The seafloor from a trait perspective. A comprehensive life history dataset of soft sediment macrozoobenthos.

Biological trait analysis (BTA) is a valuable tool for evaluating changes in community diversity and its link to ecosystem processes as well as environmental and anthropogenic perturbations. Trait-based analytical techniques like BTA rely on standardised datasets of species traits. However, there are currently only a limited number of datasets available for marine macrobenthos that contain trait data across multiple taxonomic groups. Here, we present an open-access dataset of 16 traits for 235 macrozoobenthic species recorded throughout multiple sampling campaigns of the Dutch Wadden Sea; a dynamic soft bottom system where humans have long played a substantial role in shaping the coastal environment. The trait categories included in this dataset cover a variety of life history strategies that are tightly linked to ecosystem functioning and the resilience of communities to (anthropogenic) perturbations and can advance our understanding of environmental changes and human impacts on the functioning of soft bottom systems.

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.

The importance of marshes providing soil stabilization to resist fast-flow erosion in case of a dike breach.

Salt marshes provide valuable ecosystem services including coastal protection by reducing wave loading on dikes and seawalls. If the topsoil is erosion resistant to fast-flowing water, it may also reduce breach depth if a dike fails. In this experiment, we quantified the topsoil erosion resistance from marshes and bare tidal flats with different soil types to understand the extent to which they can help reduce breach depth. Intact soil samples were collected from 11 locations in the Netherlands at different tidal elevations and then exposed for 3 h to 2.3 m/s currents. To the samples that remained stable after flow exposure, an artificial crack was made to test their stability following soil disturbance. All samples from the tidal flats were completely eroded, regardless of sediment type. In contrast, all samples from well-established marsh plateaus were stable as long as no disturbances were made, including those with sandy subsoils. After creating artificial cracks, samples with a thin cohesive top layer on top of sandy subsoil collapsed, while marshes with silty subsoils remained stable. Pioneer marshes on sandy substrate without a cohesive top layer were the only vegetated soils that completely eroded. The lower erosion of marshes with either sandy or silty soils compared to bare tidal flats was best explained by the presence of a top layer with belowground biomass, high organic content, high water content, and low bulk density. When analyzing the erodibility of marshes only, fine root density was the best predictor of erosion resistance. This study demonstrates the importance of preserving, restoring, or creating salt marshes, to obtain a topsoil that is erosion resistant under fast-flowing water, which helps reduce breach dimensions if a dike fails. The probability of topsoil erosion in established marshes with sandy subsoil is higher than in silty marshes. A silty layer of cohesive sediment on top of the sand provides extra erosion resistance as long as it does not break. Pioneer marshes that have not developed a cohesive top layer are erosion sensitive, especially in sandy soils. For future marsh creations, using fine-grained sediments or a mixture of sand with silt or clay is recommended.

Growth, maturity, and diet of the pearl whipray (Fontitrygon margaritella) from the Bijagós Archipelago, Guinea-Bissau.

The pearl whipray Fontitrygon margaritella (Compagno & Roberts, 1984) is a common elasmobranch in coastal western African waters. However, knowledge on their life-history and trophic ecology remains limited. Therefore, we aimed to determine the growth, maturity and diet of F. margaritella from the Bijagós Archipelago in Guinea-Bissau. Growth was modelled with: von Bertalanffy, Gompertz and logistic functions. Model selection revealed no model significantly outperformed another. The sampled age ranged from less than 1 to 7 years (1.8 ± 1.9 cm, mean ± standard deviation) and size (disc width) ranged from 12.2 to 30.6 cm (18.7 ± 5.2 cm). Size-at-maturity was estimated at 20.3 cm (95% CI [18.8-21.8 cm]) for males and 24.3 cm for females (95% CI [21.9-26.5 cm]), corresponding ages of 2.2 and 3.9 years. The diet differed significantly among young-of-the-year (YOY), juveniles and adults (p = 0.001). Diet of all life stages consisted mainly of crustaceans (27.4%, 28.5%, 33.3%) and polychaetes (12.5%, 26.7%, 20.3%), for YOY, juveniles and adults respectively. This study shows that F. margaritella is relatively fast-growing, matures early and experiences ontogenetic diet shifts. These results contribute to status assessments and conservation efforts of F. margaritella and closely related species.

