carbon uptake function (i.e. ability of NEE to return to pre-perturbation levels) of peat moss mixtures only slightly. These findings suggest that assemblages of co-occurring Sphagnum mosses do only marginally contribute to the stability of ecosystem functions in peatlands under drought conditions. Above all, our results highlight that predicted severe droughts can gravely affect the sink capacity of peatlands, with only a small extenuating role for peat moss mixtures.

Polishing wastewater effluent using plants: floating plants perform better than submerged plants in both nutrient removal and reduction of greenhouse gas emission.

While research on aquatic plants used in treatment wetlands is abundant, little is known about the use of plants in hydroponic ecological wastewater treatment, and its simultaneous effect on greenhouse gas (GHG) emissions. Here, we assess the effectiveness of floating and submerged plants in removing nutrients and preventing GHG emissions from wastewater effluent. We grew two species of floating plants, Azolla filiculoides and Lemna minor, and two species of submerged plants, Ceratophyllum demersum and Callitriche platycarpa, on a batch of domestic wastewater effluent without any solid substrate. In these systems, we monitored nitrogen and phosphorus removal and fluxes of CO2, CH4 and N2O, for 2 weeks. In general, floating plants produced the most biomass, whereas submerged plants were rapidly overgrown by filamentous algae. Floating plants removed nutrients most efficiently; both floating species removed 100% of the phosphate while Lemna also removed 97-100% of the inorganic nitrogen, as opposed to a removal of 81-88% in submerged plants with algae treatments. Moreover, aquaria covered by floating plants had roughly three times higher GHG uptake than the treatments with submerged plants or controls without plants. Thus, effluent polishing by floating plants can be a promising avenue for climate-smart wastewater polishing.

Overcoming establishment thresholds for peat mosses in human-made bog pools.

Globally, peatlands have been affected by drainage and peat extraction, with adverse effects on their functioning and services. To restore peat-forming vegetation, drained bogs are being rewetted on a large scale. Although this practice results in higher groundwater levels, unfortunately it often creates deep lakes in parts where peat was extracted to greater depths than the surroundings. Revegetation of these deeper waters by peat mosses appears to be challenging due to strong abiotic feedbacks that keep these systems in an undesired bare state. In this study, we theoretically explore if a floating peat mat and an open human-made bog lake can be considered two alternative stable states using a simple model, and experimentally test in the field whether stable states are present, and whether a state shift can be accomplished using floating biodegradable structures that mimic buoyant peat. We transplanted two peat moss species into these structures (pioneer sp. Sphagnum cuspidatum and later-successional sp. S. palustre) with and without additional organic substrate. Our model suggests that these open human-made bog lakes and floating peat mats can indeed be regarded as alternative stable states. Natural recovery by spontaneous peat moss growth, i.e., a state shift from open water to floating mats, is only possible when the water table is sufficiently shallow to avoid light limitation (<0.29 m at our site). Our experiment revealed that alternative stable states are present and that the floating structures facilitated the growth of pioneer S. cuspidatum and vascular plants. Organic substrate addition particularly facilitated vascular plant growth, which correlated to higher moss height. The structures remained too wet for the late-successional species S. palustre. We conclude that open water and floating peat mats in human-made bog lakes can be considered two alternative stable states, and that temporary floating establishment structures can induce a state shift from the open water state to peat-forming vegetation state. These findings imply that for successful restoration, there is a clear water depth threshold to enable peat moss growth and there is no need for addition of large amounts of donor-peat substrate. Correct species selection for restoration is crucial for success.

Resilience of beach grasses along a biogeomorphic successive gradient: resource availability vs. clonal integration.

