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Ecosystems worldwide depend on habitat-forming foundation species that engage in facultative mutualisms. Global change, however, is causing rapid declines of foundation species-structured ecosystems, often typified by sudden collapse. Although disruption of obligate mutualisms involving foundation species is known to precipitate collapse (e.g. coral bleaching), how facultative mutualisms (i.e. context-dependent, non-binding reciprocal interactions) affect ecosystem resilience is uncertain. Here, we synthesize recent advancements, and combine this with model analyses supported by real-world examples, to propose that facultative mutualisms may pose a double-edged sword for foundation species. We suggest that by amplifying self-facilitative feedbacks by foundation species, facultative mutualisms can increase foundation species resistance to global change stressors. Simultaneously, however, mutualism-dependency can generate or exacerbate bistability, implying a potential for sudden collapse when the mutualism’s buffering capacity is exceeded, while recovery requires conditions to improve beyond the initial collapse point (hysteresis). Thus, our work emphasizes the importance of acknowledging facultative mutualisms for conservation and restoration of foundation species-structured ecosystems, but highlights the potential risk of relying on mutualisms in the face of global change. We argue that significant caveats remain regarding the determination of these feedbacks, and suggest empirical manipulation across stress gradients as a way forward to identify related nonlinear responses.

1. Vegetated marine and freshwater habitats are being increasingly lost around the world. Habitat restoration is a critical step for conserving these valuable habitats, but new approaches are needed to increase restoration success and ensure their survival. 2. We investigated interactions between plants and bivalves through a review and analysis of 491 studies, determined the effects, mechanisms and key environmental variables involved in and driving positive and negative interactions, and produced guidelines for integrating positive interactions into restoration efforts in different habitats. 3. Fifty per cent of all interactions (both correlative and experimental studies) were positive. These were predominant between epifaunal bivalves and plants in all habitats, and between infaunal bivalves and plants in subtidal habitats. Plants primarily promoted bivalve survival and abundance by providing substrate and shelter, while bivalves promoted plant growth and survival by stabilizing and fertilizing the sediment, and reducing water turbidity. The prevalence of positive interactions increased with water temperature in subtidal habitats, but decreased with water temperature in intertidal habitats. The subset of studies conducted in a restoration context also showed mostly positive interactions. 4. Twenty‐five per cent of all interactions were negative, and these were predominant between plants and infaunal bivalves in intertidal habitats, except sulphide‐metabolizing bivalves, which facilitated plant survival. Interactions involving non‐native species were also mostly negative. 5. Synthesis and applications. Promoting facilitative interactions through plant–bivalve co‐restoration can increase restoration success. The prevalence of positive interactions depends on habitat and environmental conditions such as temperature, and was especially important in subtidal habitats (involving both infaunal and epifaunal bivalves) and in intertidal habitats (involving only epifaunal bivalves). Thus sites and species for co‐restoration must be carefully chosen to maximize the chances of success. If done properly, co‐restoration could increase initial survival, persistence and resilience of foundation species, and promote the recovery of associated biodiversity and ecosystem services.

Seagrasses provide multiple ecosystem services including nursery habitat, improved water quality, coastal protection, and carbon sequestration. However, seagrasses are in crisis as global coverage is declining at an accelerating rate. With increased focus on ecological restoration as a conservation strategy, methods that enhance restoration success need to be explored. Decades of work in coastal plant ecosystems, including seagrasses, has shown that positive species relationships and feedbacks are critical for ecosystem stability, expansion, and recovery from disturbance. We reviewed the restoration literature on seagrasses and found few studies have tested for the beneficial effects of including positive species interactions in seagrass restoration designs. Here we review the full suite of positive species interactions that have been documented in seagrass ecosystems, where they occur, and how they might be integrated into seagrass restoration. The few studies in marine plant communities that have explicitly incorporated positive species interactions and feedbacks have found an increase in plant growth with little additional resource investment. As oceans continue to change and stressors become more prevalent, harnessing positive interactions between species through innovative approaches will likely become key to successful seagrass restoration.

Aim: The formation of a local vegetation mosaic may be attributed to local variation in abiotic environmental conditions. Recent research, however, indicates that self-facilitating organisms and negative species interactions may be a driving factor. In this study, we explore whether heterogeneous geohydrological conditions or vegetation feedbacks and interactions could be responsible for a vegetation mosaic of rich and poor fen species. Location: Lake Aturtaun, Roundstone Bog, Ireland. Methods: In a floating fen, transects were set out to analyze the relation between vegetation type and rock–peat distance and porewater electrical conductivity. Furthermore, three distinct vegetation types were studied: rich fen, poor fen and patches of poor fen within rich fen vegetation. Biogeochemical measurements were conducted in a vertical profile to distinguish abiotic conditions of distinct vegetation types. Results: Geohydrological conditions may drive the distribution of poor and rich fen species at a larger scale in the floating fen, due to the supply of minerotrophic groundwater. Interestingly, both rich and poor fen vegetation occurred in a mosaic, when electrical conductivity values at 50 cm depth were between 300 µS/cm and 450 µS/cm. Although environmental conditions were homogeneous at 50 cm, they differed markedly between rich and poor fen vegetation at 10 cm depth. Specifically, our measurements indicate that poor fen vegetation lowered porewater alkalinity, bicarbonate concentrations and pH. No effects of rich fen vegetation at 10 cm depth on biogeochemistry was measured. However, rich fen litter had a higher mineralization rate than poor fen litter, which increases the influence of minerotrophic water in rich fen habitat. Conclusions: These results strengthen our hypothesis that species can drive formation of vegetation mosaics under environmentally homogeneous conditions in a floating fen. Positive intraspecific self-facilitating mechanisms and negative species interactions could be responsible for a stable coexistence of species, even leading to local ecosystem engineering by the species, explaining the local vegetation mosaic at the microscale level in a floating fen.

