Restoration ecology meets design-engineering:Mimicking emergent traits to restore feedback-driven ecosystems

Ecosystems shaped by habitat-modifying organisms such as reefs, vegetated coastal systems and peatlands, provide valuable ecosystem services, such as carbon storage and coastal protection. However, they are declining worldwide. Ecosystem restoration is a key tool for mitigating these losses but has proven failure-prone, because ecosystem stability often hinges on self-facilitation generated by emergent traits from habitat modifiers. Emergent traits are not expressed by the single individual, but emerge at the level of an aggregation: a minimum patch-size or density-threshold must be exceeded to generate self-facilitation. Self-facilitation has been successfully harnessed for restoration by clumping transplanted organisms, but requires large amounts of often-limiting and costly donor material. Recent advancements highlight that kickstarting self-facilitation by mimicking emergent traits can similarly increase restoration success. Here, we provide a framework for combining expertise from ecologists, engineers and industrial product designers to transition from trial-and-error to emergent trait design-based, cost-efficient approaches to support large-scale restoration.</p

Restoration ecology meets design-engineering: Mimicking emergent traits to restore feedback-driven ecosystems

Ecosystems shaped by habitat-modifying organisms such as reefs, vegetated coastal systems and peatlands, provide valuable ecosystem services, such as carbon storage and coastal protection. However, they are declining worldwide. Ecosystem restoration is a key tool for mitigating these losses but has proven failure-prone, because ecosystem stability often hinges on self-facilitation generated by emergent traits from habitat modifiers. Emergent traits are not expressed by the single individual, but emerge at the level of an aggregation: a minimum patch-size or density-threshold must be exceeded to generate self-facilitation. Self-facilitation has been successfully harnessed for restoration by clumping transplanted organisms, but requires large amounts of often-limiting and costly donor material. Recent advancements highlight that kickstarting self-facilitation by mimicking emergent traits can similarly increase restoration success. Here, we provide a framework for combining expertise from ecologists, engineers and industrial product designers to transition from trial-and-error to emergent trait design-based, cost-efficient approaches to support large-scale restoration

Water level drawdown induces a legacy effect on the seed bank and retains sediment chemistry in a eutrophic clay wetland

The lack of extreme water level fluctuations in managed, non-peat forming wetland ecosystems can result in decreased productivity through the loss of heterogeneity of these ecosystems. Stochastic disruption, such as a water level drawdown, can effectively reverse this effect and return the wetland to a more productive state, associated with higher biodiversity through new vegetation development. Yet, aside from the effect on vegetation dynamics, little is known about longer-term effects (30 years) of a water level drawdown, hereafter referred to as legacy effects, and how this may impact future water level drawdowns. Here, we aim to unravel the legacy effects of a water level drawdown, stand alone and along a water level gradient, on seed bank properties and nutrient availability in a eutrophic clay wetland. To identify these, we studied the hydrologically managed nature reserve Oostvaardersplassen in the Netherlands. Here, one section was subjected to a multi-year water level drawdown and another section was kept inundated. We determined seed bank properties in both areas, spatially and along a soil elevation gradient (20 cm). Nutrient availability was measured by taking sediment samples along the water level gradient and through experimental manipulation of the water level in an indoor mesocosm experiment. Germination was higher in locations with a water level drawdown history, especially at relatively high elevations. Additionally, the proportion of pioneer species in the seed bank was higher in the water level drawdown area. Overall, nutrient concentrations were higher compared to other aquatic systems. Nutrient availability was higher in the inundated area and did not respond to the water level gradient. We conclude that 30 years after an induced water level drawdown there is no depletion of nutrients, while we still observe a legacy effect in the number of viable seeds in the seed bank

Growth forms and life-history strategies predict the occurrence of aquatic macrophytes in relation to environmental factors in a shallow peat lake complex

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

Life cycle informed restoration:Engineering settlement substrate material characteristics and structural complexity for reef formation

