Food sources of benthic animals on intertidal and subtidal bottoms in inner Ariake Sound,southern Japan,determined by stable isotopes

To evaluate the relative importance of possible food sources, including riverine particulate organic matter, reeds, benthic microalgae, seaweeds, cultured laver (Porphyra) and coastal phytoplankton, for commercial bivalves and co-occurring benthic animals, 73 macrofaunal species were collected from intertidal and subtidal soft bottoms in the inner part of Ariake Sound, Kyushu, southern Japan, and their isotopic compositions were analyzed. The results revealed that (1) both intertidal and subtidal food webs were constituted of 3 trophic levels, (2) suspension-feeding bivalves utilize a mixture of benthic microalgae and coastal phytoplankton, and omnivores and carnivores incorporate benthic microalgae and phytoplankton through their intermediate prey, and (3) 3 bivalves (Scapharca kagoshimensis, Modiolus metcalfei and Atrina lischkeana) inhabiting both intertidal and subtidal bottoms showed similar seasonal fluctuations, suggesting no difference in the diet composition among the species and between the 2 habitats. We conclude that a large biomass of benthic microalgae which was approximately equal to that of phytoplankton and the strong tidal currents that would resuspend benthic microalgae and transport them to subtidal bottom areas account for the benthic microalgal and phytoplankton based trophic structure in the inner part of Ariake Sound.     

Hydrology of Okarito Lagoon and the inferred effects of selective logging in Okarito Forest

Okarito Lagoon (43° 11′S, 170° 14′E) is a small (20 km²) shore‐parallel, predominantly subtidal estuary, deepest near the landward end, and linked to the sea by two subtidal channels incised through shallow subtidal and intertidal flats which occupy the southern third of the lagoon. Tides at sea vary from 2.1m (spring) to 1.2 m (neap), but in the lagoon the tidal range is constant through the lunar cycle and varies from 0.80 m at the entrance to 0.17 m in the upper lagoon. Tidal water level and flow asymmetries in the subtidal channels are separated by a 1.7 h phase difference. Variations in the net discharge through the inlet result from changing flow cross‐sections rather than from variations in current velocities. Both the tidal‐averaged volume and the tidal compartment of the lagoon vary through the lunar cycle, from maxima at spring tides to minima at neap tides.

Freshwater inflows vary from less than 11 m³.s^‐1 to more than 750 m³.s^‐1. During storms, water level in the lagoon rises rapidly by 2–3 m, then declines to normal over several days. Three water masses, two with salinity and turbidity largely controlled by antecedent rainfall, normally occur in the lagoon. Suspended sediment concentrations in both freshwater inflows and lagoon waters are normally low but increase during floods. Most sediment is supplied by the Waitangi‐taona River or by erosion of tidal channel margins. The lagoon is floored with organic‐rich mud and sandy mud, deposited predominantly from suspension. Surface sediment is consistently muddier than subsurface sediment, probably reflecting an increase in the mud supply since diversion of the Waitangi‐taona River in 1967.

Comparisons of the estimated sediment yield and water inflow effects of the 1967 river diversion with short‐term observations during selective logging suggest that the effects of logging on sediment yield, water balance of the lagoon, and dissolved solids inputs will be small compared with changes caused by diversion of the Waitangi‐taona River.     

CHARACTERISTICS OF TIDAL FLATS OF THE ZHUJIANG RIVER MOUTH

The intertidal zone of the Zhujiang River mouth tidal flat is about 570,000 mu in total, and the tidal flat of the subtidal zone (shallower than -2 m of the Huanghai Sea base level) is 560,000 mu. The two types, silty mud and argillaceous silt, are predominant in the tidal flat area, which can be divided into five types in terms of origin: 1) river mouth sand spit-subaqueous natural levee type; 2) river mouth bar type; 3) bay head type; 4) around the island type; and 5) marginal flat type. The tidal flat of the Zhujiang River mouth belongs to the depositional type and is constantly accumulating silt while extending, with an average land formation of 8,940 mu/yr in 97 years (1883-1980).     