Seagrass meadows form vital ecological components of coastal zones worldwide, but are rapidly declining. Large-scale seagrass diebacks have been related to accumulation of toxic sulfide in the sediment, a phenomenon predicted to occur more frequently in the near future due to ongoing global warming and increasing organic loading of coastal systems worldwide. Recently, a facultative mutualism between seagrasses and lucinid bivalves with endosymbiotic sulfide-consuming gill bacteria was discovered that may prevent toxic sulfide accumulation in seagrass sediments. Yet, direct field-based evidence for the importance of this mutualism in alleviating sulfide stress in seagrasses is currently lacking, as well as how its role may change when sediment sulfide levels increase due to environmental change. Here, we investigated the sulfide detoxification function of this seagrass-lucinid mutualism and its resilience to organic-loading induced sulfide stress in a temperate lagoon system (Thau lagoon, France), using a correlative field survey and a full factorial field experiment. The field survey revealed a strong positive correlation between seagrass above-ground biomass and lucinid densities, and pore water sulfide concentrations close to zero at all sites. Furthermore, the field experiment revealed that addition of organic matter (starch mixed with sucrose) increased sedimentary sulfide intrusion into seagrass (Zostera noltei) leaves (a proxy for sulfide stress), while experimentally enhanced lucinid densities reduced sulfide intrusion, regardless of addition of organic matter. Moreover, addition of organic matter reduced seagrass rhizome biomass and increased pore water sulfide levels, lucinid tissue sulfur content, lucinid condition (expressed as flesh/shell dry weight ratio), and total lucinid biomass, while enhancement of lucinid densities reduced lucinid condition. These results provide the first field-based evidence that lucinid bivalves and their sulfide-oxidizing gill symbionts mitigate sulfide stress in seagrasses, and suggests that the dependence of seagrass on this seagrass-lucinid mutualism will increase under conditions of enhanced sediment sulfide production, as predicted for the near future. Therefore, we suggest that awareness of the ecological importance of the seagrass-lucinid mutualism may be instrumental for designing new measures for improving long-term restoration success and seagrass resilience to global change.

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., Coastal restoration tends to be failure-prone and expensive. Temmink and colleagues improve seagrass and cordgrass transplant survival in field experiments using biodegradable structures which temporarily mimic self-facilitation occurring in mature vegetation stands, and combine onsite and laboratory measurements on sediment stability and stem movement to test the biophysical mechanisms.

Habitat characteristics, predation and competition are known to interactively drive population dynamics. Highly complex habitats, for example, may reduce predation and competition, allowing more individuals living together in a certain area. However, the strength and direction of such interactions can differ strongly and are context dependent. Furthermore, as habitat characteristics are rapidly changing due to anthropogenic impacts, it becomes increasingly important to understand such interactions. Here, we studied the interactive effects of predation and competition on common cockle (Cerastoderma edule) recruitment, growth and survival under different habitat characteristics in the Wadden Sea, one of the world's largest intertidal ecosystems. In a predator-exclosure experiment, we manipulated cockle densities (100 vs. 1000 individuals m-2) and shorebird predation at two sites differing in habitat characteristics, namely at the wake of a blue mussel bed (Mytilus edulis) and at an adjacent sandy site. We found that recruitment was higher in the mussel-modified habitat, most likely due to reduction of hydrodynamic stress. Although bird predation strongly reduced recruit density, the combined effects still yielded more recruitment at the vicinity of the mussel bed compared to the sandy area. Furthermore, we found that high cockle densities combined with high densities of other potential prey (i.e. mussels) at the mussel-modified site, mitigated predation effects for adult cockles. Apart from these positive effects on adults, mussel-modified habitat reduced cockle growth, most likely by reducing hydrodynamics in the wake of the mussel bed and by increasing inter-specific competition for food. Our study experimentally underpins the importance of habitat characteristics, competition and predation in interactively structuring intertidal communities.

A plethora of oomycetes was isolated mostly from Zostera marina but also from other halophilic plants originating from several locations including the Wadden Sea, Limfjord (Denmark), the Dutch Delta area (the Netherlands), Thau lagoon (France), Lindholmen (Sweden) and Chesapeake Bay (Virginia, U.S.). Based on ITS sequences, seven different groups could be distinguished. The largest group was assigned to Phytophthora gemini (Germany, Sweden, the Netherlands, U.S.). The CoxI sequences of all P. gemini strains were identical indicating that P. gemini is probably an invasive species in the Wadden Sea. A second group was identified as P. inundata (the Netherlands, Denmark), that was also isolated from the halophilic plants Aster tripolium and Salicornia europaea. Four strains, originating from Chesapeake Bay clustered in a monophyletic clade with high bootstrap support at the ITS as well as the CoxI loci. They are phylogenetically closely related to P. gemini and are considered to represent a new species described here as Phytophthora chesapeakensis sp. nov. In addition, Salisapilea sapeloensis was isolated from Zostera noltii. Eleven other strains belonging to three unidentified taxa, originating from the Wadden Sea, the Dutch Delta area and Thau lagoon, clustered each in a monophyletic clade with high bootstrap support at the ITS locus, including Halophytophthora vesicula, the type species of the genus Halophytophthora. Hence, these strains were considered to belong to the Halophytophthora sensu stricto group and probably represent three new Halophytophthora species, informally designated here as Halophytophthora sp-1, Halophytophthora sp-3 and Halophytophthora sp-4 sensu Nigrelli and Thines. Halophytophthora sp-2 was not detected in this study. In addition, P. gemini and Halophytophthora sp-3 were obtained by baiting from locations in the Wadden Sea and Halophytophthora sp-1 was obtained by baiting from the Delta area.

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.

Summary Mounting evidence shows that the functioning and stability of coastal ecosystems often depends critically on habitat-forming foundation species such as seagrasses, mangroves and salt marsh grasses that engage in facultative mutualistic interactions. However, although restoration science is now gradually expanding its long-standing paradigm of minimizing competition to including intraspecific, or within species, facilitation in its designs, the potential of harnessing mutualistic interactions between species for restoration purposes remains uninvestigated. Here, we experimentally tested whether a previously documented mutualism between marsh-forming Spartina alterniflora (cordgrass) and Geukensia demissa (mussels) can increase restoration success in degraded US salt marshes. We found that co-transplanted mussels locally increased nutrients and reduced sulfide stress, thereby increasing cordgrass growth and clonal expansion by 50%. We then removed aboveground vegetation and mussels to simulate a disturbance event and discovered that cordgrass co-transplanted with mussels experienced 3-times greater survival than control transplants. Synthesis and applications. Our findings indicate that mussels amplify cordgrass re-colonization and resilience over spatial and temporal scales that exceed those of their actual mutualistic interaction. By experimentally demonstrating that mutualistic partners can enable foundation species to overcome stress barriers to establish and persist, we highlight that coastal restoration needs to evolve beyond the sole inclusion of intraspecific positive interactions. In particular, we suggest that integrating mutualisms in restoration designs may powerfully enhance long-term restoration success and ecosystem resilience in the many coastal ecosystems where mutualisms involving foundation species are important ecosystem-structuring interactions. This article is protected by copyright. All rights reserved.

Contains fulltext : 207148.pdf (Publisher’s version ) (Open Access) Abstract Foundation species are typically suggested to enhance community diversity non-trophically by increasing habitat structure and mitigating physical stress, while their trophic role is considered of minor importance. Yet, there is little experimental evidence on the relative importance of trophic and non-trophic effects and the interaction with patch size. Here, we transplanted different festoon sizes of living Tillandsia usneoides (Spanish moss) and structural mimics assessing the trophic and non-trophic roles of this habitat-forming epiphyte in mediating the invertebrate community. Compared to bare branches, mimics enhanced species and feeding guild richness and abundances, but living festoons even more so, demonstrating that trophic and non-trophic effects jointly stimulated the community. Specifically, our results show that, independent of patch size, 40% of the total species richness and 46% of total guild richness increase could be contributed to habitat structure alone, while Spanish moss trophically stimulated these metrics by another 60% and 54%. As detritivores were particularly enhanced in living festoons, our findings suggest that trophic stimulation occurred primarily through the provisioning of Spanish moss detritus. Our results highlight that foundation species can facilitate their associated communities through both trophic and non-trophic pathways, calling for studies addressing their indirect trophic role via the brown food web.

