Importance of Vascular Plant and Algal Production to Macro-invertebrate Consumers in a Southern California Salt Marsh

The dietary importance of marsh vascular plants (primarilySalicornia virginica), algae and upland particulate inputs to macro-invertebrate consumers was studied in Carpinteria Salt Marsh, southern California, using stable carbon and nitrogen isotope ratios. This marsh is predominantly a marine or hypersaline system and succulents are the most common vascular plant species. Of invertebrates collected from the vegetated marsh, tidal flats and channels, only detritivores from the vegetated marsh (Traskorchestia traskiana,Melampus olivaceus) had isotope values (δ¹³C=−20‰) that suggested some use ofSalicornia-derived carbon.T. traskianacultured in the laboratory on decomposingS. virginicaor blue-green micro-algal mat had distinctive isotopic signatures, reflecting the capability of this consumer to assimilate carbon and nitrogen derived from these sources. The δ¹³C values (generally −16‰ to −15‰) of species from tidal flats and channels (e.g.Cerithidea californica,Protothaca staminea,Mytilus galloprovincialis,Neotrypaea californiensis) were most similar to values for benthic algae and phytoplankton. Specimens ofM. galloprovincialisalong a gradient of presumed increase in marine influence had similar isotope values, suggesting little contribution to diet from upland runoff. The present results differ most noticeably from published values in the¹³C enrichment of suspension-feeders, suggesting the use of resuspended¹³C-enriched benthic microalgae in tidal channels by these consumers, and in the¹³C depletion and¹⁵N enrichment of plants and consumers along a portion of the marsh boundary receiving inputs of nutrient-enriched perched groundwater. In general, the isotopic composition of macro-invertebrates indicated the incorporation of algal production rather than ofS. virginicaor upland sources into the marsh food web.     

A Stable Carbon Isotope Study of the Food-web in a Freshwater-deprived South African Estuary, with Particular Emphasis on the Ichthyofauna

The importance of macrophytes as food sources for estuarine nekton is unclear. Previous carbon isotope investigations in the macrophyte-dominated, freshwater-deprived Kariega Estuary showed that the bivalveSolen cylindraceusdid not utilize the dominant estuarine macrophytes found within the estuary as a primary food source. This finding prompted questions as to what the nekton of this estuary utilize as primary energy sources. δ¹³C analyses of the principal autochthonous and allochthonous primary carbon sources, as well as the dominant invertebrate and fish species, indicate that there are two main carbon pathways within the Kariega Estuary. The littoral community, which incorporates the majority of crustaceans, gobies, mullet and a sparid, utilizes δ¹³C enriched primary food sources namelySpartina maritima,Zosteracapensis and epiphytes. The channel fauna, which includes the zooplankton, zooplanktivorous and piscivorous fish, utilizes a primary food source depleted in δ¹³C, which is most likely a mixture of phytoplankton, terrestrial plant debris and C₄macrophyte detritus. The C₃saltmarsh macrophytesSarcocornia perennisandChenolea diffusa, as well as benthic microalgae, appear to be less important as primary food sources to the nekton of the Kariega Estuary.     

Impacts of maintenance channel dredging in a northern Adriatic coastal lagoon. II: Effects on macrobenthic assemblages in channels and ponds

Coastal lagoons are ephemeral habitats whose conservation requires human intervention, such as maintenance dredging of inner channels. Dredging can reduce the abundance of benthic species due to the removal of individuals with the sediment, modify sediment properties, and resuspend fine sediment, nutrients and pollutants, which can lead to eutrophication, hypoxic events and increasing toxicity. Both direct effects in the dredged channel and possible indirect effects in surrounding shallow areas could be expected. This study assesses the effects of the channel maintenance dredging, performed between October 2004 and August 2005, on the invertebrate assemblages both in channels and adjacent ponds in the northern Adriatic coastal lagoon of Pialassa Baiona. The lagoon is affected by eutrophication, chemical and thermal pollution from wastewater treatment and power plants. Three impacted sites were located in the dredged channel and three in the adjacent interconnected shallow water ponds, while three non-impacted sites were located in a channel and in a pond far from the dredged area. Replicate samples were collected from each site one time before and one time after the dredging operations. Despite the extent of the intervention, effects of the dredging on macrobenthic assemblages were detected only within the dredged channel, while in the surrounding ponds no clear and unequivocal effects were found. In particular the dredging could have promoted the increase of the abundance of the polychaete Streblospio shrubsolii in the southern and central parts of the dredged channel and the increase in abundance of the amphipod Corophium insidiosum in the northern side, compared to the controls. Instead, species diversity was reduced in the central and northern parts of the dredged channel. These effects on the macrobenthic invertebrate assemblages could be related to the observed changes of sediment characteristics, contamination and toxicity. Overall, direct effects on benthic assemblages in the dredged channels were more detectable than the possible secondary effects in the surrounding shallow ponds, where the higher spatial heterogeneity can mask any relevant effects.     

