Allocation of nitrogen and carbon in an estuarine salt marsh in Portugal

Above and below-ground biomass and nitrogen and carbon composition ofSpartina maritima, Halimione portulacoides andArthrocnemum perenne, dominating species in plant communities of the lower, middle and higher salt marsh, respectively, were compared in an estuarine salt marsh in Portugal. Plant and soil nitrogen and carbon pools were estimated. For all three species root biomass was significantly higher (70–92% of total biomass) than above-ground biomass. The percentage of root biomass was related to the location of the plants in the marsh: higher values were found in plants growing in the lower salt marsh where the sediment was more unstable and subject to tidal action, which stresses the role of the roots as an anchor. For all three species nitrogen concentrations were highest in leaves, reflecting the photosynthetic role of the tissue. For carbon higher concentrations were found in the stems, with the exception ofS. maritima. In general, lower nitrogen concentrations were found in summer, which can be explained by dilution processes due to plant growth. For both nitrogen and carbon, higher concentrations were found in the soil surface layers. Higher soil nitrogen and carbon levels were associated with higher organic matter contents. Most of the nitrogen in the salt marsh occurred in the sediments (0–40 cm) and only ca. 5.7–13.3% of the total was found in the plants. The greater portion (76.5%–86%) of carbon was found in the sediment.     

Pigment composition and size distribution of phytoplankton in a confined Mediterranean salt marsh ecosystem

Pigment composition and size distribution of phytoplankton were analysed in a group of Mediterranean salt marshes, where hydrology is dominated by sudden inputs during sea storms, followed by long periods of confinement. These marshes are characterized by a low inorganic–organic nutrient ratio, and inorganic nitrogen is especially scarce due to denitrification. Nutrients were the main factor affecting phytoplankton biomass, while zooplankton grazing did not control either phytoplankton community composition, or their size distribution. The relative abundance of the different phytoplankton groups was analysed by correspondence analysis using the pigment composition measured by high-performance liquid chromatography (HPLC) and analysed with the CHEMTAX programme. In this analysis, phytoplankton pigment composition was correlated with two nutrient gradients. The first gradient was the ratio of nitrate–total nitrogen (TN), since the different phytoplankton groups were distributed according to their eco–physiological differences in nitrogen uptake. The second gradient was correlated with total nutrient loading. Biomass size distributions frequently showed a lack of intermediate sized nanophytoplankton (2.5–4 μm in diameter), and the importance of this lack of intermediate sizes correlated with dinoflagellate biomass. These results suggested that in confined environments, where nutrients are mainly in an organic form, dinoflagellates take advantage of their mixotrophy, by competing and grazing on smaller phytoplankters simultaneously.     

Nitrogen exchange between a southeastern USA salt marsh ecosystem and the coastal ocean

The salt marsh ecosystem at North Inlet, South Carolina, USA consistently exported dissolved inorganic nitrogen via tidal exchange with the coastal Atlantic Ocean. Concentrations centrations of NH ₄ ⁺ and NO ₃ ^- +NO ₂ ^- displayed distinct tidal patterns with rising values during ebb flow. These patterns suggest the importance of biogeochemical processes in the flux of material from the salt marsh. NH ₄ ⁺ export peaked during the summer (15 to 20 mg m^-2 tide^-1) during a net balance of tidal water exchange. Remineralization of NH ₄ ⁺ within the salt marsh system appears to be contributing to the estimated annual net export of bout 4.7 g NH ₄ ⁺ -N m^-2 yr^-1. NO ₃ ^- +NO ₂ ^- exports were higher in the fall and winter of 1979 (2 to 4 mg N m^-2 tide^-1). The winter export coincided with a considerable net export of water with no distinctive concentration patterns, suggesting a simple advective export. However, the fall peak of NO ₃ ^- +NO ₂ ^- export occurred during a period of net water balance in tidal exchange and an insignificant freshwater input from the western, forested boundary. During the summer and fall, tidal concentration patterns were particularly apparent, suggesting that nitrification within the salt marsh system was contributing to the estimated annual net export of ca 0.6 g NO ₃ ^- +NO ₂ ^- -N m^-2 yr^-1.Contribution No. 637 from the Belle W. Baruch Institute of Marine Biology and Coastal Research     

Diel relationships of microbial trophic groups and in situ dissolved carbohydrate dynamics in the caribbean sea

