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Seagrass meadows provide valuable ecosystem services of coastal protection and chemical habitat formation that could help mitigate the impact of sea level rise and ocean acidification. However, the intensification of hydrodynamic forces caused by sea level rise, in addition to habitat degradation threaten the provision of these ecosystem services. With quantitative field measurements of the coastal protection and chemical habitat formation services of seagrass meadows, we statistically model the relationships between hydrodynamic forces, vegetation density and the provision of these ecosystem services. Utilising a high-resolution hydrodynamic model that simulates end of the century hydrodynamic conditions and three scenarios of coral reef degradation (i.e., keep up, remain or loss) we quantify how the environmental conditions within a tropical bay will change given changes to the provision of ecosystem services. Our study shows that increasing hydrodynamic forces lead to a seafloor made up of a larger grain size that is increasingly unstable and more vulnerable to erosion. The loss of a fringing reef leads to larger hydrodynamic forces entering the bay, however, the 0.87 m increase in depth due to sea-level rise reduces the bed shear stress in shallower areas, which limits the change in the ecosystem services provided by the current benthic seagrass meadow. Loss of seagrass constitutes the greatest change in a bay ecosystem, resulting in the sediment surface where seagrass existed becoming unstable and the median sediment grain size increasing by 5-7 %. The loss of seagrass also leads to the disappearance of the unique fluctuating chemical habitat, which leaves the surrounding community vulnerable to ocean acidification. A reduction or complete loss of these ecosystem services would impact the entire community assemblage while also leaving the surrounding coastline vulnerable to erosion, thus exacerbating negative effects brought about by climate change., Civil Engineering & Geosciences, Landscape Architecture, Environmental Fluid Mechanics

Seagrass meadows provide valuable ecosystem services of coastal protection and chemical habitat formation that could help mitigate the impact of sea level rise and ocean acidification. However, the intensification of hydrodynamic forces caused by sea level rise, in addition to habitat degradation threaten the provision of these ecosystem services. With quantitative field measurements of the coastal protection and chemical habitat formation services of seagrass meadows, we statistically model the relationships between hydrodynamic forces, vegetation density and the provision of these ecosystem services. Utilising a high-resolution hydrodynamic model that simulates end of the century hydrodynamic conditions and three scenarios of coral reef degradation (i.e. keep up, remain or loss) we quantify how the environmental conditions within a tropical bay will change given changes to the provision of ecosystem services. Our study shows that increasing hydrodynamic forces lead to a seafloor made up of a larger grain size that is increasingly unstable and more vulnerable to erosion. The loss of a fringing reef leads to larger hydrodynamic forces entering the bay, however, the 0.87 m increase in depth due to sea-level rise reduces the bed shear stress in shallower areas, which limits the change in the ecosystem services provided by the current benthic seagrass meadow. Loss of seagrass constitutes the greatest change in a bay ecosystem, resulting in the sediment surface where seagrass existed becoming unstable and the median sediment grain size increasing by 5-7 %. The loss of seagrass also leads to the disappearance of the unique fluctuating chemical habitat, which leaves the surrounding community vulnerable to ocean acidification. A reduction or complete loss of these ecosystem services would impact the entire community assemblage while also leaving the surrounding coastline vulnerable to erosion, thus exacerbating negative effects brought about by climate change., SCOPUS: ar.j, info:eu-repo/semantics/published

The intensity of major storm events generated within the Atlantic Basin is projected to rise with the warming of the oceans, which is likely to exacerbate coastal erosion. Nature-based flood defence has been proposed as a sustainable and effective solution to protect coastlines. However, the ability of natural ecosystems to withstand major storms like tropical hurricanes has yet to be thoroughly tested. Seagrass meadows both stabilise sediment and attenuate waves, providing effective coastal protection services for sandy beaches. To examine the tolerance of Caribbean seagrass meadows to extreme storm events, and to investigate the extent of protection they deliver to beaches, we employed a combination of field surveys, biomechanical measurements and wave modelling simulations. Field surveys of seagrass meadows before and after a direct hit by the category 5 Hurricane Irma documented that established seagrass meadows of Thalassia testudinum remained unaltered after the extreme storm event. The flexible leaves and thalli of seagrass and calcifying macroalgae inhabiting the meadows were shown to sustain the wave forces that they are likely to experience during hurricanes. In addition, the seagrass canopy and the complex biogeomorphic landscape built by the seagrass meadows combine to significantly dissipate extreme wave forces, ensuring that erosion is minimised within sandy beach foreshores. The persistence of the Caribbean seagrass meadows and their coastal protection services during extreme storm events ensures that a stable coastal ecosystem and beach foreshore is maintained in tropical regions., info:eu-repo/semantics/published

