Bioavailability of lead in North Sea sediments

As part of an interdisciplinary research programme, lead contents were measured in the polychaeteNephtys spp. and in the sea-urchinEchinocardium cordatum as well as in the respective sediment fractions <20 μm taken from the Dogger Bank proper and the eastern coastal North Sea. A lower lead content was generally observed in the organisms taken from the German Bight than in those from the Dogger Bank, especially from its northeastern part. It is possible to divide both areas according to the slope found in the linear regression of lead versus total organic carbon contents in sediments, which is twice as steep for the Dogger Bank as for the eastern North Sea. This criterium points to a difference in sediment quality with regard to toxic metal contamination. The sediment quality of the Dogger Bank seems to be twice as bad compared with that of the eastern North Sea. This is in good agreement with the differences found in lead contamination of the sediment-dwelling polychaetes from both areas. The results indicate that lead is primarily accumulated by food ingestion.     

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     

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 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.     

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.     

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.     

Spatial variability of epifaunal communities in the North Sea in relation to sampling effort

Benthic epifauna was sampled in six areas from the German Bight towards the Norwegian Sea using a 2-m standard beam trawl. Nine replicates were taken in each area and year from 1999 to 2006. This data set (60–67 replicates per area) was used to describe the spatial variability in local species composition and to assess the effect of increasing sampling effort on species richness and community structure. Our results confirmed the importance of the 50-m depth contour for the separation of benthic fauna in the North Sea. Low species richness, sparse sessile fauna and high abundances of scavenging species such as Asterias rubens, Liocarcinus holsatus, Astropecten irregularis and Ophiura albida were characteristic of the low-water area south of the 50-m contour. Differences in community structure were less conspicuous in deeper waters north of the 50-m contour, but distribution patterns and abundances of single species such as Echinus elegans, Hyalinoecia tubicola, Ophiothrix fragilis, Scaphander lignarius as well as several hermit crabs resulted in well-defined epifaunal communities. One replicate caught 17–28% of the species found in 60–67 samples and was sufficient to separate the community in the German Bight from those in the central and northern North Sea by using multidimensional scaling. Nine replicates sampled a proportion of 53–60% and provided additional information on the spatial variability of community structure in the central and northern North Sea. Our study indicates that appropriate replication enhances the quality of the data and can partly overcome the constraints of sampling with a 2-m beam trawl. This might be helpful for future monitoring programmes.     

Effects of low salinity on settlement and strobilation of scyphozoa (Cnidaria): Is the lion’s mane <Emphasis Type="Italic">Cyanea capillata</Emphasis> (L.) able to reproduce in the brackish Baltic Sea?

Several species of scyphozoan medusae occur in river estuaries and other brackish waters but it is often unknown if the planulae settle and the scyphopolyps reproduce in those low-salinity waters. In the present study, scyphozoan species from the German Bight (North Sea) were tested in laboratory experiments to investigate their tolerance of low salinity. Planula larvae released from medusae in salinity 32 were still active after the salinity was reduced to 10 (Cyanea capillata, Cyanea lamarckii) and to 7 (Chrysaora hysoscella) in laboratory treatments. Planulae did not settle on the undersides of floating substrates when salinity was reduced to <20. By contrast, planulae released from C. capillata medusae in Kiel Bight (western Baltic Sea) in salinity 15 developed into polyps in laboratory cultures. Polyps reared from planulae in salinity 36 survived a reduction to 12 (C. capillata, C. lamarckii) and to 8 (Aurelia aurita). Polyps of all tested species strobilated and released young medusae (ephyrae) in salinity 12. These results show a high tolerance of planulae and polyps to low salinity, indicating their possible occurrence in estuaries and brackish waters. In addition to laboratory observations, young C. capillata ephyrae were collected in the western Baltic Sea (Kiel Bight) in salinity 15, which indicates that they were probably released by a local polyp population. We suggest that the polyps of the painfully stinging lion’s mane, C. capillata, may be more widespread in the Baltic Sea than previously assumed and that the occurrence of the medusae may not only depend on inflow of water masses from the North Sea.     