Incorporation of terrestrial wetland material into aquatic food webs in a tropical estuarine wetland

Carbon and nitrogen stable isotope composition of a range of organisms collected from two intermittently connected floodplain pools in the Ross River estuary were analysed to assess the extent to which carbon fixed by terrestrial wetland producers is incorporated into adjacent aquatic food webs. The two pools differed in surrounding vegetation with one surrounded by mangroves and the other by the salt couch Sporobolus virginicus. At both pools, animals showed differences in δ¹³C, indicating differences in sources of carbon. Since δ¹³C values of C₃ mangroves (−29.7 to −26.3‰) were very different from those of the C₄ salt couch (−16.3 to −15.4‰), it was possible to determine the importance of terrestrial wetland producers by comparing isotope values of consumers between sites, in a species by species approach. Most animal species collected showed lower δ¹³C at the mangrove pool than at the Sporobolus pool, which indicates a greater incorporation of mangrove carbon at the mangrove pool. However, the animals’ isotopic shifts were also similar to that shown by epiphytes, and hence the differences in animal δ¹³C could also be a result of a dependence on these producers. The IsoSource model was useful to clarify this question, indicating that mangrove and salt marsh material was a crucial contributor to the diet of several fish and invertebrate species at both sites, indicating that carbon of terrestrial origin is incorporated in the estuarine food web.     

潮沟系统水沙输运研究——以长江口崇明东滩为例

本研究以崇明东滩2015年4月实测潮间带水沙数据为基础,分析了潮沟、盐沼及光滩的水沙特征,重点研究了潮沟系统及邻近潮滩潮周期内悬沙通量情况。结果表明:（1）潮沟表层沉积物比潮滩细,二者平均中值粒径分别为21.7 μm和33.0 μm,悬沙粒径由海向陆逐渐变小;（2）大、小潮沟潮周期内潮流均以往复流为主,垂向平均流速分别为15.4 cm/s和34.6 cm/s;盐沼界和光滩则以旋转流为主,平均流速分别为11.3 cm/s和28.9 cm/s;（3）潮沟中的高悬沙浓度出现在涨潮初期,最大可达7.5 kg/m3,而潮滩高悬沙浓度则出现在潮落潮中期和高水位时刻;大、小潮沟和盐沼界站涨潮阶段平均悬沙浓度大于落潮阶段,光滩站则相反。潮沟悬沙主要来自邻近水域,而潮滩悬沙则与滩面表层沉积物密切相关;（4）潮沟在潮周期内净输沙方向均指向滩地,大潮沟潮周期单宽净输沙量可达4.0 t/m;盐沼界处垂直岸线和沿岸输沙强度相近,净输沙由海向陆,潮周期离岸输沙强度为1.0 t/m;光滩沿岸输沙强度远大于垂直岸线输沙,光滩净输沙由陆向海。研究揭示了潮间带潮沟系统的强供沙能力以及研究区域光滩冲蚀,盐沼植被带淤积的动力地貌过程。     

Nutrient dynamics and seasonal succession of phytoplankton assemblages in a Southern European Estuary: Ria de Aveiro, Portugal

The spatial and temporal dynamics of dissolved inorganic nitrogen, dissolved phosphate, dissolved silica and chlorophyll a were measured seasonally at eight stations in the Ria de Aveiro. Between December 2000 and September 2001, the seasonal succession of phytoplankton assemblages, inferred after the spatial and seasonal variation of silica and of chlorophyll a concentrations, showed that diatoms (μmol Si L⁻¹) dominated from late autumn until early spring, while chlorophytes (μg Chl a L⁻¹) bloomed during late spring and summer. The Si:N:P ratios and Si concentrations indicated no seasonal depletion in dissolved silica, as in other temperate systems, possibly because of abnormal precipitation and flood events prolonging the supply of dissolved Si to the system. The Si:N:P ratios suggested P limitation at the system level. Despite the relative proportions of available nutrients, the mean phosphorus concentration (5.3 μmol L⁻¹) was above the reported half-saturation constants for P uptake by phytoplankton. Thus, in Ria de Aveiro, the seasonal succession of phytoplankton assemblages may also be dependent on the grazing capacity of the pelagic community through top-down regulation.     

