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The recovery of top predators is thought to have cascading effects on vegetated ecosystems and their geomorphology 1,2 , but the evidence for this remains correlational and intensely debated 3,4 . Here we combine observational and experimental data to reveal that recolonization of sea otters in a US estuary generates a trophic cascade that facilitates coastal wetland plant biomass and suppresses the erosion of marsh edges-a process that otherwise leads to the severe loss of habitats and ecosystem services 5,6 . Monitoring of the Elkhorn Slough estuary over several decades suggested top-down control in the system, because the erosion of salt marsh edges has generally slowed with increasing sea otter abundance, despite the consistently increasing physical stress in the system (that is, nutrient loading, sea-level rise and tidal scour 7-9 ). Predator-exclusion experiments in five marsh creeks revealed that sea otters suppress the abundance of burrowing crabs, a top-down effect that cascades to both increase marsh edge strength and reduce marsh erosion. Multi-creek surveys comparing marsh creeks pre- and post-sea otter colonization confirmed the presence of an interaction between the keystone sea otter, burrowing crabs and marsh creeks, demonstrating the spatial generality of predator control of ecosystem edge processes: densities of burrowing crabs and edge erosion have declined markedly in creeks that have high levels of sea otter recolonization. These results show that trophic downgrading could be a strong but underappreciated contributor to the loss of coastal wetlands, and suggest that restoring top predators can help to re-establish geomorphic stability., (© 2024. The Author(s), under exclusive licence to Springer Nature Limited.)

Ecosystem restoration has traditionally focused on re-establishing vegetation and other foundation species at basal trophic levels, with mixed outcomes. Here, we show that threatened shorebirds could be important to restoring coastal wetland multifunctionality. We carried out surveys and manipulative field experiments in a region along the Yellow Sea affected by the invasive cordgrass Spartina alterniflora. We found that planting native plants alone failed to restore wetland multifunctionality in a field restoration experiment. Shorebird exclusion weakened wetland multifunctionality, whereas mimicking higher predation before shorebird population declines by excluding their key prey - crab grazers - enhanced wetland multifunctionality. The mechanism underlying these effects is a simple trophic cascade, whereby shorebirds control crab grazers that otherwise suppress native vegetation recovery and destabilize sediments (via bioturbation). Our findings suggest that harnessing the top-down effects of shorebirds - through habitat conservation, rewilding, or temporary simulation of consumptive or non-consumptive effects - should be explored as a nature-based solution to restoring the multifunctionality of degraded coastal wetlands., (© 2023. The Author(s).)

The Atlantic ribbed mussel (Geukensia demissa) is common in southeastern US salt marshes, where they form dense aggregations (mounds), that occur in the highest densities and sizes on the marsh platform close to the tidal creeks' heads. Within these marshes, mussels help build marsh elevation via their biodeposition of organic and inorganic material, stimulate the growth of the dominant foundation species cordgrass (Spartina alterniflora), and create hotspots of invertebrate biodiversity, nutrient cycling, and drought resilience. Given their powerful role, there is rising interest in assessing natural variation in the distribution of mussel mounds and using such information to guide marsh conservation and restoration strategies. However, gathering such information is challenging, because the small dimension (∼1 m) of the mounds and the presence of overlying vegetation make it difficult to quantify mound distribution on the marsh. Therefore, this study presents a new procedure to compute the distribution, height, radius, volume, and distance of mounds in marsh environments using remote sensing. A high-resolution UAV-Lidar point cloud has been collected over a highly vegetated salt marsh in Georgia, USA, using a custom-built laser scanner system. An original detection algorithm, based on a Random Forest classifier, has been implemented to identify the mounds from the point cloud. The algorithm has been trained and tested on surveyed mounds and provides their location and geometric properties. Results indicate that the classifier can distinguish mussel mounds from non-mussel mound locations with an accuracy of 95 %. The classifier identified ∼8000 mounds, which occupy 10 % of the study domain, and a volume (shells+feces/pseudofeces) of 680 m 3 . The method is highly useful in efforts to monitor mussel mounds over time and scale up to assess mounds across sites, providing invaluable data for future studies related to the geomorphic evolution of marshes to sea level rise and siting marsh conservation and enhancement projects., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

