Benthic macrofauna communities of the submersed Pleistocene Elbe valley in the southern North Sea

Macrozoobenthic community structure was studied in two surveys along a transect of 13 stations following the submersed Pleistocene Elbe valley in the south-eastern North Sea during May to June 2000 and March 2001. Two replicates of bottom samples were taken with a van Veen grab of 0.1 m² sampling size. In order to analyse the benthic macrofauna communities, the animals obtained were identified and counted, and MDS and cluster analysis were performed. Out of 200 taxa identified, 84 were polychaetes, 46 molluscs, 40 crustaceans, 15 echinoderms, and 15 belonged to other groups. Mean abundance was 4,860 individuals per m², mean biomass 32.9 g ash free dry mass per m². Mean diversity was 1.76 and mean evenness 0.54. The macrofauna of the Pleistocene Elbe valley is composed of three associations according to the cluster analysis. Each association is described by a combination of characterizing and discriminating species. An Amphiura-brachiata―Tellimya-ferruginosa-association was found in the south-eastern part of the depression, whereas a transitional association with elements of both assemblages lead to an Amphiura-filiformis―Galathowenia-oculata―Nuculoma-tenuis-association in the north-western part of the valley. In the context of the entire southern North Sea, both associations are small-scale substructures, and as such are contained in the Nucula-nitidosa-community and the Amphiura-filiformis-community, respectively. A north-westward shift of the community of the southern Elbe valley was found and discussed as a possible consequence of warm winters of the last decades.     

Neandertal mobility and large-game hunting: The exploitation of reindeer during the Quina Mousterian at Chez-Pinaud Jonzac (Charente-Maritime, France)

Neandertals were effective hunters of large ungulates throughout their geographic and temporal ranges. Equipped with this knowledge, researchers in paleoanthropology continue to seek insight on the relationships between hunting and subsistence strategies with other components of the Neandertals’ niche, such as mobility, site use, and lithic technology. The Quina Mousterian deposits from the rockshelter site of Chez Pinaud Jonzac (Charente-Maritime, France; hereafter Jonzac) offer an excellent opportunity to pursue these issues. This paper focuses on the extensive and well-preserved skeletal remains of reindeer (Rangifer tarandus) recovered from recent excavations of the site, representing at least 18 individuals that were hunted by Neandertals during the fall through winter. Our zooarchaeological results indicate that all ages of reindeer were hunted but adult individuals predominate. No bias is evident in the comparable frequencies of males and females. These prey were butchered on-site, with abundant evidence of meat filleting and marrow exploitation. In the excavated sample, the absence of hearths and the almost complete lack of burned bones or stones suggest that Neandertals were not using fire to assist with processing the reindeer carcasses. The zooarchaeological results presented here indicate that reindeer were hunted during a restricted window of time when they were seasonally abundant in the local area near Jonzac. Taken together with the lithic industry based on bifacial elements, the evidence is consistent with a pattern of site use by highly mobile hunter-gatherers making frequent, short-term visits. Ongoing research at Jonzac and other Quina Mousterian localities will contribute to a better understanding of Neandertal behavior during cold climate phases.     

Effects of the severe winter 1995/96 on the benthic macrofauna of the Wadden Sea and the coastal North Sea near the island of Sylt

The development of benthic macrofauna in the Wadden Sea and in the coastal North Sea after the severe winter of 1995/96 is compared with the preceding years with mild to moderate winters. In the intertidal of the Wadden Sea, ice-drift and low temperature caused the expected changes in species composition by increasing winter mortality in sensitive species, and by exceptionally high recruitment of some species during the succeeding summer. In the shallow subtidal (10–20 m depth), similar winter effects were observed. However, recovery of many subtidal populations was still incomplete until the summer of 1997. It is suggested that this was due to hydrographic conditions that carried many larvae or drifting juveniles into more distant offshore areas. This may have limited larval supply and may have delayed recovery at the onshore sites. Since in the eastern North Sea severe winters are accompanied by frequent easterly winds, it is not clear whether decreasing winter abundances in some species were due to increased mortality, or to a seaward dislocation of organisms. Received in revised form: 7 May 2001 Electronic Publication     

