Lead and cadmium contents in selected macrofauna species from the Dogger Bank and the eastern north Sea

Lead and cadmium concentrations were measured in the polychaetesNephtys hombergi andN. caeca, in the sea-urchinEchinocardium cordatum, and in the bivalveVenus striatula obtained from the Dogger Bank and the eastern North Sea. The cadmium concentrations determined in all four species from these areas were relatively equal, except for an increase in concentration found in those species from the northeastern part of the Dogger Bank. A lower lead content was generally observed in the individuals taken from the German Bight than in those from the Dogger Bank, especially from its northeastern part. In the case of lead, it is possible to divide the southern North Sea into three regions according to the different concentration levels by statistical treatment: the less contaminated German Bight, the more contaminated central Dogger Bank and the highly affected northeastern Dogger Bank. The results obtained contradict the prevailing opinion that offshore invertebrate populations are, in comparison to individuals from coastal regions, less affected by contaminants such as heavy metals.     

Macrofauna communities of the Dogger Bank (central North Sea) in the late 1990s: spatial distribution,species composition and trophic structure

Macrofauna samples were taken in May 1996–1998 at 28 stations on the Dogger Bank (central North Sea) to describe species composition, spatial distribution and trophic structure of the macrofaunal communities. In general, five communities could be distinguished depending on the influence of different water masses, depth, type of sediment and food availability. Except for those species which feed in the sediment, macrofauna communities did not show any correlation to the measured sediment variables. In fact, macrofauna communities of the Dogger Bank were mainly structured by lateral food supply via frontal systems. This was indicated by the dominance of interface feeders foraging in the benthic boundary layer, which are able to switch from surface deposit to suspension feeding depending on flow conditions. The importance of sandlickers in the shallowest parts of the Dogger Bank hints at an important contribution of benthic primary production to the nutritional and energetic needs of the benthic community. Electronic Publication     

The Dogger Bank: A special ecological region in the central North Sea

This paper is a review of the present state of knowledge about the ecology of the Dogger Bank, a shallow area in the central southern North Sea. The biological situation in this region is different from other regions in the North Sea; phytoplankton production occurs throughout the year, connected with low periodicity in macrofaunal abundance and condition of fish. Against prevailing opinions, the Dogger Bank, an offshore region, seems to be affected by eutrophication and pollution.     

Distribution patterns of mesozooplankton biomass in the North Sea

During spring 1986 and winter 1987, zooplankton samples were collected over the entire North Sea by means of a multi-closing net-system. Before taxonomic treatment, wet weight estimates and carbon content conversions were carried out. From this data set, 4 962 522 tons zooplankton biomass (dry weight) were estimated for the whole North Sea during the spring survey. High biomasses (more than 100 mg C/m³) were located in areas between the Orkneys and the Shetlands, off the mouth of the Firth of Forth, the Channel and the river Rhine. Considerable zooplankton biomass was also found parallel to the Danish west coast. Furthermore, a narrow tongue of high biomass (partly greater than 200 mg C/m³) intruded from the north, between 1 °E and 4 °E, into the northern North Sea, turning to the east at 56°N, and continuing into deeper water layers to form a left turning “helix” of high biomass in the central part of the North Sea. During the winter survey the carbon content of the zooplankton stock was a factor 10 lower than in summer. Altogether, 519340 tons of zooplankton biomass (dry weight) were estimated in winter. Centres of relatively high biomass were located off the mouth of the rivers Rhine, Weser and Elbe and off the British east coast moving in a cyclic way across the Dogger Bank into the central North Sea. A further maximum of zooplankton abundance was found in the Skagerrak region. However, an intrusion of zooplankton from the shelf edge into the North Sea was not observed in winter. A qualitative analysis of species composition showed that small copepods dominated the zooplankton in the southern and eastern North Sea. The “eddy” of high biomass in the northern North Sea observed in spring, however, was mostly shaped by the large copepodCalanus finmarchicus (70–90%). The distribution of zooplankton biomass in the North Sea is discussed in relation to the hydrographic conditions and to the biology of the dominant species.     

