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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., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Enhanced soil ammonium (NH4+) concentrations in wetlands often lead to graminoid dominance, but species composition is highly variable. Although NH4+ is readily taken up as a nutrient, several wetland species are known to be sensitive to high NH4+ concentrations or even suffer toxicity, particularly at low soil pH. More knowledge about differential graminoid responses to high NH4+ availability in relation to soil pH can help to better understand vegetation changes. The responses of two wetland graminoids, Juncus acutiflorus and Carex disticha, to high (2 mmol·l(-1) ) versus control (20 μmol·l(-1) ) NH4+ concentrations were tested in a controlled hydroponic set up, at two pH values (4 and 6). A high NH4+ concentration did not change total biomass for these species at either pH, but increased C allocation to shoots and increased P uptake, leading to K and Ca limitation, depending on pH treatment. More than 50% of N taken up by C. disticha was invested in N-rich amino acids with decreasing C:N ratio, but only 10% for J. acutiflorus. Although both species appeared to be well adapted to high NH4+ loadings in the short term, C. disticha showed higher classic detoxifying responses that are early warning indicators for decreased tolerance in the long term. In general, the efficient aboveground biomass allocation, P uptake and N detoxification explain the competitive strength of wetland graminoids at the expense of overall biodiversity at high NH4+ loading. In addition, differential responses to enhanced NH4+ affect interspecific competition among graminoids and lead to a shift in vegetation composition., (© 2015 German Botanical Society and The Royal Botanical Society of the Netherlands.)

In regions with intensive agriculture, water level fluctuation in wetlands has generally become constricted within narrow limits. Water authorities are, however, considering the re-establishment of fluctuating water levels as a management tool in biodiverse, base-rich fens ('rich fens'). This includes temporary inundation with surface water from ditches, which may play an important role in counteracting acidification in order to conserve and restore biodiversity. Inundation may result in an increased acid neutralizing capacity (ANC) for two reasons: infiltration of base-rich inundation water into peat soils, and microbial alkalinity generation under anaerobic conditions. The main objectives of this study were to test whether short-term (2 weeks) summer inundation is more effective than short-term winter inundation to restore the ANC in the upper 10 cm of non-floating peat soils, and to explain potential differences. Large-scale field experiments were conducted for five years in base-rich fens and Sphagnum-dominated poor fens. Winter inundation did not result in increased porewater ANC, because infiltration was inhibited in the waterlogged peat and evapotranspiration rates were relatively low. Also, low temperatures limit microbial alkalinity generation. In summer, however, when temperature and evapotranspiration rates are higher, inundation resulted in increased porewater Ca and HCO3- concentrations, but only in areas with characteristic rich fen bryophytes. This increase was not only due to stronger infiltration into the soil, but also to higher microbial alkalinity generation under anaerobic conditions. In contrast, porewater ANC did not increase in Sphagnum-plots as a result of the ability of Sphagnum spp. to acidify their environment. In both rich and poor fens, flooding-induced P-mobilization remained sufficiently low to safeguard P-limited vegetation. NO3(-) and NH4(+) dynamics showed no considerable changes either. In conclusion, short-term summer inundation with base-rich and nutrient-poor surface water is considered beneficial in the management of non-floating rich fens, and much more effective than winter inundation.

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.

Structural diversity formed by dense, floating Stratiotes aloides stands, generates hotspots of biodiversity of flora and fauna in wetlands. However, only part of the populations become emergent and provide this important facilitation. Since it has been hypothesised that its buoyancy depends on the rates of underwater photosynthesis, we investigated the role of dissolved CO2 availability and PAR on photosynthesis, biomass production and buoyancy in a controlled greenhouse experiment. Photosynthesis and growth were strongly influenced by both PAR and CO2 availability. At low PAR, plants formed less biomass and produced no emergent leaves, even when CO2 was abundant. At low CO2 levels, S. aloides switched to HCO3- use, resulting in a lower photosynthetic O2 production, decreased emergent leaf formation and increased CaCO3 precipitation on its leaves, all of which impaired buoyancy. At high PAR, low CO2 availability resulted in slower colonisation of the water layer, whereas CO2 availability did not influence PAR-limited plants. Our study shows that site conditions, rather than the sole abundance of potentially facilitating species, may strongly determine whether or not they form the structure necessary to act as a facilitator for biodiversity in aquatic environments.