Coastal ecosystems are often formed through two-way interactions between plants and their physical landscape. By expanding clonally, landscape-forming plants can colonize bare unmodified environments and stimulate vegetation-landform feedback interactions. Yet, to what degree these plants rely on clonal integration for overcoming physical stress during biogeomorphological succession remains unknown. Here, we investigated the importance of clonal integration and resource availability on the resilience of two European beach grasses (i.e. Elytrigia juncea and Ammophila arenaria) over a natural biogeomorphic dune gradient from beach (unmodified system) to foredune (biologically modified system). We found plant resilience, as measured by its ability to recover and expand following disturbance (i.e. plant clipping), to be independent on the presence of rhizomal connections between plant parts. Instead, resource availability over the gradient largely determined plant resilience. The pioneer species, Elytrigia, demonstrated a high resilience to physical stress, independent of its position on the biogeomorphic gradient (beach or embryonic dune). In contrast, the later successional species (Ammophila) proved to be highly resilient on the lower end of its distribution (embryonic dune), but it did not fully recover on the foredunes, most likely as a result of nutrient deprivation. We argue that in homogenously resource-poor environments as our beach system, overall resource availability, instead of translocation through a clonal network, determines the resilience of plant species. Hence, the formation of high coastal dunes may increase the resistance of beach grasses to the physical stresses of coastal flooding, but the reduced marine nutrient input may negatively affect the resilience of plants.

Soils in lakes: the impact of inundation and storage on surface water quality.

The large-scale storage and inundation of contaminated soils and sediments in deep waterlogged former sand pits or in lakes have become a fairly common practice in recent years. Decreasing water depth potentially promotes aquatic biodiversity, but it also poses a risk to water quality as was shown in a previous study on the impact on groundwater. To provide in the urgent need for practical and robust risk indicators for the storage of terrestrial soils in surface waters, the redistribution of metals and nutrients was studied in long-term mesocosm experiments. For a range of surface water turbidity (suspended matter concentrations ranging from 0 to 3000 mg/L), both chemical partitioning and toxicity of pollutants were tested for five distinctly different soils. Increasing turbidity in surface water showed only marginal response on concentrations of heavy metals, phosphorus (P) and nitrogen (N). Toxicity testing with bioluminescent bacteria, and biotic ligand modelling (BLM), indicated no or only minor risk of metals in the aerobic surface water during aerobic mixing under turbid conditions. Subsequent sedimentation of the suspended matter revealed the chemical speciation and transport of heavy metals and nutrients over the aerobic and anaerobic interface. Although negative fluxes occur for Cd and Cu, most soils show release of pollutants from sediment to surface waters. Large differences in fluxes occur for PO4, SO4, B, Cr, Fe, Li, Mn and Mo between soils. For an indicator of aerobic chemical availability, dilute nitric acid extraction (0.43 M HNO3; Aqua nitrosa) performed better than the conventional Aqua regia destruction. Both the equilibrium concentrations in surface waters, and fluxes from sediment, were adequately (r2 = 0.81) estimated by a 1 mM CaCl2 soil extraction procedure. This study has shown that the combination of 0.43 M HNO3 and 1 mM CaCl2 extraction procedures can be used to adequately estimate emissions from sediment to surface waters, and assess potential water quality changes, when former sand pits are being filled with soil materials.

The Water Hyacinth Microbiome: Link Between Carbon Turnover and Nutrient Cycling.

Water hyacinth (WH), a large floating plant, plays an important role in the biogeochemistry and ecology of many freshwaters globally. Its biogeochemical impact on wetland functioning is strongly mediated by the microbiome associated with its roots. However, little is known about the structure and function of this WH rhizobiome and its relation to wetland ecosystem functioning. Here, we unveil the core and transient rhizobiomes of WH and their key biogeochemical functions in two of the world's largest wetlands: the Amazon and the Pantanal. WH hosts a highly diverse microbial community shaped by spatiotemporal changes. Proteobacteria lineages were most common, followed by Actinobacteria and Planctomycetes. Deltaproteobacteria and Sphingobacteriia predominated in the core microbiome, potentially associated with polysaccharide degradation and fermentation of plant-derived carbon. Conversely, a plethora of lineages were transient, including highly abundant Acinetobacter, Acidobacteria subgroup 6, and methanotrophs, thus assuring diverse taxonomic signatures in the two different wetlands. Our findings point out that methanogenesis is a key driver of, and proxy for, community structure, especially during seasonal plant decline. We provide ecologically relevant insights into the WH microbiome, which is a key element linking plant-associated carbon turnover with other biogeochemical fluxes in tropical wetlands.

Foundation species' overlap enhances biodiversity and multifunctionality from the patch to landscape scale in southeastern United States salt marshes.

