Macrobenthic community structure of soft-bottom sediments at the Belgian Continental Shelf

Within the frame of different research projects, a large number of sites at the Belgian Continental Shelf (BCS) have been sampled for the macrobenthos between 1994 and 2000. These samples cover a diverse range of habitats: from the sandy beaches to the open sea, from the gullies between the sandbanks to the tops of the sandbanks, and from clay to coarse sandy sediments. To investigate the large-scale spatial distribution of the macrobenthos of the Belgian Continental Shelf, the data of all these research projects—728 samples—were combined and analysed. By means of several multivariate techniques, 10 sample groups with similar macrobenthic assemblage structure were distinguished. Each sample group is found in a particular physico-chemical environment and has a specific species composition. Four sample groups differ drastically, both in habitat and species composition, and are considered to represent four macrobenthic communities: (1) the muddy fine sand Abra alba–Mysella bidentata community is characterized by high densities and diversity; (2) the Nephtys cirrosa community occurs in well-sorted sandy sediments and is characterized by low densities and diversity; (3) very low densities and diversity typify the Ophelia limacina–Glycera lapidum community, which is found in coarse sandy sediments and (4) the Eurydice pulchra–Scolelepis squamata community is typical for the upper intertidal zone of sandy beaches. These macrobenthic communities are not isolated from each other, but are linked through six transitional species assemblages. The transition between the A. alba–M. bidentata community and the N. cirrosa community is characterized by a reduction in the mud content and is dominated by Magelona johnstoni. The transition between the N. cirrosa and the O. limacina–G. lapidum community is distinctive by decreasing densities and coincides with a gradual transition between medium and coarse sandy sediments. From the N. cirrosa to the E. pulchra–S. squamata community, transitional species assemblages related to the transition from the subtidal to the intertidal environment were found. Each community or transitional species assemblages was found over a specific range along the onshore–offshore gradient, four types can be discerned: (1) almost restricted to the near-shore area, but possible wider distribution; (2) distributed over the full onshore–offshore gradient; (3) restricted to the near-shore area and (4) restricted to the sandy beach environment. The diversity pattern on the BCS follows this division, with species rich and poor assemblages in the near-shore area to only species poor assemblages more offshore. The distribution and diversity patterns are linked to the habitat type, distinguished by median grain size and mud content.     

The rise of Laminaria ochroleuca in the Western English Channel (UK) and comparisons with its competitor and assemblage dominant Laminaria hyperborea

The distribution of species is shifting in response to recent climate change. Changes in the abundance and distributions of habitat‐forming species can have knock‐on effects on community structure, biodiversity patterns and ecological processes. We empirically examined temporal changes in the abundance of the warm‐water kelp Laminaria ochroleuca at its poleward range edge in the Western English Channel. Resurveys of historical sites indicated that the abundance of L. ochroleuca has increased significantly in recent decades. Moreover, examination of historical records suggested that L. ochroleuca has extended its distribution from sheltered coasts on to moderately wave‐exposed open coasts, where it now co‐exists and competes with the assemblage dominant Laminaria hyperborea. Proliferation of L. ochroleuca at its poleward range edge corresponds with a period of rapid warming in the Western English Channel. Preliminary comparisons between L. ochroleuca and L. hyperborea highlighted some subtle but ecologically significant differences in structure and function. In summer, the average biomass of epiphytic stipe assemblages on L. hyperborea was 86 times greater than on L. ochroleuca whereas, on average, L. ochroleuca had a greater stipe length and its blade supported 18 times as many gastropod grazers (Gibbula cineraria). Differences in summer growth rates were also recorded, with L. ochroleuca being more productive than L. hyperborea throughout July. Comprehensive seasonally replicated comparisons are needed to examine the wider implications of proliferation of L. ochroleuca at its poleward range edge, but our study suggests that local biodiversity patterns and ecological processes (e.g. timing of productivity and trophic pathways) on shallow subtidal reefs may be altered by shifts in the relative abundances of habitat‐forming kelp species.     