The intensity of major storm events generated within the Atlantic Basin is projected to rise with the warming of the oceans, which is likely to exacerbate coastal erosion. Nature-based flood defence has been proposed as a sustainable and effective solution to protect coastlines. However, the ability of natural ecosystems to withstand major storms like tropical hurricanes has yet to be thoroughly tested. Seagrass meadows both stabilise sediment and attenuate waves, providing effective coastal protection services for sandy beaches. To examine the tolerance of Caribbean seagrass meadows to extreme storm events, and to investigate the extent of protection they deliver to beaches, we employed a combination of field surveys, biomechanical measurements and wave modelling simulations. Field surveys of sea- grass meadows before and after a direct hit by the category 5 Hurricane Irma documented that estab- lished seagrass meadows of Thalassia testudinum re- mained unaltered after the extreme storm event. The flexible leaves and thalli of seagrass and calci- fying macroalgae inhabiting the meadows were shown to sustain the wave forces that they are likely to experience during hurricanes. In addition, the seagrass canopy and the complex biogeomorphic landscape built by the seagrass meadows combine to significantly dissipate extreme wave forces, ensuring that erosion is minimised within sandy beach fore- shores. The persistence of the Caribbean seagrass meadows and their coastal protection services dur- ing extreme storm events ensures that a stable coastal ecosystem and beach foreshore is maintained in tropical regions.

Plant species can be characterized by different growth strategies related to their inherent growth and recovery rates, which shape their responses to stress and disturbance. Ecosystem engineering, however, offers an alternative way to cope with stress: modifying the environment may reduce stress levels. Using an experimental study on two seagrass species with contrasting traits, the slow-growing Zostera marina vs. the fast-growing Zostera japonica, we explored how growth strategies versus ecosystem engineering may affect their resistance to stress (i.e. addition of organic material) and recovery from disturbance (i.e. removal of above-ground biomass). Ecosystem engineering was assessed by measuring sulphide levels in the sediment porewater, as seagrass plants can keep sulphide levels low by aerating the rhizosphere. Consistent with predictions, we observed that the fast-growing species had a high capacity to recover from disturbance. It was also more resistant to stress and still able to maintain high standing stock with increasing stress levels because of its ecosystem engineering capacity. The slow-growing species was not able to maintain its standing stock under stress, which we ascribe to a weak capacity for ecosystem engineering regarding this particular stress. Overall, our study suggests that the combination of low-cost investment in tissues with ecosystem engineering to alleviate stress creates a new path in the growth trade-off between investment in strong tissues or fast growth. It does so by being both fast in recovery and more resistant. As such low-cost ecosystem engineering may occur in more species, we argue that it should be considered in assessing plant resilience.

info:eu-repo/semantics/published

Amanipulative field experimentwas designed to investigate the effects of sediment-nutrients and sediment-organicmatterson seagrasses, Zostera japonica, using individual and population indicators. The results showed thatseagrasses quickly responded to sediment-nutrient and organicmatter loading. That is, sediment-nutrients positivelyimpacted on seagrasses by increasing N content of leaves and roots, leaf length and belowground biomass.Sediment-organic matter loading lowered N content of seagrass leaves and belowground biomass. Negative effectsof organic matter loadingwere aggravated during nutrient loading, by decreasing Ncontent of leaves, P contentof roots, leafwidth, shoot number in the middle period of the experiment, increasing C/N ratio of leaves, C/Pand N/P ratio of roots and above to belowground biomass ratio of seagrasses. Consequently, Z. japonica could beconsidered as a fast indicator to monitor seagrass ecosystem status in the eutrophic areas and facilitate tointerpreting the response of seagrasses to multiple stressors.

