Salinity is a major factor controlling the distribution of biota in aquatic systems, and most aquatic multicellular organisms are either adapted to life in saltwater or freshwater conditions. Consequently, the saltwater–freshwater mixing zones in coastal or estuarine areas are characterized by limited faunal and floral diversity. Although changes in diversity and decline in species richness in brackish waters is well documented in aquatic ecology, it is unknown to what extent this applies to bacterial communities. Here, we report a first detailed bacterial inventory from vertical profiles of 60 sampling stations distributed along the salinity gradient of the Baltic Sea, one of world's largest brackish water environments, generated using 454 pyrosequencing of partial (400 bp) 16S rRNA genes. Within the salinity gradient, bacterial community composition altered at broad and finer-scale phylogenetic levels. Analogous to faunal communities within brackish conditions, we identified a bacterial brackish water community comprising a diverse combination of freshwater and marine groups, along with populations unique to this environment. As water residence times in the Baltic Sea exceed 3 years, the observed bacterial community cannot be the result of mixing of fresh water and saltwater, but our study represents the first detailed description of an autochthonous brackish microbiome. In contrast to the decline in the diversity of multicellular organisms, reduced bacterial diversity at brackish conditions could not be established. It is possible that the rapid adaptation rate of bacteria has enabled a variety of lineages to fill what for higher organisms remains a challenging and relatively unoccupied ecological niche.

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Transitions in bacterial communities along the 2000 km salinity gradient of the Baltic Sea

Marine vegetated habitats occupy a small fraction of the ocean surface, but contribute about 50% of the carbon that is buried in marine sediments. In this Review the potential benefits of conservation, restoration and use of these habitats for coastal protection and climate change mitigation are assessed. Marine vegetated habitats (seagrasses, salt-marshes, macroalgae and mangroves) occupy 0.2% of the ocean surface, but contribute 50% of carbon burial in marine sediments. Their canopies dissipate wave energy and high burial rates raise the seafloor, buffering the impacts of rising sea level and wave action that are associated with climate change. The loss of a third of the global cover of these ecosystems involves a loss of CO2 sinks and the emission of 1 Pg CO2 annually. The conservation, restoration and use of vegetated coastal habitats in eco-engineering solutions for coastal protection provide a promising strategy, delivering significant capacity for climate change mitigation and adaption.

The role of coastal plant communities for climate change mitigation and adaptation

For more than a century, coastal wetlands have been recognized for their ability to stabilize shorelines and protect coastal communities. However, this paradigm has recently been called into question by small-scale experimental evidence. Here, we conduct a literature review and a small meta-analysis of wave attenuation data, and we find overwhelming evidence in support of established theory. Our review suggests that mangrove and salt marsh vegetation afford context-dependent protection from erosion, storm surge, and potentially small tsunami waves. In biophysical models, field tests, and natural experiments, the presence of wetlands reduces wave heights, property damage, and human deaths. Meta-analysis of wave attenuation by vegetated and unvegetated wetland sites highlights the critical role of vegetation in attenuating waves. Although we find coastal wetland vegetation to be an effective shoreline buffer, wetlands cannot protect shorelines in all locations or scenarios; indeed large-scale regional erosion, river meandering, and large tsunami waves and storm surges can overwhelm the attenuation effect of vegetation. However, due to a nonlinear relationship between wave attenuation and wetland size, even small wetlands afford substantial protection from waves. Combining man-made structures with wetlands in ways that mimic nature is likely to increase coastal protection. Oyster domes, for example, can be used in combination with natural wetlands to protect shorelines and restore critical fishery habitat. Finally, coastal wetland vegetation modifies shorelines in ways (e.g. peat accretion) that increase shoreline integrity over long timescales and thus provides a lasting coastal adaptation measure that can protect shorelines against accelerated sea level rise and more frequent storm inundation. We conclude that the shoreline protection paradigm still stands, but that gaps remain in our knowledge about the mechanistic and context-dependent aspects of shoreline protection.

The present and future role of coastal wetland vegetation in protecting shorelines: answering recent challenges to the paradigm

Salt marshes protect coastlines against waves. Wave flume experiments show that marsh vegetation causes substantial wave dissipation and prevents erosion of the underlying surface, even during extreme storm surge conditions.

