Salt marshes fronting coastal structures, such as seawalls and dikes, may offer important ecosystem-based coastal defence by reducing the wave loading and run-up levels during storms. We question (i) how the long-term salt marsh development in the Dutch Wadden Sea relates to the tidal-flat foreshore bathymetry and (ii) how the wave run-up onto dikes, which enhances the risk of dike failure, depends on foreshore bathymetry, the presence/absence of marshes, marsh vegetation properties, tidal range and wind exposure. We analysed 15 years of vegetation and bathymetry maps along the entire Dutch Wadden Sea coast, in combination with detailed process-based measurements at five locations during 3 years, to understand where salt marshes naturally form and what features determine their contribution to coastal protection. The horizontal extent of marshes along the dikes remained relatively stable over the past decade. The presence of marshes was associated with higher elevations of adjacent tidal flats (above ~0.5 m NAP), while landward-directed marsh retreat was associated with surface erosion of the fronting tidal flats. Wave run-up during storms was lower at sites with wider marshes and higher foreshore elevations. This was attributed to the marsh attenuation effect, which led to a reduction in wave heights at the dike toe. As the tidal range varies across the Dutch Wadden Sea, areas to the East with generally higher water levels experienced higher wave run-up. Synthesis and applications. We found that (i) marshes, where present, effectively protected the dikes from wave loading and (ii) the sites where marshes typically do not develop spontaneously were the most vulnerable to high wave run-up. This catch-22 problem implies that increasing reliance on nature-based coastal defences along soft-bottom coasts may require human interventions to stimulate marsh formation at the locations where it is most needed. Alternatively, ‘hard engineering’ solutions may remain necessary where implementing nature-based solutions are either too costly, unachievable, or at the expense of other ecological values, such as causing the loss of mudflats that are important for migratory birds.

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Marine reef ecosystems have degraded massively worldwide, and restoration efforts have as yet not managed to realize the scale required to reverse continued degradation. To achieve effective scales, scientific insights in restoration methods should be paired with industry-based approaches used for infrastructural development. We illustrate by five principles how long-standing experience of marine contractors with executing large-scale projects, can support reef restoration: i) utilizing industrial techniques to achieve positive impact at scale, ii) landscaping infrastructure to optimize habitat for targeted species, iii) inducing life to overcome connectivity bottle-necks and steer community composition, iv) designing nature development efforts to be self-sustainable, and v) ensuring continuity beyond project boundaries by early stakeholder engagement. Consciously connecting scientific knowledge to industry-based activities increases the likelihood that marine infrastructure development and ecosystem rehabilitation can be aligned. We plead that synergizing practices by science and industry is needed to upscale restoration efforts and truly improve marine reef ecosystems.@en

Global change processes such as sea level rise and the increasing frequency of severe storms threaten many coastlines around the world and trigger the need for interventions to make these often densely-populated areas safer. Mangroves could be implemented in Nature-Based Flood Defense, provided that we know how to conserve and restore these ecosystems at those locations where they are most needed. In this study, we investigate how best to restore mangroves along an aquaculture coast that is subject to land-subsidence, comparing two common mangrove restoration methods: 1) mangrove restoration by planting and 2) Ecological Mangrove Restoration (EMR); the assistance of natural mangrove regeneration through mangrove habitat restoration. Satellite data revealed that historically, landward mangrove expansion into the active pond zone has mainly occurred through mangrove planting on pond bunds. However, there is potential to create greenbelts along waterways by means of EMR measures, as propagule trap data from the field revealed that propagules of pioneer species were up to 21 times more abundant in creeks of the pond zone than near their source in the coastal zone. This was especially true during the prevailing onshore winds of the wet-season, suggesting that smart seasonal sluice gate management could help to efficiently trap seeds in target ponds. In the coastal zone, field experiments showed that permeable brushwood dams, aimed at expanding mangrove habitat, could not sufficiently overcome subsidence rates to increase natural mangrove expansion in the seaward direction, but did significantly increase the survival of already established (planted) seedlings compared to more wave-exposed sites. The survival and growth rate of EMR-supported plantings greatly varied between species. Out of the four planted species, Rhizophora mucronata had the highest survival (67%) but the lowest growth rate. Whereas the pioneer species Avicennia alba and Avicennia marina had lower survival rates (resp. 35 and 21%), but significantly higher growth rates, even resulting in fruiting young trees within a 16-month timeframe. Overall, we conclude that 1) EMR has potential in the pond zone, given that propagules were observed to reach well into the backwaters; and 2) that mangrove recovery in the coastal zone may be facilitated even at very challenging coastal sites by combining EMR with the planting of pioneer species.