Benthic primary production, respiration and remineralisation: in situ measurements in the soft-bottom Abra alba community of the western English Channel (North Brittany)

In situ measurements of ammonium and carbon dioxide fluxes were performed using benthic chambers at the end of spring and the end of summer in two soft-bottom Abra alba communities of the western English Channel (North Brittany): the muddy sand community (5 m, about 10% of surface irradiance) and the fine-sand community (19 m, about 1% of surface irradiance). High rates of ammonium regeneration were measured in the two communities at the end of summer (296.03±40.07 and 201.7±62.74 μmolN m⁻² h⁻¹, respectively) as well as high respiration rates (2.60±0.94 and 2.23±0.59 mmolC m⁻² h⁻¹, respectively). Significant benthic gross primary production (up to 6.11 mmolC m⁻² h⁻¹) was measured in the muddy sand community but no benthic primary production was measured in the fine-sand community. It suggests that microphytobenthic production values used in simulations previously published for these two communities were overestimated while values of community respiration were underestimated. The study confirms that this benthic system is heterotrophic and strengthens the idea that an important pelagic-benthic coupling is required for the functioning in such coastal ecosystems.     

Validation of the three-dimensional ECOHAM model in the German Bight for 2004 including population dynamics of Pseudocalanus elongatus

A three-dimensional ecosystem model for the North Sea which includes competition between Pseudocalanus elongatus and the rest of the zooplankton biomass was applied to describe the seasonal cycle of zooplankton in 2003–2004. The paper presents the comparison of simulated stage-resolved abundances with copepod counts at several stations in the German Bight during the GLOBEC-Germany project from February to October 2004. A validation of influential state variables gives confidence that the model is able to calculate reliably the stage development and abundances of P. elongatus as well as the range of bulk zooplankton biomass, and thus the ratio of population biomass to total biomass. In the German Bight, the population is below 20% in spring. The ratio increases up to 50% during summer. The number of generations was estimated from peaks in egg abundance to about 4–8 generations of P. elongatus in the southern North Sea. A mean of four generations per year were estimated in the central North Sea, six to eight generations northwest of the Dogger Bank (tails end) and five generations in the German Bight.     

Macrozoobenthic biomass in the Bay of Seine (eastern English Channel)

The benthic biomass values of various trophic groups were studied for the first time at the scale of the entire Bay of Seine (50 × 100 km) in the eastern English Channel. Sampling was carried out before and after the winter of 1999. In both cruises the suspension feeders dominated (66% of the 12 gAFDW m^− 2 in 1998 and 56% of the 10 gAFDW m^− 2 in 1999).The common European ophiuroid Ophiothrix fragilis was the most important contributor to total biomass. The repartition of its patches (sometimes > 20 gAFDW m^− 2) cannot be explained by the environmental parameters recorded (viz., granulometry, organic matter and pigment content).     

In situ Study of Bromide Tracer and Oxygen Flux in Coastal Sediments

Laboratory and in situ experiments were performed to assess the use of bromide as a tracer forin situ studies of benthic solute exchange. Bromide was used in the benthic chamber lander ‘ Elinor ’ for flux measurements in coastal sediments of the German Bight, Kiel Bight and Skagerrak (28–700 m water depth). Tracer and total oxygen uptake were monitored simultaneously in the same chamber incubation. Concurrently, in situ oxygen micro-profiles were recorded at the same locations by the profiling lander ‘ Profilur ’. Deployment in an anoxic silt (Kiel Bight) confirmed that in the absence of bioturbation and advection, tracer transport into the sediment was driven solely by molecular diffusion. This flux could be well described by a simple box model accounting for molecular diffusion only. In oxic sediments (German Bight and Skagerrak) enhanced exchange of bromide tracer due to bioirrigation parallelled enhanced oxygen uptake equivalent to a 4-fold molecular diffusive flux. Our experiments showed that incubations can be short. Depending on irrigation activity of the fauna, however, incubation length should exceed 3 h in order to provide a useful data base for flux calculations. The method demonstrating caveats is discussed and indicate possible improvements. The results show how the bromide tracer addition can be used as a tool for determining solute fluxes exceeding diffusive flux in benthic chamber incubations.     