Although seagrass-based indicators are widely used to assess coastal ecosystem status, there is littleuniversality in their application. Matching the plethora of available indicators to specific managementobjectives requires a detailed knowledge of their species-specific sensitivities and their response timeto environmental stressors. We conducted an extensive survey of experimental studies to determine thesensitivity and response time of seagrass indicators to ecosystem degradation and recovery. We iden-tified seagrass size and indicator type (i.e. level of biological organization of the measure) as the mainfactors affecting indicator sensitivity and response time to degradation and recovery. While structuraland demographic parameters (e.g. shoot density, biomass) show a high and unspecific sensitivity, bio-chemical/physiological indicators present more stressor-specific responses and are the most sensitivedetecting early phases of environmental improvement. Based on these results we present a simple deci-sion tree to assist ecosystem managers to match adequate and reliable indicators to specific managementgoals.

In coastal and estuarine systems, foundation species like seagrasses, mangroves, saltmarshes or corals provide important ecosystem services. Seagrasses are globally declining and their reintroduction has been shown to restore ecosystem functions. However, seagrass restoration is often challenging, given the dynamic and stressful environment that seagrasses often grow in. From our world-wide meta-analysis of seagrass restoration trials (1786 trials), we describe general features and best practice for seagrass restoration. We confirm that removal of threats is important prior to replanting. Reduced water quality (mainly eutrophication), and construction activities led to poorer restoration success than, for instance, dredging, local direct impact and natural causes. Proximity to and recovery of donor beds were positively correlated with trial performance. Planting techniques can influence restoration success. The meta-analysis shows that both trial survival and seagrass population growth rate in trials that survived are positively affected by the number of plants or seeds initially transplanted. This relationship between restoration scale and restoration success was not related to trial characteristics of the initial restoration. The majority of the seagrass restoration trials have been very small, which may explain the low overall trial survival rate (i.e. estimated 37%). Successful regrowth of the foundation seagrass species appears to require crossing a minimum threshold of reintroduced individuals. Our study provides the first global field evidence for the requirement of a critical mass for recovery, which may also hold for other foundation species showing strong positive feedback to a dynamic environment. Synthesis and applications. For effective restoration of seagrass foundation species in its typically dynamic, stressful environment, introduction of large numbers is seen to be beneficial and probably serves two purposes. First, a large-scale planting increases

Dynamic environments like seagrass habitats are characterised by continuous local erosion and burial processes, which may induce seagrass decline if disturbances become too intense or frequent. We aim to quantify for Zostera noltii the resilience of both the edge of a meadow and individual propagules to local-scale burial and erosion events. In a set of mesocosm and in situ manipulation experiments, we varied the frequency (sudden vs. continuous rate) and intensity (different levels; −6 cm erosion to 6 cm burial) of sediment dynamics. Our results showed that the intensity of the disturbance event (−6 up to 6 cm) was negatively correlated with plant survival. Burial frequency also had a significant effect on plant survival, with sudden events having a stronger negative impact than continuous ones. Both experiments, on individual propagules and attached rhizomes at meadow edges, demonstrated a rapid acclimatization of seagrass plants to sediment dynamics within certain levels of disturbance (−6 up to 6 cm). After erosion or burial, all surviving plants in the field and mesocosm experiments were able to relocate their rhizomes to the preferential depth (from 0.3 to 0.8 cm), which was the depth at which the rhizomes of undisturbed plants were most frequently found in the field. In situ manipulation experiments showed that at the edge of the meadow, Z. noltii invaded experimentally created hollows more easily than hills. Overall, our results demonstrate that Z. noltii meadows can rapidly recover by clonal growth after modest sediment disturbance, thereby contributing to the current understanding of seagrass recovery mechanisms following disturbances (e.g. sediment dynamics)., Dynamic environments like seagrass habitats are characterised by continuous local erosion and burial processes, which may induce seagrass decline if disturbances become too intense or frequent. We aim to quantify for Zostera noltii the resilience of both the edge of a meadow and individual propagules to local-scale burial and erosion events. In a set of mesocosm and in situ manipulation experiments, we varied the frequency (sudden vs. continuous rate) and intensity (different levels; −6 cm erosion to 6 cm burial) of sediment dynamics. Our results showed that the intensity of the disturbance event (−6 up to 6 cm) was negatively correlated with plant survival. Burial frequency also had a significant effect on plant survival, with sudden events having a stronger negative impact than continuous ones. Both experiments, on individual propagules and attached rhizomes at meadow edges, demonstrated a rapid acclimatization of seagrass plants to sediment dynamics within certain levels of disturbance (−6 up to 6 cm). After erosion or burial, all surviving plants in the field and mesocosm experiments were able to relocate their rhizomes to the preferential depth (from 0.3 to 0.8 cm), which was the depth at which the rhizomes of undisturbed plants were most frequently found in the field. In situ manipulation experiments showed that at the edge of the meadow, Z. noltii invaded experimentally created hollows more easily than hills. Overall, our results demonstrate that Z. noltii meadows can rapidly recover by clonal growth after modest sediment disturbance, thereby contributing to the current understanding of seagrass recovery mechanisms following disturbances (e.g. sediment dynamics).