渤海湾南部滩涂贝类群落结构特征及其环境因子相关性研究

本研究以渤海湾南部(滨州)滩涂为研究区域,设置6个断面共36个采样站位,覆盖潮间带(高潮带-中潮带-低潮带)和潮下带,研究该区域贝类资源群落结构特征及其环境因子相关性。结果表明:整个调查区域常见贝类总共有30种。潮间带共有23种常见贝类,优势种为光滑篮蛤(Potamocorbula laevis Hinds 1843)、四角蛤蜊(Mactra quadrangularis Reeve 1854)、短文蛤(Meretrix petechialis Lamarck 1799);潮下带共有26种常见贝类,优势种为中国蛤蜊(Mactra chinensisi Philippi 1846)、等边浅蛤(Gomphina aequilatera Sowerby 1825)、朝鲜笋螺(Terebra koreana Yoo 1976)、红带织纹螺(Nassarius succinctus A.Adams 1852)、纵肋织纹螺(Nassarius variciferus A.Adams 1851);潮间带贝类多样性指数(H'')和均匀度指数(J'')显著高于潮下带,而丰富度指数(D)则相反;潮间带贝类生物量(710.16 g/m²)和栖息密度(4 523个/m²)远高于潮下带(17.10 g/m²,35个/m²)。群落结构多元统计分析将贝类按照站位划分为5个组群,包括东侧潮下带、西侧潮下带、西侧潮间带、东侧潮间带以及D4站位。冗余分析(RDA)结果表明,对潮间带贝类影响较大的环境因子主要为沉积物的粒径以及砂、粉砂含量,而对潮下带贝类影响较大的主要为浮游植物密度、沉积物的粒径以及砂、粉砂和黏土含量。渤海湾南部(滨州)滩涂贝类资源较为丰富,但应从不同滩涂区域生态特性的角度出发,加强贝类的保护和资源修复,本研究可为该区域贝类的资源养护和可持续利用提供数据基础。     

江苏小庙洪牡蛎礁的地貌-沉积特征

利用卫星影象和野外实地勘察资料对江苏小庙洪牡蛎礁的地貌-沉积特征进行分析。结果表明，该牡蛎礁发育在强潮淤泥质潮坪上，造礁牡蛎主要是近江牡蛎和长牡蛎，表层的鲜活牡蛎为褶牡蛎。牡蛎礁区海水属盐度较高(27—30)的半成水，含沙量较大，为0．2—0．3g／L。活体牡蛎堆积体顶面高于周围潮间下带滩面1．0—1．5m。潮流较强，多在0．5—2．0m／s。海岸剖面可分为4个带，即礁后潮间带、潮沟、礁体生长带及礁前斜坡带。由于处于海岸侵蚀段，礁后潮坪缺失潮上带和大部分的潮间上带。礁后潮坪主要以粗粉砂为主，礁后潮沟冲淤变化较大，故礁体生长带时而为一沙洲，时而又与岸滩相连。礁体生长带分布在潮间下带，可看到独立的斑状礁体、带状礁体和大面积环状礁体群。环状礁的微地貌可以划分为礁塘(泻湖)、塘口和塘沟、塘口三角洲、礁墙和礁平台等。     

夏季庙岛群岛大型底栖动物群落特征

根据2022年8月大潮期间庙岛群岛岛屿的潮间带和潮下带大型底栖动物的调查结果,对其种类组成、数量分布、优势种、多样性以及群落结构特点进行了研究。结果表明:本次潮间带定量调查共鉴定大型底栖动物52种,其中甲壳动物最多(17种);定性调查共鉴定42种,软体动物最多(30种)。潮下带定量调查共鉴定大型底栖动物39种,低于潮间带定量调查,其中软体动物最多(26种)。潮间带大型底栖动物平均丰度为234个/m²,平均生物量为82.82g/m²。潮下带平均丰度为180.2个/m²,平均生物量为708.02g/m²。潮间带大型底栖动物多样性指数(H′)、丰富度指数(D)和均匀度指数(J′)平均值分别为1.187、1.723和0.540;潮下带H′、D和J′平均值均高于潮间带,分别为1.301、2.204和0.669。在本次潮间带定量调查中,大型底栖动物优势种为施氏玻璃钩虾(Hyaleschmidti)、平背蜞(Gaeticedepressus)、潮间海钩虾(Pontogeneialittorea)等。潮下带优势种与...     