Marine molluscs and fish as biomarkers of pollution stress in littoral regions of the Red Sea, Mediterranean Sea and North Sea

The intensive development of industry and urban structures along the seashores of the world, as well as the immense increase in marine transportation and other activities, has resulted in the deposition of thousands of new chemicals and organic compounds, endangering the existence of organisms and ecosystems. The conventional single biomarker methods used in ecological assessment studies cannot provide an adequate base for environmental health assessment, management and sustainability planning. The present study uses a set of novel biochemical, physiological, cytogenetic and morphological methods to characterize the state of health of selected molluscs and fish along the shores of the German North Sea, as well as the Israeli Mediterranean and Red Sea. The methods include measurement of activity of multixenobiotic resistance-mediated transporter (MXRtr) and the system of active transport of organic anions (SATOA) as indicators of antixenobiotic defence; glutathione S-transferase (GST) activity as an indicator of biotransformation of xenobiotics; DNA unwinding as a marker of genotoxicity; micronucleus test for clastogenicity; levels of phagocytosis for immunotoxicity; cholinesterase (ChE) activity and level of catecholamines as indicators of neurotoxicity; permeability of external epithelia to anionic hydrophilic probe, intralysosomal accumulation of cationic amphiphilic probe and activity of non-specific esterases as indicators of cell/tissue viability. Complete histopathological examination was used for diagnostics of environmental pathology. The obtained data show that the activity of the defensive pumps, MXRtr and SATOA in the studied organisms was significantly higher in the surface epithelia of molluscs from a polluted site than that of the same species from control, unpolluted stations, providing clear evidence of response to stress. Enhanced frequency of DNA lesions (alkaline and acidic DNA unwinding) and micronucleus-containing cells was significantly higher in samples from polluted sites in comparison to those from the clean sites that exhibited genotoxic and clastogenic activity of the pollutants. In all the studied molluscs a negative correlation was found between the MXRtr levels of activity and the frequency of micronucleus-containing hemocytes. The expression of this was in accordance with the level of pollution. The complete histopathological examination demonstrates significantly higher frequencies of pathological alterations in organs of animals from polluted sites. A strong negative correlation was found between the frequency of these alterations and MXRtr activity in the same specimens. In addition to these parameters, a decrease in the viability was noted in molluscs from the polluted sites, but ChE activities remained similar at most sites. The methods applied in our study unmasked numerous early cryptic responses and negative alterations of health in populations of marine biota sampled from the polluted sites. This demonstrates that genotoxic, clastogenic and pathogenic xenobiotics are present and act in the studied sites and this knowledge can provide a reliable base for consideration for sustainable development. Received: 2 March 1999 / Received in revised form: 2 August 1999 / Accepted: 3 August 1999     

Influence of pollution on the harpacticoid copepods of two North Sea estuaries

Seasonal monitoring of the meiobenthos in the Dutch estuaries revealed an anomaly in density and diversity of harpacticoid copepods in the Westerschelde.Another Dutch estuary, the Eems Dollard, has comparable hydrodynamical, physical and sedimentological, characteristics and a similar fauna, but even in the severely organically polluted oligohaline mudflats of this estuary, annual average density and diversity of endo-epibenthic harpacticoid communities are higher than at similar less enriched meso- to polyhaline biotopes of the Westerschelde.Besides the concentrations of inorganic pollutants, such as phosphates and nitrates, the concentrations of pesticides, cyanide, detergents, phenoles, oils, polychlorobenzenes, polycyclic aromatic hydrocarbons and heavy metals were compared in both estuaries and compared to suggested permissive levels when available. From this it appears that these pollutants are present in the Westerschelde either in too low concentrations to be considered dangerous or at concentrations comparable to those occurring in the Eems Dollard, except for heavy metals.The load of a.o. Zn, Cu and Pb is distinctly and persistently higher in sediments and suspensions of the Westerschelde than in the Eems Dollard and copper is continuously present in a concentration at which, according to bioassays, egg production and larval development of planktonic copepods are severely affected. The remarkable scarcity of harpacticoid life on nutrient rich mudflats of the Westerschelde is thus probably due to heavy metal pollution.Since no other hardbodied meio- and macrobenthic taxa nor the plankton of this estuary show such a marked impoverishment, benthic harpacticoids prove to be suitable as indicators for the first stages of ecosystem-breakdown in estuarine and coastal zones polluted by trace-metals.     

