Microphytobenthos: The ecological role of the “secret garden” of unvegetated,shallow-water marine habitats. II. role in sediment stability and shallow-water food webs

The microphytobenthos form an important component of all shallow-water ecosystems where enough light reaches the sediment surface to support appreciable primary production. Although less conspicuous than macroalgae or vascular plants, the microphytobenthos can contribute significantly to primary production and can modify habitat characteristics. The microphytobenthos alter sediment properties (e.g., erodibility) both directly, in the extreme forming a mat or scum on the sediment surface, and indirectly by modifying the activities of benthic infauna (e.g., pelletization, burrowing, tube building, and sediment tracking). Carbon dioxide fixed by the microphytobenthos supports higher, grazing trophic levels. These include deposit-feeding and suspension-feeding macrofauna as well as meiofauna and microfauna. Quantitative relations between the feeding and growth rates of macrofauna and the abundance of microphytobenthos and suspended organic matter (i.e., functional responses) are reviewed. Given the current state of knowledge of the direct and indirect interactions involving trophic dynamics, sediment properties, and benthic microalgae, we argue for reductionist studies of particular interactions as distinct entities. This is a prerequisite for the emergence of a comprehensive picture of unvegetated ecosystems and the ability to predict their responses to man’s activities. *** DIRECT SUPPORT *** A01BY074 00005     

Ciliate epibiont effects on feeding,energy reserves,and sensitivity to hydrocarbon contaminants in an estuarine harpacticoid copepod

Although epibiotic protozoans are commonly observed on the chitinous exoskeleton of aquatic crustaceans, relatively little is known about their ecological significance. The significance of protozoan epibionts on benthic copepods has never been examined.Coullana sp., a meiobenthic harpacticoid copepod, is abundant in Louisiana salt marshes and has high incidence (∼50%) of ciliate epibionts. Field and laboratory grazing experiments indicated that ciliate epibionts did not hinderCoullana feeding on benthic or planktonic algae. Contrary to expectations,Coullana with high levels of ciliate epibionts (>8 ind⁻¹) grazed at a significantly higher rate on¹⁴C-labeled benthic diatoms added to intact sediment cores than didCoullana with no epibionts.Coullana neutral lipids (examined using Nile Red, a hydrophobic fluorophore) were not significantly influenced by the presence of ciliate epibionts, suggesting that copepods are able to compensate for any additional energetic demands imposed by epibionts. Epibiont effects onCoullana susceptibility to hydrocarbon contaminants were measured by examining survivorship in diesel-spiked sediments. The presence of ciliate epibionts significantly decreased survivorship at relatively low PAH concentrations (12.8 ppm). While ciliate epibionts onCoullana do not dramatically alter total food acquisition or energy storage, they may cause stress, which in turn makesCoullana more susceptible to contaminants and possibly other natural stressors such as food limitation.     

Till mélange at Amsdorf, central Germany: sediment erosion, transport and deposition in a complex, soft-bedded subglacial system

Glacial mélange in the open-cast mine at Amsdorf, central Germany, consists of several square meters of large, sorted sediment blocks embedded in till. The blocks are composed of largely intact to slightly deformed glaciofluvial and glaciolacustrine sand, silt and clay, initially deposited in a proglacial lake (2–3 km up-ice) and subsequently overridden by a glacier. The blocks typically have cuboid to subrounded outlines, are randomly distributed in the till, and the contacts with the surrounding till are distinctly sharp. Underneath the mélange are varved clays which exhibit strong deformations occasionally intervening with entirely undisturbed areas. It is suggested that the blocks were entrained into debris-rich basal-ice by bulk freeze-on when the glacier sole was lowered onto the bottom of an overridden lake. After entrainment the blocks were transported englacially and re-deposited (with far-traveled till matrix) as a melt-out till from stagnant ice. The glacier moved mainly by sliding enhanced by low-permeability varved clays in the substratum. The glacier is believed to have been of a polythermal type. These results show that bulk freeze-on can lead to entrainment of soft sediment blocks at least 20 m² in size, and that these blocks can be englacially transported with little or no deformation for several kilometers and more. The occurrence of deformed and undeformed clays under the till mélange indicates a possible mosaic of coupled and decoupled ice, the latter caused by a thin, transient subglacial water film separating the bed from the glacier.     