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The risk of coastal storm flooding is deteriorating under global warming, especially for the heavily urbanized deltaic cities, like Shanghai. The Nature-Based Flood Defense (NBFD), as an eco-friendly design alternative for hard infrastructure against coastal flooding, is gaining attention. Nevertheless, the vulnerability of saltmarsh due to the biological instability, resulting in the uncertainties on coastal protection, is considered the bottleneck challenge that hinders the broad application of the NBFD concept. We argue that except for direct wave attenuations by the above-ground vegetation during storms, the gradual sediment trapping and consolidating during the non-storm period is a more crucial function of coastal saltmarsh, which mitigates storm waves by forming a broader and higher intertidal morphology. This benefit is an important value of saltmarsh-based coastal protection but is largely neglected in many NBFD studies. Taking Chongming Dongtan Shoal (CDS) as a case study, we demonstrated that over 2/3th wave attenuation during storms is contributed by the saltmarsh morphology, and less than 1/3th is from the saltmarsh vegetation. The relative contribution of the saltmarsh morphology on wave mitigation is even enhanced under the increasing storm grades from 100 yrs. to 5000 yrs. return levels. To promote this idea for broader application, the cost-benefit analysis of three artificial NBFD solutions (e.g., submerged breakwater, timber piles, and sand nourishment) are compared. We identified an optimal measure of the submerged breakwater for CDS, which minimizes the ecological impact and maximizes the cost-benefit. Moreover, the wave-free zone behind the breakwater increases the chance of vegetation establishment, helps suspended sediment trapping, hence fostering a beneficent cycle for saltmarsh restoration. In summary, ignoring the contribution of saltmarsh morphology on wave attenuation largely underestimated the benefits of vegetation-based coastal protection, which should be greatly emphasized to provide a solid basis for developing NBFD.

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Nature-based strategies, such as wave attenuation by tidal marshes, are increasingly proposed as a complement to mitigate the risks of failure of engineered flood defense structures such as levees. However, recent analysis of historic coastal storms revealed smaller dike breach dimensions if there were natural, high tidal marshes in front of the dikes. Since tidal marshes naturally only experience weak flow velocities (~0-0.3 ms^-1 during normal spring tides), we lack direct observations on the stability of tidal marsh sediments and vegetation under extreme flow velocities (order of several ms^-1) as may occur when a dike behind a marsh breaches. As a first approximation, the stability of a tidal marsh sediment bed and winter-state vegetation under high flow velocities were tested in a flume. Marsh monoliths were excavated from Phragmites australis marshes in front of a dike along the Scheldt estuary (Dutch-Belgian border area) and installed in a 10 m long flume test section. Both sediment bed and vegetation responses were quantified over 6 experimental runs under high flow velocities up to 1.75 ms^-1 and water depth up to 0.35 m for 2 hours. These tests showed that even after a cumulative 12 hours exposure to high flow velocities, erosion was limited to as little as a few millimeters. Manual removal of the aboveground vegetation did not enhance the erosion either. Present findings may be related to the strongly consolidated, clay- and silt-rich sediment and P. australis root system in this experiment. During the flow exposure, the P. australis stems were strongly bent by the water flow, but the majority of all shoots recovered rapidly when the flow had stopped. Although present results may not be blindly extrapolated to all other marsh types, they do provide a strong first indication that marshes can remain stable under high flow conditions, and confirm the potential of well-developed tidal marshes as a valuable extra natural barrier reducing flood discharges towards the hinterland, following a dike breach. These outcomes promote the consideration to implement tidal marshes as part of the overall flood defense and to rethink dike strengthening in the future.