Although there is mounting evidence that biodiversity is an important and widespread driver of ecosystem multifunctionality, much of this research has focused on small-scale biodiversity manipulations. Hence, which mechanisms maintain patches of enhanced biodiversity in natural systems and if these patches elevate ecosystem multifunctionality at both local and landscape scales remain outstanding questions. In a 17 month experiment conducted within southeastern United States salt marshes, we found that patches of enhanced biodiversity and multifunctionality arise only where habitat-forming foundation species overlap--i.e. where aggregations of ribbed mussels (Geukensia demissa) form around cordgrass (Spartina alterniflora) stems. By empirically scaling up our experimental results to the marsh platform at 12 sites, we further show that mussels--despite covering only approximately 1% of the marsh surface--strongly enhance five distinct ecosystem functions, including decomposition, primary production and water infiltration rate, at the landscape scale. Thus, mussels create conditions that support the co-occurrence of high densities of functionally distinct organisms within cordgrass and, in doing so, elevate salt marsh multifunctionality from the patch to landscape scale. Collectively, these findings suggest that patterns in foundation species' overlap drive variation in biodiversity and ecosystem functioning within and across natural ecosystems.We therefore argue that foundation species should be integrated in our conceptual understanding of forces that moderate biodiversity--ecosystem functioning relationships, approaches for conserving species diversity and strategies to improve the multifunctionality of degraded ecosystems.

Habitat collapse due to overgrazing threatens turtle conservation in marine protected areas.

Marine protected areas (MPAs) are key tools for combatting the global overexploitation of endangered species. The prevailing paradigm is that MPAs are beneficial in helping to restore ecosystems to more 'natural' conditions. However, MPAs may have unintended negative effects when increasing densities of protected species exert destructive effects on their habitat. Here, we report on severe seagrass degradation in a decade-old MPA where hyper-abundant green turtles adopted a previously undescribed below-ground foraging strategy. By digging for and consuming rhizomes and roots, turtles create abundant bare gaps, thereby enhancing erosion and reducing seagrass regrowth. A fully parametrized model reveals that the ecosystem is approaching a tipping point, where consumption overwhelms regrowth, which could potentially lead to complete collapse of the seagrass habitat. Seagrass recovery will not ensue unless turtle density is reduced to nearly zero, eliminating the MPA's value as a turtle reserve. Our results reveal an unrecognized, yet imminent threat to MPAs, as sea turtle densities are increasing at major nesting sites and the decline of seagrass habitat forces turtles to concentrate on the remaining meadows inside reserves. This emphasizes the need for policy and management approaches that consider the interactions of protected species with their habitat.

Addition of iron does not ameliorate sulfide toxicity by sargassum influx to mangroves but dampens methane and nitrous oxide emissions.

Sargassum spp. strandings in the tropical Atlantic harm local ecosystems due to toxic sulfide levels. We conducted a mesocosm experiment to test the efficacy of iron(III) (hydr)oxides in (a) mitigating sulfide toxicity in mangroves resulting from Sargassum and (b) reducing potentially enhanced greenhouse gas emissions. Our results show that iron addition failed to prevent mangrove mortality caused by highly toxic sulfide concentrations, which reached up to 15,000 µmol l-1 in 14 days; timely removal may potentially prevent mangrove death. Sargassum-impacted mesocosms significantly increased methane, nitrous oxide, and carbon dioxide emissions, producing approximately 1 g CO2-equivalents m-2 h-1 during daylight hours, thereby shifting mangroves from sinks to sources of greenhouse gasses. However, iron addition decreased methane emissions by 62 % and nitrous oxide emissions by 57 %. This research reveals that Sargassum strandings have multiple adverse effects related to chemical and ecological dynamics in mangrove ecosystems, including greenhouse gas emissions.

Sludge degradation, nutrient removal and reduction of greenhouse gas emission by a Chironomus-Azolla wastewater treatment cascade.

