Estimating estuarine residence times in the Westerschelde (The Netherlands) using a box model with fixed dispersion coefficients

The residence time of the water masses in the Westerschelde estuary was determined using a simple compartment-model that simulates the advective-diffusive transport of a conservative dissolved substance (chlorinity). The residence time of a water parcel in the upstream part of the estuary (i.e. the time needed for this water parcel to leave the estuary) varied from about 50 days in winter to about 70 days in summer. The most seaward compartment had residence times of about 10-15 days.Dispersive coefficients that are fixed in time were able to reproduce the observed salinity distributions very well in the Westerschelde. They were obtained by calibration on observed chlorinities. It is argued that the apparent relationship of dispersive coefficients with freshwater flow, which is observed in certain studies, could (partly) reflect the deviation from steady state conditions which are required assumptions to calculate these dispersive coefficients directly from salinity profiles.     

Composition and abundance of resident marsh-surface nekton: comparison between tidal freshwater and salt marshes in Virginia,USA

Previous research on intertidal nekton communities has identifiedimportant determinants of community structure and distribution; however, fewstudies have compared nekton utilization of disparate marsh habitats. Inthis study, abundance and distribution patterns of resident nekton werecompared between tidal freshwater marsh and salt marsh surfaces varying inflooding depth and duration. Nekton were collected in pit traps installedalong elevational transects at four marshes in coastal Virginia (twofreshwater, two saline) from April through November 1992–1993. Thedominant fish collected at all sites was the mummichog Fundulusheteroclitus. The daggerblade grass shrimp Palaemonetes pugio was thedominant nekton species collected at salt marsh sites, and was seasonallyabundant on tidal freshwater marshes. A positive correlation betweenflooding depth and nekton abundance was observed on salt marshes; anopposite pattern was observed on tidal freshwater marshes. Tidal floodingregime influences the abundance of resident nekton, however, the effect maybe confounded by other environmental variables, including variation insurface topography and seasonal presence or absence of submerged aquaticvegetation (SAV) in adjacent subtidal areas. In mid-Atlantic tidalfreshwater wetlands, SAV provides a predation refuge and forage site forearly life stages of marsh-dependent nekton, and several species utilizethis environment extensively. Salt marshes in this region generally lackdense SAV in adjacent subtidal creeks. Consequently, between-sitedifferences in species and size-specific marsh surface utilization byresident nekton were observed. Larvae and juveniles represented 79%and 59% of total fish collected at tidal freshwater and salt marshsites, respectively. The resident nekton communities of tidal freshwater andsalt marsh surfaces are characterized by a few ubiquitous species with broadenvironmental tolerances. This revised version was published online in July 2006 with corrections to the Cover Date.     

Secondary production of the brackish copepod communities and their contribution to the carbon fluxes in the Westerschelde estuary (The Netherlands)

The zooplankton community of the brackish part of the Westerschelde estuary (November 1989–October 1990) was dominated by two calanoid copepods, Eurytemora affinis and Acartia tonsa. Eurytemora was present during a longer period of the year and was much more important in terms of total abundances and biomasses than Acartia.The secondary production of these species was estimated by means of the growth rate method, using weight-specific growth rates obtained from laboratory cultures (Eurytemora) or from the literature (Acartia).Due to the substantially higher growth rates of Acartia compared to Eurytemora, total yearly productions of both communities were comparable, notwithstanding the large discrepancies in biomass. They amounted to about 5 and 6 g C m^–2 y^–1 by Acartia and Eurytemora respectively.The food needed to realise this production was estimated to be about 14 and 17 g C m^–2 y^–1 by Acartia and Eurytemora respectively. Provided that the copepods are able to selectively ingest the phytoplankton, in situ net production provides sufficient carbon for zooplankton demands for a short period of the year only. As phytoplankton standing stock is very low and net phytoplankton productivity is negative from late fall to early spring, nutritional demands of the copepods have to be fulfilled by other than algal food at least during this period of the year.Although the copepods in the brackish part can have an important impact on some food items, their contribution to total carbon fluxes in the brackish zone is negligible: each year some 6% of all consumed carbon in the brackish part of the estuary passes through the copepod food web.     