The fate of coastal ecosystems depends on their ability to keep pace with sea-level rise-yet projections of accretion widely ignore effects of engineering fauna. Here, we quantify effects of the mussel, Geukensia demissa, on southeastern US saltmarsh accretion. Multi-season and -tidal stage surveys, in combination with field experiments, reveal that deposition is 2.8-10.7-times greater on mussel aggregations than any other marsh location. Our Delft-3D-BIVALVES model further predicts that mussels drive substantial changes to both the magnitude (±<0.1 cm·yr -1 ) and spatial patterning of accretion at marsh domain scales. We explore the validity of model predictions with a multi-year creekshed mussel manipulation of >200,000 mussels and find that this faunal engineer drives far greater changes to relative marsh accretion rates than predicted (±>0.4 cm·yr -1 ). Thus, we highlight an urgent need for empirical, experimental, and modeling work to resolve the importance of faunal engineers in directly and indirectly modifying the persistence of coastal ecosystems globally., (© 2023. The Author(s).)

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.

Many wetlands around the world that occur at the base of watersheds are under threat from land-use change, hydrological alteration, nutrient pollution, and invasive species. A relevant measure of whether the ecological character of these ecosystems has changed is the species diversity of wetland-dependent waterbirds, especially those of conservation value. Here, we evaluate the potential mechanisms controlling variability over time and space in avian species diversity of the wetlands in the Palo Verde National Park, a Ramsar Site of international importance in Costa Rica. To do so, we assessed the relative importance of several key wetland condition metrics (i.e., surface water depth, wetland extent, and vegetation greenness), and temporal fluctuations in these metrics, in predicting the abundance of five waterbirds of high conservation value as well as overall waterbird diversity over a 9-yr period. Generalized additive models revealed that mean NDVI, an indicator of vegetation greenness, combined with a metric used to evaluate temporal fluctuations in the wetland extent best predicted four of the five waterbird species of high conservation value as well as overall waterbird species richness and diversity. Black-bellied Whistling-ducks, which account for over one-half of all waterbird individuals, and all waterbird species together were better predicted by including surface water depth along with wetland extent and its fluctuations. Our calibrated species distribution model confidently quantified monthly averages of the predicted total waterbird abundances in seven of the 10 sub-wetlands making up the Ramsar Site and confirmed that the biophysical diversity of this entire wetland system is important to supporting waterbird populations both as a seasonal refuge and more permanently. This work further suggests that optimizing the timing and location of ongoing efforts to reduce invasive vegetation cover may be key to avian conservation by increasing waterbird habitat., (© 2021 The Ecological Society of America.)

Formed at the confluence of marine and fresh waters, estuaries experience both the seaside pressures of rising sea levels and increasing storm severity, and watershed and precipitation changes that are shifting the quality and quantity of freshwater and sediments delivered from upstream sources. Boating, shoreline hardening, harvesting pressure, and other signatures of human activity are also increasing as populations swell in coastal regions. Given this shifting landscape of pressures, the factors most threatening to estuary health and stability are often uncertain. To identify the greatest contemporary threats to coastal wetlands and oyster reefs across the southeastern United States (Mississippi to North Carolina), we summarized recent population growth and land-cover change and surveyed estuarine management and science experts. From 1996 to 2019, human population growth in the region varied from a 17% decrease to a 171% increase (mean = +43%) with only 5 of the 72 SE US counties losing population, and nearly half growing by more than 40%. Individual counties experienced between 999 and 19,253 km 2 of new development (mean: 5725 km 2 ), with 1-5% (mean: 2.6%) of undeveloped lands undergoing development over this period across the region. Correspondingly, our survey of 169 coastal experts highlighted development, shoreline hardening, and upstream modifications to freshwater flow as the most important local threats facing coastal wetlands. Similarly, experts identified development, upstream modifications to freshwater flow, and overharvesting as the most important local threats to oyster reefs. With regards to global threats, experts categorized sea level rise as the most pressing to wetlands, and acidification and precipitation changes as the most pressing to oyster reefs. Survey respondents further identified that more research, driven by collaboration among scientists, engineers, industry professionals, and managers, is needed to assess how precipitation changes, shoreline hardening, and sea level rise are affecting coastal ecosystem stability and function. Due to the profound role of humans in shaping estuarine health, this work highlights that engaging property owners, recreators, and municipalities to implement strategies to improve estuarine health will be vital for sustaining coastal systems in the face of global change., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Consumers are often overlooked as key drivers of vegetation structure and ecosystem functioning in coastal wetlands. This oversight is particularly apparent in Asia, where much of the variation in coastal wetland plant growth and composition is attributed to physical stress gradients. To address this knowledge gap and quantify the relative importance of consumers in Asian coastal wetlands across temporal variation in environmental stress, we conducted a two-year experiment spanning relatively spring wet (2018) and spring dry (2019) years in which we manipulated the presence of the numerically dominant herbivorous crab, Helice tientsinensis, and evaluated its effects on Phragmites australis growth and structure in a Yellow River Delta salt marsh. In spring wetter 2018, Phragmites biomass and stem density were 75% and 34% higher in Crab Exclusion relative to Ambient Crab plots. In 2019 which experienced spring drought and elevated soil salinity, Phragmites biomass and stem density remained similarly high relative to 2018 in Crab Exclusion plots, but fell further, to only 16% and 39% of levels of 2018 observed in Ambient Crab plots. Phragmites' inflorescences density was also significantly reduced in Ambient Crab than Crab Exclusion plots in 2019. Together, these results highlight the significant role that crab herbivores can play in regulating Phragmites in Yellow River Delta salt marshes and suggest that the magnitude of their top-down control may be amplified, although in a non-additive manner, with spring drought stress in the region., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier B.V. All rights reserved.)