Wastewater treatment plants (WWTPs) are a point source of nutrients, emit greenhouse gases (GHGs), and produce large volumes of excess sludge. The use of aquatic organisms may be an alternative to the technical post-treatment of WWTP effluent, as they play an important role in nutrient dynamics and carbon balance in natural ecosystems. The aim of this study was therefore to assess the performance of an experimental wastewater-treatment cascade of bioturbating macroinvertebrates and floating plants in terms of sludge degradation, nutrient removal and lowering GHG emission. To this end, a full-factorial experiment was designed, using a recirculating cascade with a WWTP sludge compartment with or without bioturbating Chironomus riparius larvae, and an effluent container with or without the floating plant Azolla filiculoides, resulting in four treatments. To calculate the nitrogen (N), phosphorus (P) and carbon (C) mass balance of this system, the N, P and C concentrations in the effluent, biomass production, and sludge degradation, as well as the N, P and C content of all compartments in the cascade were measured during the 26-day experiment. The presence of Chironomus led to an increased sludge degradation of 44% compared to 25% in the control, a 1.4 times decreased transport of P from the sludge and a 2.4 times increased transport of N out of the sludge, either into Chironomus biomass or into the water column. Furthermore, Chironomus activity decreased methane emissions by 92%. The presence of Azolla resulted in a 15% lower P concentration in the effluent than in the control treatment, and a CO2 uptake of 1.13 kg ha-1 day-1. These additive effects of Chironomus and Azolla resulted in an almost two times higher sludge degradation, and an almost two times lower P concentration in the effluent. This is the first study that shows that a bio-based cascade can strongly reduce GHG and P emissions simultaneously during the combined polishing of wastewater sludge and effluent, benefitting from the additive effects of the presence of both macrophytes and invertebrates. In addition to the microbial based treatment steps already employed on WWTPs, the integration of higher organisms in the treatment process expands the WWTP based ecosystem and allows for the inclusion of macroinvertebrate and macrophyte mediated processes. Applying macroinvertebrate-plant cascades may therefore be a promising tool to tackle the present and future challenges of WWTPs.

Habitat complexity drives food web structure along a dynamic mangrove coast.

Structurally complex habitats, such as mangrove forests, allow for rich assemblages of species that benefit from the provided space, volume and substrate. Changes in habitat complexity can affect species abundance, diversity and resilience. In this study, we explored the effects of habitat complexity on food web networks in four developmental stages of mangrove forests with differing structural complexities: climax > degrading > colonizing > bare, by analyzing food web structure, stable isotopes and habitat complexity. We found that food webs became gradually more biodiverse (species richness: +119 %), complex (link density: +39 %), and robust (connectance: -35 %) in climax versus bare stages with increasing complexity of the mangrove forest (i.e., number of trees, leaf cover, and pneumatophore densities). This study shows that habitat complexity drives food web network structure in dynamic mangrove forests. We recommend restoration practitioners to use this food web network approach to quantify habitat restoration successes complementary to traditional biodiversity metrics.

Restoring gradual land-water transitions in a shallow lake improved phytoplankton quantity and quality with cascading effects on zooplankton production.

Land-water transition areas play a significant role in the functioning of aquatic ecosystems. However, anthropogenic pressures are posing severe threats on land-water transition areas, which leads to degradation of the ecological integrity of many lakes worldwide. Enhancing habitat complexity and heterogeneity by restoring land-water transition areas in lake systems is deemed a suitable method to restore lakes bottom-up by stimulating lower trophic levels. Stimulating productivity of lower trophic levels (phytoplankton, zooplankton) generates important food sources for declining higher trophic levels (fish, birds). Here, we study ecosystem restoration project Marker Wadden in Lake Markermeer, The Netherlands. This project involved the construction of a 700-ha archipelago of five islands in a degrading shallow lake, aiming to create additional sheltered land-water transition areas to stimulate food web development from its base by improving phytoplankton quantity and quality. We found that phytoplankton quantity (chlorophyll-a concentration) and quality (inversed carbon:nutrient ratio) in the shallow waters inside the Marker Wadden archipelago were significantly improved, likely due to higher nutrient availabilities, while light availability remained sufficient, compared to the surrounding lake. Higher phytoplankton quantity and quality was positively correlated with zooplankton biomass, which was higher inside the archipelago than in the surrounding lake due to improved trophic transfer efficiency between phytoplankton and zooplankton. We conclude that creating new land-water transition areas can be used to increase light and nutrient availabilities and thereby enhancing primary productivity, which in turn can stimulate higher trophic levels in degrading aquatic ecosystems.

Wetscapes: Restoring and maintaining peatland landscapes for sustainable futures.