Ecosystems are degrading world-wide, with severe ecological and economic consequences. Restoration is becoming an important tool to regain ecosystem services and preserve biodiversity. However, in harsh ecosystems dominated by habitat-modifying organisms, restoration is often expensive and failure prone. Establishment of such habitat modifiers often hinges on self-facilitation feedbacks generated by traits that emerge when individuals aggregate, causing density- or patch size-dependent establishment thresholds. To overcome these thresholds, adult or juvenile habitat-forming species are often transplanted in clumped designs, or stress-mitigating structures are deployed. However, current restoration approaches focus on introducing or facilitating a single life stage, while many habitat modifiers experience multiple bottlenecks throughout their life as they transition through sequential life stages. Here, we define and experimentally test ‘life cycle informed restoration’, a restoration concept that focuses on overcoming multiple bottlenecks throughout the target species’ lifetime. To provide proof of concept, and show its general applicability, we carried out complementary experiments in intertidal soft-sediment systems in Florida and the Netherlands where oysters and mussels act as reef-building habitat modifiers. We used biodegradable structures designed to facilitate bivalve reef recovery by both stimulating settlement with hard and fibrous substrates and post-settlement survival by reducing predation. Our trans-Atlantic experiments demonstrate that these structures enabled bivalve reef formation by: (a) facilitating larval recruitment via species-specific settlement substrates, and (b) enhancing post-settlement survival by lowering predation. In the Netherlands, structures with coir rope most strongly facilitated mussels by providing fibrous settlement substrate, and predation-lowering spatially complex hard attachment substrate. In Florida, oysters were greatly facilitated by hard substrates, while coir rope proved unbeneficial. Synthesis and applications. Our findings demonstrate that artificial biodegradable reefs can enhance bivalve reef restoration across the Atlantic by mimicking emergent traits that ameliorate multiple bottlenecks over the reef-forming organism’ life cycle. This highlights the potential of our approach as a cost-effective and practical tool for nature managers to restore systems dominated by habitat modifiers whose natural recovery is hampered by multiple life stage-dependent bottlenecks. Therefore, investment in understanding how to achieve life cycle informed restoration on larger scales and whether the method it is applicable to restore other ecosystems is now required

Coastal restoration success via emergent trait-mimicry is context dependent

Coastal ecosystems provide vital ecosystem functions and services, but have been rapidly degrading due to human impacts. Restoration is increasingly considered key to reversing these losses, but is often unsuccessful. Recent work on seagrasses and salt marsh cordgrasses highlights that restoration yields can be greatly enhanced by temporarily mimicking key emergent traits. These traits are not expressed by individual seedlings or small clones, but emerge in clumped individuals or large clones to locally suppress environmental stress, causing establishment thresholds where such density-dependent self-facilitation is important for persistence. It remains unclear, however, to what extent the efficacy of restoration via emergent trait-based mimicry depends on the intensity of stressors. We test this in a restoration experiment with the temperate seagrass Zostera marina at four sites (Finland, Sweden, UK, USA) with contrasting hydrodynamic regimes, where we simulated dense roots mats or vegetation canopies with biodegradable structural mimics. Results show that by mimicking sediment-stabilizing root mats, seagrass transplant survival, growth and expansion was strongly enhanced in hydrodynamically exposed environments. However, these positive effects decreased and turned negative under benign conditions, while mimics insufficiently mitigated physical stress in extremely exposed environments, illustrating upper and lower limits of the application. Furthermore, we found that aboveground structures, designed to mimic stiff rather than flexible vegetation canopies, underperformed compared to belowground mimics. Our findings emphasize the importance of understanding the conditions at the restoration site, species-specific growth requirements, and self-facilitating traits that organisms may express when applying emergent trait-mimicry as a tool to improve restoration success

Initiating and upscaling mussel reef establishment with life cycle informed restoration:Successes and future challenges

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. Our results 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

Mimicry of emergent traits amplifies coastal restoration success

Contains fulltext : 221224.pdf (publisher's version ) (Open Access

Impacts of shelter on the relative dominance of primary producers and trophic transfer efficiency in aquatic food webs : Implications for shallow lake restoration