Dissolved total carbohydrate (TCHO), polysaccharide (PCHO), monosaccharide (MCHO) and organic carbon (DOC) were determined at 3-h intervals over 5 diel cycles in the mixed layer of the northwestern Caribbean Sea while following a drogued buoy. These data have been compared to populations of phototrophic (PNAN) and heterotrophic (HNAN) nanoplankton (2–20 m diameter) and heterotrophic bacteria (HBAC) (0.2–2.0 m diameter) estimated by epifluorescence counts, as well as to CO₂, phosphate, chlorophyll a and phaeopigment data determined simultaneously. Two different types of apparent diel dissolved carbohydrate (CHO) patterns were found. On 3 d when no sustained net CO₂ uptake was evident, TCHO and PCHO generally declined during the afternoon and early evening while MCHO tended to increase. On two other days when apparent sustained CO₂ uptake occurred during the day, there were large evening TCHO and PCHO peaks with constant or declining MCHO levels. These accumulations probably resulted from the release of recently produced PCHO from phototrophs. As was found earlier in the Sargasso Sea, PNAN populations were inversely related to PCHO concentrations. The sample to sample fluctuations of PNAN also were inversely related to the apparent rates of change of TCHO and PCHO, possibly due to an inverse relation between the rates of PNAN cell division and CHO excretion. Fluctuations in HBAC populations were inversely correlated with PCHO dynamics and directly related to MCHO variations, possibly due to extracellular hydrolysis of PCHO to MCHO during periods of rapid bacterial growth as well as to net heterotrophic PCHO uptake. A direct relationship between HNAN and TCHO fluctuations suggests the importance of HNAN excretion in the release of dissolved organics. The combined PNAN and HBAC fluctuations accounted for a more significant fraction of the variance in the apparent rates of change of PCHO than did any single population parameter indicating that intimate interactions between the microbial plankton groups are important in the in-situ regulation of CHO dynamics. Total system net TCHO release and uptake rates for 5 d averaged 56 and 53 g C l^-1 d^-1 respectively, assuming that the observed fluctuations resulted from temporal planktonic processes in homogeneous water masses. While the data contain indications that this was the case, this assumption is not definitive.     

Root respiration and oxygen flux in salt marsh grasses from different elevational zones

Plants growing in waterlogged environments are subjected to low oxygen levels around submerged tissues. While internal oxygen transport has been postulated as an important factor governing flooding tolerance, respiration rates and abilities to take up oxygen under hypoxic conditions have been largely ignored in plant studies. In this study, physiological characteristics related to internal oxygen transport, respiration, and oxygen affinity were studied in low intertidal marsh species (Spartina alterniflora and S. anglica) and middle to high intertidal species (S. densiflora, S. patens, S. foliosa, a S. alterniflora × S. foliosa hybrid, S. spartinae, and Distichlis spicata). These marsh plants were compared to the inland species S. pectinata and the crop species rice (Oryza sativa), corn (Zea mays), and oat (Avena sativa). Plants were grown in a greenhouse under simulated estuarine conditions. The low marsh species S. anglica was found to transport oxygen internally at rates up to 2.2 μmol O₂ g fresh root weight⁻¹ h⁻¹. In contrast, marsh species from higher zones and crop species were found to transport significantly less oxygen internally, although rice plants were able to transport 1.4 μmol g⁻¹ h⁻¹. Under hypoxic conditions, low marsh species were better able to remove dissolved oxygen from the medium compared to higher marsh species and crops. The oxygen concentration at which respiration rates declined due to limited oxygen (P _crit) was significantly lower in low marsh species compared to inland and crop species; P _crit ranged from <4 μM O₂ in the low marsh species S. anglica up to 20 μM in the inland species corn. Flooding-sensitive crop species had significantly higher aerobic respiration rates compared to flooding-tolerant species in this study. Crop species took up 3.6–6.7 μmol O₂ g⁻¹ h⁻¹ while all but one marsh species took up <3.5 μmol O₂ g⁻¹ h⁻¹. We conclude that oxygen transport, aerobic demand, and oxygen affinity all play important and interrelated roles in flood tolerance and salt marsh zonation.     