Adaptive intertidal seed-based seagrass restoration in the Dutch Wadden Sea.

Seagrasses form the foundation of many coastal ecosystems but are rapidly declining on a global scale. The Dutch Wadden Sea once supported extensive subtidal seagrass meadows that have all disappeared. Here, we report on the setbacks and successes of intertidal seed-based restoration experiments in the Dutch Wadden Sea between 2014-2017. Our main goals were to 1) optimize plant densities, and 2) reduce seed losses. To achieve our goals, we conducted research-based, adaptive seagrass (Zostera marina) restoration, adjusting methods yearly based on previous results. We applied various seeding methods in three subsequent years-from Buoy Deployed Seeding (BuDS), and 'BuDS-in-frame' in fall, to a newly developed 'Dispenser Injection Seeding' (DIS) method. Our adaptive experimental approach revealed high seed losses between seeding and seedling establishment of the BuDS methods (>99.9%), which we mitigated by controlled harvest and storage of seeds throughout fall and winter, followed by DIS-seeding in spring. These iterative innovations resulted in 83 times higher plant densities in the field (0.012 to 1.00 plants m-2) and a small reduction in seed loss (99.94 to 99.75%) between 2015-2017. Although these developments have not yet resulted in self-sustaining seagrass populations, we are one step closer towards upscaling seagrass restoration in the Dutch Wadden Sea. Our outcomes suggest that an iterative, research-based restoration approach that focuses on technological advancement of precision-seeding may result in advancing knowledge and improved seed-based seagrass restoration successes.

Mimicry of emergent traits amplifies coastal restoration success.

Restoration is becoming a vital tool to counteract coastal ecosystem degradation. Modifying transplant designs of habitat-forming organisms from dispersed to clumped can amplify coastal restoration yields as it generates self-facilitation from emergent traits, i.e. traits not expressed by individuals or small clones, but that emerge in clumped individuals or large clones. Here, we advance restoration science by mimicking key emergent traits that locally suppress physical stress using biodegradable establishment structures. Experiments across (sub)tropical and temperate seagrass and salt marsh systems demonstrate greatly enhanced yields when individuals are transplanted within structures mimicking emergent traits that suppress waves or sediment mobility. Specifically, belowground mimics of dense root mats most facilitate seagrasses via sediment stabilization, while mimics of aboveground plant structures most facilitate marsh grasses by reducing stem movement. Mimicking key emergent traits may allow upscaling of restoration in many ecosystems that depend on self-facilitation for persistence, by constraining biological material requirements and implementation costs.

A Lévy expansion strategy optimizes early dune building by beach grasses.

Lifeforms ranging from bacteria to humans employ specialized random movement patterns. Although effective as optimization strategies in many scientific fields, random walk application in biology has remained focused on search optimization by mobile organisms. Here, we report on the discovery that heavy-tailed random walks underlie the ability of clonally expanding plants to self-organize and dictate the formation of biogeomorphic landscapes. Using cross-Atlantic surveys, we show that congeneric beach grasses adopt distinct heavy-tailed clonal expansion strategies. Next, we demonstrate with a spatially explicit model and a field experiment that the Lévy-type strategy of the species building the highest dunes worldwide generates a clonal network with a patchy shoot organization that optimizes sand trapping efficiency. Our findings demonstrate Lévy-like movement in plants, and emphasize the role of species-specific expansion strategies in landscape formation. This mechanistic understanding paves the way for tailor-made planting designs to successfully construct and restore biogeomorphic landscapes and their services.

North Sea demersal fisheries prefer specific benthic habitats.

