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.     

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

Atlantic cod bearing pseudobranchial tumours were investigated from both North Sea and Baltic Sea areas. Light and electron microscopic studies revealed a tumorous structure previously described in cod from Atlantic and Pacific waters. Additionally, cysts, multinucleated cells and early stages of mitosis are described. These findings support the hypothesis that pseudobranchial tumours are instigated by a protozoan infection causing xenotumours associated with the pseudobranch organ. The cells contained in these lesions showed similarities to amoebae.     

The frequency, distribution, and pathology of three diseases of demersal fishes in the Bering Sea

During September and October, 1975, bottom-dwelling fish from the Bering Sea were examined for diseases. The three most commonly observed diseases and their frequency of occurrence were epidermal papillomas of rock sole ( Lepidopsetta bilineata Ayres), 1-0%; lymphocystis of yellowfln sole ( Limanda aspera Pallas), 2-1 %; and epidermal tumours of the pseudobranch of adult Pacific cod ( Gadus macrocephalus Tilesius), 7-4%. Both the appearance and the histological characteristics of epidermal papillomas on rock sole resembled those found on several species of flatfish in Puget Sound. The frequency of tumour-bearing rock sole ranged from 0-23%, with the highest occurrence at sampling stations around the periphery of the Bering Sea in water depths of 20-30 fathoms (36-54 m). The virus-caused lymphocystis lesions were usually on the 'blind' side fins and skin surfaces of yellowfin sole, the largest lesions being found on the operculum, and the most common site of infection was the pectoral fin. The frequency of lymphocystis was 0-15 %, with a gradual increase of prevalence from the northern to southern sampling stations. The Pacific cod tumours were always bilateral, and contained remnants of normal-appearing pseudobranchial tissue on the surface or deep inside the tumours. Portions of some tumours were necrotic and liquefied. Tumour-bearing cod were most commonly found in the south and south-eastern Bering Sea and the frequency of affected cod ranged from 0-100%.     

Seasonal maturity development of Baltic cod in different spawning areas: importance of the Arkona Sea for the summer spawning stock

We investigated the seasonal maturity development of cod in four areas of the Baltic Sea. Two different spawning peaks were identified and found to be consistent over the period 1992–2005. In the Kiel Bight and Mecklenburg Bight (ICES SD 22) a spawning peak was observed from March to April (spring spawning). In the areas of the Arkona Sea (ICES SD 24) and Bornholm Sea (ICES SD 25) the spawning peak occurred during summer. In the Bornholm Sea, the main spawning activities began in June and ended in September, with a spawning peak in June–August (summer spawning). In the Arkona Sea, which is a transition area between the Mecklenburg Bight and the central Baltic Sea, spawning began in March and lasted until July, with a spawning peak in June–July (summer spawning). Seasonal maturity development and proportions of spawning cod in June in the Arkona Sea were similar to that of the Bornholm Sea. In addition, the proportion of spawning cod in the Arkona Sea was positively correlated with the size of the spawning stock in the Bornholm Sea. Our results provide evidence of a spatial expansion of spawning activities of the summer spawning stock from the eastern Baltic Sea into the Arkona Sea. Therefore, the Arkona Sea should be considered as one of the spawning habitats of the summer spawning stock of Baltic cod.     

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.     

Long-term variability of winter nitrate concentrations in the Northern Wadden Sea driven by freshwater discharge,decreasing riverine loads and denitrification

The hypothesis of a recent reversal in the eutrophication of the Wadden Sea and the potential of inshore waters in denitrification is explored. Salinity, temperature and nitrate concentrations in the List Tidal Basin (Northern Wadden Sea) have been measured about twice weekly since 1984. Salinity has a clear seasonal cycle with lowest salinities of about 27 in late winter and highest salinities of about 31 in summer. Mean annual deviations from the long-term mean salinity correlate significantly with riverine freshwater discharge. Winter nitrate concentrations are generally high (about 50 μM on average). The major part of the variability is related to salinity (∼35%). Temperature had a minor impact (∼1%). Superimposed on this, a long-term decrease of about 1 μM per year was found. Together, these processes account for about 45% of the nitrate variability. The long-term decrease of about 2% per year is similar to continental riverine trend in total nitrogen loads. In contrast to the List Tidal Basin, salinity explained more than 90% of nitrate variability in the off-shore German Bight. Salinity (30) normalised winter nitrate data of the German Bight also show a long-term decreasing trend. Most of the List Tidal Basin data are either on or below the nitrate–salinity relation found in the German Bight. This observation suggests that denitrification has a major impact on the winter nitrate concentrations in the Northern Wadden Sea compared to the German Bight. It is hypothesised that a large part of the unexplained variability is related to weather-dependent changes in residence time of tidal water masses in the Wadden Sea and circulation patterns within the German Bight.     