Fens represent a large array of ecosystem services, including the highest biodiversity found among wetlands, hydrological services, water purification and carbon sequestration. Land-use change and drainage has severely damaged or annihilated these services in many parts of North America and Europe; restoration plans are urgently needed at the landscape level. We review the major constraints on the restoration of rich fens and fen water bodies in agricultural areas in Europe and disturbed landscapes in North America: (i) habitat quality problems: drought, eutrophication, acidification, and toxicity, and (ii) recolonization problems: species pools, ecosystem fragmentation and connectivity, genetic variability, and invasive species; and here provide possible solutions. We discuss both positive and negative consequences of restoration measures, and their causes. The restoration of wetland ecosystem functioning and services has, for a long time, been based on a trial-and-error approach. By presenting research and practice on the restoration of rich fen ecosystems within agricultural areas, we demonstrate the importance of biogeochemical and ecological knowledge at different spatial scales for the management and restoration of biodiversity, water quality, carbon sequestration and other ecosystem services, especially in a changing climate. We define target processes that enable scientists, nature managers, water managers and policy makers to choose between different measures and to predict restoration prospects for different types of deteriorated fens and their starting conditions., (© 2014 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

Seagrass beds are highly productive coastal ecosystems providing a large array of ecosystem services including fisheries and carbon sequestration. As seagrasses are known to be highly sensitive to anthropogenic forcing, we evaluated the use of trace metal concentrations in seagrasses as bioindicators for trace metal pollution of coastal regions at both global and local scale. We carried out a meta-analysis based on literature data to provide a global benchmark list for trace metal accumulation in seagrasses, which was lacking in literature. We subsequently carried out a case study at the Caribbean islands of Curaçao and Bonaire to test for local-scale differences in trace metal concentrations in seagrasses, and internal metal allocation. The benchmark and local study show that trace metal concentrations in seagrass leaves, regardless of the species, can vary over a 100-1000-fold range, and are related to the level of anthropogenic pressure, making seagrasses highly valuable indicators., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Internal phosphorus (P) mobilisation from aquatic sediments is an important process adding to eutrophication problems in wetlands. Salinisation, a fast growing global problem, is thought to affect P behaviour. Although several studies have addressed the effects of salinisation, interactions between salinity changes and nutrient cycling in freshwater systems are not fully understood. To tackle eutrophication, a clear understanding of the interacting effects of sediment characteristics and surface water quality is vital. In the present study, P release from two eutrophic sediments, both characterized by high pore water P and very low pore water iron (Fe(2+)) concentrations, was studied in a long-term aquarium experiment, using three salinity levels. Sediment P release was expected to be mainly driven by diffusion, due to the eutrophic conditions and low iron availability. Unexpectedly, this only seemed to be the driving mechanism in the short term (0-10 weeks). In the long term (>80 weeks), P mobilisation was absent in most treatments. This can most likely be explained by the oxidation of the sediment-water interface where Fe(2+) immobilises P, even though it is commonly assumed that free Fe(2+) concentrations need to be higher for this. Therefore, a controlling mechanism is suggested in which the partial oxidation of iron-sulphides in the sediment plays a key role, releasing extra Fe(2+) at the sediment-water interface. Although salinisation was shown to lower short-term P mobilisation as a result of increased calcium concentrations, it may increase long-term P mobilisation by the interactions between sulphate reduction and oxygen availability. Our study showed time-dependent responses of sediment P mobilisation in relation to salinity, suggesting that sulphur plays an important role in the release of P from FeSx-rich sediments, its biogeochemical effect depending on the availability of Fe(2+) and O2.

