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Due to the major declines of seagrasses worldwide, there is an urgent need for effective restoration methods and strategies. In the Dutch Wadden Sea, intertidal seagrass restoration has proven very challenging, despite numerous restoration trials with different restoration methods. Recently, however, the first field trial performed with a newly developed “dispenser injection seeding” method (DIS) resulted in record-high plant densities and seed recruitment. Here, we present the further development of the methodology and consequently improved restoration results. During two consecutive growing seasons, we honed the seeding technique and experimentally investigated how seeding depth (2/4 cm), injection density (25/100 injects/m2), and seed amount (2/20 seeds/inject) affected restoration of intertidal annual Zostera marina. We found that all variables had a significant impact on plant establishment. Seeding deeper (4 cm) had the largest positive effect on restored plant densities, while lowered seed densities (2 seeds/inject) had the largest positive impact on seed recruitment. The optimized DIS method, combined with an altered placement of the seeding hole, resulted in a 50-fold increase in restored plant densities (from approximately 1 to 57 plants/m2) and a simultaneous increase in seed recruitment (from 0.3 to 11.4%). These improvements stem from the method's ability to counteract a recruitment bottleneck, where seeds are lost through hydrodynamic forcing. The methodological improvements described here are important steps toward restoring self-sustaining seagrass populations in the future and our study demonstrates the high potential of the seed-based DIS method for seagrass restoration.

Coastal ecosystem restoration is often ineffective and expensive in practice. As a consequence, upscaling restoration efforts to functionally relevant spatial scales remains one of the largest hurdles for coastal restoration practice. On small scales, restoration success of vegetated ecosystems (i.e. salt marshes and seagrasses) can be amplified by spatial designs that harness positive interactions. However, it remains unknown if positive interactions can be harnessed with seed-based approaches, that are considered to be more cost-effective and scalable than traditional shoot-based restoration methods.Here, we investigated with a full-factorial seeding experiment if (1) restoration scale (4, 40 and 400 m2) and (2) seeding density (10 and 50 injections/m2) affected multi-year recruitment efficiency (measured as restored plants/seed injection) of annual eelgrass Zostera marina in the Dutch Wadden Sea.We found that the largest restoration scale (400 m2) increased second-generation recruitment efficiency by suppressing a sedimentation-related negative feedback. With increased restoration scale, the inner parts of the restoration plots captured less sediment, which decreased the desiccation stress of the restored eelgrass during low tide. Due to this stress alleviation, plants grew larger and produced more seed-bearing spathes, which the following year resulted in two and three times higher recruitment efficiency at the largest restoration scale compared to the smaller scales. Moreover, lower seeding density more than doubled second-generation recruitment efficiency compared to the higher density, supporting recent work showing that the effectiveness of ‘clumped’ spatial designs is context dependent.Synthesis and applications. The efficiency of restoration efforts is seldom taken into account, but can offer restoration projects a valuable metric with which workload, donor material and cost-requirements can be reduced. We demonstrate that simple modifications to seed-based coastal restoration designs (e.g. scale and density) can have a substantial impact on recruitment efficiency and multi-year restoration yields. Thus, optimised restoration designs can strongly contribute to the upscaling potential of coastal ecosystem restoration. However, optimal restoration designs are expected to be strongly context dependent and we therefore argue that investigating optimal designs should be adopted as common practice, providing a crucial steppingstone between ‘proof-of-concepts’ and true large-scale restoration attempts (km2).

Coastal ecosystems provide vital services, including water storage, carbon sequestration, biodiversity, and coastal protection. Human disturbances, however, cause massive losses. The most direct impact is habitat destruction through infrastructure development, restricting the space available to coastal ecosystems and impeding their capacity to adapt to sea level rise by landward retreat – a phenomenon called ‘coastal squeeze’. While shoreline retreat is intensively studied, coastal congestion through infrastructure remains unquantified. Here we calculated the distance to the nearest human-made structure along 263,900 transects worldwide to show that infrastructure occurs at a 560-meter median distance from the shoreline. Moreover, we find that 18% of global sandy shores harbour less than 100 m of infrastructure-free space, and that 14-17% of the unimpacted space may drown by 2100 according to sea level rise projections. Further analyses show that population density and gross domestic product explain 40-44% of observed squeeze variation, emphasizing the intensifying pressure imposed as countries develop and populations expand. Encouragingly, we find that nature reserves relieve squeezing by 3-5 times, illustrating their effectiveness. Yet, at present only 16% of world’s sandy shores has a protected status. We therefore argue that expansion of nature reserves could be key to preserving coastal resilience to sea level rise.