Seasonal variation of Mn adsorption on to calcareous surfaces in the English Channel, and its implication on the manganese distribution coefficient

Dissolved and particulate manganese in seawater samples derived from the English Channel has been analyzed using graphite furnace atomic absorption (GFAAS), and inductively coupled plasma atomic emission (ICP—AES) spectroscopies. Because of the high contents of carbonate minerals found in the suspended matter samples, the English Channel constitutes an ideal field area for the study of the Mn²⁺/Ca²⁺/CaCO₃ system. Owing to the chemical speciation of particulate manganese and the combined use of the X-ray diffraction and electron spin resonance spectroscopy we have shown the importance of the carbonate phase in the stabilization of manganese (II). This has been confirmed by p-pH measurements in the field. All these studies have also indicated that: (1) manganese is associated with calcite in the form of a solid solution, Mn_xCa_1−xCO₃; and (2) significant increases in the concentrations of particulate manganese, especially in offshore waters, occur in summer. This seasonal phenomenon has been attributed to the proliferation of coccolithophorids, which are known to be covered with calcified skeletons at high specific surface areas. To appraise the implication of the coccolithophorid-blooms phenomenon on the Mn²⁺/Ca²⁺/CaCO₃ system, we have used the manganese distribution coefficient, D_i, between the liquid phase and CaCO₃ particles. Overall we have shown that: (1) D_i in summer (i.e. when coccoliths considered as very fine-grained calcite are abundant) is much higher than that obtained in winter; and (2) in the vicinity of the French coast, D_i does not vary significantly even in summer. This is because of the high content of chalk-derived particles found in the near-shore waters.     

Extracellular release of small and large molecules by phytoplankton in the Southern Bight of the North Sea

Extracellular release by phytoplankton of small and large metabolites (molecular weight 〈and〉 500 d) is determined kinetically in various marine ecosystems including the eutrophic Scheldt estuary, the Belgian coastal zone and the oligotrophic English Channel. Kinetic curves show that while small metabolites are immediately released in the external medium, a lag phase is always observed in the release of large metabolites. Results indicate that 9–48% of the total excreted amount ends up in the small metabolites pool with no dependence on light intensity or nutritive conditions. Relative rates of heterotrophic uptake of the phytoplankton extracellular metabolites are calculated from the time dependent distribution of ¹⁴C labelled extracellular products. The data indicate that small molecules with high turnover rates (up to 25% h^?1) may occasionally be excreted by phytoplankton cells. The turnover time of high molecular weight excreted compounds is much longer. A correlation is found between the rate of production of these large extracellular metabolites and the exoenzymatic activity of the bacterial populations.     

Mesoscale and microscale spatial variability of bacteria and viruses during a Phaeocystis globosa bloom in the Eastern English Channel

Sampling was conducted within inshore and offshore sites, characterized by highly dissimilar hydrodynamic and hydrobiological conditions, in the Eastern English Channel. The eutrophic inshore site was dominated by the influence of a dense bloom of the Prymnesiophyceae phytoplankton species Phaeocystis globosa, while the offshore site was characterized by more oceanic conditions. Within each site the microscale distributions of chlorophyll a and several flow cytometrically-defined subpopulations of heterotrophic bacteria and viruses were measured at a spatial resolution of 5 cm. The inshore site was characterized by comparatively high levels of microscale spatial variability, with concentrations of chlorophyll a, heterotrophic bacteria, and viruses varying by 8, 11 and 3.5-fold respectively across distances of several centimeters. Within the offshore site, microscale distributions of chlorophyll a and bacteria were markedly less variable than within the inshore site, although viruses exhibited slightly higher levels of heterogeneity. Significant mesoscale variability was also observed when mean microbial parameters were compared between the inshore and offshore sites. However, when the extent of change (max/min and coefficient of variation) was compared between meso- and microscales, the variability observed at the microscale, particularly in the inshore site, was substantially greater. This pattern suggests that microscale processes associated with Phaeocystis globosa bloom dynamics can generate heterogeneity amongst microbial communities to a greater degree than large scale oceanographic discontinuities.     