Larval fish assemblages,environment and circulation in a semienclosed sea (Gulf of California,Mexico)

Fish larvae and hydrographic data collected in the Gulf of California (GC) in December 2002 are used to describe larval fish assemblages (LFAs) and to explore their relationships with environmental variables (temperature, salinity, dissolved oxygen, fluorescence maximum, ?$?$ and superficial chlorophyll a). The Bray–Curtis dissimilarity index defined three LFAs, distributed in areas with distinctly different environmental conditions. The affinity of most of the species with the environmental characteristics of their areas of distribution could be interpreted as an indication that spawning occurred inside those areas. Particle tracking in current fields from a 3D numerical model and connectivity matrices are used to assess larval retention in the LFA areas. The technique is well suited for seas like the GC that have well-defined circulation patterns. On time scales around 30 days, retention (from 56% to 73% of the particles) occurred (1) for the North LFA in the Upper GC, (2) for the Channel-Center LFA in the anticyclone over the Northern GC and in Ballenas Channel, and (3) for the South LFA in the eddy over San Pedro Mártir basin and in the shallow zone off the peninsula. Therefore, the Lagrangian analysis revealed that the observed LFAs have a permanency long enough to allow fish larvae to remain in a favorable environment until they develop motility. The main particle export path (less than 26% of the particles) was from the North to the South LFA, following the anticyclonic main flow and coinciding with the gradient in species number and larval abundance.     

Colonization process by macrobenthic infauna after a managed coastal realignment in the Bidasoa estuary (Bay of Biscay, NE Atlantic)

The colonization process by macrobenthic infauna after the completion of managed realignment on the fertile plain of Jaitzubia, along the Bidasoa estuary, is assessed here. The benthic invertebrate fauna was sampled annually for three years in the newly recovered intertidal areas. The results show a rapid colonization process. Six months after the completion of the restoration works, all the stations exhibited a large number of small opportunistic benthic species, as well as some large individuals of Hediste diversicolor, that dominated the biomass of the community. One year later, a new stage of development or succession towards the settlement of the Scrobicularia plana–Cerastoderma edule community was observed, showing a reduction of both abundance and species richness, and an increase in biomass, mainly due to the presence of individuals of S. plana (which were absent previously). The following year, a further reduction in the abundance and number of species was observed, as well as an increase in biomass, but on this occasion S. plana was the main contributor to community biomass. Evidence of the importance of passive or active dispersion in the adult stage of the species H. diversicolor and S. plana in the recovery process was also observed.     

Assessment of spatial variation in carbon utilization by benthic and pelagic invertebrates in a temperate South African estuary using stable isotope signatures

Stable isotope analyses (δ¹³C and δ¹⁵N) were used to evaluate the spatial variations in carbon flow from primary producers to consumers at two sites in the temperate and permanently open Kariega Estuary on the southeastern coast of South Africa during October 2005 and February 2006. One site was located opposite a salt marsh while the second was upstream of the marsh. Except for significantly enriched δ¹³C values of Zostera capensis and surface sediments near the salt marsh, the δ¹³C and δ¹⁵N signatures of the producers were similar between sites. The invertebrates were clustered into groups roughly corresponding to the predominant feeding modes. The suspension feeders showed δ¹³C values closest to the seston, whereas the deposit feeders, detritivores and scavengers/predators had more enriched δ¹³C values reflecting primary carbon sources that were likely a combination of seston, Spartina maritima and Z. capensis at the upstream site, with an increased influence of benthic algae and Z. capensis at the salt marsh site. The δ¹⁵N signatures of the consumers showed a stepwise continuum rather than distinct levels of fractionation, indicating highly complex trophic linkages and significant dietary overlap among the species. Consumers exhibited significantly enriched δ¹³C values at the salt marsh site, an effect that was attributed to enriched Z. capensis detritus in this region in addition to increased phytoplankton biomass in their diets compared with invertebrates living upstream. The data reinforce the concept that between-site variations in the stable isotope ratios of consumers can result not only from dietary shifts, but also from alterations in the isotope ratios of primary producers.     