Reef-forming species form integral aspects of coastal ecosystems, but are rapidly degrading world-wide. To mitigate these declines, nature managers increasingly rely on the restoration of habitat-structuring, reef-forming species by, for example, introducing artificial reefs that may directly function as complex reef habitat. Since the use of biodegradable structures to restore biogenic reefs is becoming a popular technique, its effectiveness as reef habitat must be assessed. Therefore, we examine the trophic complexity on experimental large-scale biodegradable artificial reefs using food web network analysis. We placed biodegradable artificial reefs on soft-sediment intertidal flats in the Dutch Wadden Sea in a large-scale (~650 m) and 2.5-year-long experiment. We compared food web networks and biodiversity indicators between biodegradable reefs and bare controls and quantified species composition inside and near the artificial reef community to assess the expansion of the reef community. During 2.5 years, we observed that artificial reefs changed food web networks compared to bare controls: in species richness (+76%), link density (the number of interactions per species; +15%) and the fraction of basal species (species of lowest trophic level; +40%), but lowered the connectance: the realized fraction of all possible links between species (−33%). Their effects on food web networks increased over time with a higher species richness (+22%) and more complex food web (link density +13%) on the artificial reef 2.5 years after deployment compared to 1.5 years. However, the effects of the reefs did not extend beyond the reef structures; the species composition and biodiversity of macrozoobenthos near the reefs were comparable to the control. Synthesis and applications. This study shows that biodegradable artificial reefs offer an effective tool for the restoration of food web complexity and biodiversity of intertidal soft-sediment systems. However, application needs to be carefully considered as the reef-building species did not expand beyond our structures, despite the ambitious spatial extent of this experiment. Therefore, we recommend restoration practitioners to design artificial reefs in such a way that they generate ecosystem connectivity (facilitation of higher trophic levels) and biogeomorphological effects on a landscape scale (reef expansion beyond the structures).

13 pages, 5 figures.-- This is an open access article under the terms of the Creative Commons Attribution License, Coastal ecosystem functioning often hinges on habitat-forming foundation species that engage in positive interactions (e.g. facilitation and mutualism) to reduce environmental stress. Seagrasses are important foundation species in coastal zones but are rapidly declining with losses typically linked to intensifying global change-related environmental stress. There is growing evidence that loss or disruption of positive interactions can amplify coastal ecosystem degradation as it compromises its stress mitigating capacity. Multiple recent studies highlight that seagrass can engage in a facultative mutualistic relationship with lucinid bivalves that alleviate sulphide toxicity. So far, however, the generality of this mutualism, and how its strength and relative importance depend on environmental conditions, remains to be investigated. Here we study the importance of the seagrass-lucinid mutualistic interaction on a continental-scale using a field survey across Europe. We found that the lucinid bivalve Loripes orbiculatus is associated with the seagrasses Zostera noltii and Zostera marina across a large latitudinal range. At locations where the average minimum temperature was above 1 °C, L. orbiculatus was present in 79% of the Zostera meadows; whereas, it was absent below this temperature. At locations above this minimum temperature threshold, mud content was the second most important determinant explaining the presence or absence of L. orbiculatus. Further analyses suggest that the presence of the lucinids have a positive effect on seagrass biomass by mitigating sulphide stress. Finally, results of a structural equation model (SEM) support the existence of a mutualistic feedback between L. orbiculatus and Z. noltii. We argue that this seagrass-lucinid mutualism should be more solidly integrated into management practices to improve seagrass ecosystem resilience to global change as well as the success of restoration efforts, JdF was supported by NWO Open Competition #ALWOP.203 and TvdH by NWO-Vidi Career no. 16588