Unusual diatoms linked to climatic events in the northeastern English Channel

The composition of the micro-phytoplankton was investigated at two fixed stations in the northeastern English Channel from November 1997 to December 2005. In late summer-autumn 2005, proliferations of the centric diatoms Proboscia indica and Rhizosolenia hebetata f. semispina were recorded for the first time in this area. The weather in 2005 was abnormal: a cold winter and early arrival of summer conditions that extended to late autumn. This resulted in an unusually long, sustained calm and warm period. We suggest that the reduction in mixing may have allowed diatoms of more-stratified waters to be competitive in the normally highly mixed northeastern English Channel in summer-autumn 2005. In 2003 and in 2005, two warm-water diatom species were recorded for the first time. A colony of Eucampia cornuta was observed after an exceptional heat wave in September of 2003, and again in September 2005. Simultaneously at both fixed stations, Chaetoceros peruvianus was recorded in the warm October of 2005. The climatic events are associated with the northward spreading of thermophilic diatoms such as E. cornuta and Ch. peruvianus in the European Atlantic waters, and the proliferation of diatoms of stratified environments such as P. indica and R. hebetata f. semispina in the usually highly turbulent waters of the northeastern English Channel.     

Inter-season and interannual variations in fish and macrocrustacean community structure on a eastern English Channel sandy beach: Influence of environmental factors

The intertidal zone of an exposed sandy beach on the French coast of the English Channel was sampled with a 1.5 m beam-trawl over five years (2000 and 2003–2006) at weekly intervals. The fish and macrocrustacean catches were analysed to determine the inter-season and interannual variation in community structure and relate these variations to changes in the major environmental variables. Only six species (plaice Pleuronectes platessa, common goby Pomatoschistus microps, sprat Sprattus sprattus, sand eel Ammodytes tobianus, brown shrimp Crangon crangon and shore crab Carcinus maenas) from the 27 species captured can be considered as dominant species of the intertidal zone, and they accounted for >90% of total numbers. Most individuals caught were young-of-the-year or juvenile. Analysis of similarities (ANOSIM) and similarities percentage (SIMPER) indicated that inter-season variability of community structure (mean average similarity = 47%) was more pronounced than interannual variability (mean average similarity = 65%). Canonical correspondence analysis (CCA) indicates that a substantial component (32.2%) of the measured inter-season variation in community structure can be explained by environmental factors (mainly water temperature). The main inter-season changes in the abundance and community structure were due to the variation of the six key species and reflect the different times of their recruitment. During the five years of the study, the structure of the fish and macrocrustacean spring community persisted from year to year, with the dominant species reappearing consistently even though their abundances fluctuated from year to year. This interannual variation probably reflects variable recruitment success influenced by physico-chemical conditions. In spite of the considerable interannual variation (40 times) in the spring bloom of the prymnesiophyte alga Phaeocystis globosa we found no effect of this bloom on either fish and macrocrustacean species densities or diversity index.     

Distribution and feeding modes of macrobenthos within three oceanographic feature areas of the KwaZulu-Natal Bight,South Africa

The composition and distribution of macrobenthic communities was investigated in three areas in the KwaZulu- Natal Bight, a section of shelf off the east coast of South Africa. Areas were pre-selected on the basis of three known oceanographic features, posited to deliver land- or Agulhas Current-derived nutrients onto the shelf and to drive ecosystem functioning in this region. Replicate sediment samples were collected with a 0.2 m² van Veen grab, during two surveys (A, B) corresponding with normal periods of high and low rainfall, respectively. A subset of the full station array was selected across the shelf in an arrangement of increasing depths (inner-, mid- and outer shelf) through each feature area to investigate the spatial distribution and feeding modes of macrobenthic taxa. The two periods showed some differences in abundance and numbers of macrobenthic taxa, but were not statistically different. Total macrobenthic abundance from Survey A was 20 215 individuals from 642 taxa, decreasing to 18 000 individuals from 503 taxa during Survey B. Polychaeta and Crustacea were the dominant taxa sampled; abundance of the latter was attributed largely to a proliferation of Paguristes sp.1 at inner-shelf samples in the midbight (Thukela) region during Survey B. Similarity classification distinguished seven sample groups reflecting differences in feature areas and shelf positions under investigation. The Thukela River midshelf community supported the highest macrobenthic abundance, while the midshelf off the southern bight (Durban region) was most species rich. Findings were attributed to the habitat complexity of the midshelf which includes a palaeo-dune cordon at the 60 m isobath. Functionally, the community was dominated by interface- and deposit-feeding fauna, emphasising the importance of trophic plasticity in an environmentally variable and heterogeneous shelf environment.     