Contains fulltext : 207180.pdf (Publisher’s version ) (Open Access) Abstract Coastal exploitation and human-mediated modifications have markedly altered the community composition and functioning of coastal wetlands worldwide. Although recent work has shown that harnessing positive density-dependent feedbacks can greatly enhance the recovery of habitat-modifying species in degraded wetlands, the role of these intraspecific feedbacks in explaining the persistence of altered, unfavorable plant communities remains largely unexplored. Here, we experimentally tested whether intraspecific facilitation may explain the persistence of common reed (Phragmites australis) in human-modified coastal wetlands. We performed a full-factorial mesocosm experiment crossing low-density pioneer versus high-density established development stages with saline (20 psu) versus freshwater conditions. Results showed a clear shift in plant growth response from intraspecific competition under freshwater conditions to self-facilitation in saline treatments. We identified two positive feedback mechanisms enabling the established treatment to overcome salinity stress: (1) Enhanced root oxygenation of the sediment at higher plant density decreased accumulation and intrusion of phytotoxic sulfide, and (2) density-dependent rainwater infiltration into the soil lowered salinity in the dense root mat, preventing salt stress. Our study demonstrates that intraspecific facilitation can be an important factor in explaining the persistence of Phragmites australis in coastal wetlands. We emphasize the importance of integrating positive interactions in coastal restoration but argue that they should either be harnessed when restoring vegetation or broken when eradication of nontarget species is the management objective.

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., Random walk movement patterns with specific step size distributions are commonly associated with resource search optimization strategies in mobile organisms. Here, the authors show that clonal expansion of beach grasses follows a Lévy-type step size strategy that optimizes early dune building.

1. Salt marshes suffered large-scale degradation in recent decades. Extreme events such as hot and dry spells contributed significantly to this, and are predicted to increase not only in intensity, but also in frequency under future climate scenarios. Such repetitive extreme events may generate cumulative effects on ecosystem resilience. It is therefore important to elucidate how marsh vegetation responds to repetitive stress, and whether changes in key species interactions can modulate vegetation resilience.2. In this study, we investigated how moderate but repetitive desiccation events, caused by the combined effects of drought and high temperatures, affect cordgrass (Spartina alterniflora), the dominant habitat-forming grass in southeastern US salt marshes. In a 4-month field experiment, we simulated four consecutive desiccation events by periodically excluding tidal flooding and rainfall, while raising temperature. We crossed this desiccation treatment with the presence/absence of ribbed mussels (Geukensia demissa) - a mutualist of cordgrass known to enhance its desiccation resilience - and with grazing pressure by the marsh periwinkle (Littoraria irrorata) that is known to suppress cordgrass' desiccation resilience.3. We found that each subsequent desiccation event deteriorated sediment porewater conditions, resulting in high salinity (53 ppt), low pH-levels (3.7) and increased porewater Al and Fe concentrations (approximate to 800 mu mol/L and approximate to 1,500 mu mol/L) upon rewetting. No effects on porewater chemistry were found as a result of snail grazing, while ribbed mussels strongly mitigated desiccation effects almost to control levels and increased cordgrass biomass by approximately 128%. Importantly, although cordgrass generally appeared healthy above-ground at the end of the experiment, we found clear negative responses of the repetitive desiccation treatment on cordgrass below-ground biomass, on proline (osmolyte) levels in shoots and on the number of tillers (-40%), regardless of mussel and/or snail presence.4. Synthesis. Even though the mutualism with mussels strongly mitigated chemical effects in the sediment porewater throughout the experiment, mussels could not buffer the adverse ecophysiological effects observed in cordgrass tissue. Our results therefore suggest that although mussels may alleviate desiccation stress, the predicted increased frequency and intensity of hot dry spells may eventually affect saltmarsh resilience by stressing the mutualism beyond its buffering capacity.

Coastal ecosystems are often formed through two-way interactions between plants and their physical landscape. By expanding clonally, landscape-forming plants can colonize bare unmodified environments and stimulate vegetation–landform feedback interactions. Yet, to what degree these plants rely on clonal integration for overcoming physical stress during biogeomorphological succession remains unknown. Here, we investigated the importance of clonal integration and resource availability on the resilience of two European beach grasses (i.e. Elytrigia juncea and Ammophila arenaria) over a natural biogeomorphic dune gradient from beach (unmodified system) to foredune (biologically modified system). We found plant resilience, as measured by its ability to recover and expand following disturbance (i.e. plant clipping), to be independent on the presence of rhizomal connections between plant parts. Instead, resource availability over the gradient largely determined plant resilience. The pioneer species, Elytrigia, demonstrated a high resilience to physical stress, independent of its position on the biogeomorphic gradient (beach or embryonic dune). In contrast, the later successional species (Ammophila) proved to be highly resilient on the lower end of its distribution (embryonic dune), but it did not fully recover on the foredunes, most likely as a result of nutrient deprivation. We argue that in homogenously resource-poor environments as our beach system, overall resource availability, instead of translocation through a clonal network, determines the resilience of plant species. Hence, the formation of high coastal dunes may increase the resistance of beach grasses to the physical stresses of coastal flooding, but the reduced marine nutrient input may negatively affect the resilience of plants. Electronic supplementary material The online version of this article (10.1007/s00442-019-04568-w) contains supplementary material, which is available to authorized users.

Foundation species often interact with each other and co-create habitat upon which other species depend. Whether the presence of these facilitated species feeds back to mediate the growth and resilience of the foundation species themselves, and influence the strength of their interactions, remains poorly understood. In a 16-month field experiment in a southeastern US salt marsh, we tested how the overlapping presence of two foundation species, cordgrass (Spartina alterniflora) and ribbed mussels (Geukensia demissa), influences the abundance of facilitated species, specifically burrowing crabs (mainly Uca pugnax), and how crabs, in turn, affect each foundation species and their mutualistic interaction. Mussel aggregations enhanced crab abundance 3.9-fold, which in turn reduced both mussel and cordgrass growth and stifled cordgrass recovery after a simulated disturbance. Porewater and plant tissue analyses suggest crabs reduced cordgrass growth by reducing nitrogen availability, damaging roots, and potentially interfering with mussel deposition of nutrient-rich pseudofeces. A five-site field survey along 700 km southeastern US coastline revealed that cordgrass biomass and crab abundance are consistently higher in mussel aggregations. Furthermore, cordgrass biomass correlated negatively with crab abundance, supporting our experimental findings and the hypothesis that facilitated biota can negatively impact the foundation species upon which they depend. We anticipate that such negative, but non-lethal, feedbacks between foundation species and the biota they facilitate may be a common but overlooked phenomenon controlling foundation species growth and interactions in a wide range of ecosystems.