Evidence for density-dependent growth in North Sea gadoids

The classical view on density-dependent growth is that an increase in population density heightens intraspecific competition for food, leading to reduced food intake and depressed growth. This paper summarizes the analysis of English Groundfish Survey, International Young Fish Survey and ICES Roundfish Working Group data for evidence of density-dependent growth of I- and II-group cod, haddock and whiting in the North Sea.
Within Roundfish areas there was a negative relationship between growth and density in some instances, but it was small. However, there were persistent differences in both growth rate and density between Roundfish areas. In all cases, except for I-group cod, there appeared to be a tendency for fish in low-density areas to be larger than fish in high-density areas; this might be due to feeding competition but other explanations are possible. The results urge caution in interpreting commercial catch data for evidence of density-dependent growth, since changes in fishing effort in particular areas could significantly bias the conclusions.
There was evidence of enhanced survival of small fish in good year-classes; this would depress the mean size at age, thereby mimicking the effects of density-dependent growth.     

The conservation of British North Sea estuaries

Estuaries are amongst the most productive, but scarce, ecosystems in the world. Britain and the southern North Sea have many and extensive estuaries. Many British North Sea coast estuaries are nationally and internationally important for nature conservation for their variety of sand-dune, saltmarsh, sandflat and mudflat habitats, and the plants and animals that they support. The wildlife of estuaries are currently being affected by a great many of man's activities. Many, particularly land-claim, pollution and recreational activities cause loss and damage to this scarce wildlife habitat. Conservation of British estuaries is achieved mainly through site safeguard — the notification of Sites of Special Scientific Interest (SSSIs) and declaration of National Nature Reserves (NNRs) by the Nature Conservancy Council (NCC), and the designation of sites of international importance under the Ramsar Convention and the EC Directive on Wild Birds — and the amelioration of existing damage by sensitive management and pollution control. Several nationwide surveys, particularly NCC's Estuaries Review, are underway in Britain. These will provide sound comparable information on all estuarine sites, and will form the basis for developing an overall estuarine conservation strategy.     

Seasonal variability of inorganic and organic nitrogen in the North Sea

This study considers the cycling of nitrogen in the waters of the North Sea, particularly focussing on organic nitrogen. Dissolved inorganic nitrogen (DIN), dissolved organic nitrogen (DON) and particulate organic nitrogen (PON) were measured in the North Sea over a one-year period (autumn 2004–summer 2005). The surface water concentrations of nitrate, ammonium, DON and PON during the present study ranged from <0.1–7.2 μM, <0.1–2.0 μM, 1.9–11.2 μM and 0.3–5.6 μM, respectively, with DON the dominant fraction of total nitrogen at all times. These nutrients concentrations were significantly lower compared to previous studies in the southern North Sea. The seasonal variations showed high mean surface concentrations of nitrate (4.7 ± 0.6 μM) and DON (8.9 ± 0.9 μM), low ammonium (<0.1 μM) and PON (0.8 ± 0.1 μM) in winter, shifting to low nitrate (0.3 ± 0.3 μM) and DON (4.2 ± 1.2 μM) in summer, with high ammonium (0.8 ± 0.4 μM) in autumn and PON (2.5 ± 1.2 μM) in spring. Highest mean surface DON concentration was measured in winter and may be due to resuspension of the organic matter from the bottom sediments. For autumn and spring, phytoplankton DON release was likely to be the most significant source of DON as shown by high concentrations of low molecular weight (LMW) DON and its positive correlation to chlorophyll a. Low total and LMW DON concentrations during summer were likely to be due to the uptake of the LMW DON fraction by phytoplankton and bacteria and the stratification of the water column. DON is therefore shown to be a potentially important source of nitrogen in shelf seas especially after the spring bloom has depleted nitrate to limiting concentrations. Handling editor: L. Naselli-Flores     