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Conservation and restoration of salt marsh ecosystems are becoming increasingly important because of the many ecosystem services they provide. However, the processes controlling salt marsh establishment and persistence, especially on bare tidal flats in muddy areas, remain unclear. As muddy sediments typically experience a restriction of soil drainage, we expect that a surface drainage relief due to a heterogeneity topography, as might occur on the edge of tidal channels, can facilitate the establishment of salt marsh vegetation on muddy tidal flats. By means of a manipulative field experiment, using “Mega-Marsh Organ” mesocosms, we investigated the impact of surface drainage and elevation relative to mean sea level on (1) the survival of Spartina anglica seedlings from three different age classes: 1-yr, 3-month, and 1-week; and (2) the growth performance of mature S. anglica marsh tussocks. S. anglica seedling survival, especially in the establishment phase, was positively affected by better surface drainage, increases of seedling age, and higher elevation relative to mean sea level. That is, the survival rate of S. anglica seedlings at the end of 6th week increased from 0% (at surface water undrained, 1-week, 0 cm elevation) to 94.44% (at surface water drained, 1-yr, 90 cm elevation). In contrast, surface drainage did not affect the performance of large S. anglica marsh tussocks, as only increased elevation relative to mean sea level was shown to affect S. anglica tussock growth in terms of plant height, shoot numbers, and dry biomass. Based on our findings, we proposed a conceptual model to understand how surface drainage-driven feedbacks in a heterogeneous topography may be reinforced to induce salt marsh establishment in muddy systems. Further testing of present hypothesized model would be beneficial for insights into salt marsh establishment on tidal mudflats.

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Despite the well-recognized importance of plant traits for biogeomorphic development of landscapes, our understanding remains limited of how species-specific plant traits respond to and serve as drivers for the sedimentary dynamics within a biogeomorphic landscape. By manipulating a series of laboratory experiments, using mesocosms and a flume, we examined how species-specific differences in expansion strategy, i.e., clonal step-length of laterally expanding tillers, both respond to sediment type and drive cliff formation and persistence. We compared three marsh species, with contrasting clonal expansion traits, that are dominant in many estuaries worldwide: Spartina anglica, Scirpus maritimus, and Phragmites australis. Our results revealed that S. anglica tussocks tend to have high shoot density due to a short clonal expansion step-length, whereas S. maritimus tussocks were much more diffuse and tend to have a longer clonal expansion step-length. P. australis showed intermediate traits. Clonal expansion step-length did show within-species variation in response to sediment texture, but species-specific differences remained the most important. Species with smaller clonal step-lengths, such as S. anglica in this study, are more likely to induce cliffs at the marsh edge by driving formation of larger cliff heights and also having lower capacity to grow out from cliffs. Our findings thus illustrate how dynamic landscape features like cliffs at marsh edges depend on the clonal expansion traits of the dominant species. This enhances current understanding of the formation and development of marsh edges, and is instructive for understanding the role of species-specific traits in driving distinct biogeomorphic landscape dynamics.

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BACKGROUND AND AIMS: It is important to have an in-depth mechanistic understanding of tidal marsh establishment and dynamics to ensure the long-term persistence of these valuable ecosystems. As wave forcing may be expected to impact seedling establishment, we studied the effect of water-imposed drag forces on seedling survival, morphology and biomechanical properties of three marsh pioneer species that are dominant along the salinity gradient in many areas around the world: Spartina anglica (salt to brackish), Scirpus maritimus (brackish) and Phragmites australis (brackish to fresh). METHODS: Using a newly developed plant-shaking mesocosm (PSM) that mimicked water-imposed wave drag forces, the effect of wave stress on seedling survival was examined, together with impacts on morphology and biomechanical properties. KEY RESULTS: After 7 weeks of exposure to wave stress, lowered seedling survival and growth for all species was revealed. Wave treatments increased the root/shoot biomass ratio to enhance anchorage and made seedlings more flexible (i.e. reduced flexural rigidity), which might be regarded as a mixed outcome between a stress avoidance and stress tolerance strategy. CONCLUSIONS: The different biomechanical responses between the three dominant marsh pioneer species, overall, make them less resistant to external stress. Therefore, our results indicate that the likelihood of marshes becoming established is reduced if wave energy increases. Despite the different biomechanical response of these three pioneer species to waves, the seedlings of all species were found to have low resistance to external stresses.@en