The biogeography and community structure of kelp forest macroinvertebrates

Understanding species distributions and their community structure is increasingly important when taking an ecosystem‐based approach to conservation and management. However, knowledge of the distribution and community structure of species in mid‐range trophic levels (e.g. macroinvertebrates) is lacking in most marine ecosystems. Our study aimed to examine the spatial distribution and community‐level biogeographic patterns of common kelp forest–rocky reef macroinvertebrates in Southern California and to evaluate the effects of environmental gradients on these communities. Quantitative SCUBA surveys were used to estimate macroinvertebrate densities at 92 sites from 2008–2012. Non‐metric multidimensional scaling was used to evaluate spatial patterns of macroinvertebrate communities among Regions. We found that kelp forest–rocky reef macroinvertebrate communities are distinct among different island and mainland regions, and their community patterns exhibited a strong relationship with an environmental gradient (i.e. sea surface temperature) even after controlling for geographic distance between sites. High abundances of urchin species (Strongylocentrotus purpuratus and Strongylocentrotus franciscanus) were strong drivers of regional differences. Macroinvertebrate community patterns were driven by characteristic species that were typically more prevalent at warmer or colder sites. Our results provide the first quantitative analysis of macroinvertebrate community structure within the California kelp forest ecosystem. We also describe the distribution and abundance of 92 conspicuous kelp forest‐rocky reef macroinvertebrates among nine pre‐defined Regions. This study provides important preliminary information on these macroinvertebrate species that will be directly useful to inform management of invertebrate fisheries and spatial protection of marine resources.     

Spatial distribution patterns of the peppery furrow shell Scrobicularia plana (da Costa, 1778) along the European coast: A review

The bivalve Scrobicularia plana is an important species of shallow water benthic communities with a wide geographic distribution but also with a general patchy pattern, i.e. irregular in occurrence and in density. This review aims to determine the processes responsible for the species' spatial distribution pattern based on the available information on S. plana. Although several pre- and post-settlement processes are believed to influence spatial patterns of marine invertebrates, the general patchy distribution of S. plana seems to be determined by the existence of specific environmental conditions during settlement. Factors such as temperature, salinity, sediment type, hydrographic conditions and predation affect settlement and spat survival and not one but a combination of factors seems to explain the species distribution pattern. Future work should focus on determining the scale of patchiness, using hierarchical sampling, as well as the connectivity between populations by analysing the population genetic structure.     

Does the Port 2000 harbour construction have an effect on the Seine estuary suprabenthic community?

Using a Before/During/After protocol, the effects of Port 2000 (the Le Havre harbour extension) on the suprabenthic communities in the Northern Channel of the Seine estuary (English Channel) were examined from March 2001 to March 2009. Since the beginning of Port 2000 construction in 2002, there have been several changes in the hydrological and sedimentary characteristics in the North Channel, including changes in current velocity, bottoms salinity and sand accumulation, as well as in the benthic communities themselves, moving from mud to medium and coarse sand communities. For the suprabenthos, changes were moderate, corresponding to an increase in species richness in the upper-part of the North Channel and a decrease in species density in the dominant mysids species (i.e., mainly Mesopodospsis slabberi) over time. Due to hydrological changes since the beginning of the Port 2000 project (several years of low freshwater input) it has been difficult to attribute the biological changes to Port 2000 construction only. In the future, the results obtained for the suprabenthos will have to be compared to those obtained for the macrobenthos. Similarly, since the suprabenthos is known to play an important role in the estuarine trophic chain, it will also be essential to estimate the fish distribution in the North Channel and to examine their stomach contents to evaluate the probable changes since the Port 2000 project began.     