Carbonate lithofacies as paleolatitude indicators: problems and limitations

Detailed study of Miocene carbonates in the Mediterranean region and their analogues on modern carbonate shelves (in the Mediterranean Sea, Brazil and other areas) reveals at least three major types of carbonate platform lithofacies in addition to the classic tropical coral reef (chlorozoan) lithofacies: (a) chloralgal lithofacies, similar to the chlorozoan, but without hermatypic corals; (b) rhodalgal lithofacies, characterized by abundant encrusting coralline algae; and (c) molechfor lithofacies, consisting of benthic foraminifers, molluscs, echinoids, bryozoans and barnacles. These carbonate lithofacies present complex distribution patterns seemingly related primarily to latitude and depth that control water temperature, although other factors (e.g., water circulation, river discharge, suspended sediment) controlling water salinity and temperature, nutrient content, light penetration, etc., also play important roles. Chloralgal and rhodalgal lithofacies can be considered two transitional terms between the two end-members: the chlorozoan lithofacies, which characterizes shallow tropical shelves; and the molechfor lithofacies, which characterizes colder and/or deeper areas. Detailed textural and sequential analysis are required for satisfactory interpretation of these lithofacies in ancient rocks.     

The Influence of Bioturbation on Iron and Sulphur Cycling in Marine Sediments: A Model Analysis

The geochemical cycles of iron and sulphur in marine sediments are strongly intertwined and give rise to a complex network of redox and precipitation reactions. Bioturbation refers to all modes of transport of particles and solutes induced by larger organisms, and in the present-day seafloor, bioturbation is one of the most important factors controlling the biogeochemical cycling of iron and sulphur. To better understand how bioturbation controls Fe and S cycling, we developed reactive transport model of a coastal sediment impacted by faunal activity. Subsequently, we performed a model sensitivity analysis, separately investigating the two different transport modes of bioturbation, i.e. bio-mixing (solid particle transport) and bio-irrigation (enhanced solute transport). This analysis reveals that bio-mixing and bio-irrigation have distinct—and largely opposing effects on both the iron and sulphur cycles. Bio-mixing enhances transport between the oxic and suboxic zones, thus promoting the reduction of oxidised species (e.g. iron oxyhydroxides) and the oxidation of reduced species (e.g. iron sulphides). Through the re-oxidation of iron sulphides, bio-mixing strongly enhances the recycling of Fe and S between their reduced and oxidised states. Bio-irrigation on the other hand removes reduced solutes, i.e. ferrous iron and free sulphide, from the sediment pore water. These reduced species are then reoxidised in the overlying water and not recycled within the sediment column, which leads to a decrease in Fe and S recycling. Overall, our results demonstrate that the ecology of the macrofauna (inducing bio-mixing or bio-irrigation, or both) matters when assessing their impact on sediment geochemistry. This finding seems particularly relevant for sedimentary cycling across Cambrian transition, when benthic fauna started colonizing and reworking the seafloor.     