Despite active seagrass restoration gaining traction as a tool to halt and reverse worldwide seagrass losses, overall success remains limited. Restoration strategies, through seeding or transplantation, face different environmental bottlenecks that limit success. Choosing the most appropriate strategy of the two for a specific location, however, is hampered by lack of direct practical comparisons between strategies within a single system. To investigate potential life stage dependent bottlenecks, we compared seed-based and transplant-based restoration of Zostera marina in the subtidal saltwater Lake Grevelingen. Our results demonstrate that seedling recruitment was negatively impacted by bioturbation from the lugworm Arenicola marina and sediment movement due to hydrodynamic exposure. Transplant-based restoration was clearly more successful butsurprisingly best predicted by leaf gluing by the ragworm Platynereis dumerilii. This previously undescribed interaction caused seagrass leaves to clump and reduce effective photosynthetic surface and leaf movement. We suggest that the observed behavior of these worms may result from a lack of foodweb interactions, illustrating the importance of trophic control for seagrass restoration. Thus, in addition to recognizing life stage dependent environmental bottlenecks for restoration strategy selection, seagrass restoration may also require the active recovery of their associated food webs.

Vegetated coastal ecosystems such as salt marshes, dunes and seagrass meadows occur at the land–sea interface—a dynamic environment typified by harsh growing conditions. These ecosystems are known as biogeomorphic landscapes because their functioning depends on biophysical interactions by which organisms engineer landforms to their own benefit. The strength of such biogeomorphic feedbacks depends on plant traits, such as stem flexibility and shoot density. Recent work demonstrated that dune grasses with similar morphological traits can build contrasting landscapes due to differences in their spatial shoot organization. However, in contrast to dune grasses that trap and stabilize sand particles in aeolian landscapes, flow attenuation in aquatic environments can generate scouring around plant stems and cause uprooting, leading to establishment thresholds for young plants. Yet, it remains unknown how findings from aeolian landscapes translate to aquatic systems and how young clonally expanding plants in hydrodynamically exposed conditions overcome these establishment thresholds by optimizing shoot placement. Here, we measured shoot patterns of 90 establishing cordgrass patches Spartina anglica at 18 European field sites that cover a broad range of hydrodynamic conditions. Next, we carried out a field experiment to investigate how observed spatial shoot patterns affect plant–sediment feedbacks. Surprisingly, field survey analyses reveal highly consistent clustered shoot patterns, regardless of environmental conditions. Experimental results demonstrate that this clustered pattern minimizes scouring compared to densely clumped organizations typically observed in established patches. Synthesis. In contrast to earlier findings highlighting that establishing dune grasses optimize their landscape engineering capacity via a flexible shoot placement strategy, we find that cordgrass instead follows a fixed strategy that minimizes engineering effects in its early life stages. We suggest that marsh grasses avoid physical stress and associated establishment thresholds in their early life stage, and switch to an ecosystem engineering strategy once established. These findings shed new light on how plant traits interact with their environment to shape the landscape and pave the way for improved restoration designs by mimicking the natural shoot organization of establishing vegetation.

Contains fulltext : 294468.pdf (Publisher’s version ) (Open Access) 1. Coastal ecosystems are generally controlled by the combination of bottom-up (resourcedriven) and top-down (consumer-driven) trophic, and non-trophic interactions. Anthropogenic disruption of these interactions, e.g., through eutrophication or overfishing, leads to loss of the foundation species composing these ecosystems. Within degraded ecosystems, new interactions may become dominant, hampering restoration of foundation species. 2. We demonstrate this concept in saltwater Lake Grevelingen, where seagrass restoration was seriously hampered by a newly discovered non-trophic interaction between the seagrass Zostera marina and the ragworm Platynereis dumerilii. Platynereis constructs dwelling tubes between seagrass leaves by gluing and entangling them, leading to loss of biomass and increased epiphyte cover. To test the roles of epiphyte grazing (top-down) and eutrophication (bottom-up) on this newfound interaction, we executed a field experiment introducing the snail Littorina littorea under natural nutrient concentrations and Platynereis densities, and performed a full factorial mesocosm experiment applying Littorina, Platynereis and nutrient treatments. 3. Our field results show that survival of reintroduced Zostera in Lake Grevelingen is seriously hampered by Platynereis, which can be suppressed by grazer addition. 4. In our mesocosm experiment we demonstrate that top-down control by Littorina protected seagrass through two pathways: (1) mitigating bottom-up regulated epiphyte growth by trophic top-down forcing and (2) hampering the non-trophic interaction between Zostera and Platynereis by grazing epiphytes and dwelling tubes. 5. Synthesis and applications. With coastal ecosystems globally under pressure, our study emphasises the importance of understanding ecosystem-specific trophic and non-trophic key interactions when considering restoration. We show how negative interspecific interactions can become dominant in the absence of key facilitating interactions. Next, we show that negative interactions (leaf gluing), inhibiting foundation species restoration, can be overruled by the inclusion of positive interspecific interactions (epiphyte grazing by mesograzers). Finally, we provide a three-step co-introduction approach for restoration management. 10 mei 2023