Growth dynamics of eelgrass, Zostera marina, in the intertidal zone of Seomjin Estuary, Korea

To examine the growth dynamics of eelgrass, Zostera marina, in the intertidal zone of Seomjin Estuary, Korea, we surveyed environmental factors such as water temperature, underwater irradiance, tidal exposure, and nutrient concentrations in the water column and sediment pore water in relation to the shoot density, biomass, morphological characteristics, and growth of Z. marina inhabiting the upper and lower intertidal zones. The survey was conducted monthly from January 2003 to December 2004. The water temperature of the two areas displayed seasonal fluctuations. Underwater irradiance was significantly higher in the upper intertidal zone than in the lower intertidal zone. Tidal exposure was also markedly longer in the upper intertidal zone than in the lower intertidal zone, whereas tidal exposure was highest in the spring and lowest in the summer in both areas. Water column NH₄ ⁺ and sediment pore water NO₃ ^?+NO₂ ^? concentrations were significantly higher in the upper intertidal zone than the lower intertidal zone. The eelgrass shoot density, biomass, morphology, and leaf productivity were significantly higher in the lower intertidal zone than in the upper intertidal zone. Both areas displayed a clear seasonal variation depending on changes in water temperature. However, leaf turnover time was significantly shorter in the upper intertidal zone than in the lower intertidal zone, with a higher turnover rate in the upper intertidal zone. Compared to the seagrasses in the lower intertidal zone, those in the upper intertidal zone showed more effective adaptations to the stress of long tidal exposure through downsizing and increased turnover time. These results suggest that tidal exposure, coupled with desiccation stress, can be a limiting factor for seagrass growth in the intertidal zone, along with underwater irradiance, water temperature, and nutrient availability.     

Ecological classification of European transitional waters in the North-East Atlantic eco-region

A new methodology to classify European North-East Atlantic transitional waters into ecological types has been developed based on the most important hydrological and morphological features that are likely to determine the ecology of aquatic systems in transitional waters. Hydrological indicators help identifying if a transitional water area is dominated by fresh or sea water and/or by intertidal or subtidal areas, while morphological indicators allow an estimation of the complexity of the transitional water and the diversity of the habitats involved. Twelve transitional waters of the southern Bay of Biscay were classified using this methodology and the five hydro-morphological types obtained were validated with benthic macro-invertebrate data. Transitional waters with a complex morphology showed the highest values of species diversity, while those with a smaller tidal influence showed lower species diversity. The ‘Scrobicularia’ and ‘Abra’ assemblages, previously identified in the study area, were found to be related to different types of transitional waters. The ‘Abra’ assemblage only appeared in estuaries with a complex morphology and dominated by tidal influences, while the ‘Scrobicularia’ assemblage was detected in all the transitional waters except for a single coastal lagoon. This classification of transitional waters may therefore be useful to establish the biological reference conditions needed for European Directives.     

三沙湾西北部滨海湿地潮间带生物生态研究

本文报道三沙湾西北部滨海湿地潮间带生物109种,其中多毛类、软体动物和甲壳动物占总种数的89.90%,三者构成潮间带生物主要类群.断面间比较,Sch1断面的种数最少（42种）,Sch3断面的种数最多（73种）.种类季节变化,以春季种数（83种）大于秋季（59种）.三沙湾西北部滨海湿地潮间带生物平均生物量12.45g/m^2,平均栖息密度为368个/m^2.生物量以软体动物居第一位（7.34g/m^2）,甲壳动物居第二位（2.55g/m^2）;栖息密度以软体动物居第一位（196个/m^2）,多毛类居第二位（154个/m^2）.数量垂直分布,生物量以中潮区（15.20g/m^2）大于高潮区（12.37g/m^2）大于低潮区（9.78g/m^2）;栖息密度以低潮区（563个/m^2）大于中潮区（483个/m^2）大于高潮区（56个/m^2）.数量季节变化,生物量以春季（16.69g/m^2）大于秋季（8.21g/m^2）;栖息密度以春季（677个/m2）大于秋季（58个/m^2）.三沙湾西北部滨海湿地潮间带生物按断面和所处的位置可分为3个群落：Sch1泥沙滩群落,高潮区：黑口滨螺-粗糙滨螺带;中潮区：中蚓虫-鸭嘴蛤-秀丽长方蟹带;低潮区：似蛰虫-焦河篮蛤-鸭嘴蛤带.Sch2泥沙滩群落,高潮区：黑口滨螺-粗糙滨螺带;中潮区：才女虫-鸭嘴蛤-淡水泥蟹带;低潮区：似蛰虫-侧底理蛤-鸭嘴蛤带.Sch3泥沙滩群落,高潮区：黑口滨螺-粗糙滨螺带;中潮区：才女虫-鸭嘴蛤-短拟沼螺-弧边招潮带;低潮区：不倒翁虫-侧底理蛤-齿腕拟盲蟹带.结果分析表明：春季Sch1和Sch2群落不稳定,秋季Sch1、Sch2群落和Sch3群落相对稳定,总体显示,三沙湾西北部滨海湿地潮间带生物群落发生扰动,初步分析认为与优势种鸭嘴蛤和优势种才女虫季节演替有关.     