1.Populations of marine megaherbivores including green turtle (Chelonia mydas) have declined dramatically at a global scale as a result of overharvesting and habitat loss. This decline can be expected to also affect the tolerance of seagrass systems to coastal eutrophication. Until now, however, simultaneous effects of top–down control by megaherbivore grazing and bottom–up control by nutrient input have not been tested experimentally. 2.We therefore investigated the interacting effects of nutrient (N and P) addition and mimicked green turtle grazing on seagrass and epiphyte productivity, seagrass biomass and nutrient contents in exclosures at a pristine seagrass site in the Indo-Pacific region (Kalimantan, Indonesia). 3.Grazing almost doubled leaf biomass production rates, while nutrient addition (N+P, slow-release granules) did not have an effect on these rates. Rhizome biomass was, however, strongly reduced by nutrient addition. In contrast to phosphorus, tissue nitrogen contents increased after nutrient addition, showing that nitrogen was not limiting primary productivity. Epiphyte growth was, however, strongly correlated with high water column P concentrations, indicating an indirect negative effect of eutrophication when turtle grazing would be absent. We calculated that green turtle leaf grazing leads to substantial exports of N and P, at rates of at least 8% of the standing stock per day equalling the daily seagrass production, up to 13 (N) and 1.4 (P) mg m−2 day−1. 4.Synthesis. By combining our quantified effects with literature data, we propose a conceptual model of seagrass functioning under megaherbivore leaf grazing and eutrophication. In tropical seagrass systems with high green turtle grazing pressure, grazing alleviates the negative effects of eutrophication by the stimulation of seagrass production and concomitant nutrient uptake, the increased export of nutrients and the indirect prevention of low below-ground biomass. Similar to the role of terres, 1.Populations of marine megaherbivores including green turtle (Chelonia mydas) have declined dramatically at a global scale as a result of overharvesting and habitat loss. This decline can be expected to also affect the tolerance of seagrass systems to coastal eutrophication. Until now, however, simultaneous effects of top–down control by megaherbivore grazing and bottom–up control by nutrient input have not been tested experimentally. 2.We therefore investigated the interacting effects of nutrient (N and P) addition and mimicked green turtle grazing on seagrass and epiphyte productivity, seagrass biomass and nutrient contents in exclosures at a pristine seagrass site in the Indo-Pacific region (Kalimantan, Indonesia). 3.Grazing almost doubled leaf biomass production rates, while nutrient addition (N+P, slow-release granules) did not have an effect on these rates. Rhizome biomass was, however, strongly reduced by nutrient addition. In contrast to phosphorus, tissue nitrogen contents increased after nutrient addition, showing that nitrogen was not limiting primary productivity. Epiphyte growth was, however, strongly correlated with high water column P concentrations, indicating an indirect negative effect of eutrophication when turtle grazing would be absent. We calculated that green turtle leaf grazing leads to substantial exports of N and P, at rates of at least 8% of the standing stock per day equalling the daily seagrass production, up to 13 (N) and 1.4 (P) mg m−2 day−1. 4.Synthesis. By combining our quantified effects with literature data, we propose a conceptual model of seagrass functioning under megaherbivore leaf grazing and eutrophication. In tropical seagrass systems with high green turtle grazing pressure, grazing alleviates the negative effects of eutrophication by the stimulation of seagrass production and concomitant nutrient uptake, the increased export of nutrients and the indirect prevention of low below-ground biomass. Similar to the role of terres