泥沙起动概率的高速摄影测量方法

为了通过实验测量泥沙起动概率,基于封闭槽道内开展的泥沙起动实验,系统论述了基于高速摄影的起动概率测量方法所涉及的起动概率定义、相机采样频率、图像处理方法、起动比例与运动比例的关系等关键问题.根据泥沙在水流紊动猝发作用下起动的物理机制,定义实测起动概率为单个紊动猝发周期内,从一定面积床面上冲刷外移的泥沙比例.根据推移质运动概念模型,提出高速图片拍摄间隔应小于推移质运动的中间时间尺度,以捕获泥沙的每一次起动.在高速图像处理时,通过连续两帧图片之间的灰度差实现运动比例的无偏差提取.进一步分析表明,运动泥沙可处于起动、止动或滑翔状态,运动比例等于起动比例、止动比例和滑翔比例之和,起动比例与运动比例之比等于两张图片之间的时间间隔除以推移质运动的中间时间尺度.基于上述关键技术,将高速摄影测量方法用于泥沙起动概率的测量,结果表明,临界起动水流条件时,床沙处于少量起动现象,对应的临界起动概率与窦国仁所推导的2.28%接近.     

Sediment entrainment under fully developed waves as a function of water depth,boundary layer thickness,bottom slope and roughness

Many sediment entrainment equations for oscillatory waves are based on the linear (Airy) theory for deep water, but at the depth where such waves begin to transport sediments they commonly have trochoidal or cnoidal (non-linear) forms. These changes in the wave profile, together with the fact that it is displaced upward with respect to the still water level (SWL), have a profound influence on the hydrodynamics. A method is presented to determine the thickness of the boundary layer from the wave profile, which can be used to calculate the boundary velocity under the wave crest and trough, respectively, in any water depth. The critical boundary velocity can be determined from a published procedure based on laboratory experiments that takes account of the sediment and water properties as well as the wave period. An adjustment is made for the bottom slope and roughness, so that differential land- or seaward sediment entrainment can be predicted for any defined wave cycle. The results explain why sediments are normally transported landward under fair weather conditions and seaward during storms.     

On the ecology of oligochaetes: Monthly variation of community composition and environmental characteristics in two South Carolina tidal creeks

In the tidal creeks of the southeastern United States, the numerically and ecologically dominant macrobenthic organisms are typically oligochaetes. Due to their relatively small size and difficult taxonomy, little is known about the short-term and seasonal changes in the oligochaetes of tidal creeks. This study presents a report of the spatial and temporal changes of the oligochaete taxa within and between two tidal creeks in southern South Carolina, at monthly intervals over a 13-month period. These changes are framed within the reference of monthly changes in benthic chlorophylla, sediment composition, and porewater ammonia, as well as in the perspective of seasonal changes in the entire tidal creek macrobenthic community. The most abundant oligochaete found in this study was the tubificidMonopylephorus rubroniveus, followed by the naidParanais litoralis and the tubificidsTubificoides heterochaetus andT. brownae. All of the oligochaetes exhibited strong month-to-month and spatial changes, indicative of changes in water quality and sediment habitat characteristics (e.g., low dissolved oxygen, high benthic chlorophylla). There were significant correlations between the abundance of most species and either benthic chlorophylla concentration or the silt-clay fraction of the sediment. Looking at short-term changes in this rapidly changing component of the macrobenthic community provides insight not only into the ecology of the oligochaetes, but also into the changes in the tidal creek ecosystem and their potential effects on other biota.     

Relationships between benthic community condition, water quality, sediment quality, nutrient loads, and land use patterns in Chesapeake Bay