1. Grazing can significantly impact spatial heterogeneity and conservation value of ecosystems. Earlier work revealed that overgrazing may stimulate persistent vegetation collapse in low‐productivity environments where vegetation survives by concentrating scarce resources within its local environment. However, it remains unclear whether grazer fluctuations may cause persistent vegetation changes in high‐productivity systems where dense stands facilitate their own survival by hampering grazer access.2. Here, we experimentally tested how the release from grazing by greylag geese (Anser answer) affects spatial vegetation structure in a highly productive, brackish marsh in which dense reed (Phragmites australis) stands and bare roosting areas coexist. Next, we assessed the resilience of the change in vegetation patterning by reintroducing the geese after a 2‐year exclosure period. 3. During herbivore exclusion, vegetation rapidly colonized the bare areas, while reintroduction of herbivores generated a clear species‐specific response. Specifically, the pioneer species, Bolboschoenus maritimus, was immediately eradicated, while the dense and high structure of P. australis facilitated its own persistence by limiting grazer access. Surface accretion (~1 cm/year) during herbivore exclusion further amplified this herbivore‐inhibiting feedback, because greylag geese primarily rely on waterlogged conditions for grubbing. 4. Synthesis and applications. Our results indicate that temporary reductions in herbivore numbers may induce persistent unfavourable changes in the spatial structure of a high‐productivity system. It is therefore important to first assess whether vegetation changes are naturally reversible or persistent. If state shifts are indeed persistent, sufficiently high grazer densities must be maintained to warrant the favourable heterogeneous system. If changes in vegetation structure negatively impact grazer densities, active management such as sod cutting or mowing may be required to restore ecosystem structure and functions.

Seagrass meadows are vital ecosystems in coastal zones worldwide, but are also under global threat. One of the major hurdles restricting the success of seagrass conservation and restoration is our limited understanding of ecological feedback mechanisms. In these ecosystems, multiple, self-reinforcing feedbacks can undermine conservation efforts by masking environmental impacts until the decline is precipitous, or alternatively they can inhibit seagrass recovery in spite of restoration efforts. However, no clear framework yet exists for identifying or dealing with feedbacks to improve the management of seagrass ecosystems. Here we review the causes and consequences of multiple feedbacks between seagrass and biotic and/or abiotic processes. We demonstrate how feedbacks have the potential to impose or reinforce regimes of either seagrass dominance or unvegetated substrate, and how the strength and importance of these feedbacks vary across environmental gradients. Although a myriad of feedbacks have now been identified, the co-occurrence and likely interaction among feedbacks has largely been overlooked to date due to difficulties in analysis and detection. Here we take a fundamental step forward by modelling the interactions among two distinct above- and belowground feedbacks to demonstrate that interacting feedbacks are likely to be important for ecosystem resilience. On this basis, we propose a five-step adaptive management plan to address feedback dynamics for effective conservation and restoration strategies. The management plan provides guidance to aid in the identification and prioritisation of likely feedbacks in different seagrass ecosystems.

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., Intensifying drought has caused massive die-offs in ecosystems worldwide. Here, Angelini et al. use observations, experiments, and models in US salt marshes to show that a key mutualism increases ecosystem resilience by maintaining stress-resistant habitat patches that aid post-drought recovery.

Dense mats of floating vegetation form complex structures that support high biodiversity in wetlands. Since the 1960s, however, high-density stands of Stratiotes aloides have shown strong declines throughout Europe and restoration efforts have often failed, both of which have been linked to high nitrogen (N) input rates. The low success rate of reintroduction is striking, since seemingly healthy, high-density stands are observed to grow under similar environmental conditions. We therefore hypothesise that N tolerance of S. aloides increases with density due to joint N uptake and detoxification. To test our hypothesis, we set up a controlled full factorial mesocosm experiment in which we manipulated N loading and S. aloides density, and investigated interacting effects on habitat biogeochemistry and plant performance. High-density S. aloides stands strongly lowered ammonium availability through shared uptake, even at extremely high N loads. Furthermore, dense stands strongly reduced water layer oxygen concentrations, which stimulated sediment phosphorus mobilisation, and increased carbon dioxide concentrations, which enhanced underwater photosynthesis. High-density stands thus not only detoxify ammonium, but also facilitate population growth through habitat manipulation. Combined, these mechanisms enabled S. aloides to remain viable at high loads of 800 kg N ha − 1 y − 1 , whereas low-density stands already collapsed at 200 kg N ha − 1 y − 1 . Overall, our results show that under similar environmental conditions, high-density stands can thrive, while restoration or natural recruitment by a low number of individuals may be impossible. We therefore conclude that acknowledging and harnessing intraspecific facilitation can be vital for successful conservation and restoration.

Food webs are an integral part of every ecosystem on the planet, yet understanding the mechanisms shaping these complex networks remains a major challenge. Recently, several studies suggested that non-trophic species interactions such as habitat modification and mutual isms can be important determinants of food web structure. However, it remains unclear whether these findings generalize across ecosystems, and whether non-trophic interactions affect food webs randomly, or affect specific trophic levels or functional groups. Here, we combine analyses of 58 food webs from seven terrestrial, freshwater and coastal systems to test (1) the general hypothesis that non-trophic facilitation by habitat-forming foundation species enhances food web complexity, and (2) whether these enhancements have either random or targeted effects on particular trophic levels, functional groups, and linkages throughout the food web. Our empirical results demonstrate that foundation species consistently enhance food web complexity in all seven ecosystems. Further analyses reveal that 15 out of 19 food web properties can be well-approximated by assuming that foundation species randomly facilitate species throughout the trophic network. However, basal species are less strongly, and carnivores are more strongly facilitated in foundation species' food webs than predicted based on random facilitation, resulting in a higher mean trophic level and a longer average chain length. Overall, we conclude that foundation species strongly enhance food web complexity through non-trophic facilitation of species across the entire trophic network. We therefore suggest that the structure and stability of food webs often depends critically on non-trophic facilitation by foundation species.