Aquatic ecosystems provide vital services, and macrophytes play a critical role in their functioning. Conceptual models indicate that in shallow lakes, plants with different growth strategies are expected to inhabit contrasting habitats. For shallow peat lakes, characterized by incohesive sediments, roles of growth forms, life-history strategies and environmental factors in determining the occurrence of aquatic vegetation remain unknown. In a field survey, we sampled 64 points in a peat lake complex and related macrophyte occurrence to growth forms (floating-leaved rooted and submerged), life-history strategies for overwintering (turions, seeds, rhizomes) and environmental factors (water depth, fetch, and porewater nutrients). Our survey showed that macrophyte occurrence relates to water depth, wind-fetch, and nutrients, and depends on growth form and life-history strategies. Specifically, rooted floating-leaved macrophytes occur at lower wind-fetch/shallower waters. Submerged macrophytes occur from low to greater wind-fetch/water depth, depending on life-history strategies; macrophytes with rhizomes occur at greater wind-fetch/depth relative to species that overwinter with seeds or turions. We conclude that growth form and life-history strategies for overwintering predict macrophytes occurrence regarding environmental factors in peat lakes. Therefore, we propose an adapted model for macrophyte occurrence for such lakes. Altogether, these results may aid in species-selection to revegetate peat lakes depending on its environment.

Marine ecosystem engineers such as seagrasses and bivalves create important coastal habitats sustaining high biodiversity and ecosystem services. Restoring these habitats is difficult due to the importance of feedback mechanisms that can require large-scale efforts to ensure success. Incorporating facilitative interactions could increase the feasibility and success of small-scale restoration efforts, which would limit pressure on donor sites and reduce costs and time associated with restoration. Here, we tested two methods for providing facilitation in small-scale eelgrass (Zostera marina) restoration plots across northern Europe: (1) co-restoration with blue mussels (Mytilus edulis, M. trossulus); and (2) the use of biodegradable establishment structures (BESEs). Eelgrass-mussel co-restoration showed promise in aquaria, where eelgrass growth was nearly twice as high in treatments with medium and high mussel densities than in treatments without mussels. However, this did not translate to higher shoot length or shoot densities in subsequent field experiments. Rather, hydrodynamic exposure limited both eelgrass and mussel survival, especially in the most exposed sites. The use of BESEs showed more potential in enabling small-scale restoration success: they effectively enhanced eelgrass survival and reduced mussel loss, and showed potential for enabling mussel recruitment in one site. However, eelgrass planted in BESE plots along with mussels had a lower survival rate than eelgrass planted in BESE plots without mussels. Overall, we show that though co-restoration did not work at small scales, facilitation by using artificial structures (BESEs) can increase early eelgrass survival and success of small-scale eelgrass and bivalve restoration.