BACKGROUND AND AIMS: It is important to have an in-depth mechanistic understanding of tidal marsh establishment and dynamics to ensure the long-term persistence of these valuable ecosystems. As wave forcing may be expected to impact seedling establishment, we studied the effect of water-imposed drag forces on seedling survival, morphology and biomechanical properties of three marsh pioneer species that are dominant along the salinity gradient in many areas around the world: Spartina anglica (salt to brackish), Scirpus maritimus (brackish) and Phragmites australis (brackish to fresh). METHODS: Using a newly developed plant-shaking mesocosm (PSM) that mimicked water-imposed wave drag forces, the effect of wave stress on seedling survival was examined, together with impacts on morphology and biomechanical properties. KEY RESULTS: After 7 weeks of exposure to wave stress, lowered seedling survival and growth for all species was revealed. Wave treatments increased the root/shoot biomass ratio to enhance anchorage and made seedlings more flexible (i.e. reduced flexural rigidity), which might be regarded as a mixed outcome between a stress avoidance and stress tolerance strategy. CONCLUSIONS: The different biomechanical responses between the three dominant marsh pioneer species, overall, make them less resistant to external stress. Therefore, our results indicate that the likelihood of marshes becoming established is reduced if wave energy increases. Despite the different biomechanical response of these three pioneer species to waves, the seedlings of all species were found to have low resistance to external stresses.@en

Foreshores consisting of both bare tidal flats and vegetated salt marshes are found worldwide and they are well studied for their wave attenuating capacity. However, most studies only focus on the small scale: just some isolated locations in space and only up to several years in time. In order to stimulate the implementation of foreshores serving as reliable coastal defense on a large scale, we need to quantify the decadal wave attenuating capacity of the foreshore on the scale of an estuary. To study this, a unique bathymetrical dataset is analyzed, covering the geometry of the Westerschelde estuary (The Netherlands) over a time-span of 65 years. From this dataset, six study sites were extracted (both sheltered sites and exposed sites to the prevailing wind direction) and divided into transects. This resulted in 36 transects covering the entire foreshore (composed of the bare tidal flat and the vegetated salt marsh). The wave attenuation of all transects under daily conditions (with and without vegetation) and design conditions (i.e. events statistically occurring once every 10,000 years) was modelled. Overall, the spatial variability of the geometry of a single foreshore was observed to be much larger than the temporal variability. Temporal changes in salt marsh width did not follow the variability of the entire foreshore. Both under daily and design conditions, vegetation contributes to decreasing wave energy and decreases the variability of incoming wave energy, thereby decreasing the wave load on the dike. The southern foreshores, sheltered from the prevailing wind direction, were more efficient in wave attenuation than the exposed northern foreshores. A linear relation between marsh width and wave attenuation over a period of 65 years was observed at all marshes. The present study provides insights needed to calculate the length of a salt marsh to obtain a desired minimum wave attenuating capacity.

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Globally, erosion of muddy tropical coasts that are dominated by aquaculture ponds, is an increasing problem. Restoration of mangrove greenbelts may counteract such erosion, by restoring the sediment balance. Hence, we aim to unravel the processes controlling natural mangrove regeneration in both “landward” (i.e., into aquaculture ponds) and seaward direction, using the fast eroding coastline of Demak (Indonesia) as case study. Firstly, we investigated which physical and chemical factors drive landward mangrove expansion by relating them to the presence/absence of mangrove seedlings in abandoned aquaculture ponds. Secondly, we investigated which physical parameters control seaward mangrove expansion by relating them to expansion and retreat at the sea-side of mature mangrove stands. Landward mangrove expansion into abandoned aquaculture ponds was positively related to both emergence time (%) and sediment stability (i.e., shear strength), which are in turn both associated to bed level elevation and pond drainage. Surprisingly, there was no effect of soil chemistry. Seaward expansion of existing mangrove stands was strongly associated to foreshore morphology. Mangroves only expanded in the presence of an elevated mudflat, whereas the absence of a mudflat in combination with a concave (hollow) profile was associated with mangrove retreat. Our findings suggest that restoration of a mangrove greenbelt can be stimulated landward by improving drainage of abandoned aquaculture ponds. This enhances sediment stability and allows ponds to accrete. Seaward expansion can be induced by restoring foreshore morphology. Present results are discussed in the context of large-scale applications.