As a result of anthropogenic disturbances and natural stressors, seagrass beds are often patchy and heterogeneous. The effects of high loads of nutrients and organic matter in patch development and expansion in heterogeneous seagrass beds have, however, poorly been studied. We experimentally assessed the in situ effects of sediment quality on seagrass (Zostera noltii) patch dynamics by studying patch (0.35 m diameter) development and expansion for 4 sediment treatments: control, nutrient addition (NPK), organic matter addition (OM) and a combination (NPK+OM). OM addition strongly increased porewater sulfide concentrations whereas NPK increased porewater ammonium, nitrate and phosphate concentrations. As high nitrate concentrations suppressed sulfide production in NPK+OM, this treatment was biogeochemically comparable to NPK. Sulfide and ammonium concentrations differed within treatments, but over a 77 days period, seagrass patch survival and expansion were impaired by all additions compared to the control treatment. Expansion decreased at porewater ammonium concentrations >2,000 μmol L(-1). Mother patch biomass was not affected by high porewater ammonium concentrations as a result of its detoxification by higher seagrass densities. Sulfide concentrations >1,000 μmol L(-1) were toxic to both patch expansion and mother patch. We conclude that patch survival and expansion are constrained at high loads of nutrients or organic matter as a result of porewater ammonium or sulfide toxicity., (Copyright © 2014 Elsevier B.V. All rights reserved.)

The conservation of biodiverse wetland vegetation, including that of rich fens, has a high priority at a global scale. Although P-eutrophication may strongly decrease biodiversity in rich fens, some well-developed habitats do still survive in highly fertilized regions due to nutrient filtering services of large wetlands. The occurrence of such nutrient gradients is well-known, but the biogeochemical mechanisms that determine these patterns are often unclear. We therefore analyzed chemical speciation and binding of relevant nutrients and minerals in surface waters, soils and plants along such gradients in the large Ramsar nature reserve Weerribben-Wieden in the Netherlands. P-availability was lowest in relatively isolated floating rich fens, where plant N:P ratios indicated P-limitation. P-limitation can persist here despite high P-concentrations in surface waters near the peripheral entry locations, because only a small part of the P-input reaches the more isolated waters and fens. This pattern in P-availability appears to be primarily due to precipitation of Fe-phosphates, which mainly occurs close to entry locations as indicated by decreasing concentrations of Fe- and Al-bound P in the sub-aquatic sediments along this gradient. A further decrease of P-availability is caused by biological sequestration, which occurs throughout the wetland as indicated by equal concentrations of organic P in all sub-aquatic sediments. Our results clearly show that the periphery of large wetlands does indeed act as an efficient P-filter, sustaining the necessary P-limitation in more isolated parts. However, this filtering function does harm the ecological quality of the peripheral parts of the reserve. The filtering mechanisms, such as precipitation of Fe-phosphates and biological uptake of P, are crucial for the conservation and restoration of biodiverse rich fens in wetlands that receive eutrophic water from their surroundings. This seems to implicate that biodiverse wetland vegetation requires larger areas, as long as eutrophication has not been seriously tackled., (Copyright © 2014 Elsevier B.V. All rights reserved.)

Peat forming Sphagnum mosses are able to prevent the dominance of vascular plants under ombrotrophic conditions by efficiently scavenging atmospherically deposited nitrogen (N). N-uptake kinetics of these mosses are therefore expected to play a key role in differential N availability, plant competition, and carbon sequestration in Sphagnum peatlands. The interacting effects of rain N concentration and exposure time on moss N-uptake rates are, however, poorly understood. We investigated the effects of N-concentration (1, 5, 10, 50, 100, 500 µM), N-form ((15)N-ammonium or nitrate) and exposure time (0.5, 2, 72 h) on uptake kinetics for Sphagnum magellanicum from a pristine bog in Patagonia (Argentina) and from a Dutch bog exposed to decades of N-pollution. Uptake rates for ammonium were higher than for nitrate, and N-binding at adsorption sites was negligible. During the first 0.5 h, N-uptake followed saturation kinetics revealing a high affinity (Km 3.5-6.5 µM). Ammonium was taken up 8 times faster than nitrate, whereas over 72 hours this was only 2 times. Uptake rates decreased drastically with increasing exposure times, which implies that many short-term N-uptake experiments in literature may well have overestimated long-term uptake rates and ecosystem retention. Sphagnum from the polluted site (i.e. long-term N exposure) showed lower uptake rates than mosses from the pristine site, indicating an adaptive response. Sphagnum therefore appears to be highly efficient in using short N pulses (e.g. rainfall in pristine areas). This strategy has important ecological and evolutionary implications: at high N input rates, the risk of N-toxicity seems to be reduced by lower uptake rates of Sphagnum, at the expense of its long-term filter capacity and related competitive advantage over vascular plants. As shown by our conceptual model, interacting effects of N-deposition and climate change (changes in rainfall) will seriously alter the functioning of Sphagnum peatlands.

