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.

The threat of groundwater pollution for petrifying springs; defining nutrient threshold values for an endangered bryophyte dominated habitat.

Eutrophication by human activities is increasingly affecting ecosystem functioning and plant community composition. So far, studies mainly focus on the effects of atmospheric nitrogen deposition, surface water eutrophication or soil nutrient accumulation. Groundwater pollution of spring habitats, however, has received much less attention, although numerous papers report groundwater nutrient enrichment worldwide. This study presents a survey on groundwater pollution (with emphasis on nitrate and phosphate) and bryophyte composition in 51 ambient petrifying springs in 5 NW European countries, which were compared to published data from 173 other sites in 11 European countries. The reviewed dataset covers a broad range of unpolluted to heavily polluted springs with nitrate concentrations between 0.7 and 3227 µmol l-1. Most petrifying springs in the rural lowlands of NW Europe were found to have elevated concentrations of nitrate and phosphate with the most polluted springs occurring in The Netherlands. The cover of individual characteristic bryophyte species significantly correlates with groundwater nutrient concentrations indicating that nutrient pollution of spring waters affects bryophyte composition. Palustriella commutata, Eucladium verticillatum and Brachythecium rivulare prefer unpolluted petrifying springs whereas Cratoneuron filicinum and Pellia endiviifolia show a much broader tolerance to groundwater pollution. In order to sustain at least the basic conditions for the typical bryophyte composition of petrifying springs habitats, threshold values of 288 µmol (18 mg l-1) NO3- l-1 and 0.42 µmol (0.04 mg l-1) ortho-PO43- l-1 were defined. Data analysis of the spring water composition indicates that the main source for nutrient and nutrient induced base cation enrichment are nitrate losses from intensively used agricultural fields. The anthropogenically induced but regionally different chemical processes in subsoil and aquifers can result in different levels of nutrient pollution in springs. Further regulations for nitrate and phosphate application are required to conserve and restore groundwater fed ecosystems in Europe.

Trophic Transfer of Cd, Cu, Pb, Zn, P and Se in Dutch Storage Water Reservoirs.

Heavy metals are naturally omnipresent in aquatic systems. Excess amounts of heavy metals can accumulate in organisms of pollution impacted systems and transfer across a food web. Analysing the food web structure and metal contents of the organisms can help unravel the pathways of biomagnification or biodilution and gain insight in trophic linkages. We measured heavy metals and other elements in mussel bank detritus and organisms of the Biesbosch reservoirs (the Netherlands) and linked those to stable isotopic signatures. The heavy metal contents (cadmium, copper, lead, and zinc) were often lowest in benthivorous, omnivorous and piscivorous species (mainly fish); whereas, phosphorus contents were lower in the autotrophs. Mussel bank detritus contained the highest amounts of heavy metals. The heavy metals were negatively correlated with δ15N values. For selenium no clear trend was observed. Furthermore, there was a negative correlation between fish length and some heavy metals. Based on all 20 analysed elemental contents, similarities between species became apparent, related to niche or habitat. This study confirms that elemental contents of species can differ between feeding guilds and/or species, which can be attributed to metabolic and physiological processes. The organisms in higher trophic levels have adaptations preventing metal accumulation, resulting in lower contents. Within the fish species biodilution occurs, as most metal contents were lowest in bigger fish. Overall, the metals did not seem to biomagnify, but biodilute in the food web. Metal analyses combined with isotopic signatures could thus provide insights in metal transfer and possible trophic linkages within a system.

Lower nitrate leaching from dairy cattle slurry compared to synthetic fertilizer calcium ammonium nitrate applied to grassland.

Nitrate leaching from agriculture can be reduced by the choice of fertilizer and a proper timing of its application. For permanent grassland grown under temperate conditions, nitrate leaching was hypothesized to be lower from dairy cattle slurry (CS) compared to synthetic fertilizer calcium ammonium nitrate (CAN), based on differences in chemical composition, consequential effects on nitrogen (N) conversion processes in soil, and resulting differences in synchronization of (nitrate) N availability and plant N uptake. We tested the hypothesis in a two-year field experiment on cut grassland on a leaching-sensitive sandy soil, fertilized each year with 320 kg ha-1 of plant-available N from either 100% top-dressed CAN or a combination of 40% from CAN and 60% from sod-injected CS, and measured effects on grass herbage yield, herbage N uptake, and nitrate concentration in pore water at 1.0 m depth. Our results show a comparable level of herbage N uptake for both treatments, allowing for a proper comparison of nitrate leaching at a similar level of plant-available N. Average nitrate concentration in pore water in the main leaching period (over winter) was after the first (dry) growing season 44% lower for CS + CAN (41 mg l-1) compared to CAN only (73 mg l-1), and after the second (wet) growing season 35% lower for CS + CAN (32 mg l-1) compared to CAN only (49 mg l-1). Nitrogen application increased nitrate concentration at 1.0 m depth not only in winter but also in the growing season. We conclude that for permanent grasslands in temperate regions, nitrate leaching from timely applied CS may be considerably lower than from CAN, which is different from previous assumptions.