Benthic community structures in the North Sea

Coherent assemblages of marine benthic species have been recognized from the early twentieth century, and the classical papers of Petersen (1914, 1918) were based on studies of limited areas in the North Sea. In 1986, a synoptic survey of the North Sea north to 57°N was undertaken by a group of ten laboratories from seven North Sea countries. The results of this survey have recently been published (Heip et al., 1992a, b; Künitzer et al., 1992; Huys et al., 1992), and some of the results are summarized in this paper. The analysis of the macrofauna is based on slightly more than 700 taxa. In general, the North Sea macrofauna consists of northern species extending south to the northern margins of the Dogger Bank, and southern species extending north to the 100 m depth line. The central North Sea is an area of overlap of southern and northern species, especially around the 70 m depth contour. Consistent groupings of species are recognized that were summarized in seven faunal groupings. Macrofaunal body weight, density and diversity increase linearly towards the north. Macrofaunal biomass for the whole area averages 7 g adwt. m⁻² and decreases from south to north. Distribution patterns and trends within the meiofauna were very different. Nematodes, which are the dominant taxon overall, are least abundant in the sandy sediments of the Southern Bight, then increase to a maximum around 53° 30′ N and slowly decrease again towards the north. Copepod density and diversity are highest in the Southern Bight, due to the presence of many interstitial species. A large number of species new to science were recorded by the North Sea Benthos Survey and about 1500 species are expected to occur. Copepods show very distinct assemblages according to water depth and sediment type. The contrasting patterns in latitudinal gradients of body weight and number of species of macro- and meiofauna can be only partially explained. Latitude and sediment characteristics, such as grain size and content in plant pigments, and water depth, determine part of the variance in species composition, density and biomass of the benthic fauna, but the patterns that are observed are different for different benthic groups, requiring careful consideration as to their use in biological monitoring procedures. Distributions are related to current patterns in the North Sea, annual temperature variations and availability of food. However, large parts of the variance in many parameters remain unexplained.     

Mapping the spawning grounds of North Sea cod (Gadus morhua) by direct and indirect means

Despite recent evidence for sub-stock structuring, North Sea cod are assessed as a single unit. As a consequence, knowledge of sub-stock trends is poor. In particular, there are no recent evaluations of which spawning grounds are active. Here we report results from the first ichthyoplankton survey to cover the whole North Sea. Also, this survey, conducted in 2004, was the first to make extensive use of DNA-based molecular methods to unambiguously identify early developmental stage cod eggs. We compare the findings from the plankton survey with estimated egg production inferred from the distribution of mature cod in contemporaneous trawl surveys. Results from both approaches were in general agreement and showed hot spots of egg production around the southern and eastern edges of the Dogger Bank, in the German Bight, the Moray Firth and to the east of the Shetlands. These areas broadly coincide with known spawning locations from the period 1940 to 1970. We were, however, unable to directly detect significant numbers of cod eggs at the historic spawning ground off Flamborough (northeast coast of England). The results demonstrate that most of the major spawning grounds of cod in the North Sea are still active but that some localized populations may have been reduced to the point where it is now difficult to detect the presence of eggs in the plankton.     

Environmental influence on the bottom and near-bottom megafauna communities of the Dogger Bank: a long-term survey

This paper deals with climate-driven changes of the species composition of the bottom and near-bottom megafauna of the Dogger Bank (central North Sea), which was sampled each summer with a 2-m beam trawl on a yearly basis since 1991. The station grid consists of 37 stations, covering an area of approximately 17.000?km2. A selection of commoner species is analysed and correlated with temperature data gained during the research period. Temperatures are derived from our own measurements, combined with CTD data from the International Council for the Exploration of the Sea. The results show a decrease in biodiversity and a clear regime shift around the beginning of the 21st century, combined with rising mean bottom temperatures. In addition, details are given about the Dogger Bank hydrography and the climate sensitivity and abundance of the main species caught with the beam trawl. Our long-term study reveals the changes in the community structure of the megafauna of the Dogger Bank over a period of almost two decades. It suggests a link between changes in species composition/abundance and changes in the environment, especially the marine climate.     