Associations between macrobenthic communities, measures of water column and sediment exposure, and measures of anthropogenic activities throughout the watershed were examined for the Chesapeake Bay, U.S. The condition of the macrobenthic communities was indicated by a multimetric benthic index of biotic integrity (B-IBI) that compares deviation of community metrics from values at reference sites assumed to be minimally altered by anthropogenic sources of stress. Correlation analysis was used to examine associations between sites with poor benthic condition and measures of pollution exposure in the water column and sediment. Low dissolved oxygen events were spatially extensive and strongly correlated with benthic community condition, explaining 42% of the variation in the B-IBI. Sediment contamination was spatially limited to a few specific locations including Baltimore Harbor and the Southern Branch of the Elizabeth River and explained about 10% of the variation in the B-IBI. After removing the effects of low dissolved oxygen events, the residual variation in benthic community condition was weakly correlated with surrogates for eutrophication—water column concentrations of total nitrogen, total phosphorus, and chlorophylla. Associations between benthic conditions and anthropogenic inputs and activities in the watershed were also studied by correlation analysis. Benthic condition was negatively correlated with measures of urbanization (i.e., population density, point source loadings, and total nitrogen loadings) and positively correlated with watershed forestation. Significant correlations were observed with population density and nitrogen loading below the fall line, but not above it, suggesting that near-field activities have a greater effect on benthic condition than activities in the upper watershed. At the tributary level, the frequency of low dissolved oxygen events and levels of sediment contaminants were positively correlated with population density and percent of urban land use. Sediment contaminants were also positively correlated with point source nutrient loadings. Water column total nitrogen concentrations were positively correlated with nonpoint nutrient loadings and agricultural land use while total phosphorus concentrations were not correlated with land use or nutrient loadings. Chlorophylla concentrations were positively correlated with nitrogen and phosphorus concentrations in the water column and with agricultural land use but were not correlated with nutrient loads.     

Foraminifera from the Irish Sea glacigenic deposits at Aberdaron,western Lleyn,North Wales: Palaeoenvironmental implications

In a sequence of glacigenic sediments at Aberdaron, Foraminifera were obtained from samples located specifically in order to differentiate between opposing models of depositional environment. All the diamict samples yielded remarkably uniform assemblages, with similar numbers of benthic specimens and benthic species per unit weight of sediment, similar planktic : benthic ratios, and similar ratios of clearly allochthonous to possibly autochthonous elements. This is precisely as predicted by the terrestrial model of sedimentation, where all of the sediments are interpreted as being derived from the melting of glacier ice rich in marine debris entrained during passage along the Irish Sea Basin. The results lend no support to a glacial marine model, since no faunal responses to increasingly distal sedimentary environments are observed. However, the fauna is dominated by the Foraminifera Elphidium excavatum (Terquem) forma clavata Cushman, which is commonly assumed to indicate glacial marine conditions. The modern distribution of similar assemblages suggests that it is just as likely to represent the cold, reduced salinity conditions that would have prevailed in the northern Irish Sea Basin for much of the Quaternary.     

Use of annular flumes to determine the influence of current velocity and bivalves on material flux at the sediment-water interface

A benthic annular flume for both laboratory and in situ deployment on intertidal mudflats is described. The flume provides a means of quantifying material flux (i.e., biodeposition of suspended particulates, sediment resuspension, nutrients, oxygen, and contaminants) across the sediment-water interface in relation to changes in current velocity and benthic community structure and/or population density of key macrofauna species. Flume experiments have investigated the impact of the infaunal bivalveMacoma balthica and the epifaunal bivalveMytilus edulis on seston and sediment flux at the sediment-water interface. The bioturbatorMacoma was found to increase the sediment resuspension and/or erodability by 4-fold, at densities similar to those recorded at the Skeffling mudflat (Humber estuary) (i.e., >1000 individuals m^?2). There was a significant correlation between sediment resuspension andMacoma density (r=0.99; p<0.001), which supported previous in situ field observations indicating bioturbation byMacoma enhanced sediment erodability. Biodeposition rates (g m^?2 h¹) ofMytilus edulis andCerastoderma edule were quantified and related to changes in population density in a mussel bed (Cleethorpes, Humber estuary). Biodeposition rates were up to 40-times the natural sedimentation rates. At the highest mussel bed densities (i.e., 50–100% cover or >1400 mussels m^?2) the physical presence of this epifaunal bivalve on the sediment surface reduced erosion by 10-fold. The shift from net biodeposition to net erosion occurred at current velocities of 20–25 cm s^?1. These results demonstrate that infaunal and epifaunal bivalves can have a significant impact on seston flux or sediment deposition and on sediment resuspension or erodability in estuaries where there are extensive mudflats.     