N2, N2O and O2Profiles in a Tagus Estuary Salt Marsh

Vertical gas profiles of N₂, N₂O and O₂were obtained in intact sediment cores from a Tagus estuary salt marsh using membrane inlet mass spectrometry. This technique allows direct measurements of dissolved gas concentrations with minimal disturbance. O₂concentrations decreased sharply with depth, becoming undetectable below 14mm. Denitrification products (N₂and N₂O) occurred in the surface layer of the sediment where O₂was present. Diffusion of N₂and N₂O from the anaerobic zone, denitrification in anaerobic microsites and aerobic denitrification are possible explanations for this observation. N₂was the sole product of denitrification in control sediment cores probably because of the great demand for electron acceptors in this sediment. The addition of NO₃ ⁻ and CH₃CO₂ ⁻ increased the concentrations of N₂and N₂O in the sediment. Significantly higher concentrations in treated cores occurred between 1·5 and 2·0cm for N₂and between 0·5 and 1·5cm for N₂O. The peak in N₂concentration occurred in the anaerobic zone of the sediment, close to the aerobic–anaerobic interface while the peak in N₂O concentration occurred above this interface where concentrations of O₂were approximately 10μM. This is indicative that, in this sediment, production of N₂O is less sensitive to the presence of O₂than reduction of N₂O to N₂.     

The Distribution of Talitrid Amphipods (Crustacea) on a Salt Marsh in Southern Tasmania, in relation to Vegetation and Substratum

Supralittoral and terrestrial talitrid amphipods were collected from a salt marsh in Southern Tasmania by pitfall trapping along transects from the mid-tidal level to above the high-tide mark, and by hand collecting from sites chosen to represent the full range from salt marsh to terrestrial vegetation. At each site, the cover of all major plant species was recorded, and soil samples were collected from which soil moisture, organic content and salinity were measured.Eight talitrid species from four ecological groups were collected; one palustral species, one beachflea, three coastal landhoppers and three eastern forest landhoppers. There was substantial overlap in the distributions of these groups. The undescribed beachflea had the widest distribution, from the wettest, most saline sites to theSchoenus nitenstussock grassland at the extreme high-tide mark. The palustral species,Eorchestia palustris, overlapped substantially with the beachflea, but was found within a narrower band of salinities (though not in the most saline sites) and in more poorly-drained sites than the beachflea. Coastal landhoppers,Austrotroides maritimus,Keratroides rexand an undescribed species ofTasmanorchestia, were found mainly in theS. nitenstussock grassland, where they overlapped with forest landhoppers,Keratroides vulgaris,Mysticotalitrus tasmaniaeandM. cryptus, which were found mainly in non-saltmarsh terrestrial sites, well above the high-tide mark.These distributions are discussed in terms of the likelihood that salt marshes provided the route by which talitrid amphipods colonized land. There is no reason from these data to reject salt marshes as the route to land, and it is suggested that they are a more likely route than via rocky or sandy shores.     

枯季长江口南汇东滩潮间带盐沼湿地冲淤变化对大型底栖动物功能群的影响

盐沼湿地的冲淤变化是影响大型底栖动物功能群组成及其分布特征的一个重要因素。本文选择长江口南汇东滩作为研究区域,设置固定样地,分别于2015年11月和2016年2月,进行了大型底栖动物和沉积物理化因子的季节性取样分析,并在期间内每月进行滩面冲淤变化的观测,分析了枯季长江口南汇东滩潮间带盐沼湿地冲淤变化对大型底栖动物功能群的影响及作用机理。调研期间共采集到大型底栖动物37种,从功能群组成来看,以杂食者居多,共25种;植食者次之,共7种;肉食者最少,仅为5种。3种食性功能群的物种数、密度和生物量的时空变化特征有所不同。调研期间,3个功能群物种数都略有下降;植食者密度显著增加（P<0.01）,而肉食者、杂食者密度变化不显著（P>0.05）;3个功能群的生物量变化均不显著（P>0.05）。植食者密度和生物量在样地间都存在极显著差异（P<0.01）;肉食者和杂食者仅密度在2016年2月的调研中存在样地间的显著差异（P<0.05）。不同样地月际冲淤变化特征有所不同,但总体均呈淤涨态势。多数环境因子存在显著的空间差异（P<0.05）,仅不同深度的沉积物中值粒径在样地间无显著差异（P>0.05）。滩面冲淤变化对沉积物理化指标的变化具有主导作用。通过非线性回归分析筛选出的与各功能群物种数、密度、生物量变化具有显著相关性（P<0.05）的环境因子组合存在差异。其中,对植食者密度、生物量的变化和肉食者物种数的变化有显著影响的环境因子组合中都包含绝对冲淤变化量。滩面冲淤变化除对大型底栖动物产生直接影响外,还会通过改变沉积物理化因子特征,间接对大型底栖动物产生影响。对于3种食性功能群来说,产生影响作用的主要生境因子存在明显差异。虽然各食性功能群的物种数均有下降,但相对另外两种功能群而言,沉积物中的营养环境还是更利于杂食者的生存,因此杂食者在竞争过程中更具优势。未来相关研究需要在多因子系统监测的基础上,结合洪季的观测结果,进行综合分析。     