Restoration has been elevated as an important strategy to reverse the decline of coastal wetlands worldwide. Current practice in restoration science emphasizes minimizing competition between out-planted propagules to maximize planting success. This paradigm persists despite the fact that foundational theory in ecology demonstrates that positive species interactions are key to organism success under high physical stress, such as recolonization of bare substrate. As evidence of how entrenched this restoration paradigm is, our survey of 25 restoration organizations in 14 states in the United States revealed that >95% of these agencies assume minimizing negative interactions (i.e., competition) between outplants will maximize propagule growth. Restoration experiments in both Western and Eastern Atlantic salt marshes demonstrate, however, that a simple change in planting configuration (placing propagules next to, rather than at a distance from, each other) results in harnessing facilitation and increased yields by 107% on average. Thus, small adjustments in restoration design may catalyze untapped positive species interactions, resulting in significantly higher restoration success with no added cost. As positive interactions between organisms commonly occur in coastal ecosystems (especially in more physically stressful areas like uncolonized substrate) and conservation resources are limited, transformation of the coastal restoration paradigm to incorporate facilitation theory may enhance conservation efforts, shoreline defense, and provisioning of ecosystem services such as fisheries production.

By changing habitat conditions, ecosystem engineers increase niche diversity and have profound effects on the distribution and abundances of other organisms. Although many ecosystems contain several engineering species, it is still unclear how the coexistence of multiple engineers affects the physical habitat and the structure of the community on a landscape scale. Here, we investigated through a large-scale field manipulation how three coexisting engineers on intertidal flats (cockles Cerastoderma edule; lugworms Arenicola marina; blue mussels Mytilus edulis) influence the functional composition of the local macrobenthic community and what the consequences are at the landscape level. By using biological trait analysis (BTA), we show that on the local scale biogenic changes in sediment accumulation and organic matter content translated into specific shifts in the distribution of functional traits within the community. At a landscape scale, the co-occurrence of multiple ecosystem engineers resulted in the spatial separation of different functional groups, i.e. different functional groups dominated unique complementary habitats. Our results emphasize the role of co-occurring multiple engineers in shaping natural communities, thus contributing to a better knowledge of community assembly rules. Th is understanding can profitably be used to improve ecosystem-based management and conservation actions.

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.

Ecosystem engineering research has recently demonstrated the fundamental importance of non-trophic interactions for food-web structure. Particularly, by creating benign conditions in stressful environments, ecosystem engineers create hot beds of elevated levels of recruitment, growth, and survival of associated organisms; this should fuel food webs and promote production on the ecosystem scale. However, there is still limited empirical evidence of the influence of non-trophic interactions on the classical food-web processes that determine energy transfer, that is, consumer-resource interactions. On the basis of a biomanipulation experiment covering 600 m2 of an intertidal flat, we show that ecosystem engineers influence resource uptake efficiency and the accumulation of algae following nutrient enrichment in a soft-sediment food web. Nutrient additions increased chlorophyll a concentrations in the sediment by 90%, but only in plots where we also introduced high densities (2000 per m2) of a burrowing bivalve, the common cockle Cerastoderma edule. The artificial cockle beds increased the nutrient uptake efficiency of the biofilm and promoted sediment accumulation, which suggests that the cockles facilitated the sediment-living algae by increasing sediment stability. This indicates that ecological interactions, rather than the availability of nutrients per se, set the limits for production in this coastal ecosystem. Our results emphasize the need to include facilitation theory and recognize that positive interactions between species are key to understand, manage, and restore ecosystems under human influence.

Self-organized spatial patterns occur in many terrestrial, aquatic, and marine ecosystems. Theoretical models and observational studies suggest self-organization, the formation of patterns due to ecological interactions, is critical for enhanced ecosystem resilience. However, experimental tests of this cross-ecosystem theory are lacking. In this study, we experimentally test the hypothesis that self-organized pattern formation improves the persistence of mussel beds (Mytilus edulis) on intertidal flats. In natural beds, mussels generate self-organized patterns at two different spatial scales: regularly spaced clusters of mussels at centimeter scale driven by behavioral aggregation and large-scale, regularly spaced bands at meter scale driven by ecological feedback mechanisms. To test for the relative importance of these two spatial scales of self-organization on mussel bed persistence, we conducted field manipulations in which we factorially constructed small-scale and/or large-scale patterns. Our results revealed that both forms of self-organization enhanced the persistence of the constructed mussel beds in comparison to nonorganized beds. Small-scale, behaviorally driven cluster patterns were found to be crucial for persistence, and thus resistance to wave disturbance, whereas large-scale, self-organized patterns facilitated reformation of small-scale patterns if mussels were dislodged. This study provides experimental evidence that self-organization can be paramount to enhancing ecosystem persistence. We conclude that ecosystems with self-organized spatial patterns are likely to benefit greatly from conservation and restoration actions that use the emergent effects of self-organization to increase ecosystem resistance to disturbance.

Human exploitation of bivalve populations has changed intertidal landscapesworldwide.Many bivalves are ecosystemengineers thatmodify the physical environment, affecting the conditions for their survival. Herewe argue that lack of recovery of overexploited intertidal bivalve beds may be partly caused by the loss of important biological feedbacks from depleted populations. In a large-scale experiment we investigated engineering effects of cockles (Cerastoderma edule L.) and lugworms (Arenicola marina L.) on juvenile cockles by adding high densities of either species to 5 × 5 mplots in areas with different hydrodynamic and sediment conditions in the intertidal flats of the Wadden Sea. We hypothesized that cockles would facilitate the new generations by increasing sediment stability, while lugworms would have negative effects on juvenile cockles through sediment disturbance. We found that in sandy areas with high wave and current energy cockles enhanced sediment accumulationand promoted local densities of young cockles, while lugworms did not have any effect on juvenile cockles. In muddy sites sheltered from the tidal currents by mussel reefs (Mytilus edulis L.), juvenile cockle densities weregenerally high, demonstrating the general importance of biological engineering for recruitment processes in the intertidal. We suggest that the acknowledgement of positive feedbacks between bivalves and sediment stabilityis essential to achieve long-term restoration goals in coastal ecosystems. © 2014 Elsevier

In the Caribbean, green turtles graze seagrass meadows dominated by Thalassia testudinum through rotational grazing, resulting in the creation of grazed and recovering (abandoned) patches surrounded by ungrazed seagrasses. We evaluated the seagrass community and its environment along a turtle grazing gradient; with the duration of (simulated) grazing as a proxy for the level of grazing pressure. The grazing levels consisted of Short-term (4 months clipping), Medium-term (8 months clipping), Long-term grazing (8 months of clipping in previously grazed areas), 8-months recovery of previously grazed patches, and ungrazed or unclipped patches as controls. We measured biomass and density of the seagrasses and rhizophytic algae, and changes in sediment parameters. Medium- and Long-term grazing promoted a shift in community species composition. At increasing grazing pressure, the total biomass of T. testudinum declined, whereas that of early-successional increased. Ammonium concentrations were highest in the patches of Medium-term (9.2 + 0.8 μM) and Long-term grazing levels (11.0 + 2.2 μM) and were lowest in the control areas (4.6 + 1.5 μM). T. testudinum is a late-successional species that maintains sediment nutrient concentrations at levels below the requirements of early-successional species when dominant. When the abundance of this species declines due to grazing, these resources become available, likely driving a shift in community composition toward a higher abundance of early-successional species.