Microphytobenthos: The ecological role of the “secret garden” of unvegetated,shallow-water marine habitats. I. Distribution,abundance and primary production

The microphytobenthos consists of unicellular eukaryotic algae and cyanobacteria that grow within the upper several millimeters of illuminated sediments, typically appearing only as a subtle brownish or greenish shading. The surficial layer of the sediment is a zone of intense microbial and geochemical activity and of considerable physical reworking. In many shallow ecosystems, the biomass of benthic microalgae often exceeds that of the phytoplankton in the overlying waters. Direct comparison of the abundance of benthic and suspended microalgae is complicated by the means used to measure biomass and by the vertical and horizontal distribution of the microphytobenthos in the sediment. Where biomass has been estimated as chlorophyll a, there may be negligible to large (40%) error due to interference by degradation products, except where chlorophyll is measured by high-performance liquid chromatography. The vertical distribution of microphytobenthos, aside from mat-forming species, is determined by the opposing effects of their vertical migration, which tends to concentrate them near the surface, and physical mixing by overlying currents, which tends to cause an even vertical distribution through the mixed layer of sediment. Uncertainties in vertical distribution are compounded by frequently patchy horizontal distribution. Under-sampling on small (<1 m) scales can lead to errors in the estimate that are comparable to the ranges of seasonal and geographic variation. These uncertainties are compounded by biases in the techniques used to estimate production by the microphytobenthos. In most environments studied, biomass (as chlorophyll a) and light availability appear to be the principal determinants of benthic primary production. The effect of variable light intensities on integral production can be described by a functional response curve. When normalized to the chlorophyll content of the surficial sediment, the residual variation in the data described by the functional response curve is due to changes in the chlorophyll-specific response to irradiance. Production by the benthos is often a significant fraction of production in the water column and microphytobenthos may contribute directly to water column production when they are resuspended. Thus on both the basis of biomass and biogeochemical reactivity, benthic microalgae play significant roles in system productivity and trophic dynamics, as well as such habitat characteristics as sediment stability. *** DIRECT SUPPORT *** A01BY074 00003     

泥沙异重流与环境物质交换经验式对比

采用异重流层平均水沙耦合数学模型,模拟开闸式和恒定入流式泥沙异重流水槽实验,对比分析异重流与环境之间物质交换经验式的适应性和不确定性。考虑4个水卷吸经验式:e_w59、e_w86、e_w87和e_w01,5个泥沙侵蚀经验式:E_s77、E_s86、E_s87、E_s93和E_s04。数值研究表明:水卷吸对于水槽异重流影响较小,应用综合考虑底床摩擦和剪切不稳定的e_w经验式时模拟结果较好;开闸式异重流对床面侵蚀能力有限。对于恒定入流式异重流,应用E_s87和E_s93侵蚀经验式计算所得淤积厚度与实测值吻合较好,可能是率定时综合考虑了异重流实验数据。     

Long-Term Trends of Water Quality and Biotic Metrics in Chesapeake Bay: 1986 to 2008

We analyzed trends in a 23-year period of water quality and biotic data for Chesapeake Bay. Indicators were used to detect trends of improving and worsening environmental health in 15 regions and 70 segments of the bay and to assess the estuarine ecosystem’s responses to reduced nutrient loading from point (i.e., sewage treatment facilities) and non-point (e.g., agricultural and urban land use) sources. Despite extensive restoration efforts, ecological health-related water quality (chlorophyll-a, dissolved oxygen, Secchi depth) and biotic (phytoplankton and benthic indices) metrics evaluated herein have generally shown little improvement (submerged aquatic vegetation was an exception), and water clarity and chlorophyll-a have considerably worsened since 1986. Nutrient and sediment inputs from higher-than-average annual flows after 1992 combined with those from highly developed Coastal Plain areas and compromised ecosystem resiliency are important factors responsible for worsening chlorophyll-a and Secchi depth trends in mesohaline and polyhaline zones from 1986 to 2008.     