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.

Adequate biogeochemical characterization and monitoring of aquatic ecosystems, both for scientific purposes and for water management, pose high demands on spatial and temporal replication of chemical analyses. Near-infrared reflectance spectroscopy (NIRS) may offer a rapid, low-cost and reproducible alternative to standard analytical sample processing (digestion or extraction) and measuring techniques used for the chemical characterization of aquatic sediments. We analyzed a total of 191 sediment samples for total and NaCl-extractable concentrations of Al, Ca, Fe, K, Mg, Mn, N, Na, P, S, Si, and Zn as well as oxalate- extractable concentrations of Al, Fe, Mn and P. Based on the NIR spectral data and the reference values, calibration models for the prediction of element concentrations in unknown samples were developed and tested with an external validation procedure. Except Mn, all prediction models of total element concentrations were found to be acceptable to excellent (ratio of performance deviation: RPD 1.8-3.1). For extractable element fractions, viable model precision could be achieved for NaCl-extractable Ca, K, Mg, NH4 (+)-N, S and Si (RPD 1.7-2.2) and oxalate-extractable Al, Fe and P (RPD 1.9-2.3). For those elements that showed maximum total values below 3 g kg(-1) prediction models were found to become increasingly critical (RPD <2.0). Low concentrations also limited the performance of NIRS calibrations for extracted elements, with critical concentration thresholds <0.1 g kg(-1) and 3.3 g kg(-1) for NaCl and oxalate extractions, respectively. Thus, reliable NIRS measurements of trace metals are restricted to sediments with high metal content. Nevertheless, we demonstrated the suitability of NIRS measurements to determine a large array of chemical properties of aquatic sediments. The results indicate great potential of this fast technique as an analytical tool to better understand the large spatial and temporal variation of sediment characteristics in an economically viable way.

In wetland soils and underwater sediments of marine, brackish and freshwater systems, the strong phytotoxin sulfide may accumulate as a result of microbial reduction of sulfate during anaerobiosis, its level depending on prevailing edaphic conditions. In this review, we compare an extensive body of literature on phytotoxic effects of this reduced sulfur compound in different ecosystem types, and review the effects of sulfide at multiple ecosystem levels: the ecophysiological functioning of individual plants, plant-microbe associations, and community effects including competition and facilitation interactions. Recent publications on multi-species interactions in the rhizosphere show even more complex mechanisms explaining sulfide resistance. It is concluded that sulfide is a potent phytotoxin, profoundly affecting plant fitness and ecosystem functioning in the full range of wetland types including coastal systems, and at several levels. Traditional toxicity testing including hydroponic approaches generally neglect rhizospheric effects, which makes it difficult to extrapolate results to real ecosystem processes. To explain the differential effects of sulfide at the different organizational levels, profound knowledge about the biogeochemical, plant physiological and ecological rhizosphere processes is vital. This information is even more important, as anthropogenic inputs of sulfur into freshwater ecosystems and organic loads into freshwater and marine systems are still much higher than natural levels, and are steeply increasing in Asia. In addition, higher temperatures as a result of global climate change may lead to higher sulfide production rates in shallow waters.