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.     

Some issues related to reproduction of Pacific cod <Emphasis Type="Italic">Gadus macrocephalus</Emphasis> in waters of the eastern coast of the northern Kuril Islands and the southern extremity of Kamchatka

The analysis of data collected in December 1996 and 1998 on the reproduction of cod Gadus macrocephalus in Pacific waters of the northern Kuril Islands and the southern part of Kamchatka was performed. It was shown that the individual absolute fecundity of the cod varies within 0.197–9.729 million eggs and the relative fecundity, in the range of 24-1386 eggs. The fecundity of 1000 mature females comprises 2179–2449 million eggs. The low individual fecundity of fish is related to pseudobranchial tumor growth. The main role in cod reproduction is played by females of two-three size groups characterized by the highest numbers of mature females. It is suggested that different fecundity within the range of Pacific cod is related to environmental conditions, in particular, water temperature in the spawning grounds during spawning rather than to the habitation latitude.     

Comparison of trace heavy-metal levels from monitoring in the German Bight and the southwestern Baltic Sea

Since 1973, monitoring for selected trace heavy metals has been performed in the German Bight (North Sea) and the western Baltic Sea. This paper deals with a set of filtered and unfiltered samples from a network of 22 stations covering the whole German Bight based on a cruise with RV “Gauss” in May 1974, and a similar series of samples taken at different depths at 18 stations in the Baltic Sea in October 1974. The metals Cd, Cu, Fe, Mn, and Ni were determined by flameless atomic absorption spectrometry. Data comparison is facilitated since sampling and analytical methods were identical. Distributions of metal-content data have been calculated from unfiltered and filtered samples. Qualitative and semi-quantitative aspects of the major differences or similarities between the histograms obtained from both sea areas are discussed.     

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.     

Some differences in the parasitic fauna of three samples of plaice (Pleuronectesplatessa L.) from the southern North Sea

Plaice ( Pleuronectesplatessa L.) from three spawning areas in the southern North Sea were examined for six species of metazoan parasite. Differences of both incidence and level of infestation with particular parasite species were found in fish from the three areas. Neither the incidence nor the level of infestation was associated with the length of fish in the samples except in the case of the larger male fish from Flamborough, which tended to contain more larval nematodes (Anisakidae). All fish contained the intestinal nematode Cucullanus heterochrous , although fewer were found in fish from Flamborough. Southern Bight fish were more likely to contain Capillaria wickinsi than were fish from Flamborough or the German Bight. Flamborough fish were more likely to contain Anisakidae larvae than were fish from the other two areas. Fish from the three areas were similarly infested with Zoogonoides viviparus. Male fish from Flamborough were less likely to be infested with Lepeophtheirus pectoralis than male fish from the German Bight.     

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.     

Long-Term Changes in the Gonad Condition of the Sea Urchin Strongylocentrotus intermediusfrom Alekseev Bight (Amurskii Bay) under Polluted Conditions

The gonad condition of the sea urchin Strongylocentrotus intermediuscollected in August 1997 at two stations in Peter the Great Bay was examined. One of the stations was located in a polluted area (Alekseev Bight, Popov Island) and the other, in a relatively clean area (the Verkhovskii Islands). The results were compared with analogous data for 1984, 1985, and 1989. In 1997, the gonad condition of sea urchins inhabiting the two areas differed significantly. The mean value of the gonad index (GI) for sea urchins from Alekseev Bight was less than half and the maturity index was about twice that of sea urchins from the Verkhovskii Islands. The GI of sea urchins from Alekseev Bight decreased by a factor of 1.5 between 1984 and 1997. Pronounced histopathological changes were found in sea urchin gonads at this station: granular and hydropic dystrophy of oocytes, resorption and a sharp decrease in the number of gametes (in about 20% of the sea urchins, hardly any gametes were found in the gonads), changes in the morphology of accessory cells (hypertrophy, atrophy, and necrosis), and accumulation of lipofuscin in the cytoplasm of accessory cells and oocytes, in the hemal sinuses and mesentery. The suppressed gonad condition of the sea urchin S. intermediusin Alekseev Bight may be a consequence of the unfavorable environmental situation that formed in the bight in the 1980s–1990s. The main negative factor is anthropogenic pollution of Amurskii Bay.     