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.

Facilitation cascades are chains of positive interactions that occur as frequently as trophic cascades and are equally important drivers of ecosystem function, where they involve the overlap of primary and secondary, or dependent, habitat-forming foundation species [1]. Although it is well recognized that the size and configuration of secondary foundation species' patches are critical features modulating the ecological effects of facilitation cascades [2], the mechanisms governing their spatial distribution are often challenging to discern given that they operate across multiple spatial and temporal scales [1, 3]. We therefore combined regional surveys of southeastern US salt marsh geomorphology and invertebrate communities with a predator exclusion experiment to elucidate the drivers, both geomorphic and biotic, controlling the establishment, persistence, and ecosystem functioning impacts of a regionally abundant facilitation cascade involving habitat-forming marsh cordgrass and aggregations of ribbed mussels. We discovered a hierarchy of physical and biological factors predictably controlling the strength and self-organization of this facilitation cascade across creekshed, landscape, and patch scales. These results significantly enhance our capacity to spatially predict coastal ecosystem function across scales based on easily identifiable metrics of geomorphology that are mechanistically linked to ecological processes. Replication of this approach across vegetated coastal ecosystems has the potential to support management efforts by elucidating the multi-scale linkages between geomorphology and ecology that, in turn, define spatially explicit patterns in community assembly and ecosystem functioning., Competing Interests: Declaration of Interests The authors declare no competing interests., (Copyright © 2020 The Authors. Published by Elsevier Inc. All rights reserved.)

Increasing rates of sea-level rise and wave action threaten coastal populations. Defense of shorelines by protection and restoration of wetlands has been invoked as a win-win strategy for humans and nature, yet evidence from field experiments supporting the wetland protection function is uncommon, as is the understanding of its context dependency. Here we provide evidence from field manipulations showing that the loss of wetland vegetation, regardless of disturbance size, increases the rate of erosion on wave-stressed shorelines. Vegetation removal (simulated disturbance) along the edge of salt marshes reveals that loss of wetland plants elevates the rate of lateral erosion and that extensive root systems, rather than aboveground biomass, are primarily responsible for protection against edge erosion in marshes. Meta-analysis further shows that disturbances that generate plant die-off on salt marsh edges generally hasten edge erosion in coastal marshes and that the erosion protection function of wetlands relates more to lateral than vertical edge-erosional processes and is positively correlated with the amount of belowground plant biomass lost. Collectively, our findings substantiate a coastal protection paradigm that incorporates preservation of shoreline vegetation, illuminate key context dependencies in this theory, and highlight local disturbances (e.g., oil spills) that kill wetland plants as agents that can accelerate coastal erosion., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Foundation species enhance biodiversity and multifunctionality across many systems; however, whether foundation species patch configuration mediates their ecological effects is unknown. In a 6-month field experiment, we test which attributes of foundation species patch configuration - i.e. patch size, total patch area, perimeter, area-perimeter ratio, or connectivity - control biodiversity, stability and multifunctionality by adding a standardised density of mussel foundation species in patches of 1, 5, 10, 30, 60, 90 or 180 individuals to a southeastern US salt marsh. Over 67% of response variables increased with clustering of mussels, responses that were driven by increases in area-perimeter ratio (33%), decreases in perimeter (29%), or increases in patch size (5%), suggesting sensitivity to external stressors and/or dependence on foundation species-derived niche availability and segregation. Thus, mussel configuration - by controlling the relative distribution of multidimensional patch interior and edge niche space - critically modulates this foundation species' effects on ecosystem structure, stability and function., (© 2018 The Authors Ecology Letters published by CNRS and John Wiley & Sons Ltd.)