In remote, tropical areas human influences increase, potentially threatening pristine seagrass systems. We aim (i) to provide a bench-mark for a near-pristine seagrass system in an archipelago in East Kalimantan, by quantifying a large spectrum of abiotic and biotic properties in seagrass meadows and (ii) to identify early warning indicators for river sediment and nutrient loading, by comparing the seagrass meadow properties over a gradient with varying river influence. Abiotic properties of water column, pore water and sediment were less suitable indicators for increased sediment and nutrient loading than seagrass properties. Seagrass meadows strongly responded to higher sediment and nutrient loads and proximity to the coast by decreasing seagrass cover, standing stock, number of seagrass species, changing species composition and shifts in tissue contents. Our study confirms that nutrient loads are more important than water nutrient concentrations. We identify seagrass system variables that are suitable indicators for sediment and nutrient loading, also in rapid survey scenarios with once-only measurements., In remote, tropical areas human influences increase, potentially threatening pristine seagrass systems. We aim (i) to provide a bench-mark for a near-pristine seagrass system in an archipelago in East Kalimantan, by quantifying a large spectrum of abiotic and biotic properties in seagrass meadows and (ii) to identify early warning indicators for river sediment and nutrient loading, by comparing the seagrass meadow properties over a gradient with varying river influence. Abiotic properties of water column, pore water and sediment were less suitable indicators for increased sediment and nutrient loading than seagrass properties. Seagrass meadows strongly responded to higher sediment and nutrient loads and proximity to the coast by decreasing seagrass cover, standing stock, number of seagrass species, changing species composition and shifts in tissue contents. Our study confirms that nutrient loads are more important than water nutrient concentrations. We identify seagrass system variables that are suitable indicators for sediment and nutrient loading, also in rapid survey scenarios with once-only measurements.

Traditionally, protection of the coastal area from flooding is approached from an engineering perspective. This approach has often resulted in negative or unforeseen impacts on local ecology and is even known to impact surrounding ecosystems on larger scales. In this paper, the utilization of ecosystem engineering species for achieving civil-engineering objectives or the facilitation of multiple use of limited space in coastal protection is focused upon, either by using ecosystem engineering species that trap sediment and damp waves (oyster beds, mussel beds, willow floodplains and marram grass), or by adjusting hard substrates to enhance ecological functioning. Translating desired coastal protection functionality into designs that make use of the capability of appropriate ecosystem engineering species is, however, hampered by lack of a generic framework to decide which ecosystem engineering species or what type of hard-substrate adaptations may be used where and when. In this paper we review successful implementation of ecosystem engineering species in coastal protection for a sandy shore and propose a framework to select the appropriate measures based on the spatial and temporal scale of coastal protection, resulting in a dynamic interaction between engineering and ecology. Modeling and monitoring the bio-physical interactions is needed, as it allows to upscale successful implementations and predict otherwise unforeseen impacts., Traditionally, protection of the coastal area from flooding is approached from an engineering perspective. This approach has often resulted in negative or unforeseen impacts on local ecology and is even known to impact surrounding ecosystems on larger scales. In this paper, the utilization of ecosystem engineering species for achieving civil-engineering objectives or the facilitation of multiple use of limited space in coastal protection is focused upon, either by using ecosystem engineering species that trap sediment and damp waves (oyster beds, mussel beds, willow floodplains and marram grass), or by adjusting hard substrates to enhance ecological functioning. Translating desired coastal protection functionality into designs that make use of the capability of appropriate ecosystem engineering species is, however, hampered by lack of a generic framework to decide which ecosystem engineering species or what type of hard-substrate adaptations may be used where and when. In this paper we review successful implementation of ecosystem engineering species in coastal protection for a sandy shore and propose a framework to select the appropriate measures based on the spatial and temporal scale of coastal protection, resulting in a dynamic interaction between engineering and ecology. Modeling and monitoring the bio-physical interactions is needed, as it allows to upscale successful implementations and predict otherwise unforeseen impacts.