Wind-induced turbulence can strongly impact ecological processes in shallow lake ecosystems. The creation of shelter against wind can be expected to affect both primary producers and herbivores in aquatic food webs. Shelter may benefit particular primary producers more than others by changing relative resource availabilities for different primary producers. Herbivore community compositions may be affected either directly or indirectly as a consequence of changes in their food quantity and quality that, in turn, may affect the transfer efficiency between primary producers and herbivores. A reduction in trophic transfer efficiency resulting from wind-induced turbulence potentially can lead to declines of higher trophic levels, but is generally understudied. Here, we focus on the impact of wind on aquatic primary producers and trophic transfer efficiency. We hypothesised that reducing wind-induced turbulence will stimulate higher trophic production in shallow lakes. However, the multitude of impacts of wind-induced turbulence on aquatic food webs make it challenging to predict the direction of change when creating sheltered conditions. We tested our hypothesis in the shallow waters of a newly constructed archipelago named Marker Wadden in lake Markermeer in the Netherlands. Lake Markermeer has experienced declining numbers of birds and fish. These declines have been related to wind-induced sediment resuspension that potentially limits primary production and trophic transfer efficiency. Marker Wadden is a large-scale restoration project that aims to add sheltered and heterogeneous habitat to the otherwise mostly homogeneous lake, thus targeting the potential problems associated with wind-induced turbulence. We executed a 2-month manipulative field mesocosm experiment in the shallow waters of Marker Wadden to study the effect of reduced wind-induced turbulence (i.e., shelter) on aquatic food webs. Specifically, we studied the effects on primary producers, trophic transfer efficiency between phytoplankton and zooplankton (using zooplankton biomass divided by phytoplankton Chl a as a proxy), and benthic fauna. The experiment consisted of three treatments: no shelter, shelter without macrophytes and shelter with submerged macrophytes (Myriophyllum spicatum) present at the start of the experiment. Our results clearly showed that under unsheltered conditions phytoplankton was the dominant primary producer, whereas in sheltered conditions submerged macrophytes became dominant. Interestingly, submerged macrophytes appeared rapidly in the sheltered treatment where first no macrophytes were visibly present; hence, at the end of the experiment, there was little difference among the sheltered treatments with and without initial presence of submerged macrophytes. Despite that phytoplankton concentrations were 23-fold higher under the unsheltered conditions, this did not result in higher zooplankton biomass. This can be explained by a five-fold greater trophic transfer efficiency between phytoplankton and zooplankton under the sheltered conditions. Furthermore, under the sheltered conditions the Gastropoda density reached 746 individuals m−2, whereas no Gastropoda were found under the no shelter treatment. These findings indicate that for shallow lakes that are negatively affected by wind-induced turbulence, measures aimed at ameliorating this stressor can be effective in facilitating submerged macrophyte recovery, increasing Gastropoda densities and restoring trophic transfer efficiency between phytoplankton and zooplankton. Ultimately, this may support higher trophic levels such as fish and water birds by increasing their food availability in shallow lake ecosystems

Impacts of shelter on the relative dominance of primary producers and trophic transfer efficiency in aquatic food webs: Implications for shallow lake restoration

Wind-induced turbulence can strongly impact ecological processes in shallow lake ecosystems. The creation of shelter against wind can be expected to affect both primary producers and herbivores in aquatic food webs. Shelter may benefit particular primary producers more than others by changing relative resource availabilities for different primary producers. Herbivore community compositions may be affected either directly or indirectly as a consequence of changes in their food quantity and quality that, in turn, may affect the transfer efficiency between primary producers and herbivores. A reduction in trophic transfer efficiency resulting from wind-induced turbulence potentially can lead to declines of higher trophic levels, but is generally understudied. Here, we focus on the impact of wind on aquatic primary producers and trophic transfer efficiency. We hypothesised that reducing wind-induced turbulence will stimulate higher trophic production in shallow lakes. However, the multitude of impacts of wind-induced turbulence on aquatic food webs make it challenging to predict the direction of change when creating sheltered conditions. We tested our hypothesis in the shallow waters of a newly constructed archipelago named Marker Wadden in lake Markermeer in the Netherlands. Lake Markermeer has experienced declining numbers of birds and fish. These declines have been related to wind-induced sediment resuspension that potentially limits primary production and trophic transfer efficiency. Marker Wadden is a large-scale restoration project that aims to add sheltered and heterogeneous habitat to the otherwise mostly homogeneous lake, thus targeting the potential problems associated with wind-induced turbulence. We executed a 2-month manipulative field mesocosm experiment in the shallow waters of Marker Wadden to study the effect of reduced wind-induced turbulence (i.e., shelter) on aquatic food webs. Specifically, we studied the effects on primary producers, trophic transfer efficiency between phytoplankton and zooplankton (using zooplankton biomass divided by phytoplankton Chl a as a proxy), and benthic fauna. The experiment consisted of three treatments: no shelter, shelter without macrophytes and shelter with submerged macrophytes (Myriophyllum spicatum) present at the start of the experiment. Our results clearly showed that under unsheltered conditions phytoplankton was the dominant primary producer, whereas in sheltered conditions submerged macrophytes became dominant. Interestingly, submerged macrophytes appeared rapidly in the sheltered treatment where first no macrophytes were visibly present; hence, at the end of the experiment, there was little difference among the sheltered treatments with and without initial presence of submerged macrophytes. Despite that phytoplankton concentrations were 23-fold higher under the unsheltered conditions, this did not result in higher zooplankton biomass. This can be explained by a five-fold greater trophic transfer efficiency between phytoplankton and zooplankton under the sheltered conditions. Furthermore, under the sheltered conditions the Gastropoda density reached 746 individuals m−2, whereas no Gastropoda were found under the no shelter treatment. These findings indicate that for shallow lakes that are negatively affected by wind-induced turbulence, measures aimed at ameliorating this stressor can be effective in facilitating submerged macrophyte recovery, increasing Gastropoda densities and restoring trophic transfer efficiency between phytoplankton and zooplankton. Ultimately, this may support higher trophic levels such as fish and water birds by increasing their food availability in shallow lake ecosystems