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Wave attenuation over coastal salt marshes under storm surge conditions

Akaike information criterion statistics

The objectives of the present study were twofold: (1) to identify spatial sedimentation and erosion patterns developing within patches of epibenthic structures (i.e. physical structures that protrude from the sediments, originating either from animals or plants) as a consequence of biophysical interactions; and (2) to assess the relevance of hydrodynamic flume studies for the long-term sediment dynamics in the field. We addressed these objectives by using patches of well-defined artificial structures (bamboo canes) for which we could easily monitor the long-term sediment dynamics in the field, measure the hydrodynamic effects in detail in the flume, and simulate the field and flume set-up with a commercially available hydrodynamic model. Two-year monitoring in the field showed that sedimentation was much larger in the high-density patches than the low-density ones. Within the high-density patches, comparable spatial patterns emerged at different field sites: erosion at the front and the side of the patches, and sedimentation more down-stream within the patches. The low-density patches showed no such patterns, and were generally characterised by some small-scale erosion directly around individual bamboo canes. Sedimentation and erosion in the field was well explained by the patterns in bed shear stress that were derived from our flume measurements. The 3D hydrodynamic modelling facilitated up-scaling of the flume results to the field, but failed to simulate accurately the effects at the leading edge. We conclude that: (A) field observations on sedimentation revealed interesting spatial patterns, but could not elucidate underlying processes; (B) detailed hydrodynamic measurements in a flume can elucidate these underlying processes, provided that appropriate scaling is being used; (C) flume studies are by definition not able to capture all spatial scales that are relevant for estuarine landscape formation and will always cause some flow artefacts; (D) hydrodynamic modelling offers a valuable tool to upscale flume observations, even though present models are not yet capable of fully reproducing all detailed spatial patterns; and (E) spatial heterogeneity is very important when looking at small-scale patches. There is a need for more spatially explicit and scale-dependent knowledge on bio-physical interactions.

http://www.vliz.be/imisdocs/publications/125805.pdf

Spatial flow and sedimentation patterns within patches of epibenthic structures: Combining field, flume and modelling experiments

We quantified the distribution, abundance, biomass and assemblage structure of ben- thic macrofauna at different spatio-temporal scales in a temperate estuarine, intertidal soft-sediment environment in the Schelde estuary, The Netherlands. Hierarchically scaled surveys were conducted yearly between 1994 and 2000, covering 4 spatial scales: region (10 4 m), transect (10 3 m), station (10 2 m), and replicate samples (10 -1 m). Our approach provided a powerful framework for quantify- ing the proportion of the variation among samples that was attributable to each spatial or temporal scale, and we explicitly aimed at identifying the role of environmental variables in explaining the observed variability. Variance components calculated for 11 dominant macrobenthic species revealed that variations at the scale of stations and year × station interactions were the most im- portant components of variability. Regional and transect differences were only apparent for a few species, although multivariate analysis revealed clear inter-regional differences for the macrobenthic assemblage structure. Only a few species displayed significant variability associated with the factor year solely. Both spatial and temporal components explained > 70% of the total variance. A sub- stantial part of the total variation in abundance of individual species was explained by the observed environmental variables (27 to 56%). Multiple regression with subdivision of the environmental variables into a long-term average and a temporal component showed that the long-term averages were much more important than the short-term deviations from this average, with local environ- mental variables (mud content, chlorophyll a, bed level height) explaining the largest part of the observed variation. For a few species and total biomass, salinity also explained a large part of the observed variation. Canonical correspondence analysis (CCA) with forward selection revealed that salinity and mud content, and to a lesser extent chlorophyll a and bed-level height, accounted for most of the variance in the macrobenthic species data. Partial CCA indicated that of the variation in the species data a large part was spatially structured (56.7%), with about half of this variation being explained by the environmental variables used. Only 4.1% of the species variation was temporally structured. Our results have important consequences for both the interpretation of monitoring programmes and the enhancement of sampling designs.

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Spatial and temporal variation in benthic macrofauna and relationships with environmental variables in an estuarine, intertidal soft-sediment environment

Estuaries are naturally highly dynamic and rapidly changing systems, forming a complex mixture of many different habitat types. They are very productive biomes and support many important ecosystem functions: biogeochemical cycling and movement of nutrients, mitigation of floods, maintenance of biodiversity and biological production. Human pressure on estuaries is very high. On the other hand, it is recognized that estuaries have a unique functional and structural biodiversity. Therefore, these ecosystems are particularly important for integrating sound ecological management with sustainable economics. These opportunities are explored for the Scheldt estuary, a well-documented system with an exceptional tidal freshwater area. In this article a description of the Scheldt estuary is presented, illustrating that human influence is intertwined with natural dynamics. Hydrology, geomorphology, trophic status and diversity are discussed, and possible future trends in both natural evolution and management are argued.