Seagrasses have declined at a global scale due to light reduction and toxicity events, caused by eutrophication and increased sediment loading. Although several studies have tested effects of light reduction and toxicants on seagrasses, there is at present no information available on their interacting effects. In a full-factorial 5-day laboratory experiment, we studied short-term interactive effects of light conditions, pH and reduced nitrogen (NHx) in the water layer, mimicking pulses of river discharge, on the tropical early successional species Halodule uninervis and the late successional species Thalassia hemprichii. In contrast to recent results reported for the temperate species Zostera marina, increased NHx supply did not affect leaf mortality or photochemical efficiency in H. uninervis and in 7 out of 8 treatments for T. hemprichii. However, both tropical species demonstrated striking differences in nitrogen accumulation, free amino acid composition and free NH3 accumulation. The increase in tissue nitrogen content was two times higher for H. uninervis than for T. hemprichii. Nitrogen stored as free amino acids (especially asparagine) only increased in H. uninervis. High pH only affected T. hemprichii, but only when not shaded, by doubling its free NH3 concentrations, concomitantly decreasing its photosynthetic efficiency. Our results indicate that the early successional H. uninervis has higher tolerance to high NHx loads as compared to the late successional T. hemprichii. H. uninervis was better able to avoid toxic internal NHx levels by further assimilating glutamine into asparagine in contrast to T. hemprichii. Moreover, both tropical species seem to cope much better with high NHx than the temperate Z. marina. The implications for the distribution and succession of seagrass species under high nutrient loads are discussed., Seagrasses have declined at a global scale due to light reduction and toxicity events, caused by eutrophication and increased sediment loading. Although several studies have tested effects of light reduction and toxicants on seagrasses, there is at present no information available on their interacting effects. In a full-factorial 5-day laboratory experiment, we studied short-term interactive effects of light conditions, pH and reduced nitrogen (NHx) in the water layer, mimicking pulses of river discharge, on the tropical early successional species Halodule uninervis and the late successional species Thalassia hemprichii. In contrast to recent results reported for the temperate species Zostera marina, increased NHx supply did not affect leaf mortality or photochemical efficiency in H. uninervis and in 7 out of 8 treatments for T. hemprichii. However, both tropical species demonstrated striking differences in nitrogen accumulation, free amino acid composition and free NH3 accumulation. The increase in tissue nitrogen content was two times higher for H. uninervis than for T. hemprichii. Nitrogen stored as free amino acids (especially asparagine) only increased in H. uninervis. High pH only affected T. hemprichii, but only when not shaded, by doubling its free NH3 concentrations, concomitantly decreasing its photosynthetic efficiency. Our results indicate that the early successional H. uninervis has higher tolerance to high NHx loads as compared to the late successional T. hemprichii. H. uninervis was better able to avoid toxic internal NHx levels by further assimilating glutamine into asparagine in contrast to T. hemprichii. Moreover, both tropical species seem to cope much better with high NHx than the temperate Z. marina. The implications for the distribution and succession of seagrass species under high nutrient loads are discussed.

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

Successful management and restoration of coastal vegetation requires a quantitative process-based understanding of thresholds hampering (re-)establishment of pioneer vegetation. We expect scouring to be important in explaining the disappearance of seedlings and/or small propagules of intertidal plant species, and therefore quantify the dependence of scouring depend on plant traits (flexibility, size) and physical forcing by current velocity. Flume studies with unidirectional flow revealed that scouring around seedlings increased exponentially with current velocity and according to a power relationship with plant size. Basal stem diameter rather than shoot length was found to be the factor controlling scouring volume. Flexible shoots caused far less scouring than stiff shoots, provided that the bending occurred near the sediment surface as was the case for Zostera, and not on top of a solid tussock base as we observed for Puccinellia. Therefore, shoot stiffness is likely to strongly affect the chances for initial establishment in hydrodynamically exposed areas. Plant traits such as shoot stiffness are subject to a trade-off between advantages and disadvantages, the outcome of which depends on the physical settings., Successful management and restoration of coastal vegetation requires a quantitative process-based understanding of thresholds hampering (re-)establishment of pioneer vegetation. We expect scouring to be important in explaining the disappearance of seedlings and/or small propagules of intertidal plant species, and therefore quantify the dependence of scouring depend on plant traits (flexibility, size) and physical forcing by current velocity. Flume studies with unidirectional flow revealed that scouring around seedlings increased exponentially with current velocity and according to a power relationship with plant size. Basal stem diameter rather than shoot length was found to be the factor controlling scouring volume. Flexible shoots caused far less scouring than stiff shoots, provided that the bending occurred near the sediment surface as was the case for Zostera, and not on top of a solid tussock base as we observed for Puccinellia. Therefore, shoot stiffness is likely to strongly affect the chances for initial establishment in hydrodynamically exposed areas. Plant traits such as shoot stiffness are subject to a trade-off between advantages and disadvantages, the outcome of which depends on the physical settings.