Peatlands are among the world's most carbon-dense ecosystems and hotspots of carbon storage. Although peatland drainage causes strong carbon emissions, land subsidence, fires and biodiversity loss, drainage-based agriculture and forestry on peatland is still expanding on a global scale. To maintain and restore their vital carbon sequestration and storage function and to reach the goals of the Paris Agreement, rewetting and restoration of all drained and degraded peatlands is urgently required. However, socio-economic conditions and hydrological constraints hitherto prevent rewetting and restoration on large scale, which calls for rethinking landscape use. We here argue that creating integrated wetscapes (wet peatland landscapes), including nature preserve cores, buffer zones and paludiculture areas (for wet productive land use), will enable sustainable and complementary land-use functions on the landscape level. As such, transforming landscapes into wetscapes presents an inevitable, novel, ecologically and socio-economically sound alternative for drainage-based peatland use.

Recovering wetland biogeomorphic feedbacks to restore the world's biotic carbon hotspots.

Biogeomorphic wetlands cover 1% of Earth's surface but store 20% of ecosystem organic carbon. This disproportional share is fueled by high carbon sequestration rates and effective storage in peatlands, mangroves, salt marshes, and seagrass meadows, which greatly exceed those of oceanic and forest ecosystems. Here, we review how feedbacks between geomorphology and landscape-building vegetation underlie these qualities and how feedback disruption can switch wetlands from carbon sinks into sources. Currently, human activities are driving rapid declines in the area of major carbon-storing wetlands (1% annually). Our findings highlight the urgency to stop through conservation ongoing losses and to reestablish landscape-forming feedbacks through restoration innovations that recover the role of biogeomorphic wetlands as the world's biotic carbon hotspots.

Temperature response of aquatic greenhouse gas emissions differs between dominant plant types.

Greenhouse gas (GHG) emissions from small inland waters are disproportionately large. Climate warming is expected to favor dominance of algae and free-floating plants at the expense of submerged plants. Through different routes these functional plant types may have far-reaching impacts on freshwater GHG emissions in future warmer waters, which are yet unknown. We conducted a 1,000 L mesocosm experiment testing the effects of plant type and warming on GHG emissions from temperate inland waters dominated by either algae, free-floating or submerged plants in controls and warmed (+4 °C) treatments for one year each. Our results show that the effect of experimental warming on GHG fluxes differs between dominance of different functional plant types, mainly by modulating methane ebullition, an often-dominant GHG emission pathway. Specifically, we demonstrate that the response to experimental warming was strongest for free-floating and lowest for submerged plant-dominated systems. Importantly, our results suggest that anticipated shifts in plant type from submerged plants to a dominance of algae or free-floating plants with warming may increase total GHG emissions from shallow waters. This, together with a warming-induced emission response, represents a so far overlooked positive climate feedback. Management strategies aimed at favouring submerged plant dominance may thus substantially mitigate GHG emissions.

Methanotrophic symbionts provide carbon for photosynthesis in peat bogs.

Wetlands are the largest natural source of atmospheric methane, the second most important greenhouse gas. Methane flux to the atmosphere depends strongly on the climate; however, by far the largest part of the methane formed in wetland ecosystems is recycled and does not reach the atmosphere. The biogeochemical controls on the efficient oxidation of methane are still poorly understood. Here we show that submerged Sphagnum mosses, the dominant plants in some of these habitats, consume methane through symbiosis with partly endophytic methanotrophic bacteria, leading to highly effective in situ methane recycling. Molecular probes revealed the presence of the bacteria in the hyaline cells of the plant and on stem leaves. Incubation with (13)C-methane showed rapid in situ oxidation by these bacteria to carbon dioxide, which was subsequently fixed by Sphagnum, as shown by incorporation of (13)C-methane into plant sterols. In this way, methane acts as a significant (10-15%) carbon source for Sphagnum. The symbiosis explains both the efficient recycling of methane and the high organic carbon burial in these wetland ecosystems.

Warming and eutrophication interactively drive changes in the methane-oxidizing community of shallow lakes.