Intertidal macroalgae and macroinvertebrates: Seasonal and spatial abundance patterns along an estuarine gradient

Quantitative sampling of the dominant intertidal epibiota was conducted seasonally along an estuarine gradient within the Great Bay Estuary System, New Hampshire, U.S.A. The abundance and zonation of the dominant macroorganisms varied with distance into the estuary. Replacement of marine by estuarine species occurred, and overall abundance and species richness decreased along the estuarine gradient. Zonation patterns within the inner estuary were primarily allied with substrata. Maximum abundance of invertebrates occurred in the mid-intertidal zone where a dense fucoid canopy provided habitat heterogeneity. Densities of epibiotic organisms decreased toward low water, especially in the inner estuary where hard substratum was limiting. Settlement blocks, introduced into the low intertidal zone, were dominated by barnacles and fucoid algae; after 16 months, the species composition on the settlement blocks resembled the adjacent community. Semibalanus balanoides settled in the spring, while Fucus vesiculosus var. spiralis exhibited low but constant settlement. Despite the physical rigors of the estuarine environment, only Semibalanus balanoides, Ilyanassa obsoleta and Spartina alterniflora showed significant seasonal changes in density. Thus, there are predictable and persistent epibiotic species assemblages within the intertidal zone of the Great Bay Estuary System.     

Macrofaunal spatial distribution and community structure in a lagoon without a river discharge (the Oualidia lagoon,NW Morocco)

The assemblage structure and spatial distribution patterns of macrobenthic communities according to environmental conditions were analysed through five samplings carried out at 14 intertidal soft sediment stations in the Oualidia lagoon (NW Morocco). A total of 89 macrobenthic taxa were collected, amounting to 55.188 specimens. Molluscs, polychaetes and arthropods were the most species‐rich groups. The snail Peringia ulvae was the most dominant species, especially abundant in the mid‐ and upper lagoon sections (nearly 64% of the total specimens gathered), followed by the mollusc Scrobicularia plana (10%). Diversity values were generally low (<3 bit) pointing out the high dominance by few species. Multivariate analyses clearly distinguished three distinct macrobenthic assemblages, successively distributed from lagoon inlets towards inner parts. Nevertheless, each assemblage was found in particular habitat types and had a specific species composition in response to different environmental characteristics. (a) The Scolelepis fuliginosa community occupied sandy substrates at the lower section of the lagoon with stronger marine influence and, (b) the P. ulvae community lived in sandy‐muddy substrates, including two sub‐communities dominated, respectively, by Cerastoderma edule, S. plana and Tritia pfeffeiri inhabiting the middle section and S. plana the upper section. The combination of salinity, temperature, pH and silt content was identified as the major environmental variable subset with the highest correlation with the observed macrofaunal patterns and silt fraction as the single best correlated variable. The results demonstrated significant spatial differences in macrobenthic assemblage structure and composition along environmental gradients in spite of the absence of direct freshwater river inflow.     