We examined the original manuscripts of a French national survey conducted in 1933 on the state of common eelgrass Zostera marina beds along the French Atlantic coasts during the period when wasting disease struck the entire North Atlantic population in the 1930s. Based on GIS related techniques and old sets of aerial photographs, we present the first accurate mapping of the Z. marina beds before wasting disease occurred and assess their spatial recolonization since the 1950s in the Chausey Archipelago (France), which contains large Z. marina beds. The national survey confirmed that the Z. marina beds almost totally disappeared from the French coasts during the 1930s. However, the disease symptoms seem to have begun locally a few years before. On the study site, we found that the Z. marina beds were more than twice as extended than as they are today, and covered both subtidal and intertidal areas. By the 1950s, 20 yr after the onset of the disease, the beds had hardly recolonized, and contrary to the recolonization patterns reported elsewhere in Europe, they were mainly restricted to subtidal areas. The subtidal and intertidal Z. marina beds on the site are now rapidly expanding.

Seagrasses are generally known as ecosystem engineers, as they reduce flow velocities in their canopies. In perennial subtidal meadows, this usually leads to increased net sedimentation rates and reduction of the grain size. The present study aims to describe the contribution of annual seagrass populations to these processes and elucidate the temporal dynamics. Sediment accretion and grain size modification were experimentally tested by transplanting seedlings of an annual intertidal eelgrass population to an unvegetated tidal flat. Within the planting units (79 shoots m−2) 4.7 mm of sediment accreted, whereas in the most dense parts of these units (199 shoots m−2) accretion amounted to 7.1 mm. The silt fraction (, Seagrasses are generally known as ecosystem engineers, as they reduce flow velocities in their canopies. In perennial subtidal meadows, this usually leads to increased net sedimentation rates and reduction of the grain size. The present study aims to describe the contribution of annual seagrass populations to these processes and elucidate the temporal dynamics. Sediment accretion and grain size modification were experimentally tested by transplanting seedlings of an annual intertidal eelgrass population to an unvegetated tidal flat. Within the planting units (79 shoots m−2) 4.7 mm of sediment accreted, whereas in the most dense parts of these units (199 shoots m−2) accretion amounted to 7.1 mm. The silt fraction (

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

Prof. Dr. Pieter Hendrik (Piet) Nienhuis worked for almost 40 years in all aspects of aquatic ecology and environmental science and retired on 31 October 2003. He can be characterised as a distinguished scientist, shaped in an applied estuarine and aquatic research ambience of the former Delta Institute for Hydrobiological Research (DIHO) in Yerseke in the Netherlands. His appointment as a full professor at the Radboud University Nijmegen offered him a challenging step from monodisciplinarity in ecology, via multidisciplinarity in the application of ecological knowledge in river science to interdisciplinarity in environmental science and management. This paper describes his education, teaching activities, research, scientific publications, science management, and significance for various scientific disciplines. He made important contributions to biosystematics of angiosperms and algae, the ecology of seagrasses, nutrient cycling and eutrophication in estuarine ecosystems, and the integrated modelling of the ecological functioning of estuaries. Subsequently, he paid much attention to environmental problems in river basins, ecological rehabilitation and sustainable development. His work influenced the view of ecologists, aquatic scientists and water managers in the Netherlands as well as abroad, in particular regarding the drawbacks of compartmentalization of the estuaries and the importance of connectivity and morphodynamics in river systems. In hindsight, it appears as a logical line that he gradually moved from estuarine ecological research that became increasingly driven by societal and environmental problems to the field of environmental science and management. [KEYWORDS: coastal zone management ; estuarine ecology ; environmental sciences ; interdisciplinary research ; river basin management ; river ecology], Prof. Dr. Pieter Hendrik (Piet) Nienhuis worked for almost 40 years in all aspects of aquatic ecology and environmental science and retired on 31 October 2003. He can be characterised as a distinguished scientist, shaped in an applied estuarine and aquatic research ambience of the former Delta Institute for Hydrobiological Research (DIHO) in Yerseke in the Netherlands. His appointment as a full professor at the Radboud University Nijmegen offered him a challenging step from monodisciplinarity in ecology, via multidisciplinarity in the application of ecological knowledge in river science to interdisciplinarity in environmental science and management. This paper describes his education, teaching activities, research, scientific publications, science management, and significance for various scientific disciplines. He made important contributions to biosystematics of angiosperms and algae, the ecology of seagrasses, nutrient cycling and eutrophication in estuarine ecosystems, and the integrated modelling of the ecological functioning of estuaries. Subsequently, he paid much attention to environmental problems in river basins, ecological rehabilitation and sustainable development. His work influenced the view of ecologists, aquatic scientists and water managers in the Netherlands as well as abroad, in particular regarding the drawbacks of compartmentalization of the estuaries and the importance of connectivity and morphodynamics in river systems. In hindsight, it appears as a logical line that he gradually moved from estuarine ecological research that became increasingly driven by societal and environmental problems to the field of environmental science and management. [KEYWORDS: coastal zone management ; estuarine ecology ; environmental sciences ; interdisciplinary research ; river basin management ; river ecology]