Flax Pond ecosystem study: Exchanges of phosphorus between a salt marsh and the coastal waters of Long Island Sound

The exchanges of phosphorus between the Flax Pond, a tidal Spartina alterniflora marsh on the north shore of Long Island (USA) and Long Island Sound were measured over 18 months. Phosphate was exported from the marsh from May through December and imported during the remainder of the year. Organic phosphorus appears to be accumulated in all seasons, but the yearly phosphorus budget of the marsh is approximately balanced despite the accumulation of about 6 mm of sediment annually.     

滨海盐沼及其植物群落的分布与多样性

滨海盐沼是广泛存在于世界中、高纬度地区的一种湿地生态系统,具有抵御风暴潮灾害、净化污染物和为珍稀濒危生物提供适宜生境等重要的生态和经济价值。滨海盐沼因随高程变化而急剧变化的环境梯度和植物带状分布现象而为生态学者阐释自然界物种的分布机制提供了理想系统。主要概述了滨海盐沼的定义、特点、类型、全球分布以及潮汐作用、土壤盐度等环境因子特征;阐述了不同尺度下滨海盐沼的植物群落分布和多样性特征。在滨海盐沼植物群落的分布特征上,重点阐述了中尺度下的植物带状分布,即植物群落往往在白海向陆渐高的不同高程梯度上表现出显著的分带分布,不同植物各自占据该梯度上的一定区域。通常认为,带状分布是植物竞争和物理性胁迫共同调控的结果,但其在不同地理区域的普适性仍存争议。滨海植物群落多样性往往较低,在中、小尺度上盐沼植物多样性受控于盐度、潮汐等物理性胁迫、植物间相互作用等因子的作用;在大尺度上盐沼植物多样性可能随纬度增大而增加。系统深入地认识滨海盐沼植物群落生态格局和过程,将为气候变化、生物入侵等人类影响下的滨海盐沼生态系统的管理和恢复提供有益经验。     

Nutrient fluxes within a small north temperate salt marsh

The water exchange between a small (4.1 hectare) salt marsh adjoining the Great Bay Estuary system of New Hampshire, USA was sampled during 16 tidal cycles between July, 1976 and November, 1977. Tidal amplitude, temperature, salinity, nutrient concentrations (ammonia-N, nitrate-N, nitrite-N, orthophosphate-P, total-P, silicates) and suspended particulates were measured. Conspicuous tidal hydrographic patterns were observed. Mean concentrations of nitrate-N and silicates varied with season. The tidal information, combined with volume determinations, was extrapolated to determine the net flux of hydrographic parameters on monthly and yearly bases. Ammonia-N showed a pronounced seasonality of net exchange by regression analyses. Ammonia-N and suspended particulates had a statistically significant import into the marsh. However, the net fluxes of the other materials were not statistically different from O.Published with the approval of the Director of the New Hampshire Agricultural Experiment Station as Scientific Contribution No. 930Scientific Contribution No. 69 of the Jackson Estuarine Laboratory     

Primary production and heterotrophy of a pennate and a centric salt marsh diatom

Radionuclide labeled sodium bicarbonate and glucose were used to measure the rates of primary production and glucose uptake by Nitzschia obtusa var. undulata and Cymbella pusilla. The presence of glucose in the medium enhanced primary production in both diatom species. Uptake of glucose in C. pusilla was photoinhibited. A maximum of 9.3 nmol C mg dry wt^-1 h^-1 of glucose was taken up in the light without the presence of NaHCO₃ in the medium and with NaHCO₃ present the value was 18 nmol C mg dry wt^-1 h^-1. The ratio of inorganic carbon fixed to the total carbon budget (autotrophy and heterotrophy) was 0.51 for N. obtusa var. undulata and 0.62 for C. pusilla.     

Latitudinal variation in top-down and bottom-up control of a salt marsh food web

The shrub Iva frutescens, which occupies the terrestrial border of U.S. Atlantic Coast salt marshes, supports a food web that varies strongly across latitude. We tested whether latitudinal variation in plant quality (higher at high latitudes), consumption by omnivores (a crab, present only at low latitudes), consumption by mesopredators (ladybugs, present at all latitudes), or the life history stage of an herbivorous beetle could explain continental-scale field patterns of herbivore density. In a mesocosm experiment, crabs exerted strong top-down control on herbivorous beetles, ladybugs exerted strong top-down control on aphids, and both predators benefited plants through trophic cascades. Latitude of plant origin had no effect on consumers. Herbivorous beetle density was greater if mesocosms were stocked with beetle adults rather than larvae, and aphid densities were reduced in the "adult beetle" treatment. Treatment combinations representing high and low latitudes produced patterns of herbivore density similar to those in the field. We conclude that latitudinal variation in plant quality is less important than latitudinal variation in top consumers and competition in mediating food web structure. Climate may also play a strong role in structuring high-latitude salt marshes by limiting the number of herbivore generations per growing season and causing high overwintering mortality.     