Recent insights suggest that predators should include (mildly) toxic prey when non-toxic food is scarce. However, the assumption that toxic prey is energetically as profitable as non-toxic prey misses the possibility that non-toxic prey have other ways to avoid being eaten, such as the formation of an indigestible armature. In that case, predators face a trade-off between avoiding toxins and minimizing indigestible ballast intake. Here, we report on the trophic interactions between a shorebird (red knot, Calidris canutus canutus ) and its two main bivalve prey, one being mildly toxic but easily digestible, and the other being non-toxic but harder to digest. A novel toxin-based optimal diet model is developed and tested against an existing one that ignores toxin constraints on the basis of data on prey abundance, diet choice, local survival and numbers of red knots at Banc d'Arguin (Mauritania) over 8 years. Observed diet and annual survival rates closely fit the predictions of the toxin-based model, with survival and population size being highest in years when the non-toxic prey is abundant. In the 6 of 8 years when the non-toxic prey is not abundant enough to satisfy the energy requirements, red knots must rely on the toxic alternative.

Increasing evidence shows that spatial interactions between sedentary organisms can structure communities and promote landscape complexity in many ecosystems. Here we tested the hypothesis that reef-forming mussels (Mytilus edulis L.), a dominant intertidal ecosystem engineer in the Wadden Sea, promote abundances of the burrowing bivalve Cerastoderma edule L. (cockle) in neighboring habitats at relatively long distances coastward from mussel beds. Field surveys within and around three mussel beds showed a peak in cockle densities at 50-100 m toward the coast from the mussel bed, while cockle abundances elsewhere in the study area were very low. Field transplantation of cockles showed higher survival of young cockles (2-3 years old) and increased spat fall coastward of the mussel bed compared to within the bed and to areas without mussels, whereas growth decreased within and coastward of the mussel bed. Our measurements suggest that the observed spatial patterns in cockle numbers resulted from (1) inhibition effects by the mussels close to the beds due to preemptive algal depletion and deteriorated sediment conditions and (2) facilitation effects by the mussels farther away from the beds due to reduction of wave energy. Our results imply that these spatial, scale-dependent interactions between reef-forming ecosystem engineers and surrounding communities of sedentary benthic organisms can be an important determinant of the large-scale community structure in intertidal ecosystems. Understanding this interplay between neighboring communities of sedentary species is therefore essential for effective conservation and restoration of soft-bottom intertidal communities.

International audience; 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.

The construction and restoration of riparian (temporarily flooded) wetlands as water storage and flood protection areas plays a central role in climate-adaptive water management. In general, arable and ex-arable lands are used for this type of water storage. However, inundation may lead to problems, as excess phosphorus (P) stored in these soils may be released and result in the eutrophication of the overlying surface waters. Clearly, water and nature managers need to be able to determine for which areas temporary water storage would be a feasible option without causing eutrophication problems. Here, using a controlled experimental approach, a simple predictive tool for the P mobilization rates from soils upon short-term inundation has been developed. A large suite of soil characteristics and P mobilization rates were determined during flooding for different soil types (peat and sand), at two different depths to mimic topsoil removal (topsoils and soils from −30 to −60 cm below ground level), and at two temperatures to test seasonal influence (8 °C and 18 °C). Increasing the temperature from 8 to 18 °C almost tripled P mobilization rates, but the variation could not be linked to any of the soil characteristics measured – average Q10 (temperature coefficient) values were 2.8 (2.9 for peaty soils, 2.6 for sandy soils). Although P mobilization was related to P saturation of amorphic Fe, water-extractable P was found to be by far the best predictor for short-term P mobilization rates, explaining 86.9% of the variation. The predictive tool for P mobilization after short-term rewetting is simple, low-cost and widely applicable, and can support water managers during their decision-making processes concerning the optimal location for the construction of water storage areas, the restoration of riparian wetlands, and the combinational use of different ecosystem services.

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.

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.

In the last decades, eutrophication has become a major cause for concern in aquatic ecosystems worldwide. Apart from external nutrient loading, release of dissolved nutrients from sediment pore water into the water layer contributes importantly to eutrophication. Theory implies that pore water HCO 3 - and nutrient mobility are linked, because anaerobic organic matter breakdown stimulates release of HCO 3 - and nutrients, while HCO 3 - stimulates decay rates by increasing buffer capacity and pH. An investigation was performed on how HCO 3 - relates to commonly measured eutrophication-related variables in freshwater and marine sediments, by analyzing a database of 71 marine and 206 freshwater samples, using linear and non-linear regression. Results demonstrate that HCO 3 - is indeed strongly related with NH 4 + and PO 4 3 - in both freshwater and marine pore waters. Moreover, HCO 3 - also correlated with sulfide levels in marine sediments. Contrastingly, no relationship was found with variables describing total nutrient content (i.e. organic matter, total P and N), suggesting that these do not reveal information on nutrient availability in aquatic sediments.

Shallow soft-sediment systems are mostly dominated by species that, by strongly affecting sediment dynamics, modify their local environment. Such ecosystem engineering species can have either sediment-stabilizing or sediment-destabilizing effects on tidal flats. They interplay with abiotic forcing conditions (wind, tide, nutrient inputs) in driving the community structure and generating spatial heterogeneity, determining the composition of different communities of associated species, and thereby affecting the channelling of energy through different compartments in the food web. This suggests that, depending on local species composition, tidal flats may have conspicuously different geomorphology and biological functions under similar external conditions. Here we use a historical reconstruction of benthic production in the Wadden Sea to construct a framework for the relationships between human impacts, ecosystem engineering and sediment dynamics. We propose that increased sediment disturbances by human exploitation interfere with biological controls of sediment dynamics, and thereby have shifted the dominant compartments of both primary and secondary production in the Wadden Sea, transforming the intertidal from an internally regulated and spatially heterogeneous, to an externally regulated and spatially homogenous system. This framework contributes to the general understanding of the interaction between biological and environmental control of ecosystem functioning, and suggests a general framework for predicting effects of human impacts on soft-bottom ecosystems.