Relationships Among Exceedences of Sediment Guidelines,the Results of Ambient Sediment Toxicity Tests,and Community Metrics in Estuarine Systems

To use bioassessments to help diagnose or identify the specific environmental stressors affecting estuaries, we need a better understanding of the relationships among sediment chemistry guidelines, ambient toxicity tests, and community metrics. However, this relationship is not simple because metrics generally assess the responses at the community level of biological organization whereas sediment guidelines and ambient toxicity tests generally assess or are based on the responses at the organism level. The relationship may be further complicated by the influence of other chemical and physical variables that affect the bioavailability and toxicity of chemical contaminants in the environment. Between 1990 and 1993, the U.S. Environmental Protection Agency (USEPA) conducted an Environmental Monitoring and Assessment Program (EMAP) survey of estuarine sites in the Virginian Province of the eastern United States. The surveys collected data on benthic assemblages, physical and chemical habitat characteristics, and sediment chemistry and toxicity. We characterized these estuarine sites as affected by sediment contamination based on the exceedence of sediment guidelines or on ambient sediment toxicity tests (i.e., 10-day Ampelisca abdita survival). Then, benthic invertebrate metrics were compared among affected and unaffected sites to identify metrics sensitive to the contamination. A number of benthic invertebrate metrics differed between groups of sites segregated using the organism-level measures whereas other metrics did not. The difference among metrics appears to depend on the sensitivity of the individual metrics to the stressor gradient represented by metals or persistent organic toxics in sediments because the insensitive metrics do not effectively quantify the changes in the benthic invertebrate assemblage associated with these stressors. The significant relationships suggest that a relationship exists between the organism-level effects assessed by chemistry or ambient toxicity tests and the community-level effects assessed by community metrics and that the organism-level effects are predictive, to some extent, of community-level effects.     

Early to mid-Holocene Atlantic water influx and deglacial meltwater events, Beaufort Sea slope, Arctic Ocean

Holocene high-resolution cores from the margin of the Arctic Ocean are rare. Core P189AR-P45 collected in 405-m water depth on the Beaufort Sea slope, west of the Mackenzie River delta (70°33.03′N and 141°52.08′W), is in close vertical proximity to the present-day upper limit of modified Atlantic water. The 5.11-m core spans the interval between ∼6800 and 10,400 ¹⁴C yr B.P. (with an 800-yr ocean reservoir correction). The sediment is primarily silty clay with an average grain-size of 9 φ. The chronology is constrained by seven radiocarbon dates. The rate of sediment accumulation averaged 1.35 mm/yr. Stable isotopic data (δ¹⁸O and δ¹³C) were obtained on the polar planktonic foraminifera Neogloboquadrina pachyderma (s) and the benthic infaunal species Cassidulina neoteretis. A distinct low-δ¹⁸O event is captured in both the benthic and planktonic data at ∼10,000 ¹⁴C yr B.P.—probably recording the glacial Lake Agassiz outburst flood associated with the North Atlantic preboreal cold event. The benthic foraminifera are dominated in the earliest Holocene by C. neoteretis, a species associated with modified Atlantic water masses. This species decreases toward the core top with a marked environmental reversal occurring ∼7800 ¹⁴C yr B.P., possibly coincident with the northern hemisphere 8200 cal yr B.P. cold event.     

海洋沉积物中色素生物标志物研究进展

海洋沉积物中的光合色素包含着水体、沉积物中浮游和底栖植物以及微生物群落的丰富信息,能表征特定生物来源,在埋藏到沉积物甚至发生某些改变之后仍然保留其源信息,是一类重要的化学生物标志物.结合总有机碳、总氮等其他海洋地球化学参数,沉积色素可用来研究海洋浮游植物和光合细菌的群落组成和丰度,反演海洋初级生产、水体富营养化水平及其历史趋势,指示水体和沉积物氧化还原条件,揭示海域气候条件等现状及其历史变化.沉积色素的研究,对于掌握海洋中碳的生物地球化学循环过程,回溯古环境、古海洋、古生态以及古气候记录,制定合理的海洋管理政策具有十分重要的意义.阐述了沉积物中色素的分类、来源、性质和分析方法,分析了色素在沉积物中的保存和变化规律,探讨总结了沉积色素作为化学生物标志物在海洋学研究中的应用.     