Nitrogen dynamics in the Westerschelde estuary (SW Netherlands) estimated by means of the ecosystem model MOSES

A tentative nitrogen budget for the Westerschelde (SW Netherlands) is constructed by means of a simulation model with thirteen spatial compartments. Biochemical and chemical processes in the water column are dynamically modeled; fluxes of dissolved constituents across the water-bottom interface are expressed by means of diagenetic equations.The model is calibrated on a large amount of observed variables in the estuary (1980–1986) with relatively fine temporal and spatial detail. Additional constraints are imposed by the stoichiometric coupling of carbon, nitrogen and oxygen flows and the required conservation of mass. The model is able to reproduce rather well the observed distributions of nitrate, ammonium, oxygen and Kjeldahl nitrogen both in time and space. Also, model output of biochemical oxygen demand and total organic carbon falls within observed ranges.By far the most pervasive process in the nitrogen cycle of the estuary is nitrification which mainly takes place in the water column of the upper estuarine part. On average about three times as much nitrate is leaving the estuary at the sea side compared to what enters from the river and from waste discharges. Ammonium on the other hand is consumed much faster (nitrification) than it is regenerated and only about one third of the total import leaves the estuary at the sea side. The budget for detrital nitrogen reveals import from the river, from wastes and from the sea. Phytoplankton uptake of inorganic nitrogen is negligible in the model.About 21% of total nitrogen, 33% of inorganic nitrogen, is removed from the estuary (mainly to the atmosphere through denitrification) and the load of nitrogen net exported to the sea amounts to about 51 000 tonnes per year. Total denitrification in our model is lower than what was estimated in the literature from the late seventies, where a nitrogen removal up to 40–50% of the total inorganic load was reported. Part of the differences could be methodological, but inspection of the nutrient profiles that led to these conclusions show them to be different to the ones used in our study. The oxygen deficient zone has moved upstream since the late seventies, entrailing the zone of denitrification into the riverine part of the Schelde. The nitrification process now starts immediately upon entering the estuary.     

Exotic oniscideans (Crustacea: Isopoda) in coastal salt marshes: first record of the families Halophilosciidae and Platyarthridae in Continental Chile

We report two new exotic introduced isopods in Chile: Halophiloscia couchii of the family Halophilosciidae and Niambia capensis of the family Platyarthridae. This report provides first evidence of the presence of these species in the Southeastern Pacific. These species were found in several salt marshes and beaches spanning nearly 2000?km of the Chilean coast. With the data reported here, the percentage of exotic oniscidean isopods in Chile is 28%.     

The influence of tidal marshes on upland groundwater discharge to estuaries

We investigated subsurface hydrology in two fringing tidal marshes and in underlying aquifers in the coastal plain of Virginia. Vertical distributions of hydraulic conductivity, hydraulic head and salinity were measured in each marsh and a nearby subtidal sediment. Discharge of hillslope groundwater into the base of the marshes and subtidal sediment was calculated using Darcy's law. In the marshes, fluxes of pore water across the sediment surface were measured or estimated by water balance methods. The vertical distribution of salt in shoreline sediments was modeled to assess transport and mixing conditions at depth. Hydraulic gradients were upward beneath shoreline sediments; indicating that groundwater was passing through marsh and subtidal deposits before reaching the estuary. Calculated discharge (6 to 10 liters per meter of shoreline per day) was small relative to fluxes of pore water across the marsh surface at those sites; even where discharge was maximal (at the upland border) it was 10 to 50 times less than infiltration into marsh soils. Pore water turnover in our marshes was therefore dominated by exchange with estuarine surface water. In contrast, new interstitial water entering subtidal sediments appeared to be primarily groundwater, discharged from below. The presence of fringing tidal marshes delayed transport and increased mixing of groundwater and solute as it traveled towards the estuaries. Soil-contact times of discharged groundwater were up to 100% longer in marshes than in subtidal shoreline sediments. Measured and modeled salinity profiles indicated that, prior to export to estuaries, the solutes of groundwater, marsh pore water and estuarine surface water were more thoroughly mixed in marsh soils compared to subtidal shoreline sediments. These findings suggest that transport of reactive solutes in groundwater may be strongly influenced by shoreline type. Longer soil-contact times in marshes provide greater opportunity for immobilization of excess nutrients by plants, microbes and by adsorption on sediment. Also, the greater dispersive mixing of groundwater and pore water in marshes should lead to increased availability of labile, dissolved organic carbon at depth which could in turn enhance microbial activity and increase the rate of denitrification in situations where groundwater nitrate is high.     