One of the most frequently quoted ecosystem services of seagrass meadows is their value for coastal protection. Many studies emphasize the role of above-ground shoots in attenuating waves, enhancing sedimentation and preventing erosion. This raises the question if short-leaved, low density (grazed) seagrass meadows with most of their biomass in belowground tissues can also stabilize sediments. We examined this by combining manipulative field experiments and wave measurements along a typical tropical reef flat where green turtles intensively graze upon the seagrass canopy. We experimentally manipulated wave energy and grazing intensity along a transect perpendicular to the beach, and compared sediment bed level change between vegetated and experimentally created bare plots at three distances from the beach. Our experiments showed that i) even the short-leaved, low-biomass and heavily-grazed seagrass vegetation reduced wave-induced sediment erosion up to threefold, and ii) that erosion was a function of location along the vegetated reef flat. Where other studies stress the importance of the seagrass canopy for shoreline protection, our study on open, low-biomass and heavily grazed seagrass beds strongly suggests that belowground biomass also has a major effect on the immobilization of sediment. These results imply that, compared to shallow unvegetated nearshore reef flats, the presence of a short, low-biomass seagrass meadow maintains a higher bed level, attenuating waves before reaching the beach and hence lowering beach erosion rates. We propose that the sole use of aboveground biomass as a proxy for valuing coastal protection services should be reconsidered.

The majority of studies on rhizospheric interactions focus on pathogens, mycorrhizal symbiosis, or carbon transformations. Although the biogeochemical transformations of N, S, and Fe have profound effects on vegetation, these effects have received far less attention. This review, meant for microbiologists, biogeochemists, and plant scientists includes a call for interdisciplinary research by providing a number of challenging topics for future ecosystem research. Firstly, all three elements are plant nutrients, and microbial activity significantly changes their availability. Secondly, microbial oxidation with oxygen supplied by radial oxygen loss from roots in wetlands causes acidification, while reduction using alternative electron acceptors leads to generation of alkalinity, affecting pH in the rhizosphere, and hence plant composition. Thirdly, reduced species of all three elements may become phytotoxic. In addition, Fe cycling is tightly linked to that of S and P. As water level fluctuations are very common in wetlands, rapid changes in the availability of oxygen and alternative terminal electron acceptors will result in strong changes in the prevalent microbial redox reactions, with significant effects on plant growth. Depending on geological and hydrological settings, these interacting microbial transformations change the conditions and resource availability for plants, which are both strong drivers of vegetation development and composition by changing relative competitive strengths. Conversely, microbial composition is strongly driven by vegetation composition. Therefore, the combination of microbiological and plant ecological knowledge is essential to understand the biogeochemical and biological key factors driving heterogeneity and total (i.e., microorganisms and vegetation) community composition at different spatial and temporal scales.

In order to understand the physicochemical mechanisms that could explain the massive growth of Azolla arctica in the Eocene Arctic Ocean, we carried out a laboratory experiment in which we studied the interacting effects of rain and wind on the development of salinity stratification, both in the presence and in the absence of a dense Azolla cover. Additionally, we carried out a mesocosm experiment to get a better understanding of the nutrient cycling within and beneath a dense Azolla cover in both freshwater and brackish water environments. Here we show that Azolla is able to create a windproof, small-scale salinity gradient in brackish waters, which allows for efficient recycling of nutrients. We suggest that this mechanism ensures the maintenance of a large standing biomass in which additional input of nutrients ultimately result in a further expansion of an Azolla cover. As such, it may not only explain the extent of the Azolla event during the Eocene, but also the absence of intact vegetative Azolla remains and the relatively low burial efficiency of organic carbon during this interval.

Seagrasses have declined at a global scale due to light reduction and toxicity events, caused by eutrophication and increased sediment loading. Although several studies have tested effects of light reduction and toxicants on seagrasses, there is at present no information available on their interacting effects. In a full-factorial 5-day laboratory experiment, we studied short-term interactive effects of light conditions, pH and reduced nitrogen (NH(x)) in the water layer, mimicking pulses of river discharge, on the tropical early successional species Halodule uninervis and the late successional species Thalassia hemprichii. In contrast to recent results reported for the temperate species Zostera marina, increased NH(x) supply did not affect leaf mortality or photochemical efficiency in H. uninervis and in 7 out of 8 treatments for T. hemprichii. However, both tropical species demonstrated striking differences in nitrogen accumulation, free amino acid composition and free NH₃ accumulation. The increase in tissue nitrogen content was two times higher for H. uninervis than for T. hemprichii. Nitrogen stored as free amino acids (especially asparagine) only increased in H. uninervis. High pH only affected T. hemprichii, but only when not shaded, by doubling its free NH₃ concentrations, concomitantly decreasing its photosynthetic efficiency. Our results indicate that the early successional H. uninervis has higher tolerance to high NH(x) loads as compared to the late successional T. hemprichii. H. uninervis was better able to avoid toxic internal NH(x) levels by further assimilating glutamine into asparagine in contrast to T. hemprichii. Moreover, both tropical species seem to cope much better with high NH(x) than the temperate Z. marina. The implications for the distribution and succession of seagrass species under high nutrient loads are discussed., (Copyright © 2011 Elsevier B.V. All rights reserved.)