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Many organisms rely on chemical signals and cues to determine habitat suitability and safety. Chemical signals can mediate many interactions, including those between predators and their prey. Altering prey behaviour, these non-consumptive effects (NCEs) can influence population and community dynamics. Understanding how NCEs influence early life history stages, such as ‘decisions’ of benthic species with planktonic larvae about where to settle, can provide useful information on the ecological functioning of these systems as well as the management for commercial usage, although most studies have so far focused on intertidal systems which are already subject to a set of stressful conditions. With a shallow subtidal field experiment we investigated NCEs of the common starfish Asterias rubens on one of its main preys, the blue mussels Mytilus edulis. We tested the hypotheses that (1) the presence of starfish reduces mussels settlement and that (2) the mussels that settle will invest more energy towards induced defences than to growth, and will thus remain smaller than mussels settling in an area without starfish. Two independent trials revealed a significant reduction of mussel spat on the collectors in the presence of starfish after a two-week deployment period. There was however no effect of starfish on the size distribution of the mussel spat. The delayed observation of effects of starfish, absent after the first week but evident afterwards, suggests a time dependency of NCE's on spat settlement. Harnessing this ecologically important information has the potential to increase yield of mussel seeds available for fisheries by either removing starfish from the ground-based settling areas at the onset and for the duration of spatfall or by using floating substrates that are away from the bottom-bound starfish. Moreover, these results also underlines the potential of using predator cues in the application for sustainable natural antifouling compounds in situations with low recruitment pressures.@en

The Yucatan Channel connects the Caribbean Sea with the Gulf of Mexico and is the main outflow region of the Caribbean Sea. Moorings in the Yucatan Channel show high-frequent variability in kinetic energy (50–100 days) and transport (20–40 days), but the physical mechanisms controlling this variability are poorly understood. In this study, we show that the short-term variability in the Yucatan Channel transport has an upstream origin and arises from processes in the North Brazil Current. To establish this connection, we use data from altimetry and model output from several high resolution global models. A significant 40–70 day variability is found in the sea surface height in the North Brazil Current retroflection region with a propagation toward the Lesser Antilles. The frequency of variability is generated by intrinsic processes associated with the shedding of eddies, rather than by atmospheric forcing. This sea surface height variability is able to pass the Lesser Antilles, it propagates westward with the background ocean flow in the Caribbean Sea and finally affects the variability in the Yucatan Channel volume transport.@en

The natural coastal hydrodynamics and morphology worldwide is altered by human interventions such as embankments, shipping and dredging, which may have consequences for ecosystem functionality. To ensure long-term ecological sustainability, requires capability to predict long-term large-scale ecological effects of altered hydromorphology. As empirical data sets at relevant scales are missing, there is need for integrating ecological modeling with physical modeling. This paper presents a case study showing the long-term, large-scale macrozoobenthic community response to two contrasting human alterations of the hydromorphological habitat: deepening of estuarine channels to enhance navigability (Westerschelde) vs. realization of a storm surge barrier to enhance coastal safety (Oosterschelde). A multidisciplinary integration of empirical data and modeling of estuarine morphology, hydrodynamics and benthic ecology was used to reconstruct the hydrological evolution and resulting long-term (50 years) large-scale ecological trends for both estuaries over the last. Our model indicated that hydrodynamic alterations following the deepening of the Westerschelde had negative implications for benthic life, while the realization of the Oosterschelde storm surge barriers had mixed and habitat-dependent responses, that also include unexpected improvement of environmental quality. Our analysis illustrates long-term trends in the natural community caused by opposing management strategies. The divergent human pressures on the Oosterschelde and Westerschelde are examples of what could happen in a near future for many global coastal ecosystems. The comparative analysis of the two basins is a valuable source of information to understand (and communicate) the future ecological consequences of human coastal development.