外源氮对沼泽湿地CH4和N2O通量的影响

三江平原沼泽湿地受到大气沉降、地表径流、农业排水等外源氮素的输入,对湿地生态系统CH4和N2O通量有重要影响。采用野外原位施肥试验模拟外源氮输入,设0,60,120,240kgN·hm^-24种试验处理,探讨外源氮对沼泽湿地CH4和N2O通量的影响。结果表明,外源氮促进了CH4和N2O排放。与对照处理比较,各施氮水平CH4平均排放通量分别增加了181％,254％和155％,N2O排放通量分别增加了21％,100％和533％。外源氮输入对CH4排放的季节变化形式影响不大,而N2O的季节变化形式随着氮输入表现出波动变化的趋势。不同施氮水平对CH4排放的促进作用与植物生长阶段和产CH4的微生物过程密切相关,N2O排放通量随氮输入量呈指数增加（R^2=0．97,P〈0．01）。外源氮通过影响湿地微生物过程来进一步影响CH4和N2O的排放。     

Physicochemical environments and tolerances of cyprinodontoid fishes found in estuaries and salt marshes of eastern North America

Individuals of 28 species of cyprinodontoid fishes have been reported from estuaries/salt marshes of the Atlantic and Gulf coasts of North America. Some species show limited latitudinal distributions and/or occupy a limited range of habitats; others are widely distributed and/or occupy a wide range of habitats.A literature survey was made of conditions of water temperature, dissolved-oxygen (DO) concentrations, and salinities at sites where individuals of each species had been collected, and of laboratory-determined tolerances or lethal limits and other responses to those abiotic conditions. Individuals of Cyprinodon variegatus showed the widest overall range of tolerance of environmental temperatures, −1.9–45.4°C, with Gambusia rhizophorae showing the highest lower temperature-tolerance limit, 17°C. The only species highly sensitive to hypoxia was Floridichthys carpio, which showed “stress” at DO levels of 6–8 mg kg⁻¹. All showed use of aquatic surface respiration, except for Kryptolebias marmoratus, which uses aerial respiration in the presence of H₂S, and/or under hypoxic conditions. Individuals of C. variegatus were found to tolerate ambient salinities ranging from < 0.5 to 125.2, or higher, and several species of the genus Fundulus were found to tolerate concentrations ranging from <0.5 to ≥100. However, some of the species discussed cannot tolerate salinities beyond those of dilute brackish waters. In most instances, laboratory-determined tolerance limits of temperature and salinity were wider than conditions under which individuals of these species had been found in nature. The majority of available information related to adult individuals, with few studies focused on immature stages; however, existing information permitted a brief review of spawning, incubation, and early development features in Fundulus heteroclitus.Suggestions were made, based on existing information, as to species that would be most likely to show altered population distributions resulting from continued global warming. These included five species that have tropical/subtropical, or subtropical/temperate distributions. Also, a few others were included that show extensive latitudinal distributions, most extending northward into cooler temperate regions of the Atlantic coast. At present, none of these species has shown a range alteration that can be attributed to global warming.     

Actual litter decomposition rates in salt marshes measured using near-infrared reflectance spectroscopy

Near-infrared reflectance spectroscopy (NIRS) has been widely applied as a holistic tool to investigate decomposition processes in terrestrial ecosystems. The objectives of this research were to determine the potential of NIRS to predict (1) the halophytic litter chemistry (i.e., carbon and nitrogen content) during decomposition, and (2) the stage of decomposition of halophytic litter. Decomposition experiments were conducted in the laboratory with microcosms placed under a wide range of physical characteristics and in the field with litterbags located along the elevation gradient (i.e., low to upper marsh). Microcosm experiments were used to calibrate the predictive equations. These calibration equations were then applied to the field data to test their capacity to predict %C, %N, and litter mass loss (LML). NIRS can be successfully applied to predict chemical composition of halophyte litter during decomposition processes. We hypothesized that the use of litterbags in the field might lead to a 20–40% overestimation of the decay rate as fine organic debris are lost through the meshes of the litterbags. NIRS can be used as a fast and nondestructive method to more accurately predict decay rates, and thus microbial consumption in aquatic environments.     