Existence of a deep subtidal stock of drifting Ulva in relation to intertidal algal mat developments

Blooms of drifting green algae often develop in shallow coastal zones that receive significant nutrient inputs. Each spring and summer, some fifty bays and coves in Brittany (France) are affected in this way. Until now, in this region, only the algae present in the surf zone or stranded ashore, constituting an intertidal stock, have been taken into account. Another stock of algae, which was subtidal and of the same species (Ulva spp.), was found in the Bay of Douarnenez, one of the ten areas most affected by these algal blooms. This subtidal Ulva stock was located beyond the surf zone, at depths reaching 15 m. It was about the same size as the intertidal stock, viz., a few thousand tons on average. Subtidal Ulva stocks were generally found lying on the sandy bottom in a distribution showing no particular pattern. Biomass ranged from almost zero to 1.547 kg m⁻² of fresh and spun-dried algae. However, at depths from 3 to 7 m they were often arranged in strips a few dm wide, due to the swell's effect. The bottom conditions of temperature, salinity, irradiance and dissolved inorganic nitrogen measured during spring and summer are suitable for the growth of Ulva in the subtidal zone. Both intertidal and subtidal drifting Ulva stocks are mobile and capable of exchanging material. In spring, the intertidal stock's inoculum is likely to come from the subtidal. Later in the season, the subtidal stock could be supplied, at least partially, by the intertidal.     

Structure of subtidal and intertidal communities of psammophilous ciliates of the Pechora Sea

A rich and diverse bottom ciliate fauna (168 species) was observed in the course of the studies of the subtidal and intertidal zones of the Pechora Sea; 81 of them were encountered for the first time in the Barents Sea region. As compared to the intertidal fauna, the sublittoral ciliofauna featured a higher species diversity but a lower total abundance (32.6 and 20.7 species per station and 10–100 and 2.4–443 ind./cm² in the subtidal and intertidal zones, respectively). With respect to trophic strategies, raptorial feeders were most abundant in the sublittoral zone (up to 77% of the total biomass), while both raptorial feeders (45%) and upstream filter feeders (36%) dominated in the littoral sediments. In the subtidal zone, two different types of ciliate communities were distinguished according to their total abundance and species composition. The first type is represented by abundant and species-rich communities mainly dominated by large specific interstitial ciliates developed over fine-grained well-sorted sands. The second type of communities joins mainly nonspecific eurybiotic ciliates dwelling over heterogeneous silty sediments; they feature lower abundance and species richness values. The high species diversity, the low abundance or absolute absence of meso-and polysaprobic groups indicating organic pollution, and the high values of the ABC size structure index suggested a good condition of the sediments and biota in the region studied.     

Sea-level fluctuations in a coastal lagoon

Sea-level observations made during December, 1979, at six stations in Great South Bay (which is a coastal lagoon on the south shore of Long Island, New York) reveal that there were significant subtidal fluctuations in addition to the tidal oscillations. Harmonic analysis of the tidal oscillations of sea level indicates that M₂ is the dominant tidal constituent. The M₂ amplitude, however, suffered a more than 50% reduction in the interior of the Bay due largely to the narrow inlet. The subtidal sea level fluctuations within the Bay were forced primarily by the low-frequency fluctuations of the adjacent shelf water. The active subtidal exchange induced by this Bay-shelf coupling appeared to have suffered only minor attenuation within the Bay. As a consequence, the variance associated with subtidal sea level fluctuations was greater than that associated with the tidal oscillations over most of Great South Bay.     

A numerical study of tidal asymmetry in Okatee Creek,South Carolina

The Okatee River, South Carolina is characterized by a narrow tidal channel and an extensive area of intertidal salt marshes. Current measurements in the upstream portion Okatee Creek show that tidal flow features an asymmetric pattern: ebb current is stronger than flood current. The ebb dominance is mainly caused by deformation of the dominant astronomical tidal constituent M₂. An unstructured grid, finite volume coastal ocean model (FVCOM) with wet-dry point treatment method is applied to examine physical mechanisms of M₄ overtide generation. Model experiments show that mean absolute amplitude and phase errors are 3.1 cm and 1.7^° for M₂ elevation, 2.4 cm s⁻¹ and 0.8^° for M₂ current major axis, 2.1 cm and 1.8^° for M₄ elevation, and 2.1 cm s⁻¹ and 24.6^° for M₄ current major axis. The overall pattern of tidal asymmetry is qualitatively reproduced. Various sensitivity experiments suggest that the generation of M₄ overtide is a result of nonlinear interaction of tidal currents with irregular creek geometry and bottom topography. Consistent with the classical view, the large volume of intertidal water storage is the major reason for ebb dominance in the creek. However, the zero-inertia assumption (i.e., negligible advective terms) is probably not valid for the entire tidal cycle. Besides the pressure gradient force and the bottom friction force, terms related to lateral shear of the along-estuary velocity (i.e., advective inertia and horizontal eddy viscosity) may also contribute in horizontal momentum balance. Exclusion of the flooding-draining processes over the intertidal zone will severely underestimate tidal currents in the river channel and make the tidal asymmetry less prominent.     