Effects of the large-scale uncontrolled fertilisation process along the continental coastal North Sea

In this paper, effects of eutrophication in selected compartments of the North Sea ecosystem are discussed, encompassing the possibly positive effects of nutrient enrichment. Based on a variety of studies, impacts on biomass of phytoplankton, macrozoobenthos, microzooplankton, shrimps and fishes and productivity are presented. Enhanced nutrient concentrations and loadings can be observed in several coastal areas of the North Sea. As a result, increases in the concentration, production and changes in the species composition was observed in the phytoplankton. In addition, there are some indications for an increased biomass of macrozoobenthos, whereas an increase in microzooplankton can only be assumed from mesocosm experiments. A concomitant increase of higher trophic levels such as shrimps and fishes, as observed in some coastal regions of the North Sea, is difficult to link directly to eutrophication due to a lack of conclusive field observations showing the causality of the changes. That the large fertilisation process in the North Sea has led to a series of changes is, however, without doubt. The answer, to what extent these can be claimed as being harmless, positive or negative from the anthropogenic point of view, is hampered by the lack of good assessment criteria for marine ecosystems and requires a thorough analysis of all compartments involved by means of long-term-series long enough to discriminate between man-made and natural variability.     

Mesocosm experiments: mimicking seasonal developments of microbial variables in North Sea sediments

This study investigated the suitability of mesocosms for studying the seasonal development of microbial variables in the benthic system of the North Sea. Undisturbed sediment cores were taken from two locations in the North Sea, one with sandy sediment (28 m depth) and the other with silty sediment (38 m depth) and installed in mesocosms in January–April 1989. Cores were kept as in situ temperature in the dark until December 1989. One set of sandy and silty sediments was starved and the other set received a supply of organic matter in May–June, simulating the settlement of the spring bloom of Phaeocystis pouchetii. Seasonal developments in bacterial production (methyl ³H-thymidine incorporation), abundance and biomass of bacteria and nanoflagellates and oxygen consumption were compared between the mesocosms and the field in surface sediments every 1.5 to 2.5 months. Effects of seasonal temperature variations (range 6–17.5 °C) on microbial variables in starved mesocosms were limited, which possibly indicates a subordinate role of temperature in microbial processes in North Sea sediments. Organic matter produced a direct response in bacterial production and oxygen consumption in mesocosms. Bacterial and protozoan abundance also increased. The effect of the organic input disappeared within 2 months and values of enhanced variables declined to initial levels. The organic matter enrichment in mesocosms apparently did not provide sufficient energy to keep the microbenthos active at field levels through summer.These results suggest that in the silty sediments in the field, organic matter is available for bacterial production throughout summer. In sandy sediments, the major organic matter input, which sets the seasonal pattern, appears to be in June. Apparently the seasonal development of microbial variables can be mimicked in mesocosms with organic matter supplies. Differences between the field and mesocosms are further illustrated by carbon budgets. Recycling of bacterial biomass was required to meet the bacterial carbon demand in the budget.Publication No. 22 of the project Applied Scientific Research Neth. Inst. for Sea Res. (BEWON).     