Sediment grain size effect on benthic microalgal biomass in shallow aquatic ecosystems

Benthic microalgal biomass is an important fraction of the primary producer community in shallow water ecosystems, and the factors controlling benthic microalgal biomass are complex. One possible controlling factor is sediment grain-size distribution. Benthic microalgal biomass was sampled in sediments collected from two sets of North Carolina estuaries Massachusetts and Cape Cod bays, and Manukau Harbour in New Zealand. Comparisons of benthic microalgal biomass and sediment grain-size distributions in these coastal and estuarine ecosystems frequently showed a negative relationship between the proportion of fine-grained sediments and benthic microalgal biomass measured as chlorophylla. The highest sedimentary chlorophylla levels generally occurred in sediments with lower percentages of fine particles (diameter <125 mm). A negative relationship between the proportion of fine sediments and benthic microalgal biomass suggests anthropogenic loadings of fine sediment may reduce the biological productivity of shallow-water ecosystems.     

Small-scale spatial variation of macrobenthic community structure

Examination of small-scale spatial variation in essential to understanding the relationships between environmental factors and benthic community structure in estuaries. A sampling experiment was performed in October 1993 to measure infauna association with sediment composition and salinity gradients in Nueces Bay, Texas, USA. The bay was partitioned into four salinity zones and three sediment types. Higher densities of macrofaua, were found in sediments with greater sand content and in areas with higher salinity. High diversity was also associated with high homogeneous salinity (31–33‰) and greater sand content. Macrofauna biomass and diversity were positively correlated with bottom salinity, porewater salinity, and bottom dissolved inorganic nitrogen (DIN). Furthermore, species dominance shifted along the estuarine gradient.Streblospio benedicti dominated at lower salinity, but,Mediomatsus ambiseta andMulinia lateralis were the dominant species at higher salinity. Statistical analyses revealed significant correlations for sediment characteristics (i.e., increased fine sediments, water content, and total organic carbon) with decreased total abundance and diversity. Increased salinity and DIN were correlated with increased total biomass, diversity, and macrofauma community structure. These physico-chemical variables are regulated by freshwater inflow, so inflow is an important factor influencing macrofauna community structure by indirectly influencing the physico-chemical environment.     

The environments of production and deposition of calcareous sediments on the shelf west of Scotland

The dominant calcareous organisms and sediment characteristics are described for eight different physical settings on the shelf west of Scotland, each having a different depth, substrate and degree of hydrodynamic exposure.

The principal sites of carbonate production are on shallow rocky substrates where barnacles, molluscs, echinoderms and serpulids are the dominant calcareous organisms. In sheltered shallow sandy zones, molluscs, echinoderms and benthic foraminiferans are the active producers, though the sediments are commonly barnacle-rich. Where tidal currents are enhanced between islands and the waves are suppressed, calcareous red algae (Phymatolithon calcareum) and mussel shells build localised banks. In deep, open-shelf water molluscs are the major skeletal contributor to the sediment, though on rocky sea beds bryozoans, serpulids and echinoderms are important.

The major sites of deposition are where persistent hydro- (and aero-) dynamic conditions sweep together grains from active production sites (e.g., sand ribbons or beaches and dunes adjacent to shallow rocky platforms) or in sinks where the physiographic configuration favours the deposition and retention of locally produced sediment or sediment derived from suspension. The well-sorted, cross-bedded, beach and dune sands commonly contain > 75% CaCO₃. In sheltered depressions, bioturbated muds accumulate with up to 30% calcite silt, which is probably the breakdown product of barnacles and benthic foraminiferans.