The relative importance of flood pulse dynamics and megafan behaviour for the Sábalo (Prochilodus lineatus) catches in the neotropical Pilcomayo River is studied. The Sábalo catches can mainly be explained by decreased river discharges in the preceding years resulting in smaller inundated areas during rainy season floods and thereby in a decreased area of feeding grounds for the fishes. The decreased river discharges and the related decline of Sábalo catches in the 1990's can be linked to the 90-95 El Niño event. In 2007 the Sábalo catches were comparable to the catches before the "El Niño" event. The connectivity (continuity) between the main river and flood plain areas, which is influenced by sedimentation processes, is also of great importance and very probably plays a more important role since the late 1990's.Se ha estudiado la importancia relativa de la dinámica del pulso de inundación y el comportamiento del sistema megafan para las capturas del Sábalo (Prochilodus lineatus) en el río Pilcomayo. Las capturas del Sábalo puede explicarse fundamentalmente por los bajos caudales del río en los años anteriores, resultando en menores superficies de las zonas inundadas durante la temporada de lluvia y por lo tanto en una área menor de alimentación para los peces. La disminución del caudal del río y la declinación relativa de capturas de Sábalo en los años 1990 pueden estar relacionadas con el 90-95 evento de El Niño. En 2007, la captura de Sábalo fue comparable a las capturas antes del evento de "El Niño". La conectividad (continuidad) entre el río principal y las llanuras aluviales, que son influenciadas por los procesos de sedimentación, son también de gran importancia y muy probablemente juegan un papel muy importante desde finales de los años '90.

Many populations are exposed to naturally occurring or synthetic toxicants. An increasing number of studies demonstrate that the toxicity of such compounds is not only dependent on the concentration or load, but also on the biomass or density of exposed organisms. At high biomass, organisms may be able to alleviate adverse effects of the toxicant by actively lowering ambient concentrations through either a joint detoxification mechanism or growth dilution. We show in a conceptual model that this mechanism may potentially lead to alternative stable states if the toxicant is lethal at low densities of organisms, whereas a high density is able to reduce the toxicant concentrations to sub-lethal levels. We show in an example that this effect may be relevant in real ecosystems. In an earlier published experimental laboratory study, we demonstrated that ammonia toxicity in eelgrass is highly dependent on the eelgrass shoot density. Here, we used the results of these experiments to construct a model describing the complex interactions between the temperate seagrass Zostera marina and potentially lethal ammonia. Analyses of the model show that alternative stable states are indeed present over wide ranges of key-parameter settings, suggesting that the mechanism might be important especially in sheltered, eutrophicated estuaries where mixing of the water layer is poor. We argue that the same mechanism could cause alternative stable states in other biological systems as well.

The spatial structure of seagrass landscapes is typically ascribed to the direct influence of physical factors such as hydrodynamics, light, and sediment transport. We studied regularly interspaced banded patterns, formed by elongated patches of seagrass, in a small-scale intertidal ecosystem. We investigated (1) whether the observed spatial patterns may arise from feedback interactions between seagrass and its abiotic environment and (2) whether changes in abiotic conditions may lead to predictable changes in these spatial patterns. Field measurements, experiments, and a spatially explicit computer model identified a ‘‘scale-dependent feedback’’ (a mechanism for spatial self-organization) as a possible cause for the banded patterns. Increased protection from uprooting by improved anchoring with increasing seagrass density caused a local positive feedback. Sediment erosion around seagrass shoots increased with distance through the seagrass bands, hence causing a long-range negative feedback. Measurements across the depth gradient of the intertidal, together with model simulations, demonstrated that seagrass cover and mean patch size were predictably influenced by additional external stress caused by light limitation and desiccation. Thus, our study provides direct empirical evidence for a consistent response of spatial self-organized patterns to changing abiotic conditions, suggesting a potential use for self-organized spatial patterns as stress indicators in ecosystems.

Seagrasses are threatened by human activity in many locations around the world. Their decline is often characterized by sudden ecosystem collapse from a vegetated to a bare state. In the 1930s, such a dramatic event happened in the Dutch Wadden Sea. Before the shift, large seagrass beds (Zostera marina) were present in this area. After the construction of a large dam and an incidence of the “wasting disease” in the early 1930s, these meadows became virtually extinct and never recovered despite restoration attempts. We investigated whether this shift could be explained as a critical transition between alternative stable states, and whether the lack of recovery could be due to the high resilience of the new turbid state. We analyzed the depth distribution of the historical meadows, a long-term dataset of key factors determining turbidity and a minimal model based on these data. Results demonstrate that recovery was impossible because turbidity related to suspended sediment was too high, probably because turbidity was no longer reduced by seagrass itself. Model simulations on the positive feedback suggest indeed the robust occurrence of alternative stable states and a high resilience of the current turbid state. As positive feedbacks are common in seagrasses, our findings may explain both the worldwide observed collapses and the low success rate of restoration attempts of seagrass habitats. Therefore, appreciation of ecosystem resilience may be crucial in seagrass ecosystem management.

Seagrass meadows are vital ecosystems in coastal zones worldwide, but are also under global threat. One of the major hurdles restricting the success of seagrass conservation and restoration is our limited understanding of ecological feedback mechanisms. In these ecosystems, multiple, self-reinforcing feedbacks can undermine conservation efforts by masking environmental impacts until the decline is precipitous, or alternatively they can inhibit seagrass recovery in spite of restoration efforts. However, no clear framework yet exists for identifying or dealing with feedbacks to improve the management of seagrass ecosystems. Here we review the causes and consequences of multiple feedbacks between seagrass and biotic and/or abiotic processes. We demonstrate how feedbacks have the potential to impose or reinforce regimes of either seagrass dominance or unvegetated substrate, and how the strength and importance of these feedbacks vary across environmental gradients. Although a myriad of feedbacks have now been identified, the co-occurrence and likely interaction among feedbacks has largely been overlooked to date due to difficulties in analysis and detection. Here we take a fundamental step forward by modelling the interactions among two distinct above- and belowground feedbacks to demonstrate that interacting feedbacks are likely to be important for ecosystem resilience. On this basis, we propose a five-step adaptive management plan to address feedback dynamics for effective conservation and restoration strategies. The management plan provides guidance to aid in the identification and prioritisation of likely feedbacks in different seagrass ecosystems.

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.

Mounting evidence indicates that spatial interactions are important in structuring coastal ecosystems. Until recently, however, most of this work has been focused on seemingly exceptional systems that are characterized by regular, self-organized patterns. In this review, we document that interactions that operate at long distances, beyond the direct neighborhood of individual organisms, are more common and have much more far-reaching implications for coastal ecosystems than was previously realized. We review studies from a variety of ecosystem types—including cobble beaches, mussel beds, coral reefs, seagrass meadows, and mangrove forests—that reveal a startling interplay of positive and negative interactions between habitats across distances of up to a kilometer. In addition to classical feeding relations, alterations of physical conditions constitute an important part of these long-distance interactions. This entanglement of habitats has crucial implications for how humans manage coastal ecosystems, and evaluations of anthropogenic impact should explicitly address long-distance and system-wide effects before we deem these human activities to be causing little harm.