In degraded landscapes, recolonization by pioneer vegetation is often halted by the presence of persistent environmental stress. When natural expansion does occur, it is commonly due to the momentary alleviation of a key environmental variable previously limiting new growth. Thus, studying the circumstances in which expansion occurs can inspire new restoration techniques, wherein vegetation establishment is provoked by emulating natural events through artificial means. Using the salt‐marsh pioneer zone on tidal flats as a biogeomorphic model system, we explore how locally raised sediment bed forms, which are the result of natural (bio)geomorphic processes, enhance seedling establishment in an observational study. We then conduct a manipulative experiment designed to emulate these facilitative conditions in order to enable establishment on an uncolonized tidal flat. Here, we attempt to generate raised growth‐promoting sediment bed forms using porous artificial structures. Flume experiments demonstrate how these structures produce a sheltered hydrodynamic environment in which suspended sediment and seeds preferentially settle. The application of these structures in the field led to the formation of stable, raised sediment platforms and the spontaneous recruitment of salt‐marsh pioneers in the following growing season. These recruits were composed primarily of the annual pioneering Salicornia genus, with densities of up to 140 individuals/m2 within the structures, a 60‐fold increase over ambient densities. Lower abundances of five other perennial species were found within structures that did not appear elsewhere in the pioneer zone. Furthermore, recruits grew to be on average three times greater in mass inside of the structures than in the neighboring ambient environment. The success of this restoration design may be attributed to the combination of three factors: (1) enhanced seed retention, (2) suppressed mortality, and (3) accelerated growth rates on the elevated surfaces generated by the artificial structures. We argue that restoration approaches similar to the one shown here, wherein the conditions for natural establishment are actively mimicked to promote vegetation development, may serve as promising tools in many biogeomorphic ecosystems, ranging from coastal to arid ecosystems.

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

Whether pristine ecosystems with intact trophic structures are more resilient to anthropogenic pressures than ecosystems facing human exploitation is a pressing question to ecological theory and management. For coastal vegetated ecosystems such as kelp forests and seagrass meadows, a number of studies recently highlighted that under eutrophic conditions, trophic cascades are particularly important in buffering or reducing negative effects of nutrient enrichment. Yet, it currently remains unclear how nutrient enrichment and trophic downgrading interact in oligotrophic coastal ecosystems with an intact trophic structure. Here, we factorially manipulated nutrient loading and the interactions within a tri-trophic food chain within a pristine, oligotrophic seagrass ecosystem to investigate how trophic downgrading affects its ability to buffer against eutrophication. Results revealed that nutrient addition stimulates seagrass production, while reducing the growth of benthic microalgae, presumably because the thicker seagrass canopy reduced light availability. Trophic downgrading by excluding predators, however, almost completely negated these nutrient effects, as the release of herbivores from predation strongly enhanced grazing pressure on seagrass. Exclusion of grazers in turn restored seagrass biomass by allowing the nutrient addition treatment to regain its effect, confirming that a tri-trophic cascade mediated how enhanced nutrient loading affected our system. Our results highlight that a healthy trophic structure is vital for the ability of pristine coastal ecosystems to buffer eutrophication, as trophic downgrading may degrade their ability to absorb enhanced nutrient loading. Hence, our findings emphasize the potential for multiple anthropogenic impacts to interact synergistically in degrading natural ecosystems.

1. Facilitation of foundation species is critical to the structure, function and persistence of ecosystems. Understanding the dependence of the strength of this facilitation on environmental conditions is important for informed ecosystem management and for predicting the impacts of global change. In coastal seagrass habitats, chemosymbiotic lucinid bivalves can facilitate seagrasses by decreasing potentially toxic levels of sulphide in sediment porewater. However, variation in the strength of lucinid–seagrass facilitation with environmental context has not been experimentally investigated.2. We tested the hypothesis that the presence of the tiger lucine Codakia orbicularis becomes more important to the growth and survival of the seagrass Thalassia testudinum under decreased light availability and increased sulphide stress. In a mesocosm experiment, we reduced average ambient‐light to T. testudinum by 64% and/or increased sediment porewater sulphide concentrations by ~200% and comparedgrowth and tissue chemistry of T. testudinum with and without C. orbicularis. 3. We found that T. testudinum was better able to maintain growthunder shading and sulphide stress when C. orbicularis was present. C. orbicularis strongly decreased sediment porewater sulphide, an effect that minimized sulphur build‐up in seagrass tissue and was likely achieved through bioirrigation as well as chemoautotrophy. The relative effects of C. orbicularis on T. testudinum growth were strongest in the presence of environmental stressors. 4. Synthesis. The strength of lucinid–seagrass facilitation increases under environmental conditions that hinder the ability of seagrass to detoxify sulphide. Our results provide evidence of a potential mechanism by which the spatiotemporal association between lucinids and seagrasses is maintained and support the incorporation of interspecific facilitation into conservation and restoration strategies for foundation species in the face of increasing anthropogenic impact and global change.