INTRODUCTION: The future protection of marine biodiversity through good conservation planning requires both the identification of key habitats with unique ecological characteristics and detailed knowledge of their human utilization through fisheries. Demersal fisheries are important disturbers of benthic habitats. They often have a heterogeneous spatial distribution, pressurizing particular habitats with high abundances of target species. For the North Sea, we quantified the commonness/rarity of habitats in relation to the environmental determinants of so-called fishing hotspots, to support better-informed conservation planning of benthic habitats in this intensively used continental shelf. METHODS: We first distinguished 9 main seascapes in the study area based on seabed morphology. Secondly, we determined average fishing intensity and fishing hotspots using VMS-data for the three dominant Dutch fisheries from 2008 to 2015: beam-trawlers targeting sole Solea solea (Beam-Sole), beam-trawlers targeting plaice Pleuronectes platessa (Beam-Plaice), and otter-trawlers targeting Norway lobster Nephrops norvegicus and demersal fish (Otter-Mix). Within the seascapes subjected to >80% of the fishing activity, nineteen environmental factors (summarized by PCA) were used to ecologically characterize fishing hotspot locations using MaxEnt response modelling. RESULTS: We found that all three fisheries target highly specific, uncommon habitats. Beam-Sole fishers targeted warmer, shallow, dynamic, nearshore habitats, and within these specifically the depressions between sand ridges. Beam-Plaice fishers mainly targeted the exposed, non-muddy flanks of the Dogger Bank and similar large-scale elevations (50-75 km) where especially the ridges of smaller sand banks are used. Otter-Mix fisheries concentrated in areas with low bed shear stress, located in muddy, relatively deeper areas. IMPLICATIONS: This study is the first to provide insight in benthic habitat types that are frequently targeted by fishers in the North Sea. We demonstrated unequal exploitation pressure between seabed habitats, with the majority of hotspots in the less common habitats. Our results hence contribute to a more effective, ecologically informed planning for the protection and monitoring of all seabed habitats and biodiversity of the North Sea.

Evidence for 'critical slowing down' in seagrass: a stress gradient experiment at the southern limit of its range.

The theory of critical slowing down, i.e. the increasing recovery times of complex systems close to tipping points, has been proposed as an early warning signal for collapse. Empirical evidence for the reality of such warning signals is still rare in ecology. We studied this on Zostera noltii intertidal seagrass meadows at their southern range limit, the Banc d'Arguin, Mauritania. We analyse the environmental covariates of recovery rates using structural equation modelling (SEM), based on an experiment in which we assessed whether recovery after disturbances (i.e. seagrass & infauna removal) depends on stress intensity (increasing with elevation) and disturbance patch size (1 m2 vs. 9 m2). The SEM analyses revealed that higher biofilm density and sediment accretion best explained seagrass recovery rates. Experimental disturbances were followed by slow rates of recovery, regrowth occurring mainly in the coolest months of the year. Macrofauna recolonisation lagged behind seagrass recovery. Overall, the recovery rate was six times slower in the high intertidal zone than in the low zone. The large disturbances in the low zone recovered faster than the small ones in the high zone. This provides empirical evidence for critical slowing down with increasing desiccation stress in an intertidal seagrass system.

A facultative mutualistic feedback enhances the stability of tropical intertidal seagrass beds.

Marine foundation species such as corals, seagrasses, salt marsh plants, and mangrove trees are increasingly found to engage in mutualistic interactions. Because mutualisms by their very nature generate a positive feedback between the species, subtle environmental impacts on one of the species involved may trigger mutualism breakdown, potentially leading to ecosystem regime shifts. Using an empirically parameterized model, we investigate a facultative mutualism between seagrass and lucinid bivalves with endosymbiotic sulfide-oxidizing gill bacteria in a tropical intertidal ecosystem. Model predictions for our system show that, by alleviating the build-up of toxic sulfide, this mutualism maintains an otherwise intrinsically unstable seagrass ecosystem. However, an increase in seagrass mortality above natural levels, due to e.g. desiccation stress, triggers mutualism breakdown. This pushes the system in collapse-and-recovery dynamics ('slow-fast cycles') characterized by long-term persistent states of bare and seagrass-dominated, with rapid transitions in between. Model results were consistent with remote sensing analyses that suggest feedback-mediated state shifts induced by desiccation. Overall, our combined theoretical and empirical results illustrate the potential of mutualistic feedbacks to stabilize ecosystems, but also reveal an important drawback as small environmental changes may trigger shifts. We therefore suggest that mutualisms should be considered for marine conservation and restoration of seagrass beds.

Living in the intertidal: desiccation and shading reduce seagrass growth, but high salinity or population of origin have no additional effect.