Community structure and functional roles of meiofauna in the North Sea

Knowiedge on community structure of North Sea meiofauna has greatly increased recently. A quasisynoptic picture of meiofauna densities and copepod community structure from 171 stations of the southern North Sea, sampled in April–May 1986, has been obtained during the North Sea Benthos Survey. Latitudinal patterns in meiofauna abundance and copepod weight, abundance and diversity exist in an area between 51°30′N and 58°30′N. Using TWINSPAN-classification five major groups of copepod species can be recognized which are related to sediment type, latitude and depth. The part of the meiofauna in total benthic energy flow, their role in the benthic food web and in biogeochemical cycles is discussed based on existing literature. There are still considerable gaps in knowledge and the field is not progressing rapidly. Publication no. 599 Netherlands Institute of Ecology, Centre for Estuarine and Coastal Ecology, Yerseke, The Netherlands.     

Short-term variations in the fluxes and composition of seston in near-bottom traps in the southern North Sea

In this study we attempted to measure the potential food availability for macrobenthic interface feeders using a new device (sediment recorder) which traps bed load particles at a height of 10 cm above the sea floor. The recorder is positioned flat on the seabed and alternately exposes a shallow collector mounted in a smooth surface mimicking the sediment surface. The samples were analyzed for their content of chlorophyll a (Chl) and dry weight (DM). We compared the sediment recorder data with the results obtained with a sediment trap suspended 3.2 m above the sea floor. Measurements were carried out in spring and fall at two stations in the southern North Sea with different sediment type, maximum current velocities and biomass (sand, station B vs silty sand, station F). The recorder deployments in November yielded higher Chl fluxes and a higher quality of particles (Chl:DM ratio) than the trap at both stations. Unlike the trap samples, in which Chl and DM were always positively and strongly correlated, maximum Chl and DM fluxes in the recorder samples were out of phase, particularly at station B, pointing to a highly variable quality during the tidal cycle. Unexpectedly, Chl fluxes and Chl:DM ratios in the recorder samples at station B were higher than at station F in both seasons, although the benthic biomass at the latter station was five times higher. The highest Chl fluxes and particle quality (Chl:DM) at station B were measured during slack tides when interface feeding would be least hampered by drag forces. Station B is nevertheless characterized by a low macrobenthic biomass and a scarcity of interface feeders, specifically of suspension-feeding bivalves. Possible reasons for this apparent discrepancy are discussed. Electronic Publication     

Introduction into the ecosystem of the North Sea: Hydrography,biota, and food web relationships

The North Sea, one of the most productive of the earth's seas and oceans, is also surrounded by some of earth's most densely populated and heavily industrialized regions. A growing number of signals are being received which indicate that this valuable ecosystem is increasingly under stress. This has generated a corresponding increase in concern over the steps to be taken to protect the North Sea. While there are divergent views on what constitutes an ‘ideal’ North Sea, there is a general recognition that any decisions that are made should be based on a good understanding of this ecosystem. The intention of this paper is to give an overview of what is presently known, and to identify areas where more studies are needed. A brief summary of the hydrography and the biota of the North Sea is given. Biotic and abiotic structure justify partitioning the North Sea into three ecologically different regions: southern, central, and northern. For the most part, neither the top predators,e.g. marine birds and mammals, nor the macroalgae and sea grasses are included in this overview.     

Variations in spatial and temporal distribution of Archaea in the North Sea in relation to environmental variables

The spatial and temporal distribution of pelagic Archaea was studied in the southern North Sea by rRNA hybridization, sequencing and quantification of 16S rRNA gene and membrane lipid analyses and related to physical, chemical and biological parameters to determine the factors influencing archaeal biogeography. A clear temporal variability was observed, with marine Crenarchaeota (Group I.1a) being relatively more abundant in winter and Euryarchaeota dominating the archaeal assemblage in spring and summer. Spatial differences in the lateral distribution of Crenarchaeota were also evident. In fact, their abundance was positively correlated with the copy number of the gene encoding the alpha subunit of crenarchaeotal ammonia monooxygenase (amoA) and with concentrations of ammonia, nitrate, nitrite and phosphorus. This suggests that most Crenarchaeota in the North Sea are nitrifiers and that their distribution is determined by nutrient concentrations. However, Crenarchaeota were not abundant when larger phytoplankton (>3 microm) dominated the algal population. It is hypothesized that together with nutrient concentration, phytoplankton biomass and community structure can predict crenarchaeotal abundance in the southern North Sea. Euryarchaeotal abundance was positively correlated with chlorophyll a concentrations, but not with phytoplankton community structure. Whether this is related to the potential of Euryarchaeota to perform aerobic anoxygenic phototrophy remains to be shown, but the conspicuous seasonal distribution pattern of Crenarchaeota and Euryarchaeota suggests that they occupy a different ecological niche.     