Diel variation in chlorophyll a,carbohydrate and protein content of the marine diatom Skeletonema costatum

Skeletonema costatum (Greville) Cleve isolated from Narragansett Bay, USA, was incubated at 3 light intensities (ca. 0.008, 0.040 and 0.075 ly min^-1) under a 12 h light: 12 h dark (12L:12D) photoperiod at 2°, 10° and 20°C. Cellular chlorophyll a increased at intensities less than ca. 0.040 ly min^-1; increases occured within one photoperiod at temperatures above 10°C. Cellular carbohydrate increased with light intensity at all temperatures; increases during the photophase were due to net production of the dilute acid-soluble fraction. Cellular protein increased during the photoperiod at 10° and 20°C; there was little difference in cellular protein among all cultures after one photoperiod. The rate at which cellular chlorophyll a increased in response to a decrease in light suggests that diel variation in cellular chlorophyll a is temperature-dependent in S. costatum. Protein: carbohydrate ratios ranged from ca. 0.5 to 2.0 over a diel cycle; ratios increased at lower intensities and higher temperatures. The diel range in protein:carbohydrate ratios equals that in cultures developing nitrogen deficiency; thus, use of this ratio as an index to phytoplankton physiological state must account for diel light effects.     

Consequences of omnivory for trophic interactions on a salt marsh shrub

Although omnivory is common in nature, its impact on trophic interactions is variable. Predicting the food web consequences of omnivory is complicated because omnivores can simultaneously produce conflicting direct and indirect effects on the same species or trophic level. We conducted field and laboratory experiments testing the top-down impacts of an omnivorous salt marsh crab, Armases cinereum, on the shrub Iva frutescens and its herbivorous and predatory arthropod fauna. Armases is a "true omnivore," consuming both Iva and arthropods living on Iva. We hypothesized that Armases would benefit Iva through a top-down trophic cascade, and that this benefit would be stronger than the direct negative effect of Armases on Iva. A field experiment on Sapelo Island, Georgia (USA), supported this hypothesis. Although Armases suppressed predators (spiders), it also suppressed herbivores (aphids), and benefited Iva, increasing leaf number, and reducing the proportion of dead shoots. A one-month laboratory experiment, focusing on the most common species in the food web, also supported this hypothesis. Armases strongly suppressed aphids and consumed fewer Iva leaves if aphids were available as an alternate diet. Armases gained more body mass if they could feed on aphids as well as on Iva. Although Armases had a negative effect on Iva when aphids were not present, Armases benefited Iva if aphids were present, because Armases controlled aphid populations, releasing Iva from herbivory. Although Armases is an omnivore, it produced strong top-down forces and a trophic cascade because it fed preferentially on herbivores rather than plants when both were available. At the same time, the ability of Armases to subsist on a plant diet allows it to persist in the food web when animal food is not available. Because omnivores feed on multiple trophic levels, their effects on food webs may differ from those predicted by standard trophic models that assume that each species feeds only on a single trophic level. To better understand the complexity of real food webs, the variable feeding habits and feeding preferences of different omnivorous species must be taken into consideration.     

Behaviour of horses and cattle at two stocking densities in a coastal salt marsh

Livestock grazing has been practiced in salt marshes in the Wadden Sea area since 600 B.C. Currently livestock grazing is also applied for conservation management. However, effects of such grazing management on salt marshes are likely to vary depending on the species of livestock and stocking density due to differences in the behaviour of the animals. Yet, little is known about the behaviour of different livestock species and stocking densities grazing in salt marshes. We studied the grazing behaviour of horses and cattle by focal observation in an experiment with four different grazing treatments on a coastal salt marsh. In all treatments we recorded diet choice, movement and grazing activity, and spatial distribution. Livestock species shared an overlap in diet choice. Yet, horses more often foraged on the short grass Puccinellia maritima, while the cattle diet contained a higher amount of Aster tripolium. Horses travelled longer distances per day and spent more time grazing than cattle. Spatial distribution of cattle was significantly clustered, while horses showed a random distribution utilizing the whole area. Animal behaviour differs between livestock species and stocking densities with respect to diet choice, activity and spatial distribution.     