This paper reports on experimental restoration of mussel beds in the Wadden Sea and the processes that might limit successful restoration of this foundation species (i.e. substrate, predation, hydrodynamics). The importance of substrate, predation, hydrodynamic conditions and location on mussel restoration success was studied using artificially created mussel beds. Experimental beds established on a stable substrate (coir net) were compared with control beds established on sand, at three locations in the Wadden Sea. Their persistence was followed over time. The results revealed a near disappearance of all experimental beds in just over 7 months. Providing a stable substrate did not improve mussel survival. Predation could not explain the disappearance of the beds, as the maximal predation rate by birds was found to be insufficient to have a significant effect on mussel cover. Differences in wave conditions alone could also not explain the variation in decline of mussel cover between the locations. However, the gradual disappearance of mussels from the seaward side of the bed strongly suggested that hydrodynamic conditions (i.e. combined effects of waves and current) played an important role in the poor persistence of the artificial beds. Our results highlight the fact that restoration of mussel beds in dynamic areas cannot simply be implemented by mussel transplantation, particularly if additional measures to prevent wave losses are not taken, even when artificial substrate is provided to facilitate mussel adhesion. (C) 2015 Elsevier B.V. All rights reserved.

Temperature is one of the most important factors determining growth rates of free-floating macrophytes in the field. To analyse and predict temperature dependent growth rates of these pleustophytes, modelling may play an important role. Several equations have been published for describing temperature responses of macrophytes and algae. But they are often complex or are only applicable in a limited range of temperatures. In this paper, we present a simple three-parameter equation for describing the temperature dependent growth rates of pleustophytes. The equation that we developed is tested using results from laboratory growth experiments conducted with three different species of pleustophytes ( Lemna minor , Salvinia molesta and Azolla filiculoides ). The equation is simple and demonstrates reliable fits (adjusted R 2 reaching from 0.89 to 0.95). Additionally, our equation primarily uses parameters of biological significance, resulting in estimates of useful cardinal temperatures (minimum and maximum).

1. Wilf & Labandeira (1999)suggested that increased temperatures because of global warming will cause an increase in herbivory by insects. This conclusion was based on the supposed effect of temperature on herbivores but did not consider an effect of temperature on plant growth. 2. We studied the effect of temperature on grazing pressure by the small China-mark moth (Cataclysta lemnata L.) on Lemna minor L. in laboratory experiments. 3. Between temperatures of 15 and 24 degrees C we found a sigmoidal increase in C. lemnata grazing rates, and an approximately linear increase in L. minor growth rates. Therefore, an increase in temperature did not always result in higher grazing pressure by this insect as the regrowth of Lemna changes also. 4. At temperatures below 18.7 degrees C, Lemna benefited more than Cataclysta from an increase in temperature, causing a decrease in grazing pressure. 5. In the context of global warming, we conclude that rising temperatures will not necessarily increase grazing pressure by herbivorous insects.

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.

International audience; Ecosystem engineering, the physical modification of the environment by organisms, is a common and often influential process whose significance to food web structure and dynamics is largely unknown. In the light of recent calls to expand food web studies to include non-trophic interactions, we explore how we might best integrate ecosystem engineering and food webs. We provide rationales justifying their integration and present a provisional framework identifying how ecosystem engineering can affect the nodes and links of food webs and overall organization; how trophic interactions with the engineer can affect the engineering; and how feedbacks between engineering and trophic interactions can affect food web structure and dynamics. We use a simple integrative food chain model to illustrate how feedbacks between the engineer and the food web can alter 1) engineering effects on food web dynamics, and 2) food web responses to extrinsic environmental perturbations. We identify four general challenges to integration that we argue can readily be met, and call for studies that can achieve this integration and help pave the way to a more general understanding of interaction webs in nature. Synthesis All species are affected by their physical environment. Because ecosystem engineering species modify the physical environment and belong to food webs, such species are potentially one of the most important bridges between the trophic and non-trophic. We examine how to integrate the so far, largely independent research areas of ecosystem engineering and food webs. We present a conceptual framework for understanding how engineering can affect food webs and vice versa, and how feedbacks between the two alter ecosystem dynamics. With appropriate empirical studies and models, integration is achievable, paving the way to a more general understanding of interaction webs in nature.

Bivalves are key components of coastal ecosystems because they link pelagic and benthic food webs, and shape the landscape through habitat modification. Nevertheless, many bivalve stocks have dramatically declined, and recruitment failure due to (anthropogenically-) increased predation by mesopredators and loss of facilitation mechanisms have been separately hypothesized as underlying causes. Here, we tested the interactive effects of predation and habitat modification on bivalve recruitment in a large-scale experiment in the Wadden Sea, one of the world's largest intertidal soft-sediment ecosystems. We applied anti-erosion mats to simulate biotic attachment and substrate stabilization by commonly found tubeworm beds, crossed this with addition of adult mussels, and manipulated shrimp and crab predation using exclosures within these treatments. Epibenthic mussel recruits were only found in treatments with manipulated substrates, attached to either the anti-erosion mat or adult mussels. Three out of four endobenthic species were facilitated by the mat, but were inhibited by adult mussels. In contrast, invasive surf-dwelling American razor clams were inhibited by both substrate manipulations, indicating a preference for unstable sediments. These facilitation and inhibition effects, however, only clearly emerged when predators were excluded, demonstrating strong synergistic effects between predation and habitat modification. Our findings suggest that disturbance of trophic interactions and loss of habitat modifying species interactively affect bivalve recruitment dynamics in coastal ecosystems. We conclude that conservation and restoration of bivalves should focus on protecting and restoring internal facilitation mechanisms, and should simultaneously reduce excessive mesopredator predation by restoring natural food web dynamics, including the role of top-predators. (C) 2014 Elsevier Ltd. All rights reserved.

The importance of positive effects of ecosystem engineers on associated communities is predicted to increase with environmental stress. However, incorporating such non-trophic interactions into ecological theory is not trivial because facilitation of associated species is conditional on both the type of engineer and the type of abiotic stress. We tested the influence of two allogenic ecosystem engineers (lugworms, Arenicola marina L. and cockles, Cerastoderma edule L.) on the main primary producers (microphytobenthos) of the tidal flats, under different abiotic stresses controlled by reefs of blue mussels (Mytilus edulis L.). We added 25,000 cockles or 2,000 lugworms to 5 x 5 m plots, both in a muddy site with high sedimentation rates located coastward of a mussel bed, and in a sandy site without mussels and characterized by high hydrodynamic stress. After a year, cockles increased algal biomass in the sandy area, but not in the mussel bed site, where high values were measured in all plots. However, lugworms did not affect algal biomass in any of the sites. Field measurements suggest that cockles outweighed negative effects of water currents in the site without mussels by locally increasing sediment stability, whereas mussels overruled the effects of cockles in the wake of the reefs through hydrodynamic stress alleviation and/or biodeposition. Our results suggest that non-trophic interactions by ecosystem engineering bivalves control primary production of intertidal areas, and that the sediment-stabilizing effect of cockles plays a crucial role where the overruling effects of mussel beds are not present.