CO2, CH4, and N2O emissions from dredged material exposed to drying and zeolite addition under field and laboratory conditions.

Dredging, the removal of sediment from water courses, is generally conducted to maintain their navigability and to improve water quality. Recent studies indicate that dredging can significantly reduce aquatic greenhouse gas (GHG) emissions. These studies, however, do not consider the potential emission from the dredged material (sludge) in the depot. In addition, it is unknown if and how GHG emissions from sludge depots can be reduced. Here we present spatiotemporal variations of carbon dioxide (CO2), methane (CH4), and nitrous oxide (N2O) fluxes, as well as environmental variables from a sludge depot located in the Netherlands. Measurements were conducted monthly from the time the depot was filled until the sludge was dry and the depot was abolished. We also experimentally assessed the GHG mitigation potential of 1) keeping the sludge permanently inundated, and 2) the addition of different amounts of zeolite to increase sludge nitrogen binding capacity to reduce N2O emissions. In the depot and in the laboratory, a decrease in moisture content coincided with increased CO2 and N2O emissions while CH4 emissions decreased. We observed that permanent inundation reduced emissions (∼4 times less CO2-eq than in drying sludge). Adding zeolite lowered N2O fluxes from permanently inundated sludge but did not reduce total GHG emissions. During the depot's operational period, average CO2, CH4, and N2O fluxes were 5078, 27, and 5 mg m-2 d-1, respectively. GHG emissions from drying sludge occurred mainly in the form of CO2 (73% of the total CO2-eq emissions), with average GHG emission rates comparable to those reported for ditches and ponds. We estimate that approximately 14 tons of CO2-eq were emitted from the 0.011 km2 depot, which contained ∼20,000 m3 of sludge, during its entire operational period, and we argue that more studies are needed, considering different sludge origins, to expand our understanding of sludge depots.

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.

Nitrate-dependent anaerobic methane oxidation (N-DAMO) as a bioremediation strategy for waters affected by agricultural runoff.

Agricultural drainage ditches are subjected to high anthropogenic nitrogen input, leading to eutrophication and greenhouse gas emissions. Nitrate-dependent anaerobic methane oxidation (N-DAMO) could be a promising remediation strategy to remove methane (CH4) and nitrate (NO3-) simultaneously. Therefore, we aimed to evaluate the potential of N-DAMO to remove excess NO3- and decrease CH4 release from agricultural drainage ditches. Microcosm experiments were conducted using sediment and surface water collected from three different sites: a sandy-clay ditch (SCD), a freshwater-fed peatland ditch (FPD), and a brackish peatland ditch (BPD). The microcosms were inoculated with an N-DAMO enrichment culture dominated by Candidatus Methanoperedens and Candidatus Methylomirabilis and supplemented with 13CH4 and 15NO3-. A significant decrease in CH4 and NO3- concentration was only observed in the BPD sediment. In freshwater sediments (FPD and SCD), the effect of N-DAMO inoculation on CH4 and NO3- removal was negligible, likely because N-DAMO microorganisms were outcompeted by heterotrophic denitrifiers consuming NO3- much faster. Overall, our results suggest that bioaugmentation with N-DAMO might be a potential strategy for decreasing NO3- concentrations and CH4 emission in brackish ecosystems with increasing agricultural activities where the native microbial community is incapable of efficient denitrification.

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.

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.

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.

Nutrient dynamics of Sphagnum farming on rewetted bog grassland in NW Germany.