Macroinfauna of a Southern New England salt marsh: seasonal dynamics and production

The animal-habitat relationships and seasonal dynamics of the benthic macroinfauna were investigated from November 1986 to October 1988 in the Great Sippe-wissett salt marsh (Massachusetts, USA). Total macrofaunal abundance varied seasonally, displaying a peak in late spring and early summer, then declining sharply during late summer and recovering briefly in fall before collapsing in winter. Three macroinfaunal assemblages were found in the marsh, distributed along gradients of environmental factors. These included a sandy non-organic sediment assemblage, a sandy organic sediment assemblage and a muddy sediment assemblage. The species groups characteristic of unstable sandy non-organic sediments included the polychaetes Leitoscoloplos fragilis, Aricidea jefreyssi, Magelona rosea and Streptosyllis verrilli, the oligochaete Paranais litoralis, and the crustacean Acanthohaustorius millsi. Sandy organic sediments were characterized by the polychaetes Marenzelleria viridis, Capitella capitata, Neanthes succinea, N. arenaceodonta, Polydora ligni and Heteromastus filiformis, the oligochaete Lumbricillus sp., and the mollusc Gemma gemma. In muddy sites, the polychaete Streblospio benedicti and the oligochaetes Paranais litoralis and Monopylephorus evertus were the dominant species. Secondary production of benthic macroinfauna in each of these habitats was estimated. The highest values of biomass and production were recorded in the sandy organic sediments. Secondary production was estimated to be 1850 kJ m^-2 yr^-1 in sandy organic areas, but only 281 kJ m^-2 yr^-1 in sandy non-organic areas and 113 kJ m^-2 yr^-1 in muddy areas. This results in an area-weighted average production of 505 kJ m^-2 yr^-1 for the unvegetated areas of the marsh. The Great Sippewissett salt marsh has an area of 483800 m², the total secondary production of the macroinfauna for the whole unvegetated area of the marsh was estimated as 4651 kg dry wt yr^-1, expressed as somatic growth. This production value seems consistent with production data obtained for other intertidal North Atlantic environments.     

Secondary succession on a high salt marsh at different grazing intensities

Succession occurs on a large part of German salt marshes following abandonment or reduction of grazing. Its speed and effect on the biodiversity of salt marshes has been discussed in the literature. Permanent plot studies show site-dependent differences in successional outcome. If grazing is to be continued, there is uncertainty about the stocking rates that are optimal for the conservation of plant diversity. We present an eight year permanent plot study with different grazing treatments on a high salt marsh at the marsh island of Hallig Langeness, Germany, which is enclosed by a summer dike. The study site is owned by WWF. Successional outcomes on the permanent plots depend on grazing intensity and depth of the groundwater table which is correlated with soil salinity. Intermediate stocking rates support a mixture of halophytes and glycophytes, the relative proportion of which depends on the depth of the groundwater table. Cattle grazing at stocking rates of 0.6 livestock units per ha were found to be optimal for plant diversity conservation. At sites with deeper groundwater tables and no grazing, dominantElymus spp. stands develop and diversity is strongly reduced. If the groundwater table is high, succession following grazing abandonment is retarded and small halophytes prevail.     

Phytoplankton in a temperate-zone salt marsh: Net production and exchanges with coastal waters

Phytoplankton production and associated variables were measured in Flax Pond, a 52 ha salt marsh on the north shore of Long Island, New York, from July 1972 to October 1973. Measurements made up to five times per day, once per week, yielded a mean annual net primary production, determined by the ¹⁴C technique, of 20.5 mg C/m³/h; daily means were as high as 60.0 mg C/m³/h. However, when productivity was calculated for the entire marsh ecosystem, the shallow water in the salt marsh produced only 11.7 g C/m² of marsh/year. There was a net flux of phytoplankton from the coastal waters into the marsh; during the summer up to 0.2 g chlorophy 11/m² of marsh was carried in with the tides daily and remained in the marsh. Analysis of the productivity data, as well as variables associated with productivity (pH, standing crop, nutrients, extinction coefficient), indicated that the aquatic portion of the marsh behaved more as a net consumer rather than a net producer of phytoplankton.Research carried out at Brookhaven National Laboratory, supported by the U.S. Atomic Energy Commission and the National Science Foundation under Grant No. AG-375.     