Shell‐shape and morphometric variability in Mytilus galloprovincialis from micro‐tidal environments: responses to different hydrodynamic drivers

Tidal conditions differently influence inter‐tidal organisms in terms of general physiological and metabolic responses. In this study we investigated the morphological response in shells of Mytilus galloprovincialis native to different micro‐tidal coastal environments in the Northern Adriatic Sea. Our purpose was to highlight the ecophenotypic variability across tidal levels and to elucidate how tidal currents and waves produced by anthropogenic activities may play a part in modulating shell morphology. Three sampling sites were selected: an open‐sea area 15 km off‐shore and two sites within the lagoon of Venice, the first near one of its three inlets, and the other one in the proximity of Venice city centre. At each sampling site, organisms were seasonally collected at different depths within their vertical zonation, either in the inter‐tidal zone – i.e. at both the highest and lowest tide zonation limits, and subtidally. The mussel shells were analysed by investigation of their morphometric relationships (height/length and width/length ratios) and by elliptic Fourier analysis of the shell contours. Shell thickness and condition index were also evaluated for a better comprehension of energy allocation/partitioning. Estimates based on long‐term measurements, visual observation, wind statistics and wave growth laws allowed an evaluation of the forces acting on shells. At the open‐sea site, the observed phenotypic variability of both shell shape and thickness was clearly related to the tidal vertical zonation. At the two lagoon sites, the currents generated by tidal flow through the inlet and the waves caused by the frequent passage of boats influenced both shell shape and thickness. A trade‐off between protection and growth was apparent along the tide gradient, as emphasized by the differences in the partitioning and allocation of energy between shell and flesh production.     

Fish fauna recovery in a newly re-flooded Mediterranean coastal lagoon

Drana Lagoon, located at the NW site of Evros River Delta, was drained in 1987 and re-flooded in 2004 within the framework of an integrated wetland restoration project. This study presents the results of a monitoring program of the lagoon's oceanographic, water quality and fish fauna characteristics, during the pre- and post-restoration period. Results depict the presence of high salinity water (up to 41) due to seawater intrusion, strong evaporation in its interior and inadequate freshwater inflows. Overall, nutrient levels were low depicting local changes. Tidal variability at the mouth was approximately 0.2 m, producing high velocity tidal currents (up to 0.75 m/s). Eleven fish fauna species were collected; seven species were caught in both the inlet channel and the lagoon during the pre-restoration period and nine species in the post-restoration period. Atherina boyeri (37.6%) and Pomatoschistus marmoratus (31.7%) dominated the lagoon during the post-restoration period. Most of the A. boyeri specimens (88.5%) were caught inside the lagoon, while P. marmoratus had an almost equal distribution in the inlet channel and the lagoon (56.3% and 43.7% respectively). The presence of species of the Mugilidae family (5.2% total average catches after lagoon re-flooding) was mainly in the inlet channel (12.6% of the average catches) and not inside the lagoon (only 1.3% of the average catches). The small number of fish species inhabiting the lagoon might be the result of the recent restoration or it could be related with the increased water flow observed at the lagoon mouth during the flood and ebb tidal phases, and also in the presence of a smooth bank in the concrete waterspout that connects the entrance channel with the lagoon. The limited presence of the Mugilidae juveniles inside the lagoon could be related to the prevailing tidal inlet dynamics (i.e. strong ebb flow at lagoon inlet), thus preventing the species to enter the lagoon. In order to restore the lagoon environment, careful and gradual steps should be undertaken under the basis of continuous monitoring of hydrologic, environmental and fisheries system's status.     