Annual changes in the nutritive state of North Sea dab

The nutritive state of dab Limanda limanda was investigated over a 2-year period at a fixed sampling site northwest of Helgoland (German Bight, North Sea), with respect to feeding habits and the accumulation of biochemical storage products. Ophiuroids formed the main weight of food organisms (50%) while polychaetes (10%), molluscs and crustaceans (<5% each) were less frequent. Feeding activity in males varied between summer and winter, while females fed more constantly. The condition factor and the hepatosomatic index showed characteristic seasonal cycles in both sexes. The glycogen content in the liver reached 40–60 mg g⁻¹ FW in summer and fell to about 10–20 mg g⁻¹ FW in late winter. Total lipids of the liver showed a distinct seasonal cycle with 400 mg g⁻¹ FW in summer and a minimum of 50–100 mg g⁻¹ FW in spring. The lipid content of the muscle ranged from 5 to 6 mg g⁻¹ FW and did not vary significantly between seasons.     

The human impact in the German Bight: Eutrophication during three decades (1962–1991)

The human impact in the German Bight, in the form of anthropogenic eutrophication, has been documented by a 30-year time-series measurement near the island of Helgoland. Since 1962, the Biologische Anstalt Helgoland has measured inorganic nutrients and phytoplankton abundance from daily samples at Helgoland Roads, a position 60 km off the main source of eutrophication, the River Elbe. Since the early sixties, phosphate concentrations rose for about a decade, levelling off to about twice the former concentrations for another decade, and then decreasing (since 1982) as a result of phosphate-reducing measures. Nitrate concentrations, however, have only increased since 1980/81, following Elbe river flood events. In 1987, three times the former concentrations were reached. A decrease has been observed only since 1991. This different development of phosphorus and nitrogen eutrophication led to a shift of inorganic N/P-ratios in the German Bight. The phosphate increase was more pronounced in the late summer “regeneration mode” conditions, the nitrate increase in the winter months. The eutrophication is not restricted to the inner German Bight and coastal waters of a salinity of <33, but has also occurred in more saline waters at S>33 psu (practical salinity unit), as characteristic for the outer German Bight. In this more saline water, phosphate and nitrate maximum levels occurred three years later, compared with the average Helgoland data, which are more representative of the inner German Bight. It is suggested that suspended particulate organic matter, as a long-distance carrier of nutrients, might have caused this delayed eutrophication in the outer German Bight waters. While the human impact is obvious as to nutrient concentrations, it is less obvious in phytoplankton stock enhancement. A general increase in phytoplankton biomass (about 3–4 times) was found, but this was mainly due to unidentified nanoflagellates of unknown trophic state, and subject to methodological errors. The causal relationships of phytoplankton stocks and eutrophication are not clearly understood, as natural variability is large and hydrographical factors possibly dominate. Additional nutrient input by Elbe river floods did not always result in elevated phytoplankton stocks near Helgoland, while extended periods of vertical density stratification of the German Bight water caused large plankton blooms.     

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.     