Assessment of some innate immune responses in dab (Limanda limanda L.) from the North Sea as part of an integrated biological effects monitoring

The marine flatfish dab (Limanda limanda), which lives in direct contact with contaminated sediments, is frequently used as a sentinel species in international monitoring programmes on the biological effects of contaminants. In this study, immune responses were recorded as indicators of sublethal chronic effects of contaminants, in addition to measurement of the induction of mono-oxygenase ethoxyresorufin O-deethylase (EROD) in liver cells, the inhibition of acetylcholin esterase (AChE) in muscle and a quantification of grossly visible diseases and parasites. In total, 336 dab were analysed from five sampling areas in the North Sea, including the German Bight, the Dogger Bank, the Firth of Forth, and two locations close to oil and gas platforms (Ekofisk and Danfield). When considering plasma lysozyme levels, pinocytosis and respiratory burst activity of head kidney leucocytes, a clear gradient could be observed with decreased levels in individuals collected from the Firth of Forth and locations near the oil or gas platforms compared with dab from the Dogger Bank or the German Bight. Individuals with induced EROD activity displayed reduced lysozyme and respiratory burst activities. Lysozyme levels were also reduced in dab with lymphocystis or with nematodes. The data obtained indicate that the assessment of innate immune parameters in a monitoring programme provides supplementary information about immunomodulatory effects associated with the exposure of fish to contaminants. In particular, concentrations of plasma lysozyme, which can be analysed in an easy and inexpensive assay, are considered to be an appropriate parameter for use in a battery of other bioindicators. Communicated by H. von Westernhagen and A. Diamant     

The megaepifauna of the Dogger Bank (North Sea): species composition and faunal characteristics 1991–2008

During a long-term study in the summer months of the years 1991–2008, 176 megaepifaunal species were recorded through a series of beam trawl surveys on a grid of fixed stations on the Dogger Bank (central North Sea). This paper gives a qualitative overview on species composition throughout the research period, determined from samples collected during 15 cruises. In recent years, a number of species with more oceanic distribution patterns (e.g. species from SW British coasts) has been collected. In spite of these newcomers, there was a slight decrease in total species numbers during the research period.     

SPI-ing on the seafloor: characterising benthic systems with traditional and in situ observations

This work aimed to show that the sea bed of two environmentally-different regions of the North Sea varies both spatially and temporally with respect to their biological communities and bioturbation characteristics. The two contrasting sites studied were north of the Dogger Bank (ND) (85 m) and the Oyster Grounds (OG) (45 m). The physical environment varied between and within sites, mainly influenced by sediment chlorophyll a content and water temperature. Our data revealed that the depth of the apparent Redox Potential Discontinuity (aRPD) layer at OG varied between 2.2 cm in February and 6.5 cm in October; evidence of bioturbation activity (e.g., feeding voids) was observed within the sediment profile. In contrast, at the ND site the aRPD values ranged from 1.7 cm in February to 2.5 cm in May and feeding voids and infaunal burrows were restricted to sediment depths far shallower than those observed at OG. Communities at ND were dominated by a number of surficial-sediment dwelling polychaete species (e.g., Notomastus latericeus, capitellids) while those of OG were dominated by the brittlestar Amphiura filiformis, together with significant numbers of deeper-dwelling taxa such as the ghost shrimp Callianassa subterranea and the bivalve mollusc Corbula gibba. Our data imply that regions of the North Sea which experience dissimilar environmental conditions not only possess different infaunal communities but also contrasting seasonal fluctuations and bioturbation capacities. The ecological implications of these findings, including inferences for carbon and nutrient cycling, are discussed in relation to the wider North Sea ecosystem.     

Spatial and temporal variation in mercury contamination of seabirds in the North Sea

Once the moult patterns have been taken into account, feather methylmercury levels can be used to accurately measure the mercury burdens of seabirds. We used body feathers from live seabirds and from museum collections to examine geographical and temporal patterns of mercury contamination in the North Sea. This approach identifies an increase in mercury concentrations in seabirds of the German North Sea coast during the last 100 years, especially high levels during the 1940s, and reduced contamination in the last few years. Comparisons among populations suggest that some increases in mercury levels are predominantly due to local pollution inputs, as on the German coast, while in other areas deposition from jet stream circulation of global contamination may be the major contributor. Mercury levels are far higher in seabirds from the German North Sea coast than in populations from the north and west North Sea or from most areas of the North Atlantic. We advocate the use of museum collections of birds for studies of long-term changes in levels of mercury contamination.     