The agricultural use of drained peatlands leads to huge emissions of greenhouse gases and nutrients. A land-use alternative that allows rewetting of drained peatland while maintaining agricultural production is the cultivation of Sphagnum biomass as a renewable substitute for fossil peat in horticultural growing media (Sphagnum farming). We studied Sphagnum productivity and nutrient dynamics during two years in two Sphagnum farming sites in NW Germany, which were established on drained bog grassland by sod removal, rewetting, and the introduction of Sphagnum fragments in 2011 and 2016, respectively. We found a considerable and homogeneous production of Sphagnum biomass (>3.6 ton DW ha--1 yr-1), attributable to the high nutrient levels, low alkalinity, and even distribution of the irrigation water. The ammonium legacy from former drainage-based agriculture rapidly declined after rewetting, while nutrient mobilization was negligible. CH4 concentrations in the rewetted soil quickly decreased to very low levels. The Sphagnum biomass sequestered high loads of nutrients (46.0 and 47.4 kg N, 3.9 and 4.9 kg P, and 9.8 and 16.1 kg K ha-1 yr-1 in the 7.5 y and 2.5 y old sites, respectively), preventing off-site eutrophication. We conclude that Sphagnum farming as an alternative for drainage-based peatland agriculture may contribute effectively to tackling environmental challenges such as local and regional downstream pollution and global climate change.

Effects of guanotrophication and warming on the abundance of green algae, cyanobacteria and microcystins in Lake Lesser Prespa, Greece.

Lake Lesser Prespa in Greece is a vital breeding habitat for the Dalmatian and Great White Pelican and a shelter for numerous rare and endemic species. However, eutrophication processes are distressing the lake system and the outbreaks of cyanobacterial blooms during the warm months may pose a threat to aquatic organisms due to the presence of microcystins (MCs). In this study we hypothesize that nutrients (eutrophication), nutrient-rich pelican droppings (guanotrophication) and warming (climate change) can affect the algal growth and MCs production in the water layer of Lake Lesser Prespa. Seston collected from three lake sites was incubated at ambient (20°C) and high (30°C) temperature with or without the addition of nutrients (nitrogen (N), phosphorus (P)), or pelican droppings. Results showed increased chlorophyll-a at higher temperature (30°C). N addition yielded higher chlorophyll-a levels than the non-treated water or when only P was added. The addition of both N and P as well as the addition of pelican dropping resulted in the highest chlorophyll-a at both temperatures. Notably, in the dropping-treatments, cyanobacteria and MCs were promoted while changes were evoked in the relative contribution of toxic MC-variants. Guanotrophication may thus influence the cyanobacterial dynamics and most likely their toxicity profile at Lesser Prespa.

Hidden treasures: Human-made aquatic ecosystems harbour unexplored opportunities.

Artificial water bodies like ditches, fish ponds, weirs, reservoirs, fish ladders, and irrigation channels are usually constructed and managed to optimize their intended purposes. However, human-made aquatic systems also have unintended consequences on ecosystem services and biogeochemical cycles. Knowledge about their functioning and possible additional ecosystem services is poor, especially compared to natural ecosystems. A GIS analysis indicates that currently only ~ 10% of European surface waters are covered by the European Water Framework directive, and that a considerable fraction of the excluded systems are likely human-made aquatic systems. There is a clear mismatch between the high possible significance of human-made water bodies and their low representation in scientific research and policy. We propose a research agenda to build an inventory of human-made aquatic ecosystems, support and advance research to further our understanding of the role of these systems in local and global biogeochemical cycles as well as to identify other benefits for society. We stress the need for studies that aim to optimize management of human-made aquatic systems considering all their functions and to support programs designed to overcome barriers of the adoption of optimized management strategies.

The Impact of Warming and Nutrients on Algae Production and Microcystins in Seston from the Iconic Lake Lesser Prespa, Greece.

Lake Lesser Prespa and its adjacent pond, Vromolimni in Greece, is a shallow freshwater system and a highly protected area hosting an exceptional biodiversity. The occurrence of microcystins (MCs) producing cyanobacterial blooms in these waters during recent years can be harmful to the wildlife. We tested the hypothesis that both cyanobacterial biomass and MCs are strongly influenced by nutrients (eutrophication) and warming (climate change). Lake and pond water was collected from two sites in each water body in 2013 and incubated at three temperatures (20 °C, 25 °C, 30 °C) with or without additional nutrients (nitrogen +N, phosphorus +P and both +N and +P). Based on both biovolume and chlorophyll-a concentrations, cyanobacteria in water from Lesser Prespa were promoted primarily by combined N and P additions and to a lesser extent by N alone. Warming seemed to yield more cyanobacteria biomass in these treatments. In water from Vromolimni, both N alone and N+P additions increased cyanobacteria and a warming effect was hardly discernible. MC concentrations were strongly increased by N and N+P additions in water from all four sites, which also promoted the more toxic variant MC-LR. Hence, both water bodies seem particularly vulnerable to further N-loading enhancing MC related risks.