In remote, tropical areas human influences increase, potentially threatening pristine seagrass systems. We aim (i) to provide a bench-mark for a near-pristine seagrass system in an archipelago in East Kalimantan, by quantifying a large spectrum of abiotic and biotic properties in seagrass meadows and (ii) to identify early warning indicators for river sediment and nutrient loading, by comparing the seagrass meadow properties over a gradient with varying river influence. Abiotic properties of water column, pore water and sediment were less suitable indicators for increased sediment and nutrient loading than seagrass properties. Seagrass meadows strongly responded to higher sediment and nutrient loads and proximity to the coast by decreasing seagrass cover, standing stock, number of seagrass species, changing species composition and shifts in tissue contents. Our study confirms that nutrient loads are more important than water nutrient concentrations. We identify seagrass system variables that are suitable indicators for sediment and nutrient loading, also in rapid survey scenarios with once-only measurements., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

While it is well established that ecosystems display strong responses to elevated nitrogen deposition, the importance of the ratio between the dominant forms of deposited nitrogen (NH(x) and NO(y)) in determining ecosystem response is poorly understood. As large changes in the ratio of oxidised and reduced nitrogen inputs are occurring, this oversight requires attention. One reason for this knowledge gap is that plants experience a different NH(x):NO(y) ratio in soil to that seen in atmospheric deposits because atmospheric inputs are modified by soil transformations, mediated by soil pH. Consequently species of neutral and alkaline habitats are less likely to encounter high NH(4)(+) concentrations than species from acid soils. We suggest that the response of vascular plant species to changing ratios of NH(x):NO(y) deposits will be driven primarily by a combination of soil pH and nitrification rates. Testing this hypothesis requires a combination of experimental and survey work in a range of systems., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

Worldwide, fens and peat lakes that used to be peat-forming systems have become a significant source of C, N and P due to increased peat decomposition. To test the hypothesis that net nutrient mineralization rates may be uncoupled from decomposition rates, we investigated decomposition and net mineralization rates of nutrients in relation to sediment and pore water characteristics. We incubated 28 non-calcareous peat sediments and floating fen soils under aerobic and anaerobic conditions. We also tried to find a simple indicator to estimate the potential nutrient mobilization rates from peat sediments to the water layer by studying their relation with sediment and pore water characteristics in 44 Dutch non-calcareous peat lakes and ditches. Decomposition rates were primarily determined by the organic matter content, and were higher under aerobic conditions. However, highly decomposed peat sediments with low C:P and C:N ratios still showed high net nutrient mineralization rates. At Fe:PO(4) ratios below 1molmol(-1), PO(4) mobilization from the sediment to the water layer was considerable and linearly related to the pore water PO(4) concentration. At higher ratios, there was a strong linear correlation between the Fe:PO(4) ratio and PO(4) mobilization. Hence, measuring Fe and PO(4) in anaerobic sediment pore water provides a powerful tool for a quick assessment of internal PO(4) fluxes. Mobilization of mineral N was largely determined by diffusion. Total sediment Fe:S ratios gave an important indication of the amount of Fe that is available to immobilize PO(4). Pore water Fe concentrations decreased at ratios <1molmol(-1), whereas pore water PO(4) concentrations and PO(4) mobilization to the water layer increased. As PO(4) mobilization rates from the sediment to the water layer contribute to almost half of the total P load in Dutch peat lakes and fens, it is of pivotal importance to examine the magnitude of internal fluxes. Dredging of the nutrient-rich upper sediment layer will only be a useful restoration measure if both the influx of P-rich water and its internal mobilization from the newly exposed, potentially more reactive peat layer are sufficiently low., (Copyright 2010 Elsevier Ltd. All rights reserved.)