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The natural coastal hydrodynamics and morphology worldwide is altered by human interventions such as embankments, shipping and dredging, which may have consequences for ecosystem functionality. To ensure long-term ecological sustainability, requires capability to predict long-term large-scale ecological effects of altered hydromorphology. As empirical data sets at relevant scales are missing, there is need for integrating ecological modeling with physical modeling. This paper presents a case study showing the long-term, large-scale macrozoobenthic community response to two contrasting human alterations of the hydromorphological habitat: deepening of estuarine channels to enhance navigability (Westerschelde) vs. realization of a storm surge barrier to enhance coastal safety (Oosterschelde). A multidisciplinary integration of empirical data and modeling of estuarine morphology, hydrodynamics and benthic ecology was used to reconstruct the hydrological evolution and resulting long-term (50 years) large-scale ecological trends for both estuaries over the last. Our model indicated that hydrodynamic alterations following the deepening of the Westerschelde had negative implications for benthic life, while the realization of the Oosterschelde storm surge barriers had mixed and habitat-dependent responses, that also include unexpected improvement of environmental quality. Our analysis illustrates long-term trends in the natural community caused by opposing management strategies. The divergent human pressures on the Oosterschelde and Westerschelde are examples of what could happen in a near future for many global coastal ecosystems. The comparative analysis of the two basins is a valuable source of information to understand (and communicate) the future ecological consequences of human coastal development.

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Moving water exerts drag forces on vegetation. The susceptibility of vegetation to bending and breakage determines its flow resistance, and chances of survival, under hydrodynamic loading. To evaluate the role of individual vegetation parameters in this water-vegetation interaction, we conducted drag force measurements under a wide range of wave loadings in a large wave flume. Artificial vegetation elements were used to manipulate stiffness, frontal area in still water and material volume as a proxy for biomass. The aim was to compare: (i) identical volume but different still frontal area, (ii) identical stiffness but different still frontal area, and (iii) identical still frontal area but different volume. Comparison of mimic arrangements showed that stiffness and the dynamic frontal area (i.e., frontal area resulting from bending which depends on stiffness and hydrodynamic forcing) determine drag forces. Only at low orbital-flow velocities did the still frontal area dominate the force-velocity relationship and it is hypothesised that no mimic bending took place under these conditions. Mimic arrangements with identical stiffness but different overall material volume and still frontal area showed that forces do not increase linearly with increasing material volume and it is proposed that short distances between mimics cause their interaction and result in additional drag forces. A model, based on effective leaf length and characteristic plant width developed for unidirectional flow, performed well for the force time series under both regular and irregular waves. However, its uncertainty increased with increasing interaction of neighbouring mimics.

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Moving water exerts drag forces on vegetation. The susceptibility of vegetation to bending and breakage determines its flow resistance, and chances of survival, under hydrodynamic loading. To evaluate the role of individual vegetation parameters in this water-vegetation interaction, we conducted drag force measurements under a wide range of wave loadings in a large wave flume. Artificial vegetation elements were used to manipulate stiffness, frontal area in still water and material volume as a proxy for biomass. The aim was to compare: (i) identical volume but different still frontal area, (ii) identical stiffness but different still frontal area, and (iii) identical still frontal area but different volume. Comparison of mimic arrangements showed that stiffness and the dynamic frontal area (i.e., frontal area resulting from bending which depends on stiffness and hydrodynamic forcing) determine drag forces. Only at low orbital-flow velocities did the still frontal area dominate the force-velocity relationship and it is hypothesised that no mimic bending took place under these conditions. Mimic arrangements with identical stiffness but different overall material volume and still frontal area showed that forces do not increase linearly with increasing material volume and it is proposed that short distances between mimics cause their interaction and result in additional drag forces. A model, based on effective leaf length and characteristic plant width developed for unidirectional flow, performed well for the force time series under both regular and irregular waves. However, its uncertainty increased with increasing interaction of neighbouring mimics.