Short-term fluctuations in benthic diatom numbers on an intertidal sandflat in the Westerschelde estuary (Zeeland,The Netherlands)

During the period March–May 1991, sediment samples were taken every two or three days at one intertidal station in the brackish part of the Westerschelde estuary. Quantitative cell counts were made in order to investigate the short-term temporal changes in diatom numbers and assemblage structure.Throughout the whole sampling period, the diatom assemblage was dominated by epipsammic diatoms. Three species, Achnanthes delicatula, Opephora cf. perminuta and Catenula adhaerens on average accounted for almost 67% of all valves counted. The epipsammic diatom fraction showed no significant changes in absolute numbers; its species composition appeared relatively stable. In contrast, epipelic diatom densities significantly increased towards the end of the study period. Species composition within this fraction was less stable. Multivariate analysis (Principal Components Analysis), in combination with multiple regression, indicated that total sky irradiance (on the second and third day preceding sampling) and percentage organic matter were related to the short-term fluctuations of the epipelic diatom fraction.     

Restoration of salt marshes in the Netherlands

The conquest of land from the sea has been a long tradition in the Netherlands. When salt marshes were high enough, they were embanked when it was economically feasible, and transformed into intensively exploited agricultural land. This resulted in the transformation of halophytic communities to glycophytic communities. Often as an alternative, a low levee, a summerdike was built, which greatly reduced the flooding frequency of the landward summerpolder, hence creating a sedimentation deficit therein. Such summerpolders now cover 1200 ha in the Netherlands, 2100 ha in NW-Germany and small areas in England. Due to continuous embankments, the present salt-marsh area is relatively small with respect to the tidal basins. Discussions have been started how to increase the salt-marsh area. Two options will be discussed, firstly de-embankment of summerpolders and maintenance of the protective seawall, secondly increase of the effects of saline seepage behind the seawall by top soil removal. Both options include the restoration of salt-marsh communities (target communities) in intensively agriculturally exploited sites that have been salt marshes before. From the few examples abroad and experiments it is discussed (1) to which extent the sedimentation deficit in summerpolders could be compensated for, (2) if the soil seed bank is likely to contribute to re-establishment of salt-marsh communities, (3) if the dispersal of propagules of halophytic plants will be possible by hydrochory when the summerdike is breached, (4) to what extent is dispersal by endozoochory through waterfowl important in case re-establishment in a saline seepage area behind the seawall without open connection to the sea is envisaged. Two case studies of de-embanked summerpolders in the Netherlands revealed that the sedimentation deficit can be counteracted by rapid sedimentation, provided enough transport is possible from the foreshore. Dispersal by incoming tidal water from the nearby salt-marsh source area into the target area is possible for many salt-marsh plant species. The rate of success seems to depend on the relative position of source area and target area. A case study in a saline seepage area after top soil removal in the Netherlands, showed that the number of viable seeds dispersed by droppings from waterfowl is limited. Hence the possibilities for restoration of inland halophytic plant communities seem much lower than after de-embankment of summerpolders.     

The concomitant existence of a typical coastal and a detritus food chain in the Westerschelde estuary

Until now studies of the aquatic ecology of the Westerschelde estuary have been erratic. Yet, from the results of a literature research some structural aspects of the ecosystems in the Westerschelde could be derived.Clear gradients could be found from the river to the sea for the abiotic as well as for the biotic features of the estuary. On the basis of the available quantitative data 2 tentative food chains were distinguished; a mainly detritus-based food chain in the upstream brackish part and a coastal food chain in the downstream seaward part. The centre of the area of the detritus food chain coincided with the zone of a turbidity maximum at the interface of salt and fresh water. This food chain is characterized by a low primary production but a year-round high concentration of suspended organic matter. The suspended organic matter consists of aggregates of detritus and bacteria. The zooplankton as well as the zoobenthos can reach high biomasses. The coastal food chain is mainly based on a seasonally chainging high primary production. The diversity of flora and fauna in this system is much higher than in the detritus food chain, although the biomass can be lower.Communication no. 404 of the Delta Institute, Yerseke.     