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.     

Mass occurrence of an introduced crustacean (Caprella cf. mutica) in the south-eastern North Sea

The caprellid amphipod Caprella mutica is indigenous to coastal waters of north-east Asia and was first recorded in European waters in 1995. We now detected mass occurrences of > 3000 individuals per m² in harbours of the two islands of Sylt and Helgoland in the German Bight, North Sea. Currently, Caprella mutica seems to be restricted to artificial hard substrata but we expect it to become a new species in natural hard bottom assemblages of that region as well.     

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.     

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     

Different zooplankton structures in the German Bight

In August 1982, a net of 48 stations with altogether 208 samples was investigated in the eastern German Bight with respect to temperature, salinity, as well as the amount and species composition of the mesozooplankton (>80 μm). The data were arranged into different structures by means of a cluster analysis. Four different clusters were found: (a) a “Wadden sea water” with few holoplankton organisms but a higher amount of spionid larvae; (b) a “German Bight water” with a maximum occurrence of turbellaria (Alaurina composita) and medium concentrations of copepods; (c) a mixing area between these two water masses with highest amounts ofOikopleura dioica, Temora longicornis, Acartia sp., mussel larvae and larvae of the spionid worms; (d) a “North Sea water” mass with highest concentrations ofPseudocalanus elongatus, Paracalanus parvus undOithona similis. The differences in the concentrations of the species mentioned between the four clusters were significant on the 0.1%-level.     

Gymnodinium catenatum Graham (Dinophyceae)in Europe: a growing problem?

The microreticulate resting cyst of the potentially toxic, chain-forming,unarmoured neritic dinoflagellate Gymnodinium catenalum Graham1943. the planktonic stage of which is not known from NorthEuropean waters, is reported for the first time from recentGerman coastal sediments of the North Sea and Baltic Sea. Insandy mud sediments of the German Bight, a maximum of 8 5 livingcysts cm^–3 were found. In Kiel Bight sediments G.catenalumwas found in maximum concentrations of 17.0 living cysts cm^–3.In surface waters of the German Bight resuspended G catenatumcysts were observed at concentrations of up to 3.6 cysts l^–1.Successful germination experiments conducted with natural seawatershow that the occurrence of a vegetative form of G.catenatumin northern Europe is very likely. The present study highlightsthat cyst surveys provide an important tool for the evaluationof areas with potential toxicity problems, as they may indicatethe presence of hitherto overlooked species in the water column.     

Predation on barnacles of intertidal and subtidal mussel beds in the Wadden Sea

Balanids are the numerically dominant epibionts on mussel beds in the Wadden Sea. Near the island of Sylt (German Bight, North Sea), Semibalanus balanoides dominated intertidally and Balanus crenatus subtidally. Field experiments were conducted to test the effects of predation on the density of barnacle recruits. Subtidally, predator exclusion resulted in significantly increased abundances of B. crenatus, while predator exclusion had no significant effects on the density of S. balanoides intertidally. It is suggested that recruitment of B. crenatus to subtidal mussel beds is strongly affected by adult shore crabs (Carcinus maenas) and juvenile starfish (Asterias rubens), whereas recruits of S. balanoides in the intertidal zone are mainly influenced by grazing and bulldozing of the very abundant periwinkle Littorina littorea, which is rare subtidally. Thus, not only do the barnacle species differ between intertidal and subtidal mussel beds, but the biotic control factors do so as well. Electronic Publication     

The recent arrival of the oceanic isopodIdotea metallica Bosc off Helgoland (German Bight, North Sea): an indication of a warming trend in the North Sea?