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Moving water exerts drag forces on vegetation. The susceptibility of vegetation to bending and breakage determines its flow resistance, and chances of survival, under hydrodynamic loading. To evaluate the role of individual vegetation parameters in this water-vegetation interaction, we conducted drag force measurements under a wide range of wave loadings in a large wave flume. Artificial vegetation elements were used to manipulate stiffness, frontal area in still water and material volume as a proxy for biomass. The aim was to compare: (i) identical volume but different still frontal area, (ii) identical stiffness but different still frontal area, and (iii) identical still frontal area but different volume. Comparison of mimic arrangements showed that stiffness and the dynamic frontal area (i.e., frontal area resulting from bending which depends on stiffness and hydrodynamic forcing) determine drag forces. Only at low orbital-flow velocities did the still frontal area dominate the force-velocity relationship and it is hypothesised that no mimic bending took place under these conditions. Mimic arrangements with identical stiffness but different overall material volume and still frontal area showed that forces do not increase linearly with increasing material volume and it is proposed that short distances between mimics cause their interaction and result in additional drag forces. A model, based on effective leaf length and characteristic plant width developed for unidirectional flow, performed well for the force time series under both regular and irregular waves. However, its uncertainty increased with increasing interaction of neighbouring mimics.

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Thorough understanding of the potential for threshold dynamics and catastrophic shifts to occur in natural systems is of great importance for ecosystem conservation and restoration. However, verifying the presence of alternative stable states, one of the theoretical explanations for sudden shifts in natural systems, has proven to be a major challenge. We examine processes on local and landscape scales in salt-marsh pioneer zones, to assess the presence of alternative stable states in this system. To that end, we investigated the presence of typical characteristics of alternative stable states: bimodality and threshold dynamics. We also studied whether vegetation patches remained stable over long time periods. Analysis of false-color aerial photographs revealed clear bimodality in plant biomass distribution. By transplanting Spartina anglica plants of three different biomass classes on three geographically different marshes, we showed that a biomass threshold limits the establishment of Spartina patches, potentially explaining their patchy distribution. The presence of bimodality and biomass thresholds points to the presence of alternative stable states and the potential for sudden shifts, at small, within-patch scales and on short time scales. However, overlay analysis of aerial photographs from a salt marsh in The Netherlands, covering a time span of 22 years, revealed that there was little long-term stability of patches, as vegetation cover in this area is slowly increasing. Our results suggest that the concept of alternative stable states is applicable to the salt-marsh pioneer vegetation on small spatio-temporal scales. However, the concept does not apply to long-term dynamics of decades or centuries of heterogeneous salt-marsh pioneer zones, as landscape-scale processes may determine the large-scale dynamics of salt marshes. Hence, our results provide the interesting perspective that threshold dynamics may occur in systems with, on the long term, only a single stable state.@en

Thorough understanding of the potential for threshold dynamics and catastrophic shifts to occur in natural systems is of great importance for ecosystem conservation and restoration. However, verifying the presence of alternative stable states, one of the theoretical explanations for sudden shifts in natural systems, has proven to be a major challenge. We examine processes on local and landscape scales in salt-marsh pioneer zones, to assess the presence of alternative stable states in this system. To that end, we investigated the presence of typical characteristics of alternative stable states: bimodality and threshold dynamics. We also studied whether vegetation patches remained stable over long time periods. Analysis of false-color aerial photographs revealed clear bimodality in plant biomass distribution. By transplanting Spartina anglica plants of three different biomass classes on three geographically different marshes, we showed that a biomass threshold limits the establishment of Spartina patches, potentially explaining their patchy distribution. The presence of bimodality and biomass thresholds points to the presence of alternative stable states and the potential for sudden shifts, at small, within-patch scales and on short time scales. However, overlay analysis of aerial photographs from a salt marsh in The Netherlands, covering a time span of 22 years, revealed that there was little long-term stability of patches, as vegetation cover in this area is slowly increasing. Our results suggest that the concept of alternative stable states is applicable to the salt-marsh pioneer vegetation on small spatio-temporal scales. However, the concept does not apply to long-term dynamics of decades or centuries of heterogeneous salt-marsh pioneer zones, as landscape-scale processes may determine the large-scale dynamics of salt marshes. Hence, our results provide the interesting perspective that threshold dynamics may occur in systems with, on the long term, only a single stable state.@en