Freshwater ecosystems are the largest natural source of the greenhouse gas methane (CH4), with shallow lakes a particular hot spot. Eutrophication and warming generally increase lake CH4 emissions but their impacts on the sole biological methane sink-methane oxidation-and methane-oxidizer community dynamics are poorly understood. We used the world's longest-running freshwater climate-change mesocosm experiment to determine how methane-oxidizing bacterial (MOB) abundance and composition, and methane oxidation potential in the sediment respond to eutrophication, short-term nitrogen addition and warming. After nitrogen addition, MOB abundance and methane oxidation potential increased, while warming increased MOB abundance without altering methane oxidation potential. MOB community composition was driven by both temperature and nutrient availability. Eutrophication increased relative abundance of type I MOB Methyloparacoccus. Warming favoured type II MOB Methylocystis over type I MOB Methylomonadaceae, shifting the MOB community from type I dominance to type I and II co-dominance, thereby altering MOB community traits involved in growth and stress-responses. This shift to slower-growing MOB may explain why higher MOB abundance in warmed mesocosms did not coincide with higher methane oxidation potential. Overall, we show that eutrophication and warming differentially change the MOB community, resulting in an altered ability to mitigate CH4 emissions from shallow lakes.

A Novel Laboratory-Scale Mesocosm Setup to Study Methane Emission Mitigation by Sphagnum Mosses and Associated Methanotrophs.

Degraded peatlands are often rewetted to prevent oxidation of the peat, which reduces CO2 emission. However, the created anoxic conditions will boost methane (CH4) production and thus emission. Here, we show that submerged Sphagnum peat mosses in rewetted-submerged peatlands can reduce CH4 emission from peatlands with 93%. We were able to mimic the field situation in the laboratory by using a novel mesocosm set-up. By combining these with 16S rRNA gene amplicon sequencing and qPCR analysis of the pmoA and mmoX genes, we showed that submerged Sphagnum mosses act as a niche for CH4 oxidizing bacteria. The tight association between Sphagnum peat mosses and methane oxidizing bacteria (MOB) significantly reduces CH4 emissions by peatlands and can be studied in more detail in the mesocosm setup developed in this study.

Variation in symbiotic N2 fixation rates among Sphagnum mosses.

Biological nitrogen (N) fixation is an important process supporting primary production in ecosystems, especially in those where N availability is limiting growth, such as peatlands and boreal forests. In many peatlands, peat mosses (genus Sphagnum) are the prime ecosystem engineers, and like feather mosses in boreal forests, they are associated with a diverse community of diazotrophs (N2-fixing microorganisms) that live in and on their tissue. The large variation in N2 fixation rates reported in literature remains, however, to be explained. To assess the potential roles of habitat (including nutrient concentration) and species traits (in particular litter decomposability and photosynthetic capacity) on the variability in N2 fixation rates, we compared rates associated with various Sphagnum moss species in a bog, the surrounding forest and a fen in Sweden. We found appreciable variation in N2 fixation rates among moss species and habitats, and showed that both species and habitat conditions strongly influenced N2 fixation. We here show that higher decomposition rates, as explained by lower levels of decomposition-inhibiting compounds, and higher phosphorous (P) levels, are related with higher diazotrophic activity. Combining our findings with those of other studies, we propose a conceptual model in which both species-specific traits of mosses (as related to the trade-off between rapid photosynthesis and resistance to decomposition) and P availability, explain N2 fixation rates. This is expected to result in a tight coupling between P and N cycling in peatlands.

Contribution of plant-induced pressurized flow to CH4 emission from a Phragmites fen.

The widespread wetland species Phragmites australis (Cav.) Trin. ex Steud. has the ability to transport gases through its stems via a pressurized flow. This results in a high oxygen (O2) transport to the rhizosphere, suppressing methane (CH4) production and stimulating CH4 oxidation. Simultaneously CH4 is transported in the opposite direction to the atmosphere, bypassing the oxic surface layer. This raises the question how this plant-mediated gas transport in Phragmites affects the net CH4 emission. A field experiment was set-up in a Phragmites-dominated fen in Germany, to determine the contribution of all three gas transport pathways (plant-mediated, diffusive and ebullition) during the growth stage of Phragmites from intact vegetation (control), from clipped stems (CR) to exclude the pressurized flow, and from clipped and sealed stems (CSR) to exclude any plant-transport. Clipping resulted in a 60% reduced diffusive + plant-mediated flux (control: 517, CR: 217, CSR: 279 mg CH4 m-2 day-1). Simultaneously, ebullition strongly increased by a factor of 7-13 (control: 10, CR: 71, CSR: 126 mg CH4 m-2 day-1). This increase of ebullition did, however, not compensate for the exclusion of pressurized flow. Total CH4 emission from the control was 2.3 and 1.3 times higher than from CR and CSR respectively, demonstrating the significant role of pressurized gas transport in Phragmites-stands.