The limiting effects of stressors like desiccation, light and salinity on seagrass growth and distribution are well-studied. However, little is known about their interactive effects, and whether such effects might differ among populations that are adapted to different local conditions. In two laboratory experiments we tested (a) if growth and development of intertidal, temperate Zostera noltii is affected by emergence time (experiment 1 and 2), and (b) how this is affected by an additional, second stressor, namely shading (experiment 1) or high salinity (25, 30 and 35, experiment 2). In addition, we tested (c) whether the effects of emergence time and salinity varied between three different European seagrass populations (Saint-Jacut/France, Oosterschelde/The Netherlands, and Sylt/Germany), which are likely adapted to different salinity levels (experiment 2). In both experiments, emergence of 8 h per tidal cycle (of 12 h) had a negative effect on seagrass relative growth rate (RGR), and aboveground biomass. Emergence furthermore reduced either rhizome length (experiment 1) or belowground biomass (experiment 2). Shading (experiment 1) resulted in lower RGR and a two-fold higher aboveground/belowground ratio. We found no interactive effects of emergence and shading stress. Salinity (experiment 2) did not affect seagrass growth or morphology of any of the three populations. The three tested populations differed greatly in morphology but showed no differential response to emergence or salinity level (experiment 2). Our results indicate that emergence time and shading show an additive negative effect (no synergistic or antagonistic effect), making the plants still vulnerable to such combination, a combination that may occur as a consequence of self-shading during emergence or resulting from algal cover. Emergence time likely determines the upper limit of Z. noltii and such shading will likely lower the upper limit. Shading resulted in higher aboveground/belowground ratios as is a general response in seagrass. Z. noltii of different populations originating from salinity 30 and 35 seem tolerant to variations in salinity within the tested range. Our results indicate that the three tested populations show morphotypic rather than ecotypic variation, at least regarding the salinity and emergence, as there were no interactive effects with origin. For restoration, this implies that the salinity regime of the donor and receptor site of Z. noltii is of no concern within the salinity range 25-35.

Review: Host-pathogen dynamics of seagrass diseases under future global change.

Human-induced global change is expected to amplify the disease risk for marine biota. However, the role of disease in the rapid global decline of seagrass is largely unknown. Global change may enhance seagrass susceptibility to disease through enhanced physiological stress, while simultaneously promoting pathogen development. This review outlines the characteristics of disease-forming organisms and potential impacts of global change on three groups of known seagrass pathogens: labyrinthulids, oomycetes and Phytomyxea. We propose that hypersalinity, climate warming and eutrophication pose the greatest risk for increasing frequency of disease outbreaks in seagrasses by increasing seagrass stress and lowering seagrass resilience. In some instances, global change may also promote pathogen development. However, there is currently a paucity of information on these seagrass pathosystems. We emphasise the need to expand current research to better understand the seagrass-pathogen relationships, serving to inform predicative modelling and management of seagrass disease under future global change scenarios.

Seagrass leaf element content: A global overview.

Knowledge on the role of seagrass leaf elements and in particular micronutrients and their ranges is limited. We present a global database, consisting of 1126 unique leaf values for ten elements, obtained from literature and unpublished data, spanning 25 different seagrass species from 28 countries. The overall order of average element values in seagrass leaves was Na>K>Ca>Mg>S>Fe>Al>Si>Mn>Zn. Although we observed differences in leaf element content between seagrass families, high intraspecific variation indicated that leaf element content was more strongly determined by environmental factors than by evolutionary history. Early successional species had high leaf Al and Fe content. In addition, seagrass leaf element content also showed correlations with macronutrients (N and P), indicating that productivity also depends on other elements. Expected genomes of additional seagrass species in combination with experiments manipulating (micro)nutrients and environmental drivers might enable us to unravel the importance of various elements to sustain productive and flourishing meadows.

Copper treatment during storage reduces Phytophthora and Halophytophthora infection of Zostera marina seeds used for restoration.