Species assemblages of pelagic fish embryos in the southern North Sea between 1984 and 2000

The occurrence and abundance of the pelagic eggs of southern North Sea spring-spawning fish were analysed between 1984 and 2000. Species number varied between six (1986) and 14 (1999) and was positively correlated with sea surface temperature. With one exception, dab eggs were always the most abundant and usually highly dominant. Ranking of species depended on temperature, but no significant differences in ranking between years was discernible. Although with the increase in temperature in the 1990s a change in species assemblage was evident [species belonging to the boreal-Mediterranean (Lusitanian) group became more apparent in the species assemblage] this did not lead to an increased species diversity (Shannon Index) or a change in other community parameters. It appears that the recent developments regarding spawning stock biomass of commercial North Sea fish is reflected in the declining egg abundance of the respective commercial and larger species (i.e. cod, flounder, plaice) and an increase in abundance of the eggs of small species (i.e. long rough dab, rockling) over the years. Received in revised form: 5 November 2001 Electronic Publication     

The houting Coregonus oxyrinchus(L.)(Salmoniformes: Coregonidae), a globally extinct species from the North Sea basin

Coregonus oxyrinchus is redescribed and a neotype is designated (BMNH 1862.11.20.1.). It was restricted to south England and the lower parts of the Rivers Rhine, Meuse and Schelde. It is now globally extinct; the last individual was caught in 1940. Coregonus oxyrinchus is distinguished from other coregonids by having 38–46 gill rakers and a long, pointed snout.     

Native Parasites Adopt Introduced Bivalves of the North Sea

Introduced species may have a competitive advantage over native species due to a lack of predators or pathogens. In the North Sea region, it has been assumed that no metazoan parasites are to be found in marine introduced species. In an attempt to test this assumption, we found native parasites in the introduced bivalves Crassostrea gigas and Ensis americanus with a prevalence of 35% and 80%, respectively, dominated by the trematode Renicola roscovita. When comparing these introduced species with native bivalves from the same localities, Mytilus edulis and Cerastoderma edule, trematode intensity was always lower in the introduced species. These findings have three major implications: (1) introduced bivalves are not free of detrimental parasites which raises the question whether introduced species have an advantage over native species after invasion, (2) introduced bivalves may divert parasite burdens providing a relief for native species and (3) they may affect parasite populations by influencing the fate of infectious stages, ending either in dead end hosts, not being consumed by potential final hosts or by adding new hosts. Future studies should consider these implications to arrive at a better understanding of the interplay between native parasites and introduced hosts.     

Year-to-year variation in larval fish assemblages of the Southern North Sea

In order to test the temporal stability within and the reproducibility of larval fish assemblages between years, the larval fish assemblage at Helgoland Roads, North Sea (NE Atlantic) was quantitatively sampled almost daily from January 2003 to December 2005. The survey resulted in a total of 462 samples containing 50,632 larval fish of at least 42 taxa. In winter the larval fish assemblage was mainly dominated by larvae emerging from demersal eggs. This changed gradually to larvae hatching from pelagic eggs. Larvae from pelagic eggs dominated the ichthyoplankton assemblage in summer. A remarkably stable seasonality in terms of dominance patterns with recurring, season-specific fish assemblages was observed over the 3 years, despite substantial variation in environmental conditions such as a temperature difference of almost 20°C between summer and winter. The lesser sandeel (Ammodytes marinus), was the only species which showed significant fluctuations in abundance between the years. After removal of this species from the analysis, the dominance patterns of the remaining fish species were almost identical between years.