Generally, seagrass Zostera marina L. distribution in the Wadden Sea and south-west Netherlands is Limited to waters with low to moderate nutrient concentrations. However, it is known that Z. marina also occurs at high nutrient concentrations when growing in low salinity environments. In this study, we investigated the separate and interactive effects of nutrients and salinity on Z, marina plants in a 5 wk experiment. Two populations were tested; one originating from a relatively marine habitat and the other from an estuarine habitat. Supplied salinities were 23, 26 and 30 parts per thousand S, and sup plied water nutrient levels were nitrate:ammonium:phosphate, 1:3:2, 3:9:4.5 and 60:9:9 mu M at a refreshment rate of 1 d(-1), corresponding with a load of 20, 95 and 625 kg N ha(-1) yr(-1). Z. marina was negatively influenced by high salinity. The estuarine plants showed a decreased 'vitality' (calculated from 6 plant response parameters), whereas the marine plants showed a lesser number of shoots at high salinity. The negative effect acted on the estuarine plants at 26 and 30 parts per thousand S, and on the marine plants at 30 parts per thousand S. At these high salinities, a high nutrient load had no detectable effect on the marine plants, whereas the estuarine plants were negatively influenced by high nutrient loads. At low salinity levels, i.e. marine plants at 23 and 26 parts per thousand S and estuarine plants at 23 parts per thousand S, plants from both populations were positively influenced by higher nutrient loads. It is argued that these results may explain the distribution and decline of Z. marina in many areas of the northern hemisphere. Examples from both sides of the Atlantic Ocean are presented. [KEYWORDS: eelgrass; eutrophication; interaction; nitrogen; phosphorus population dynamics; salinity stress; seagrass; Wadden sea Column nitrate enrichment; coastal lagoon; mesocosm experiments; halodule-wrightii; plant-communities; wasting disease; amino-acids, Generally, seagrass Zostera marina L. distribution in the Wadden Sea and south-west Netherlands is Limited to waters with low to moderate nutrient concentrations. However, it is known that Z. marina also occurs at high nutrient concentrations when growing in low salinity environments. In this study, we investigated the separate and interactive effects of nutrients and salinity on Z, marina plants in a 5 wk experiment. Two populations were tested; one originating from a relatively marine habitat and the other from an estuarine habitat. Supplied salinities were 23, 26 and 30 parts per thousand S, and sup plied water nutrient levels were nitrate:ammonium:phosphate, 1:3:2, 3:9:4.5 and 60:9:9 mu M at a refreshment rate of 1 d(-1), corresponding with a load of 20, 95 and 625 kg N ha(-1) yr(-1). Z. marina was negatively influenced by high salinity. The estuarine plants showed a decreased 'vitality' (calculated from 6 plant response parameters), whereas the marine plants showed a lesser number of shoots at high salinity. The negative effect acted on the estuarine plants at 26 and 30 parts per thousand S, and on the marine plants at 30 parts per thousand S. At these high salinities, a high nutrient load had no detectable effect on the marine plants, whereas the estuarine plants were negatively influenced by high nutrient loads. At low salinity levels, i.e. marine plants at 23 and 26 parts per thousand S and estuarine plants at 23 parts per thousand S, plants from both populations were positively influenced by higher nutrient loads. It is argued that these results may explain the distribution and decline of Z. marina in many areas of the northern hemisphere. Examples from both sides of the Atlantic Ocean are presented. [KEYWORDS: eelgrass; eutrophication; interaction; nitrogen; phosphorus population dynamics; salinity stress; seagrass; Wadden sea Column nitrate enrichment; coastal lagoon; mesocosm experiments; halodule-wrightii; plant-communities; wasting disease; amino-acids