Climate change and consumer outbreaks are driving ecosystem collapse worldwide. Although much research has demonstrated that these factors can interact, how heterogeneity in top-down control intensity and physical forcing modulates ecosystem resilience to climate stress remains poorly understood. Here, we explore whether the nocturnal herbivorous crab Sesarma reticulatum can control spatially dominant cordgrass (Spartina alterniflora) growth and how its top-down effects vary with crab density, drought stress, and large-scale disturbance in southeastern US salt marshes. In multiple field experiments and surveys, we show that Sesarma depresses cordgrass growth and that its effects increase in a saturating manner with increasing crab density, such that the highest naturally occurring densities of this consumer can trigger local cordgrass die-off. This top-down effect of Sesarma is similar in magnitude to what is thought to be the dominant grazer in the system, the marsh periwinkle snail Littoraria irrorata. In a drought stress by Sesarma density experiment, we further show that salinity stress and intensive crab herbivory additively suppress cordgrass drought resistance. After drought subsides, surveys and experiments reveal that Sesarma also stifles cordgrass re-growth into existing die-off areas. Together, these results show that multiple grazers powerfully regulate the productivity and drought resilience of these intertidal grasslands and that heterogeneity in physical stress and consumer density can dictate when and where top-down forcing is important. More generally, this work provides a rare, experimental demonstration of the critical role top-down control can play across the initiation and recovery stages of ecosystem die-off.

Natural history gave birth to ecology and evolutionary biology, but today its importance is sometimes marginalized. Natural history provides context for ecological research, a concept that we illustrate using a consumer-driven vegetation die-off case study. For three decades, local predator depletion promoted the formation of high-density crab (Sesarma reticulatum) grazing and burrowing fronts, resulting in the spread of vegetation die-off through southern New England and Long Island marshes. We review results from a decade of research on this phenomenon and synthesize these findings with new field surveys, experiments, and historical reconstructions to test the hypothesis that the locations and processes of vegetation die-off and recovery are spatially predictable. We discovered that crab-driven die-off consistently begins on marsh creek heads, where peat and high flow conditions overlap, before spreading to inner creeks following peat availability, stunted cordgrass, and flow. Eventually, die-off eliminates most low marsh vegetation, leaving behind unvegetated substrate too soft to support burrows. Vegetation recovery exhibits the reverse patterns of die-off; it consistently begins in the low marsh within inner creeks, where soft substrate and low flow conditions overlap, before spreading to creek heads. This spatially explicit, substrate-dependent recovery eventually leads to ungrazed cordgrass abutting grazed cordgrass on the high marsh border. We present a conceptual model of die-off through recovery progression to provide managers and landowners with a diagnostic tool for identifying marsh die-off and recovery status. Collectively, this work illustrates the fundamental importance of long-term, natural history-based investigations of ecosystem dynamics in informing ecology, conservation, and management practices.

Climate change and other anthropogenic stressors are converging on coastal ecosystems worldwide. Understanding how these stressors interact to affect ecosystem structure and function has immediate implications for coastal planning, however few studies quantify stressor interactions. We examined past and potential future interactions between two leading stressors on New England salt marshes: sea-level rise and marsh crab (Sesarma reticulatum) grazing driven low marsh die-off. Geospatial analyses reveal that crab-driven die-off has led to an order of magnitude more marsh loss than sea-level rise between 2005 and 2013. However, field transplant experimental results suggest that sea-level rise will facilitate crab expansion into higher elevation marsh platforms by inundating and gradually softening now-tough high marsh peat, exposing large areas to crab-driven die-off. Taking interactive effects of marsh softening and concomitant overgrazing into account, we estimate that even modest levels of sea-level rise will lead to levels of salt marsh habitat loss that are 3x greater than the additive effects of sea-level rise and crab-driven die-off would predict. These findings highlight the importance of multiple stressor studies in enhancing mechanistic understanding of ecosystem vulnerabilities to future stress scenarios and encourage managers to focus on ameliorating local stressors to break detrimental synergisms, reduce future ecosystem loss, and enhance ecosystem resilience to global change.