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The Scheldt estuary: a description of a changing ecosystem

Few macrobenthic studies have dealt simultaneously with the two major gradients in estuarine benthic habitats: the salinity gradient along the estuary (longitudinal) and the gradients from high intertidal to deep subtidal sites (vertical gradient). In this broad-scale study, a large data set (3112 samples) of the Schelde estuary allowed a thorough analysis of these gradients, and to relate macrobenthic species distributions and community structure to salinity, depth, current velocities and sediment characteristics. Univariate analyses clearly revealed distinct gradients in diversity, abundance, and biomass along the vertical and longitudinal gradients. In general, highest diversity and biomass were observed in the intertidal, polyhaline zone and decreased with decreasing salinity. Abundance did not show clear trends and varied between spring and autumn. In all regions, very low values for all measures were observed in the subtidal depth strata. Abundance in all regions was dominated by both surface deposit feeders and sub-surface deposit feeders. In contrast, the biomass of the different feeding guilds showed clear gradients in the intertidal zone. Suspension feeders dominated in the polyhaline zone and showed a significant decrease with decreasing salinity. Surface deposit feeders and sub-surface deposit feeders showed significantly higher biomass values in the polyhaline zone as compared with the mesohaline zone. Omnivores showed an opposite trend. Multivariate analyses showed a strong relationship between the macrobenthic assemblages and the predominant environmental gradients in the Schelde estuary. The most important environmental factor was depth, which reflected also the hydrodynamic conditions (current velocities). A second gradient was related to salinity and confirms the observations from the univariate analyses. Additionally, sediment characteristics (mud content) explained a significant part of the macrobenthic community structure not yet explained by the two other main gradients. The different assemblages are further described in terms of indicator species and abiotic characteristics. The results showed that at a large, estuarine scale a considerable fraction of the variation in abundance and biomass of the benthic macrofauna correlated very well with environmental factors (depth, salinity, tidal current velocity, sediment composition).

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Large-scale spatial patterns in estuaries: estuarine macrobenthic communities in the Schelde estuary, NW Europe

To examine the spatial and temporal variability of sediment grain size in exposed tidal wetlands with ample sediment supply, we sampled sediments and measured hydrodynamics, accretion/erosion rates, and vegetation characteristics in the Yangtze Delta. Sediment grain size exhibited a landward/upward decreasing trend. This trend is mainly attributed to attenuation of hydrodynamics. A 630-day series of daily surface sediment sampling at a fixed site on an unvegetated intertidal flat revealed significant seasonal and storm-cyclic changes in grain size. This temporal variability was related to alternating accretion/erosion events, with erosion associated with coarser grain size. Such temporal dynamics were not present in vegetation, where sediment remained fine grained throughout the year. In the marsh, vegetation cover enables the trapping of fine-grained sediments in the following ways: (a) adherence of suspended sediments onto plants; (b) deposition of suspended sediments stimulated by attenuation of hydrodynamics through plant obstruction; and (c) prevention of resuspension of fine-grained deposits due to the protection of the plant canopy. The influence of vegetation on sediment grain size was clearly seen when comparing sediment trapped by different vegetation types and seasonal patterns of trapped sediment on different vegetation canopy densities. The relatively high plant biomass of the recently introduced Spartina alterniflora enhanced the trapping effect, whereas plant degradation due to buffalo grazing reduced the trapping effect. We conclude that for exposed tidal wetlands with ample sediment supply such as the Yangtze Delta, the spatial and temporal variability of sediment grain size is governed predominantly by physical controls on the unvegetated flat and predominantly by biophysical interaction of hydrodynamics and vegetation in the salt marsh, rather than by sediment supply.

T.J. Ysebaert

Papers

Spatial flow and sedimentation patterns within patches of epibenthic structures: Combining field, flume and modelling experiments

Spatial and temporal variation in benthic macrofauna and relationships with environmental variables in an estuarine, intertidal soft-sediment environment

The Scheldt estuary: a description of a changing ecosystem

Large-scale spatial patterns in estuaries: estuarine macrobenthic communities in the Schelde estuary, NW Europe

Spatial and temporal variations in sediment grain size in tidal wetlands, Yangtze Delta: On the role of physical and biotic controls