Use of different intertidal habitats by faunal communities in a temperate coastal lagoon

The faunal communities of four intertidal habitats namely sand, mud, seagrass (Zostera noltii) and seagrass patches (mixSM) of a temperate coastal lagoon, Ria Formosa (southern Portugal), were sampled. A total of 47 species were taken in 428 bottomless drop sampler samples, with the highest number of species and the more commonly occurring species belonging to the Mollusca phylum. The dominance of these gastropod species underlines the importance of the grazing food chain in these habitats. Bittium reticulatum was the most abundant species, being especially abundant in the seagrass habitat. The most frequent and highest biomass species in the community was Carcinus maenas, a predator that makes use of the available resources and that is adapted to the highly variable intertidal environment. Pomatoschistus microps was the most abundant fish species, with highest densities in the mud habitat, which demonstrates an ability to occupy a low depth area. The seagrass habitat had the highest diversity, abundance and biomass, followed by the mixSM habitat and was different from all the others. Assemblages were highly influenced by the presence of vegetation, providing forage and refuge from predation. A well defined summer group was identified in all habitats. These results highlight the importance of seagrass beds and the idea that their decrease implies the decrease of lagoon production through the impoverishment of the trophic structure of the lagoon.     

Keeping things in order: multivariate direct gradient analysis of a strongly fluctuating benthic community

Two spatial surveys of the macrobenthos of an estuarine intertidal area, the Oosterschelde in the southwestern part of The Netherlands, were conducted in different years. In the period between the surveys changes in the hydrodynamics of the estuary took place as a result of the construction of a storm-surge barrier. The works reduced the tidal volume in the basin and as a consequence the altitude of the tidal flats decreased by approximately 0.1 to 0.2 m. Considerable differences in species densities were found between the two surveys, but these large differences could not be explained by changes in the ‘spatial' environmental variables (such as altitude) that were taken into account in a direct gradient analysis. Unknown and ‘only-to-time-related' (e.g. weather-related) factors had considerably influenced species abundance. This implies that the impact of the works on the macrobenthos could not be evaluated on the basis of observed species densities in the before- and after-the-works surveys. Generally, it implies that even if the levels of future ‘spatial' environmental variables are known, direct gradient analysis based on a single survey cannot be used for prediction of future densities in absolute terms in such strongly fluctuating communities. Yet, the gradient analyses for the two years revealed rather similar estimates of the major environmental gradients. The same was true for the parameters that described the relative change in species densities in response to a change in the environmental gradient levels. Hence the most appropriate way of evaluating the changes that have occurred in the benthic community as a result of the works was in terms of the observed changes in the ‘spatial' environmental gradient levels. The levels of the first environmental gradient, which was strongly related to the sediment structure, did not change between years. Levels of the second gradient, which could be interpreted as a tidal zone gradient, decreased. This suggests that the works particularly affected taxa that are strongly related to this second gradient, such as Corophium living in the upper tidal areas and Nephthys living in the lower tidal zone.     

Understanding the association between the non-geniculate coralline red alga Spongites discoidea and the mollusc Oxystele sinensis

Nearly all adult winkles Oxystele sinensis encountered in the Kalk Bay (Western Cape province, South Africa) shallow subtidal zone were observed to bear a thick, convoluted form of the non-geniculate coralline red alga Spongites discoidea. To understand this association, various data were collected. The results show that O. sinensis is for the most part restricted to the shallow subtidal zone and rockpools, whereas the morphologically similar O. tigrina is largely restricted to the intertidal zone. Fleshy macroalgae were absent from all habitats sampled, with much of the primary substratum being occupied by encrusting algae. Spongites discoidea was restricted to intertidal rockpools and the shallow subtidal zone where O. sinensis was the most abundant winkle. Subtidal specimens of S. discoidea were generally thicker (7.24?mm [SE 0.55] on shells, 6.02?mm [SE 0.67] on boulders) than intertidal specimens (2.39?mm [SE 0.21] on boulders, 1.02?mm [SE 0.09] on bedrock), were more heavily burrowed into (44% [SE 3.32] for subtidal shells, 36% [SE 3.94] for subtidal boulders), and were more weakly attached (2.2?kg [SE 0.58] for subtidal shells, 8.4?kg [SE 0.77] for subtidal boulders). Experimental evidence showed that the coralline has no preference for shells of O. sinensis and that, contrary to our assumption, there is no apparent advantage to the winkle bearing the coralline, other than perhaps for camouflage to avoid predation. The high frequency of the association is more likely due to the substantial overlap in the niches of the coralline and the winkle.