Restoration is increasingly considered an essential tool to halt and reverse the rapid decline of vital coastal ecosystems dominated by habitat-forming foundation species such as seagrasses. However, two recently discovered pathogens of marine plants, Phytophthora gemini and Halophytophthora sp. Zostera, can seriously hamper restoration efforts by dramatically reducing seed germination. Here, we report on a novel method that strongly reduces Phytophthora and Halophytophthora infection of eelgrass (Zostera marina) seeds. Seeds were stored in seawater with three different copper sulphate concentrations (0.0, 0.2, 2.0 ppm) crossed with three salinities (0.5, 10.0, 25.0 ppt). Next to reducing seed germination, infection significantly affected cotyledon colour: 90% of the germinated infected seeds displayed a brown cotyledon upon germination that did not continue development into the seedling stage, in contrast to only 13% of the germinated non-infected seeds. Copper successfully reduced infection up to 86% and the 0.2 ppm copper sulphate treatment was just as successful as the 2.0 ppm treatment. Infection was completely eliminated at low salinities, but green seed germination was also dramatically lowered by 10 times. We conclude that copper sulphate treatment is a suitable treatment for disinfecting Phytophthora or Halophytophthora infected eelgrass seeds, thereby potentially enhancing seed-based restoration success.

Marine Phytophthora species can hamper conservation and restoration of vegetated coastal ecosystems.

Phytophthora species are potent pathogens that can devastate terrestrial plants, causing billions of dollars of damage yearly to agricultural crops and harming fragile ecosystems worldwide. Yet, virtually nothing is known about the distribution and pathogenicity of their marine relatives. This is surprising, as marine plants form vital habitats in coastal zones worldwide (i.e. mangrove forests, salt marshes, seagrass beds), and disease may be an important bottleneck for the conservation and restoration of these rapidly declining ecosystems. We are the first to report on widespread infection of Phytophthora and Halophytophthora species on a common seagrass species, Zostera marina (eelgrass), across the northern Atlantic and Mediterranean. In addition, we tested the effects of Halophytophthora sp. Zostera and Phytophthora gemini on Z. marina seed germination in a full-factorial laboratory experiment under various environmental conditions. Results suggest that Phytophthora species are widespread as we found these oomycetes in eelgrass beds in six countries across the North Atlantic and Mediterranean. Infection by Halophytophthora sp. Zostera, P. gemini, or both, strongly affected sexual reproduction by reducing seed germination sixfold. Our findings have important implications for seagrass ecology, because these putative pathogens probably negatively affect ecosystem functioning, as well as current restoration and conservation efforts.

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.

How habitat-modifying organisms structure the food web of two coastal ecosystems.

The diversity and structure of ecosystems has been found to depend both on trophic interactions in food webs and on other species interactions such as habitat modification and mutualism that form non-trophic interaction networks. However, quantification of the dependencies between these two main interaction networks has remained elusive. In this study, we assessed how habitat-modifying organisms affect basic food web properties by conducting in-depth empirical investigations of two ecosystems: North American temperate fringing marshes and West African tropical seagrass meadows. Results reveal that habitat-modifying species, through non-trophic facilitation rather than their trophic role, enhance species richness across multiple trophic levels, increase the number of interactions per species (link density), but decrease the realized fraction of all possible links within the food web (connectance). Compared to the trophic role of the most highly connected species, we found this non-trophic effects to be more important for species richness and of more or similar importance for link density and connectance. Our findings demonstrate that food webs can be fundamentally shaped by interactions outside the trophic network, yet intrinsic to the species participating in it. Better integration of non-trophic interactions in food web analyses may therefore strongly contribute to their explanatory and predictive capacity.

Seagrasses as indicators for coastal trace metal pollution: a global meta-analysis serving as a benchmark, and a Caribbean case study.

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.

Eutrophication threatens Caribbean seagrasses - An example from Curaçao and Bonaire.

Seagrass beds are globally declining due to human activities in coastal areas. We here aimed to identify threats from eutrophication to the valuable seagrass beds of Curaçao and Bonaire in the Caribbean, which function as nursery habitats for commercial fish species. We documented surface- and porewater nutrient concentrations, and seagrass nutrient concentrations in 6 bays varying in nutrient loads. Water measurements only provided a momentary snapshot, due to timing, tidal stage, etc., but Thalassia testudinum nutrient concentrations indicated long-term nutrient loads. Nutrient levels in most bays did not raise any concern, but high leaf % P values of Thalassia in Piscadera Bay (∼0.31%) and Spanish Water Bay (∼0.21%) showed that seagrasses may be threatened by eutrophication, due to emergency overflow of waste water and coastal housing. We thus showed that seagrasses may be threatened and measures should be taken to prevent loss of these important nursery areas due to eutrophication.

Toxic effects of increased sediment nutrient and organic matter loading on the seagrass Zostera noltii.

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.