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., (© 2016 The Author(s).)

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.

Overexploitation of predators has been linked to the collapse of a growing number of shallow-water marine ecosystems. However, salt-marsh ecosystems are often viewed and managed as systems controlled by physical processes, despite recent evidence for herbivore-driven die-off of marsh vegetation. Here we use field observations, experiments, and historical records at 14 sites to examine whether the recently reported die-off of northwestern Atlantic salt marshes is associated with the cascading effects of predator dynamics and intensive recreational fishing activity. We found that the localized depletion of top predators at sites accessible to recreational anglers has triggered the proliferation of herbivorous crabs, which in turn results in runaway consumption of marsh vegetation. This suggests that overfishing may be a general mechanism underlying the consumer-driven die-off of salt marshes spreading throughout the western Atlantic. Our findings support the emerging realization that consumers play a dominant role in regulating marine plant communities and can lead to ecosystem collapse when their impacts are amplified by human activities, including recreational fishing.

Massive anthropogenic and climate-related disturbances are now common in ecosystems worldwide, generating widespread die-off and subsequent community recovery dominated by remnant-patch dynamics rather than open-gap dynamics. Whether communities can recover and, if so, which factors mediate recolonization rate and extent remain unresolved. Here we evaluate recolonization dynamics of southern U.S. salt marshes that experienced extensive, drought-induced die-off of the foundation species Spartina alterniflora over the previous decade. Surveys of Georgia (USA) salt marshes showed little seedling recruitment in die-off areas but persistence of Spartina particularly in large, rather than small, remnant patches. Given this natural variation in remnant patch size, we conducted field experiments to test whether key plant-controlling biotic (grazing, plant neighbor presence) and abiotic (water availability) factors differentially impact Spartina recolonization at small and large-patch scales. In the small-patch (< 1 m2) experiment in 2009, removing grazers and plant neighbors prompted dramatically higher expansion and growth of Spartina relative to controls, while adding freshwater to reduce water limitation had little effect. In contrast, large-patch (> 20 m2 borders advanced significantly over the same time period regardless of grazer or neighbor removal. We continued the large-patch experiments in 2010, a year that experienced drought, and also added freshwater or salt to borders to modify ambient drought stress; overall, borders advanced less than the previous year but significantly more where neighbors were removed or freshwater added. Thus, water availability appears to mediate Spartina recovery by fueling large-patch expansion during wet summers and intensifying interspecific competition during drought. Combined, these findings suggest ecosystems can recover from massive disturbance if remnant foundation species' patches are large enough to overcome biotic inhibition and successfully expand during periods of relaxed abiotic stress.

Foundation species provide habitat to other organisms and enhance ecosystem functions, such as nutrient cycling, carbon storage and sequestration, and erosion control. We focus on freshwater wetlands because these ecosystems are often characterized by foundation species; eutrophication and other environmental changes may cause the loss of some of these species, thus severely damaging wetland ecosystems. To better understand how wetland primary producer foundation species support other species and ecosystem functions across environmental gradients, we reviewed ~150 studies in subtropical, boreal, and temperate freshwater wetlands. We look at how the relative dominance of conspicuous and well-documented species (i.e., sawgrass, benthic diatoms and cyanobacteria, Sphagnum mosses, and bald cypress) and the foundational roles they play interact with hydrology, nutrient availability, and exposure to fire and salinity in representative wetlands. Based on the evidence analyzed, we argue that the foundation species concept should be more broadly applied to include organisms that regulate ecosystems at different spatial scales, notably the microscopic benthic algae that critically support associated communities and mediate freshwater wetlands' ecosystem functioning. We give recommendations on how further research efforts can be prioritized to best inform the conservation of foundation species and of the freshwater wetlands they support. [ABSTRACT FROM AUTHOR]