Hydrodynamic and chemical conditions during onset of a red-tide assemblage in an estuarine upwelling ecosystem

The hydrodynamics and nitrogen/silicon biogeochemistry accompanying the development of a red-tide assemblage were examined in the Ría de Vigo (northwest Spain), a coastal embayment affected by upwelling, during an in situ diel experiment in September 1991. Despite a low N:Si molar ratio (0.5) of nutrients entering the surface layer, which was favourable for diatom growth, the diatom population began to decline. Limited N-nutrient input, arising from moderate coastal upwelling in a stratified water column, restricted net community production (NCP = 630 mg C m⁻² d⁻¹). In addition, light-limitation of gross primary production (GPP = 1525 mg C m⁻² d⁻¹) was observed. The relatively high f-ratio (= NCP:GPP) recorded (0.41, characteristic of intense upwelling conditions) would have been as low as 0.15 had not GPP been limited by light intensity. Temporal separation of carbohydrate synthesis during the photoperiod from protein synthesis in the dark could be inferred from the time-course of the C:N ratio of particulate organic matter. Severe light-limitation would lead to diatom collapse were the diatoms not able to meet all their energy requirements during the hours of darkness. Under the hydrodynamic, nutrient and light conditions of the experiment, an assemblage of red-tide-forming species began to develop, aided by their ability to migrate vertically and to synthesize carbohydrates during the light in surface waters and protein during the dark at the 4 m-deep pycnocline. Thermal stratification, reduced turbulence, intense nutrient mineralization, and the limited nitrogen input through moderate upwelling were all favourable to the onset of a red-tide assemblage. Received: 15 February 1997 / Accepted: 26 September 1997     

城市生态系统-大气间的碳通量研究进展

城市生态系统对全球碳收支具有显著的贡献,城市化进程促进城市向大气排放碳。随着观测手段和仪器的发展,涡动相关法已成为陆地生态系统碳通量观测的主要手段,并广泛应用于中心城市碳通量观测;城市CO2浓度和通量变化具有日、周、季节变化特征,不同城市之间CO2浓度和通量也有很大差异,其主要原因是由于城市生态系统的下垫面类型不均一,土壤、植被、路面、交通、人呼吸等对碳通量都有贡献,绿地（树木、草坪）对燃料燃烧、人呼吸等的碳排放起着一定的抵消作用;由于多种因素对碳通量起作用,在对城市通量数据进行分析时必须开展贡献区即碳足迹分析。本文综述了城市生态系统-大气间碳通量研究的历程;分析了城市生态系统碳通量的时空变化特征及其主要控制因子;探讨了复杂下垫面条件下城市生态系统-大气间碳通量研究的特点和难点,并提出了未来研究的重点。     

Diel and tidal cycles regulate larval dynamics in salt marshes and mangrove forests

Variation in the release and recruitment of larvae of estuarine invertebrates affects the distribution and abundance of adults, as well as trophic interactions in both the plankton and the benthos. Larval release and supply are often timed to environmental cycles such as the diel and tidal cycles. Here, we determined using plankton tows whether the abundance of larvae spanning salt marsh and mangrove habitats across the intertidal landscape varied with diel and tidal cycles. Using three different sampling designs across two sites and within each of two estuaries over a 12-month period, we covered a range of spatial and temporal scales. This allowed us to test the general prediction that densities of meroplankton in the water column would be greater during nocturnal ebb tides than during other phases of the diel or tidal cycle. As predicted, nocturnal ebb tides yielded the highest densities of meroplanktonic larvae and were dominated by first-stage crab zoeae and this finding was most pronounced in the salt marsh. Throughout the course of the year, greater numbers of meroplankters consistently occurred during the ebb tide compared with the flood tide. The densities of other taxa (e.g. gastropods and polychaetes) showed no clear trends with diel or tidal cycles. This study highlights the effects of these pervasive physical cycles on the timing of larval release and supply in the salt marsh–mangrove complex, and emphasises their contribution to the trophic interactions and the dynamics of benthic populations within estuaries.