The summertime net transport of dissolved oxygen,salt and heat in a salt marsh basin,North Inlet,S.C.

This study addresses the impact marshes have on the dissolved oxygen content of tidal waters, particularly during summer when respiratory demand for oxygen in adjacent coastal waters is at a maximum and the solubility of oxygen is lowest. The net transports of dissolved oxygen, salt and heat have been measured for 65 tidal cycles during late spring and summer for a small (0·14 km²) salt marsh basin near North Inlet, S.C. The results indicate that export of dissolved oxygen occurs only on tidal cycles that begin between 2:00 am and 10:00 am such that high tide occurs within 4 h of noon. The largest exports of oxygen and heat are produced by spring tides beginning near sunrise. Although the time window for oxygen export is only about 8 h in duration, there is a more or less overall long-term balance between export and import because the magnitude of oxygen export is about 25% greater than import. The magnitude of heat export similarly exceeds heat import but because the time windows for heat export and import are equal, there is an overall export of heat. This study thus suggests that in summer salt marshes of the Atlantic coast export heat and are in balance with respect to the export and import of dissolved oxygen. However, because of the interaction of the diurnal tide with the daily cycle of solar radiation, transient dissolved oxygen concentrations in tidal waters can range from 1.5 to 10.0 ppm. Thus loading of additional oxygen consuming materials to these waters possibly could lead to significant periods of anoxia.     

我国苏北建川潮汐盐沼有孔虫垂直分带的发现和特征

通过三个断面加密有孔虫表层采样和精确高程测量,发现建川开敞型盐沼存在特征性盐沼有孔虫组合,这些组合与盐沼植被带相应,受高程控制,有明显的垂直分带,垂直带顶面即最高高潮位以上有孔虫突然消失.这个有孔虫垂直分布模式可高精度地恢复古海面(精度最高可达±5cm).该研究为我国高分辨恢复古海面提供了一个地区性的现代有孔虫标尺,同时该工作也填补了国外仅局限于研究隐蔽型海岸盐沼有孔虫垂直分带的不足.     

Daily and inter-tidal variations of Fe,Mn and Hg in the water column of a contaminated salt marsh: Halophytes effect

It has been shown that salt marshes may function as efficient sinks for contaminants, namely for mercury. At the rhizo-sediment Hg may be associated with Mn and Fe oxyhydroxides, precipitated as sulphides or incorporated into organic matter. However, to our knowledge, in situ studies have not focused on the related processes at a daily or tidal cycle scales. Thus, the present work aims to study the effect of a common salt marsh halophyte in temperate latitudes (Sarcocornia perennis) on dissolved Fe, Mn and Hg concentrations in the water column. The in situ approach was carried out at a mercury-contaminated salt marsh and at the adjacent non-vegetated area (distance ≤ 4 m), covering two consecutive tidal cycles in order to include the photosynthetic active period and the night processes. During high tide no daily or spatial effects were observed on the concentrations of Mn, Fe and Hg in the water column, due to the dilution effect of the incoming seawater. During low tide the concentrations of Mn, Fe and Hg were significantly higher in the overlaying water column of the salt marsh. At S. perennis mats the concentration of dissolved total Hg was significantly related with the concentration of Mn (r = 0.459, p = 0.028, n = 23), but not with that of Fe (r = 0.367, p = 0.085, n = 23) while no significant relations were found at the adjacent non-vegetated sediments.     