Microbial Diversity during Biodegradation of Crude Oil in Seawater from the North Sea

Microbial communities were characterized during biodegradation of immobilized oil in seawater from the Statfjord field and the German Bight in the North Sea. Seawater samples were collected at different distances from pollution sources at the two locations. A Statfjord oil was immobilized on hydrophobic synthetic Fluortex fabrics and submerged in closed flasks (no headspace) with natural or sterile seawater and incubated at 13°C for 56 days. Biodegradation of immobilized n-alkanes was measured by gas chromatography, total microbes were enumerated by epifluorescence microscopy, and culturable heterotrophic and oil-degrading microorganisms were quantified by most probable number (MPN) analysis. Polymerase chain reaction (PCR) amplification of bacterial 16S rDNA in water samples was conducted during biodegradation experiments. The amplified 16S rDNA fragments were characterized by denaturing gradient gel electrophoresis (DGGE), and by sequence analysis of cloned inserts. Biodegradation rates of alkanes in seawater collected at different distances from the pollution sources did not differ significantly (P > 0.05). Concentrations of oil-degrading microorganisms showed a temporary peak after 7 days of degradation, with a subsequent decline later in the period. DGGE analysis of 16S rRNA genes showed that community diversity decreased during the first 2–3 weeks of biodegradation, with the emergence of a few dominant bands. Cloning, restriction analysis, and sequence analysis of the 16S rDNA fragments revealed >30 different phylotypes. Abundant types during biodegradation belonged to the -Proteobacteria, in waters from both Statfjord and the German Bight. Cloning and sequencing studies indicated that the most abundant bacteria during biodegradation belonged to the family Rhodobacteraceae, with the closest relationship to the genera Sulfitobacter and Roseobacter.     

Larvae of uncommon caridean decapods in the German Bight: Species composition,distribution and abundance

Typically, the most abundant group of shrimp larvae in the German Bight is formed by representatives of the family Crangonidae. Larvae of the remaining species have been largely ignored, and only scarce information concerning their ecology is available. Thus, the purpose of the present study was to determine the species composition, distribution and abundance of noncrangonid shrimp larvae in the German Bight in July 1990, after the mildest winter of the century. The material is based upon plankton samples collected at 77 stations, covering the entire German Bight. Eight species were identified, as well as larvae of Palaemonidae andProcessa-juveniles.Processa nouveli holthuisi (53.0%) andP. modica (31.3%) were predominant in the collection. The distribution of the two species was clearly separated: the main concentration ofP. nouveli holthuisi (peak concentration of 1.94 larvae per m³) was confined to the northwest corner of the German Bight, while a majority ofP. modica larvae (peak concentration of 0.54 larvae per m³) occurred at the southwesterly stations. The spatial distribution ofCaridion steveni andEualus occultus around Helgoland indicates the presence of an adult population at the only rocky island in the study area. Other taxa, such as larvae of Palaemonidae and juvenilePandalina brevirostris were collected exclusively in estuarine habitats. Based upon both the results of the present study and comparable data, we conclude that developmental stages of ten non-crangonid species, as well as representatives of Palaemonidae, can be expected to occur in the plankton of the German Bight. The extremely mild temperatures of the preceding winter may have been, in part, responsible for the relatively high densities of some taxa encountered during our plankton survey. We assume that warm winter temperatures favour the immigration, reproduction and survival of cold-sensitive species.     

Nitrogen removal in coastal sediments of the German Wadden Sea

Although sediments of the German Wadden Sea are suspected to eliminate a considerable share of nitrate delivered to the SE North Sea, their denitrification rates have not been systematically assessed. We determined N₂ production rates over seasonal cycles (February 2009–April 2010) at two locations with two sediments types each, the first site (Meldorf Bight) receiving nitrate during all seasons from the Elbe river plume, and a second site on the island of Sylt, where nitrate is depleted during summer months. In sediments from the Sylt site, N₂ production ranged from 15 to 32 μmol N₂ m^?2 h^?1 in the fine sand station and from 7 to 13 μmol N₂ m^?2 h^?1 in the coarse sand station; N₂ production was not detected when nitrate was depleted in May and July of 2009. N₂ production in the Meldorf Bight sediments were consistently detected at higher rates (58–130 μmol N₂ m^?2 h^?1 in the very fine sand station and between 14 and 30 μmol N₂ m^?2 h^?1 in the medium sand station). Analysis of ancillary parameters suggests that major factors controlling N₂ production in coastal sediments of the German Wadden Sea are the nitrate concentrations in the overlying water, the ambient temperature, and the organic matter content of the sediment. Extrapolating our spot measurements to the zone of nitrate availability and sediment types, we estimate an annual nitrogen removal rate around 16 kt N year^?1 for the entire northern sector of the German Wadden Sea area. This corresponds to 14% of the annual Elbe river nitrogen load.     

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.