Seagrass meadows are threatened biodiversity hot spots that provide essential ecosystem services. Green sea turtles may overgraze meadows, further enhancing seagrass decline. However, we observed an unexpected, remarkable recovery of seagrasses in a previously overgrazed meadow with abundant unattached branched coralline algae, suggesting that turtle grazing had ceased. We hypothesize that this recovery is due to an effective grazing-protection mutualism, in which the spiny coralline algae structures protect the seagrass meadows from overgrazing, while the seagrasses protect the algae from removal by currents and waves. Removing coralline algae from recovered seagrass plots allowed the turtles to resume grazing, while addition of coralline algae to grazed plots caused cessation of grazing. Coralline algae that were placed on bare sand were quickly displaced by wave action, whereas those placed in grazed or ungrazed seagrass remained. Our experiments demonstrate a grazing-protection mutualism, which likely explains the witnessed recovery of an overgrazed seagrass meadow. To our knowledge, this is the first account of a plant–plant grazing-protection mutualism in an aquatic environment. Our findings show that grazing-protection mutualisms can be vital for the maintenance and recovery of ecosystems shaped by habitat-structuring foundation species, and highlight the importance of mutualisms in coastal ecosystems. As seagrasses, sea turtles and coralline algae share habitats along tropical shores worldwide, the mutualism may be a global phenomenon. Overgrazing is expected to increase, and this mutualism adds a new perspective to the conservation and restoration of these valuable ecosystems.

In soft-bottom marine ecosystems, bedform variation is induced by wind- and tidal-driven hydrodynamics. The resulting megaripples, sand waves and sandbanks form a spatially and temporally heterogeneous seafloor landscape. The strong physical forces imposed by the migration of these bedforms are important determinants for the occurrence of different macrozoobenthic species. Quantifying the effect of these forces can help in differentiating natural- and anthropogenically induced physical stressors. However, large-scale mapping of seabed morphology at high resolution using multibeam echosounder is challenging, costly and time-consuming, especially in shallow seas, prohibiting wide swaths. Instead, their bathymetry is typically studied using single-beam transects that are interpolated to bathymetric grids with a relatively coarse resolution (20 m). However, this leaves out information on smaller scale (versus anthropogenic disturbances such as dredging and bottom trawling activities. We discuss that by repurposing already available single-beam data in this way, bedforms can be characterized at high resolution without the need for additional equipment or mapping campaigns, yielding novel input to decision-making on marine management and conservation.

Worldwide, coastal ecosystems are rapidly degrading in quality and extent. While novel restoration designs include facilitation to enhance restoration success in stressful environments, they typically focus on a single life-stage, even though many organisms go through multiple life-stages accompanied by different bottlenecks. A new approach – life cycle informed restoration – was designed to ameliorate multiple bottlenecks throughout an organism's life cycle. It has successfully been tested on a small scale to facilitate intertidal bivalve reef formation in the Netherlands and Florida. Yet, it remains unknown whether this approach can be scaled to ecosystem-relevant scales. To test whether life cycle informed restoration is upscalable, we conducted a large-scale restoration experiment using blue mussel reefs as a model system. In our experiment, we used biodegradable structures to temporarily facilitate mussel reef formation by providing early-life settlement substrates, and subsequently, reduce post-settlement predation on an intertidal flat in the Wadden Sea, the Netherlands. The structures were placed in 10 × 20 m plots, mimicking bands found in natural mussel beds, spread out across 650 m, and were followed for two years. Ourresults show that the structures enhance mussel biomass (0.7 ± 0.2 kg DW m −2), as mussels were absent in bare plots. However, biomass varied within plots; in intact structures it was 60 times higher (1.2 ± 0.2 kg DW m−2) than in those that became buried (0.02 ± 0.009 kg DW m−2). Next to burial, 18–46% of the structures were lost due to technical failure, especially during winters at this exposed site. We show that the life cycle informed restoration principle works, but we encountered technical challenges due to larger scale processes (e.g. sedimentation). Furthermore, environmental information is essential for site selection, and for restoration, the functioning of such structures should be tested under extreme conditions before upscaling.