Epidemiology of pseudobranchial tumours in Atlantic cod(Gadus morhua) from the North Sea and the Baltic Sea

Investigations during several cruises (1979–1981) showed that Atlantic cod with pseudobranchial tumours occurred in all areas investigated. Statistical analysis of data revealed that there was a marked prevalence of the condition in the centre of the German Bight, whereas a lower incidence of the condition was observed on the Dogger Bank, off the Dutch coast and in the western Baltic Sea. The higher prevalence of pseudobranchial tumours in the German Bight coincides with a dense population of cod and impacts of pollution.     

High abundance of small zoobenthos around biogenic structures in tidal sediments of the Wadden Sea

On the tidal flats of the island of Sylt (eastern part of the North Sea) the quantity of micro- and meiofauna associated with shoots of seagrass(Zostera noltii), with infaunal bivalves(Macoma balthica), and with tubes and burrows of polychaetes(Pygospio elegans, Pectinaria koreni, Nereis diversicolor, Nereis virens, Arenicola marina) was found to add up to 5 to 33 % of the overall abundance. These structures, taken together, account for 10 to 50 % of the faunal abundance on an average tidal flat at Sylt. The quantitative effect of biogenic structures at the sediment surface (casts and funnels) is small compared to that of tubes and burrows penetrating the anaerobic subsurface layer. In providing stable oxic microenvironments these elite structures frequently bring together more individuals than occur in the entire reducing sediment below surface. Faunal composition of irrigated dwellings of large infauna is different from that of the oxic surface sediment. The common denominator of all elite structures of the subsurface is an oxic halo. Burrows without such a halo are unattractive. There is no evidence that owners of burrows prey on their smaller inmates.     

Seagrass recovery in the Northern Wadden Sea?

Aerial surveys on seagrass (Zostera spp.) indicate a three to fourfold increase in bed area from 1994 to 2006 with up to 100 km² or 11% of intertidal flats in the Northfrisian Wadden Sea (coastal eastern North Sea), observed at seasonal maximum in August when flying during low tide exposure 300 to 500 m above ground. When viewed from the air, difficulties in distinguishing between seagrass and green algae and a lack of contrast on dark-coloured mudflats are sources of error in areal estimates. Particularly the positioning of beds remote from shores was imprecise. However, the consistency in method over time gives confidence to the inferred positive trend which is opposite to the global pattern. Both, the spatial pattern and a recent decrease in storminess suggest that sediment stability is the key factor for seagrass dynamics in this tidal area. On exposed sand flats, high sediment mobility may be limiting and along the sheltered mainland shore land claim activities with high accretion rates may cause a scarcity of seagrass. The potential area of seagrass beds may be twice as large as the realized maximum in 2006 but eventually the rising sea level will reverse the observed seagrass expansion.     

Regional patterns in prevalence of principal external diseases of dab Limanda limanda in the North Sea and adjacent areas 1992-1997

The prevalence and spatial distribution of major diseases of dab Limanda limanda in the North Sea and adjacent areas were studied in the summers 1992 to 1997. Areas covered were the North Sea, Irish Sea, northern and northeastern British Waters and the English Channel. The diseases studied were lymphocystis, epidermal hyperplasia/papilloma and skin ulceration. To standardise data, results were analysed for females >15 cm (>3 yr old). Data were subjected to median polish, and additive, extended and additive plus multiplicative models were applied to best account for effects of region and year. Annual differences in disease prevalence were low whilst differences between areas were pronounced. For lymphocystis higher prevalence was observed in the northwestern sector of the North Sea, at the northern tip of Scotland and in an area south of Iceland. Prevalence was low in the Irish Sea, the English Channel and the southern North Sea, and intermediate in the German Bight. For epidermal hyperplasia/papilloma, levels were low at Icelandic stations, in the northern Irish Sea, in the southern North Sea and the English Channel, whilst levels were high in the northwestern part of the North Sea and the German Bight. Elevated levels of skin ulceration were found on the Dogger, at 1 station in the Irish Sea (off Sellafield) and at 1 station to the south of Iceland. Lower levels were detected west of Iceland. Prevalence in all other areas was intermediate. It is concluded that a detailed analysis of available data on disease prevalence and putative causative factors is desirable and, given the good availability of data, would be a promising step forward toward elucidating possible cause and effect relationships between diseases and anthropogenic factors.     