Below-ground biomass in healthy and impaired salt marshes

Twelve salt marshes in south Louisiana (USA) were classified as either impaired or healthy before a summer sample collection of above- and below-ground biomass and determination of sediment accretion rates. The above-ground biomass of plant tissues was the same at both impaired and healthy salt marshes and was not a good predictor of marsh health. However, below-ground root biomass in the upper 30cm was much lower in the impaired marshes compared to the healthy marshes. Compromises to root production apparently occur before there is an obvious consequence to the above-ground biomass, which may quickly collapse before remedial action can be taken. The subsequent change in vertical position of the marsh surface may be equivalent to many years of accretion, and be irreversible within decades without considerable effort. These results are consistent with the hypothesis that it is the plants below-ground accumulation of organic matter, not inorganic matter that governs the maintenance of salt marsh ecosystem in the vertical plane. Reversing the precursor conditions leading to marsh stress before the collapse of the above-ground biomass occurs is therefore a prudent management objective and could be easier than restoration.     

Changes in phytoplankton biomass and primary production between 1991 and 2001 in the Westerschelde estuary (Belgium/The Netherlands)

The Westerschelde estuary is a very polluted and turbid estuary, but the last decade the waterquality improved. Dredging activity also increased in 1997 to allow bigger ships to enter the port of Antwerpen. This could potentially decrease the light conditions for the phytoplankton. Because of all these recent changes in the estuary we studied primary productivity in 2001 and compared it to values in 1991. The results show that due to a decrease in discharge in particulate and dissolved organic carbon the oxygen concentrations in general have increased in the upstream region, although in spring and summer low oxygen concentrations (10–30% saturation) can still be found. Phosphate and ammonia concentrations have decreased and the zone of nitrification which was very large in 1991 has become very small and is now located in the uppermost upstream region of the estuary. Si-concentrations have remained the same. All nutrient concentrations are still high enough not to limit phytoplankton growth. Turbidity remained unaltered as a result of the dredging works, and as a result phytoplankton biomass in most of the estuary did not show a decrease, although there were signs that in the upstream region phytoplankton biomass decreased, possible caused by increased grazing pressure. The relationship between phytoplankton biomass and primary productivity did not change, and from the data it can be concluded that the dredging activity will not influence the gross and net primary productivity of the phytoplankton.     

Freshwater marshes as dissolved silica recyclers in an estuarine environment (Schelde estuary, Belgium)

Compared to knowledge about N and P processing in the aquatic continuum of lakes, wetlands and estuaries, knowledge concerning transport and cycling of Si is only fragmentary. Furthermore, Si research in estuaries has mainly been focused on subtidal benthic sediments and uptake and recycling by diatom communities. The biogeochemical cycling of Si in tidal wetlands, which can contain large amounts of Si, has thus far been neglected. We have conducted several whole ecosystem Si mass-balances on a freshwater marsh located in the Schelde estuary (6 tidal cycles, 2 with BSi included). Our measurements show that the freshwater marsh acts as an important source of dissolved Si to the main river (1–18% more export than import, on average 0.114 g m^–2). This export is compensated by import of amorphous silica into the marsh (19–55% more import than export). The marsh was shown to act as silica recycler, resupplying biologically available dissolved Si to the estuarine ecosystem. Extrapolations show that during summer and spring months, when dissolved silica is depleted due to diatom growth, almost half of the total dissolved silica load in the main river channel could result from marsh recycling.     

Comparison of physical characteristics between created and natural estuarine marshes in Galveston Bay,Texas

Five natural and ten created Spartinaalternifloramarshes in the Lower Galveston BaySystem were compared to determine if there weresignificantly different physical characteristicsassociated with each type of marsh. The saltmarsheswere compared on the basis of microhabitats,length-width ratio, area-perimeter ratio, marsh-wateredge ratio, total size of S. alternifloraplantcommunities, fetch distances, angle of exposure,orientation, and elevation. All physicalmeasurements, except for elevation, were obtained fromphotography analyzed with the use of a GeographicInformation System with digital image processingcapabilities. Differences existed between natural andcreated marshes. The natural marsh sites in this studywere characterized by highly undulant marsh-wateredges, island-like S. alternifloraplant stands,concave shorelines, and low elevations. Createdmarshes were characterized by relatively smoothmarsh-water edges, an unbroken shoreline morphology,convex to straight shoreline configurations, andelevations on the edge and inner portions of the marshhigher than those of natural marshes. The lowelevations of the natural marsh appear to be due tocoastal subsidence in the Galveston Bay area alongwith rising sea level. Reticulated marshes andundulant shorelines appear to be caused by consequentdrowning of the natural marshes. High elevations insome of the created marshes are related to erosion ofthe low elevation marsh or deposition of coarsesediments at the marsh-water edge.     