The sources and accumulation rate of sedimentary organic matter in the Pearl River Estuary and adjacent coastal area, Southern China

In this study, the contents, sources and accumulation rate of sedimentary organic matter (OM) in the Pearl River Estuary (PRE) and adjacent coastal area were investigated. The stable carbon isotopic composition (δ¹³C) is a reliable geochemical proxy and was used to indicate the OM origin here. Nevertheless, the organic carbon and nitrogen molar ratios (TOC/TN) and the stable nitrogen isotopic composition (δ¹⁵N) were affected by diagenesis and could be the supplementary indicators. The sources of OM were estimated based on the two end-member model. The results showed that in the estuary, sedimentary OM originated from terrestrial and aquatic mixing origins, whereas, OM in coastal sediments was dominantly algae-derived. The accumulation rate of sedimentary OM was analyzed based on ²¹⁰Pb dating. Due to the sampling sites and the distinct hydraulic environments, the accumulation rates of TOC, aquatic and terrestrial OC were obviously higher in the estuary than in coastal area. TOC accumulation rates were 18–27 mg cm⁻² y⁻¹ in the estuary, and 0.84–3.6 mg cm⁻² y⁻¹ in coastal area. Aquatic OC accumulation rates were 7.9–11.3, 0.8–1.3, and 2.6–3.1 mg cm⁻² y⁻¹, and terrestrial OC accumulation rates were 9.7–16.3, 0.02–0.14, 0.16–0.42 mg cm⁻² y⁻¹ in cores 2, 5, 6, respectively. It could be seen from the high accumulation rate of organic matter in the estuary that, when nutrients increased in the river, phytoplankton biomass and productivity would also have increased. As a result, phytoplankton sinking and organic matter sedimentation usually increased with primary productivity, resulting in the observed accumulation rate of aquatic OC in the estuary. Furthermore, terrestrial OC accumulation rates in the estuary and coastal area showed an increasing trend with the age.     

Vertical distribution of barnacle larvae at a fixed nearshore station in southern California: Stage-specific and diel patterns

Patterns of vertical distribution in marine invertebrate larvae interact with coastal hydrodynamics to determine cross-shore distributions, dispersal ranges, and scales of connectivity among populations. We present observations on the vertical distribution of barnacle larvae from southern California, collected from 3 depth intervals every 2 h and over a 48-h period at a fixed nearshore station. Larvae were identified to species and stage of development, and their vertical distributions were tested for day/night and between-stage differences within each species, as well as for correlations with environmental variability. Stage-specific patterns of vertical distribution were distinct and consistent across species. Nauplii were most abundant near the surface at all times, and were numerically dominated by Chthamalus spp. (78% of N_n = 1835). Cyprids were substantially less abundant than nauplii in our samples (N_c = 146) and corresponded mostly to the subtidal barnacle Balanus nubilus (84%). Despite the lower counts and different species composition, our data suggest that cyprids remain within the mid-depth and bottom layers, and that changes in their mean depth of distribution (MDD) may track changes in pycnocline depth. Temporal changes in nauplii MDD were not correlated with water-column variability, wind forcing, or tidal height. The apparent vertical segregation of nauplii and cyprids may be related to previously documented differences in the cross-shore distribution of these stages.     

Observed fluxes of water,salt and suspended sediment in a partly mixed estuary

Observations of the residual fluxes of water, salt and suspended sediment are presented for seven stations along the Tamar Estuary. The data include measurements over single spring and neap tidal cycles, and are generally applicable to medium or high run-off conditions.Surface to bed differences in salinity are typically of the order of several parts per thousand. Gravitational circulation is an important component of residual flow in the deep, lower reaches of the estuary. Here, Stokes drift is insignificant. In the shallow upper reaches, the major residual currents are generated by Stokes drift and freshwater inputs. Data are compared with predictions from Hansen and Rattray's (1966) model of estuarine circulation.Salt fluxes due to tidal pumping and vertical shear are directed up-estuary at spring tides, tidal pumping being dominant. Tidal pumping of salt is also directed up-estuary at neap tides, although it is insignificant in the lower reaches, where vertical shear dominates.Tidal pumping of suspended sediment is directed up-estuary near the head at spring tides, and probably contributes to the formation of the turbidity maximum. The existence of the turbidity maximum is predicted using a simplified model of the transport of water and sediment. The model shows that an additional mechanism for the existence of the turbidity maximum is an up-estuary maximum in the tidal current speeds (and thus resuspension). In the lower reaches, transport of suspended sediment is directed down-estuary at both spring and neap tides, and sediment is essentially flushed to sea with the fresh water.