Review on the effects of pollution on marine fish life and fisheries in the North Sea

Information on possible effects of pollution on fish life is reviewed with special emphasis to the North Sea. Concentrations of heavy metals are high in onshore waters, sediments and organisms. Offshore regions are considered to be not endangered. In estuaries concentrations of certain heavy metals are often surpassing thresholds, which have been experimentally demonstrated to produce sublethal effects on marine organisms. Also mussel culture areas had to be closed due to excess of cadmium contamination. Mercury in fillets of cod in the southern North Sea sometimes reaches 0.3 ppm representing 1/3 of the legal limit. Reductions of inputs are necessary due to the bioaccumulative and toxic potential of heavy metals. Organochlorine contamination of water, sediments and organisms is higher in the southern than in the northern North Sea. In offshore areas clearly elevated levels of PCB's, which are the dominating organochlorine compound in marine organisms, have been analyzed. At present only a minor fraction of total organochlorine residues accumulated in marine biota is analyzed. Margins between experimentally derived lethal levels and those occurring in marine biota are narrow suggesting that sublethal effects might occur. One of the special concerns in the context of organochlorine contamination has to be seen in the potential of these substances affecting the reproductive capacity of marine organisms. Reduction measures are urgently necessary and should include substances with bioaccumulative and toxicological potential, which are not presently included in Marine Monitoring Programmes. Impact of oil pollution is considered to be a regional problem, although especially in estuaries concentrations in the water column might be high. Input through oil platforms has led to local reductions in benthic communities. Measures have to be taken, to further reduce these inputs also in view to avoid contamination of fish in the vicinity of oil fields. Oxygen depletion in German and Danish coastal waters has been shown to be a severe degradation in coastal water quality resulting in significant impact on fish populations. Indications for large-scale avoidance reactions of low DO-areas and mortalities of fish and benthos organisms have been demonstrated. It is suspected that the low DO-situations are related to increased nutrients input and high phytoplankton production. Improvement of the situation can only be expected, if input of nutrients is reduced. Reasons for the occurrence of toxic plankton blooms are not understood. An intensified research is necessary, to avoid damage to fisheries and mariculture. Although activity of radionucleids in North Sea waters is increased due to input of radioactive material from La Hague and Sellafield reprocessing units, the present levels of activity are not considered to be a threat for marine life. The same holds for thermal pollution, which is, depending on the recipient area, expected to have only local effects. Sewage sludge dumping and discharge of untreated sewage introduce significant amounts of accompanying heavy metals, which can be expected to have a negative influence on the total metal budget of recipient waters. Nutrient input through sewage will exhibit only local effects. Dumping of wastes from titanium dioxide industry are suspected to be causally linked to the occurrence of increased prevalences of certain diseases of dab in the German Bight. Present management strategies should take account of the fact that through dumping of waste from titanium dioxide production significant quantities of heavy metal are introduced into the southern North Sea. Also the incineration of chemical wastes at sea should be reduced due to uncertainties concerning quantities and toxicity of incineration products and their potential impact on marine life. It is concluded that clear evidence exists for pollution impact in estuarine regions. Most of the major estuaries at the North Sea coast show benthos reductions, which can be related to water quality and it has to be assumed that areas of a sublethal effect are located in the vicinity of those areas of well demonstrated effects. They are of unknown size and attempts have to be undertaken to quantify areas of sublethal impact. The occurrence of fish diseases is interpreted as an expression of degraded water quality. Some of the diseases detected can produce mortality, thereby having an as yet unquantified impact on fish populations. The presence of fish diseases also reduces the marketability of catches. It cannot be excluded that changes in species composition of fish populations in German Wadden Sea areas are related to pollution. Concentrations of organochlorine substances in fishes of the southern North Sea give rise to the concern that their reproductive capability might be impaired. Future work has to be directed to this problem. Impact of pollution on populations of marine organisms has so far not been demonstrated.     