Nutrients,light and primary production by phytoplankton and microphytobenthos in the eutrophic,turbid Westerschelde estuary (The Netherlands)

Abiotic factors and primary production by phytoplankton and microphytobenthos was studied in the turbid Westeschelde estuary. Because of the high turbidity and high nutrient concentrations primary production by phytoplankton is light-limited. In the inner and central parts of the estuary maximum rates of primary production were therefore measured during the summer, whereas in the more marine part spring and autumn bloom were observed. Organic loading is high, causing near anaerobic conditions upstream in the river Schelde. Because of this there were no important phytoplankton grazers in this part of the estuary and hence the grazing pressure on phytoplankton was minimal. As this reduced losses, biomass is maximal in the river Schelde, despite the very low growth rates.On a number of occasions, primary production by benthic micro-algae on intertidal flats was studied. Comparison of their rates of primary production to phytoplankton production in the same period led to the conclusion that the contribution to total primary production by benthic algae was small. The main reason for this is that the photosynthetic activity declines rapidly after the flats emerged from the water. It is argued that CO₂-limitation could only be partially responsible for the noticed decrease in activity.     

The role of salt marshes in the Mira estuary (Portugal)

The Mira estuary is a narrow entrenched pristine estuary of the Ria type, about 30 km long. It comprises an area of 285 ha of salt marsh, of which250 ha have been proposed for reclamation for aquaculture. Dredging, village and recreation development menace the yet undisturbed estuarine ecosystem. To assess the biological importance of this wetland, a multidisciplinary study was conducted in apart of the salt marsh, considered as being representative of the whole area. Halophytic vegetation covering 75% of the total salt marsh site is dominated by Spartina maritima (28% of total vegetation area). Total primary production attains63,766 kg/yr (dw). A net export of 1541 kg/yr of COM to the relatively oligotrophic adjacent waters was also found. Insects and birds are described for the first time in the saltmarsh. Macrobenthic communities are dominated by Hediste diversicolor, Nepthys caeca and Scrobicularia plana. The fiddler crabUca tangeri attains here its north distribution limit. The mud flats and creeks associated with the salt marsh act as a nursery for 40.8% of the fish species present. The food web is dominated by detritivorous species like the grey mullets. The results obtained in this study support the need for an effective conservation of this area. This revised version was published online in July 2006 with corrections to the Cover Date.     

The burrowing crab Neohelice granulata affects the root strategies of the cordgrass Spartina densiflora in SW Atlantic salt marshes

Salt marshes are among the most productive systems of the world, with plant primary production limited by soil oxygen deficiency and nutrient availability. Nevertheless, root adaptations to anoxia and nutrient acquisition are different and often incompatible. The SW Atlantic salt marshes are characterized by high densities of the deep (up to 1 m) burrowing crab Neohelice granulata (Dana, 1851) that may change soil physical and chemical characteristics by burrow construction. In this work, we experimentally evaluated the hypothesis that crab burrowing can enhance soil oxygenation, causing changes in Spartina densiflora Brongniart below ground tissues from structures adapted to anoxia to systems adapted for nutrient acquisition. This response, in turn, would enhance plant productivity. Results from field observations show that oxygen availability is higher in zones with high burrow densities. As burrow densities increased, the plant root distribution changed from shallow (associated to low oxygen availability) to deeper and vertically homogeneous, with a positive correlation between burrow density and plant aboveground biomass. Experimental exclusion of crabs shows that they induce changes of root strategies from anoxia toleration to nutrient efficient acquisition, with increasing plant productivity. The invasive success that this plant shows in other parts of the world is likely to be due to their ability to tolerate harsh environmental conditions. Our results suggest that the morphological plasticity of S. densiflora is also important in their native zone given the characteristics of their specific habitat.