Azolla along a phosphorus gradient: biphasic growth response linked to diazotroph traits and phosphorus-induced iron chlorosis.

Azolla spp., a water fern often used for phytoremediation, is a strong phosphorus (P) accumulator due to its high growth rate and N2 fixing symbionts (diazotrophs). It is known that plant growth is stimulated by P, but the nature of the interactive response of both symbionts along a P gradient, and related changes in growth-limiting factors, are unclear. We determined growth, and N and P sequestration rates of Azolla filiculoides in N-free water at different P concentrations. The growth response appeared to be biphasic and highest at levels ≥10 P µmol l-1. Diazotrophic N sequestration increased upon P addition, and rates were three times higher at high P than at low P. At 10 µmol P l-1, N sequestration rates reached its maximum and A. filiculoides growth became saturated. Due to luxury consumption, P sequestration rates increased until 50 µmol P l-1. At higher P concentrations (≥50 µmol l-1), however, chlorosis occurred that seems to be caused by iron- (Fe-), and not by N-deficiency. We demonstrate that traits of the complete symbiosis in relation to P and Fe availability determine plant performance, stressing the role of nutrient stoichiometry. The results are discussed regarding Azolla's potential use in a bio-based economy.

A keystone mutualism underpins resilience of a coastal ecosystem to drought.

Droughts are increasing in severity and frequency, yet the mechanisms that strengthen ecosystem resilience to this stress remain poorly understood. Here, we test whether positive interactions in the form of a mutualism between mussels and dominant cordgrass in salt marshes enhance ecosystem resistance to and recovery from drought. Surveys spanning 250 km of southeastern US coastline reveal spatially dispersed mussel mounds increased cordgrass survival during severe drought by 5- to 25-times. Surveys and mussel addition experiments indicate this positive effect of mussels on cordgrass was due to mounds enhancing water storage and reducing soil salinity stress. Observations and models then demonstrate that surviving cordgrass patches associated with mussels function as nuclei for vegetative re-growth and, despite covering only 0.1-12% of die-offs, markedly shorten marsh recovery periods. These results indicate that mutualisms, in supporting stress-resistant patches, can play a disproportionately large, keystone role in enhancing ecosystem resilience to climatic extremes.

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

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

Drought, Mutualism Breakdown, and Landscape-Scale Degradation of Seagrass Beds.

In many marine ecosystems, biodiversity critically depends on foundation species such as corals and seagrasses that engage in mutualistic interactions [1-3]. Concerns grow that environmental disruption of marine mutualisms exacerbates ecosystem degradation, with breakdown of the obligate coral mutualism ("coral bleaching") being an iconic example [2, 4, 5]. However, as these mutualisms are mostly facultative rather than obligate, it remains unclear whether mutualism breakdown is a common risk in marine ecosystems, and thus a potential accelerator of ecosystem degradation. Here, we provide evidence that drought triggered landscape-scale seagrass degradation and show the consequent failure of a facultative mutualistic feedback between seagrass and sulfide-consuming lucinid bivalves that in turn appeared to exacerbate the observed collapse. Local climate and remote sensing analyses revealed seagrass collapse after a summer with intense low-tide drought stress. Potential analysis-a novel approach to detect feedback-mediated state shifts-revealed two attractors (healthy and degraded states) during the collapse, suggesting that the drought disrupted internal feedbacks to cause abrupt, patch-wise degradation. Field measurements comparing degraded patches that were healthy before the collapse with patches that remained healthy demonstrated that bivalves declined dramatically in degrading patches with associated high sediment sulfide concentrations, confirming the breakdown of the mutualistic seagrass-lucinid feedback. Our findings indicate that drought triggered mutualism breakdown, resulting in toxic sulfide concentrations that aggravated seagrass degradation. We conclude that external disturbances can cause sudden breakdown of facultative marine mutualistic feedbacks. As this may amplify ecosystem degradation, we suggest including mutualisms in marine conservation and restoration approaches.