Restored shellfish reefs provide valuable habitat for fish, but it is not clear how different approaches affect performance, and either promote the development of new fish populations (i.e. “production”) or simply attract individuals from the broader seascape (i.e. “attraction”). We measured the effects of a 1.5 ha shellfish reef restoration site on fish assemblages in Pumicestone Passage in eastern Australia, which contains replicates of six different restoration units: shell patch reefs, crates of shells, and biodegradable matrix, and each had replicates with and without live oysters. Fish were surveyed before restoration and then every 6 months for 30 months with baited (at restoration and control sites) and unbaited (at 106 sites across the seascape to detect potential fish attraction, and at the different restoration units) underwater cameras. Shellfish reef restoration represents an addition to the carrying capacity of Pumicestone Passage for fish for two key reasons. First, restoration significantly enhanced the diversity and abundance of fish assemblages and the density of harvestable fish at the restoration site by 3.8, 10.7, and 16.4 times, respectively. Second, fish distributions across the broader seascape did not change in response to succession at the restoration site. Fish assemblages did not differ between restoration units or the presence or absence of oysters. These findings further support the notion that restored shellfish reefs significantly enhance fish abundance and diversity and that restored reefs can enhance the overall carrying capacity of seascapes for fish, rather than simply centralizing them at restoration sites.

Habitat fragmentation into small patches is regarded as a vital cause of biodiversity loss. Fragmentation of habitat-forming species is especially harmful, as patchiness of such species often controls ecosystem stability and resilience by density and patch size-dependent self-reinforcing feedbacks. Although fragmentation are expected to weaken or even break such feedbacks, it remains unclear how the resulting patchiness of habitat-forming species affect ecosystem resilience to environmental stresses. Here, using Spartian alterniflora, the habitat-forming species in saltmarshes as a model, we investigate how patch size, plant density, and shell aggregation interactively control the persistence of a degrading salt marsh that suffered from erosion induced by hydrodynamics. Our results demonstrate that large patches can trap more shells along the patch edge than the smaller ones, therefore significantly facilitating plant re-growth within the patch. Shell removal experiments further reveal that large patches trapping more shells along patch edges reinforce their own persistence by decreasing erosion and thus facilitating plant recovery. By contrast, small patches with lesser plants cannot persist as they trap less shells along patch edges but are able to accumulate more shells at interior locations where they hinder plant re-growth, indicating a critical threshold of patch size ~20 m2 below which ecosystem collapses. The current study highlights the importance to identify critical threshold of stress-resistant patch sizes in transition-prone ecosystems as early-warning to alert undesired ecosystem collapse and restoration practice.

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

Ecosystems worldwide depend on habitat‐forming foundation species that often facilitate themselves with increasing density and patch size, while also engaging in facultative mutualisms. Anthropogenic global change (e.g., climate change, eutrophication, overharvest, land‐use 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, nonbinding reciprocal interactions) affect ecosystem resilience is uncertain. Here, we synthesize recent advancements and combine these 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 stress from anthropogenic impact. 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., 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. Here, we synthesize recent advancements and combine this with model analyses to propose that facultative mutualisms can increase foundation species resistance to global change stressors, but simultaneously increase the risk of sudden collapse by generating or exacerbating bistability.

In biogeomorphic landscapes, plant traits can steer landscape development through plant‐mediated feedback interactions. Interspecific differences in clonal expansion strategy can therefore lead to the emergence of different landscape organisations. Yet, whether landscape‐forming plants adopt different clonal expansion strategies depending on their physical environment remains to be tested. Here, we use a field survey and a complementary mesocosm approach to investigate whether sediment deposition affects the clonal expansion strategy employed by dune‐building marram grass individuals. Our results reveal a consistent shift in expansion pattern from more clumped, Brownian‐like, movement in sediment‐poor conditions, to patchier, Lévy‐like, movement under high sediment supply rates. Additional model simulations illustrate that the sediment‐dependent shift in movement strategies induces a shift in optimisation of the cost–benefit relation between landscape engineering (i.e. dune formation) and expansion. Plasticity in expansion strategy may therefore allow landscape‐forming plants to optimise their engineering ability depending on their physical landscape., Clonal dune grasses can express different movement strategies that determine their ecosystem engineering ability. Here we show that dune grasses can switch between local movement patterns – indicating low engineering strength – to multi‐scale movement patterns – signifying high engineering efficiency – when sediment deposition is low or high respectively. This environment‐dependent trait expression has important consequences for our understanding of coastal landscape formation and resilience under global change.

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.