Both eutrophication and SO4 pollution can lead to higher availability of nutrients and potentially toxic compounds in wetlands. To unravel the interaction between the level of eutrophication and toxicity at species and community level, effects of SO4 were tested in nutrient-poor and nutrient-rich fen mesocosms. Biomass production of aquatic and semi-aquatic macrophytes and colonization of the water layer increased after fertilization, leading to dominance of highly competitive species. SO4 addition increased alkalinity and sulphide concentrations, leading to decomposition and additional eutrophication. SO4 pollution and concomitant sulphide production considerably reduced biomass production and colonization, but macrophytes were less vulnerable in fertilized conditions. The experiment shows that competition between species, vegetation succession and terrestrialization are not only influenced by nutrient availability, but also by toxicity, which strongly interacts with the level of eutrophication. This implies that previously neutralized toxicity effects in eutrophied fens may appear after nutrient reduction measures have been taken.

Background and Aims: Plants need different survival strategies in habitats differing in hydrological regimes. This probably has consequences for vegetation development when former floodplain areas that are currently confronted with soil flooding only, will be reconnected to the highly dynamical river bed. Such changes in river management are increasingly important, especially at locations where increased water retention can prevent flooding events in developed areas. It is therefore crucial to determine the responses of plant species from relatively low-dynamic wetlands to complete submergence, and to compare these with those of species from river forelands, in order to find out what the effects of such landscape-scale changes on vegetation would be., Methods: To compare the species' tolerance to complete submergence and their acclimation patterns, a greenhouse experiment was designed with a selection of 19 species from two contrasting sites: permanently wet meadows in a former river foreland, and frequently submerged grasslands in a current river foreland. The plants were treated with short (3 weeks) and long (6 weeks) periods of complete submergence, to evaluate if survival, morphological responses, and changes in biomass differed between species of the two habitats., Key Results: All tested species inhabiting river forelands were classified as tolerant to complete submergence, whereas species from wet meadows showed either relatively intolerant, intermediate or tolerant responses. Species from floodplains showed in all treatments stronger shoot elongation, as well as higher production of biomass of leaves, stems, fine roots and taproots, compared with meadow species., Conclusions: There is a strong need for the creation of temporary water retention basins during high levels of river discharge. However, based on the data presented, it is concluded that such reconnection of former wetlands (currently serving as meadows) to the main river bed will strongly influence plant species composition and abundance.

Although sulphur deposition rates in Europe have considerably decreased over the last decades, sulphate concentrations in freshwater wetlands are still high, as a result of drainage, nitrate pollution, and increased sulphur loads in rivers. High sulphur fluxes may cause sulphide toxicity and eutrophication, and strongly interfere with the biogeochemical cycling of iron and phosphorus. In the present study the ecotoxicological interactions between sulphur, phosphate, iron, and trace metals in freshwater wetlands are reviewed.

After flooding, iron reduction in riverine wetlands may cause the release of large quantities of phosphorus. As phosphorus is an important nutrient causing eutrophication in aquatic systems, it is important to have a tool to predict this potential release. In this study we examined the P release to the soil pore water in soil cores from floodplains in the Netherlands and from less anthropogenically influenced floodplains from Poland. During the inundation experiment, concentrations of P in the pore water rose to 2-90 times the initial concentrations. P release was not directly related to the geographic origin of the soils. An important predictor variable of P release was found in the ratio between the concentration of iron-bound P and amorphous iron. This ratio may provide a practical tool for the selection of new areas for wetland creation, and for impact assessment of plans for riverine wetland restoration and floodwater storage.

Sulfur pollution can lead to serious problems in freshwater wetlands, including phosphorus eutrophication and sulfide toxicity. We tested the effects of anaerobic iron-rich groundwater discharge in fens, simulated by iron injection, on two characteristic species (Juncus effusus and Caltha palustris) in a sulfidic environment. Biomass production of C. palustris roots showed an optimum response to the combined addition of iron and sulfide, with highest values at intermediate concentrations of both substances. Iron deficiency apparently occurred at low iron concentrations, while at high iron concentrations, growth was decreased. For J. effusus, in contrast, no toxic effects were found of both iron and sulfide. This could be explained by larger radial oxygen loss (ROL) of J. effusus and could not be explained by differences in phosphorous concentrations. The results of our experiments confirm that iron-rich groundwater discharge has the potential to affect vegetation composition through toxicity modification in sulfidic environments.