In 1988 a long-term study was started of the isopod fauna associated with surface drift material off Helgoland (German Bight, North Sea). In the summer of 1994 specimens ofIdotea metallica Bosc were recorded for the first time. There is no evidence that this species has ever been present in the German Bight before. The samples contained males, both gravid and non-gravid females, and juveniles, indicating that the species reproduced successfully in the Helgoland region. Interbreeding of specimens from Helgoland and the western Mediterranean produced fertile off-spring. As a neustonic species,I. metallica shows a high natural capacity for dispersal. It thus seems unlikely that the arrival of the species in the North Sea resulted from an accidental introduction by man. We are probably witnessing an extension of the species’ geographical range by natural means of dispersal, as a response to recent changes in the ecological conditions of the German Bight. Temperature data measured by the Biologische Anstalt Helgoland since 1962 show that the last decade (except 1996) was characterized by unusually mild winters. Following the severe winter of 1996,I. metallica was again absent from the Helgoland region. After the subsequent mild winters (1997 and 1998), however, the species reappeared in the summer of 1998 with higher numbers than ever before. This suggests that the observed phenomena are closely connected with the recent temperature anomalies.I. metallica can be regarded as a potential immigrant to a warmer North Sea, and may be useful as a sensitive indicator of the predicted long-term warming trend.     

Species-specific PCR discrimination of species of the calanoid copepod Pseudocalanus, P. acuspes and P. elongatus, in the Baltic and North Seas

Two species of the calanoid copepod Pseudocalanus, P. acuspes and P. elongatus, have been reported from the Baltic and North Seas. Morphometric analysis based on Frost (Can J Zool 67:525–551, 1989) did not identify the species in samples collected during the German GLOBEC program; the majority of measurements fell outside reported ranges for Baltic and North Sea specimens. A multiplexed species-specific PCR (SS-PCR) protocol was designed from a ~700 bp sequence of mitochondrial cytochrome oxidase I (mtCOI), with primers designed to amplify regions of different sizes for the two species to allow detection by gel electrophoresis. The accuracy and reliability of the multiplexed SS-PCR reaction was confirmed by sequencing the mtCOI region. Preliminary SS-PCR analysis of geographic distributions of the two species indicated that P. acuspes occurred exclusively at sampling sites in the Baltic Sea, whereas P. elongatus was found in the southern North Sea and at one site in the Baltic. The SS-PCR protocol can be used for routine identification of the two species for studies of population, community and ecosystem dynamics.     

Diseases in North Sea fishes

Prior to the studies reviewed here, only lymphocystis and skeletal deformities of a variety of fish species and certain diseases of eel were known to occur in the German Bight (North Sea). From 1977 until now, 9 externally visible lesions on North Sea fishes were observed; in addition to those mentioned before, they comprise: fin rot, ulcerations, epidermal papilloma, hyperplasia, pseudobranchial tumour, eye diseases and gill swellings. With the exception of information on changes in frequencies of vertebral deformities of herring from the 1950's to the 1970's, there are no long-term data characterizing changes in frequencies of the diseases under study. For pseudobranchial tumours of cod and epidermal papilloma of dab, information is provided on occurrence and abundance. The distribution pattern of cod afflicted with pseudobranchial tumours is strongly influenced by the migratory behaviour of the fish. Epidermal papillomas of dab were more frequently found at stations within the inner German Bight than in neighbouring areas. The Bight is used for dumping of wastes from titaniumdioxide production. Further disease hot spots are areas off the Humber estuary and the British coast. Analysis of chromium in dab from the German Bight revealed elevated concentrations in epidermal tissues of specimens from the dumping area compared with that found in dab from neighbouring localities. Particulate iron was demonstrated to occur in mucous cells of dab from the dumping area. From increased levels of heavy metals with cancerogenic potential in sensitive target tissues and from increased prevalences of diseased fish in the dumping area it is concluded that these phenomena are possibly causally linked. In the vicinity of the Humber estuary high disease rates were encountered and areas with high prevalences of dab afflicted with epidermal papilloma extended over regions shown to be transport routes for persistent pollutants such as radioactive materials. It is therefore suggested that the long-range distribution of fish diseases in the southern North Sea might reflect the long-range transport of persistent pollutants.     

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.