凡纳滨对虾-缢蛏生态循环养殖池塘中浮游植物群落结构与水质因子相关性研究

为研究凡纳滨对虾(Litoperaaeus vannamei)与缢蛏(Siaoraovacula constricta)生态循环养殖过程中浮游植物群落结构及其与水质因子变化特征之间的相关性,本实验于2018年6月至11月在宁波市鄞州区椿霖养殖场对虾蛏循环养殖池塘中水质及浮游植物的动态开展了逐月采样监测,分析了养虾塘与养蛏塘中的浮游植物群落结构变化及水质因子变动。结果显示:(1)对虾养殖期间共鉴定出6个门101种浮游植物(包含9个未定种),从种的数量上来看,硅藻门>甲藻门>绿藻门>蓝藻门>裸藻门>隐藻门,其中包含优势种14种,养殖初期优势种为硅藻门的新月菱形藻(Nitzschia closteriu)、牟氏角毛藻(Chaetoceros muelleri)等,随后甲藻门的海洋原甲藻(Prorocentrum micans)和绿藻门的小球藻(Chlorella vulgaris)等逐渐占据优势,后期仍以硅藻门为主要优势种。(2)浮游植物的丰度介于6.8×10⁵-2.5×10⁸cell/L,生物量为2.04-65.72...     

Variabilité spatio-temporelle de la biomasse microphytobenthique en rade de Brest et en Manche occidentale

Microphytobenthic biomass was estimated in the Bay of Brest (1994) and the western English Channel (Trezen Vraz, 1993) using spectrophotometry. Best results (42 % difference) were obtained with an instantaneous extraction procedure at room temperature, compared with the cold extraction procedure. Chlorophyll biomass in sediment was higher in the western English Channel (65–215 mg Chl a m⁻²) than in the Bay of Brest (10–113 mg Chl a m⁻²), in contrast to total pigment biomass (Chl a + pheo a : 88–254 mg m⁻² at Trezen Vraz vs. 131–934 mg m⁻² in the Bay of Brest). This study emphasized decreasing pigment biomass from the estuary to the open sea. Low benthic pigment biomass in the Bay, compared to eutrophic ecosystems, agrees with previous studies on the pelagic system, suggesting that there is no eutrophication in the Bay of Brest. Microphytobenthic blooms were strongly correlated with environmental factors, such as river floods, light intensity, water temperature and the spring/neap tidal cycle. Turbidity and light could be limiting factors at Roscanvel, i.e. the deepest site facing gyre circulation. The latter would favour sedimentation of silt particles at the sediment-water interface. Microphytobenthos population dynamics would be also dependent on the spring/neap tidal cycle, since maximum chlorophyll biomass occurred at neap tides at all three sites in the Bay of Brest, in contrast to the western English Channel. Lower temporal variations of pigment biomass in sediment were observed in the Channel, compared with the Bay of Brest; maximum chlorophyll biomass in sediment was found to occur three to four weeks after the phytoplankton blooms.     

Zooplankton communities and oceanographic structures in a high-resolution grid in the south-eastern corner of the Bay of Biscay

A one year long monthly sampling series at a fixed station was combined with a high-spatial resolution survey (May 2003) to investigate factors controlling zooplankton distribution in the south-eastern corner of the Bay of Biscay. Species abundance and gonad maturity for Calanoides carinatus and Calanus helgolandicus were estimated to clarify whether the observed spatial patterns were attributable to increased population growth or to mechanical accumulation. The two studies included distinct oceanographic regimes. The eastern corner of the grid was characterized by Adour river plume waters and the Cap Breton canyon, and the rest of the area by an alternating pattern of physical convergences and divergences. In the river plume, the zooplankton community diversity and abundances were highest in response to the nutrient enriched river plume waters and the submarine canyon. In the rest of the area, distribution and abundance patterns were the result of a combination of behaviour and physical transport.