Cold winter effects on benthic macrofauna communities in near- and offshore regions of the North Sea

In order to investigate the effect of severe winters on the long-term variability of benthic macrofauna communities four stations from the inner German Bight towards the Dogger Bank (North Sea) were sampled in March 1990 and from May 1995 to 2002. The stations were chosen to reflect a gradient in the hydrographic regime, temperature and organic matter supply. We initially hypothesized that the effect of severe winters on benthic communities will be more pronounced in the offshore more stable environment than in the nearshore regions. Our results clearly showed the opposite pattern. The benthic communities at the nearshore stations in the German Bight changed dramatically in species abundance and community structure after the severe winter 1995/1996. But the community structure in the German Bight returned to the previous stage within 2 years. In contrast, the offshore stations in the Oyster Ground and at the Dogger Bank remained rather unaffected by the severe winter and changed gradually during the study period. The results are discussed concerning the ecological importance of severe winters for benthic communities and the utility of our results for interpreting long-term changes.     

The potential effect of sustained hypoxia on nitrogen cycling in sediment from the southern North Sea: a mesocosm experiment

The potential effect of sustained hypoxia (up to 70 days) on the production of N₂ gas through denitrification and anammox, as well as sediment–water exchange of nitrite, nitrate and ammonia, oxygen consumption and penetration, were measured in mesocosms using sediment collected from the southern North Sea (north of Dogger Bank). As expected, both the penetration of oxygen into, and consumption of oxygen by, the sediment decreased by 42 and 46 %, respectively, once hypoxia was established. Importantly, the oxygen regime did not change significantly (P > 0.05) during the experiment, suggesting that organic carbon was not depleted. During the first 10 days, the exchange of NO₃ ^?, NO₂ ^? and NH₄ ⁺ between the sediment and water was erratic but once a steady state was established the sediment acted as either a sink for fixed nitrogen under hypoxia or as a source in the controls. Over the course of the mesocosm experiment the rate of both anammox and denitrification increased, with anammox increasing disproportionately under hypoxia relative to the controls, whereas the rate of increase in denitrification was the same for both. Under sustained hypoxia the production of N₂ gas increased by 72 % relative to the controls, with this increase in N₂ production remaining constant regardless of the duration of hypoxia. Longer periods of stratification and oxygen depletion are predicted to occur more regularly in the bottom waters of shallow coastal seas as one manifestation of climate change. Under sustained hypoxia the potential for nitrogen removal by the production of N₂ gas in this region of the southern North Sea was estimated to increase from 2.1 kt N 150 days^?1 to 3.6 kt 150 days^?1, while the efflux of dissolved inorganic nitrogen ceased altogether; both of which could down regulate the productivity of this region as a whole.     

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.     

Stomach contents of the Antarctic Silverfish Pleuragramma antarcticum from the southern and eastern Weddell Sea (Antarctica)

Summary The stomach contents of 142 Pleuragramma antarcticum from the southernmost part of the Weddell Sea (Gould Bay: 77°19S) and from the eastern coast (72°–74°S) were analyzed. The fish were collected in February 1983 and stored frozen. Size range of the investigated specimens was from 6.4 cm SL (7.3 cm TL) to 21.9 cm SL (24.1 cm TL). The fish were caught by bottom trawl (Gould Bay) and Agassiz Trawl and pelagic RMT-m net in the upper 300 m layer at the eastern coast, respectively. Thirty-six stomachs were empty or contained sand only. The most important prey in terms of biomass were euphausiids, which occurred in 49 stomachs at a mean number of 2. Their dry weight was estimated to be 15–50 times that of the next important food items, which were copepods, gastropods and gammarids. Krill (Euphausia superba) constituted up to 14% of the euphausiids in the diet of fish caught in the eastern Weddell Sea. The rest was usually made up by Euphausia crystallorophias. This species was taken with preference even when the abundance of E. superba in accompanying plankton catches was 16 times higher. In the Gould Bay, gastropods were frequently eaten despite high numbers of copepods in the plankton, whereas in the eastern Weddell Sea, copepods were abundant both in plankton and in the stomach content. The almost complete absence of the early stages of fish in the diet of Pleuragramma antarcticum in the southern and eastern Weddell Sea is due to a pronounced vertical segregation of fish of different sizes. This distribution pattern is thought to be an adaptive mechanism to avoid intraspecific predation, as 90% of the ichthyoplankton in that area is Pleuragramma antarcticum.