Metal pollution can be a serious threat to ecosystems at a global scale. Although the bioavailability of potentially toxic metals is determined by many biotic and abiotic factors, including pH and redox potential, total metal concentrations in the soil are used widely to assess or predict toxicity. In the present study we tested the effect of desiccation of soils differing in acidification potential and total heavy metal contamination on the growth and metal uptake of three typical, common wetland species: Caltha palustris, Juncus effusus, and Rumex hydrolapathum. We found that plant growth in wet soils mainly was determined by nutrient availability, though in dry soils the combined effects of acidification and increased metal availability prevailed. Metal uptake under anaerobic conditions was best predicted by the acidification potential (sediment S/[Ca + Mg] ratio), not by total metal concentrations. We propose that this is related to radial oxygen loss by wetland plant roots, which leads to acidification of the rhizosphere. Under aerobic conditions, plant metal uptake was best predicted by the amount of CaCl2-extractable metals. We conclude that total metal concentrations are not suitable for predicting bioavailability and that the above diagnostic parameters will provide insight into biogeochemical processes involved in toxicity assessment and soil policy.

The Thiobacilli are an important group of autotrophic bacteria occurring in nature linking the biogeochemical cycles of sulfur and nitrogen. Betaproteobacterial Thiobacilli are very likely candidates for mediating the process of nitrate-dependent anoxic iron sulfide mineral oxidation in freshwater wetlands. A Thiobacillus denitrificans-like bacterium was present in an enrichment on thiosulfate and nitrate, derived from an iron-sulfide- and nitrate-rich freshwater environment. Preliminary FISH analysis showed that the 16S rRNA gene-based bacterial probe mix showed great variation in intensity under different culture conditions. Furthermore, the widely applied 23S rRNA gene-based probe set BET42a/GAM42a incorrectly identified the T. denitrificans-like bacterium as a member of the Gammaproteobacteria. To circumvent these problems, the 23S rRNA genes of two T. denitrificans strains were partially sequenced and a new 23S rRNA gene-based probe (Betthio 1001) specific for betaproteobacterial Thiobacilli was designed. Use of this new probe Betthio 1001, combined with field measurements, indicates the involvement of Thiobacilli in the process of nitrate-dependent iron sulfide mineral oxidation.

In many Dutch freshwater wetlands, concentrations of sulfate in the surface water and groundwater have increased. It is especially in peaty areas that this can lead to problems, including the reduction of sulfate to toxic sulfide. Our aquarium experiments showed that even low sulfide concentrations of 50 micromol/L are toxic to the freshwater macrophyte Nitella flexilis and the freshwater oligochaete Ophidonais serpentina. Sulfide toxicity can be modified by the availability of free iron in soil moisture or sediment from iron-rich groundwater discharge. The iron precipitates the sulfide, thereby immobilizing it and decreasing its toxicity. However, iron itself can be a toxicant as well. We found a detoxifying effect of moderate iron concentrations on sulfide toxicity, as well as a toxic effect of high iron concentrations on the growth of Potamogeton acutifolius. At the intermediate range, the formation of metal sulfides can simultaneously decrease metal and sulfide toxicity. Results of the experiments are discussed in relation to hydrological changes in freshwater wetlands.

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.

Sexually mature female tilapia were exposed to sublethal concentrations of waterborne Cu and/or Cd over 6 days, and subsequent body concentrations of these metals were determined in several organs. The results show that the distribution of Cu and Cd was metal and organ specific. This is demonstrated, for example, by the observation that in tilapia, Cu exposure did not result in Cu accumulation in the liver, whereas in the intestinal wall, notably high concentrations of Cu and Cd were measured in metal exposed fish. In addition to single metal exposed fish, we also determined Cu and Cd body distribution in Cu?Cd co-exposed fish. The observed interactions in metal accumulation were most pronounced in the organs of fish exposed to low, environmentally realistic, metal concentrations.