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.     

Evidence of a hybrid-zone in Atlantic cod (Gadus morhua) in the Baltic and the Danish Belt Sea revealed by individual admixture analysis

The study of hybrid zones is central to our understanding of the genetic basis of reproductive isolation and speciation, yet very little is known about the extent and significance of hybrid zones in marine fishes. We examined the population structure of cod in the transition area between the North Sea and the Baltic Sea employing nine microsatellite loci. Genetic differentiation between the North Sea sample and the rest increased along a transect to the Baltic proper, with a large increase in level of differentiation occurring in the Western Baltic area. Our objective was to determine whether this pattern was caused purely by varying degrees of mechanical mixing of North Sea and Baltic Sea cod or by interbreeding and formation of a hybrid swarm. Simulation studies revealed that traditional Hardy-Weinberg analysis did not have sufficient power for detection of a Wahlund effect. However, using a model-based clustering method for individual admixture analysis, we were able to demonstrate the existence of intermediate genotypes in all samples from the transition area. Accordingly, our data were explained best by a model of a hybrid swarm flanked by pure nonadmixed populations in the North Sea and the Baltic Sea proper. Significant correlation of gene identities across loci (gametic phase disequilibrium) was found only in a sample from the Western Baltic, suggesting this area as the centre of the apparent hybrid zone. A hybrid zone for cod in the ecotone between the high-saline North Sea and the low-saline Baltic Sea is discussed in relation to its possible origin and maintenance, and in relation to a classical study of haemoglobin variation in cod from the Baltic Sea/Danish Belt Sea, suggesting mixing of two divergent populations without interbreeding.     

Warm-water decapods and the trophic amplification of climate in the North Sea

A long-term time series of plankton and benthic records in the North Sea indicates an increase in decapods and a decline in their prey species that include bivalves and flatfish recruits. Here, we show that in the southern North Sea the proportion of decapods to bivalves doubled following a temperature-driven, abrupt ecosystem shift during the 1980s. Analysis of decapod larvae in the plankton reveals a greater presence and spatial extent of warm-water species where the increase in decapods is greatest. These changes paralleled the arrival of new species such as the warm-water swimming crab Polybius henslowii now found in the southern North Sea. We suggest that climate-induced changes among North Sea decapods have played an important role in the trophic amplification of a climate signal and in the development of the new North Sea dynamic regime.     

An environmental gradient dominates ecological and genetic differentiation of marine invertebrates between the North and Baltic Sea

Environmental gradients have emerged as important barriers to structuring populations and species distributions. We set out to test whether the strong salinity gradient from the marine North Sea to the brackish Baltic Sea in northern Europe represents an ecological and genetic break, and to identify life history traits that correlate with the strength of this break. We accumulated mitochondrial cytochrome oxidase subunit 1 sequence data, and data on the distribution, salinity tolerance, and life history for 28 species belonging to the Cnidaria, Crustacea, Echinodermata, Mollusca, Polychaeta, and Gastrotricha. We included seven non‐native species covering a broad range of times since introduction, in order to gain insight into the pace of adaptation and differentiation. We calculated measures of genetic diversity and differentiation across the environmental gradient, coalescent times, and migration rates between North and Baltic Sea populations, and analyzed correlations between genetic and life history data. The majority of investigated species is either genetically differentiated and/or adapted to the lower salinity conditions of the Baltic Sea. Species exhibiting population structure have a range of patterns of genetic diversity in comparison with the North Sea, from lower in the Baltic Sea to higher in the Baltic Sea, or equally diverse in North and Baltic Sea. Two of the non‐native species showed signs of genetic differentiation, their times since introduction to the Baltic Sea being about 80 and >700 years, respectively. Our results indicate that the transition from North Sea to Baltic Sea represents a genetic and ecological break: The diversity of genetic patterns points toward independent trajectories in the Baltic compared with the North Sea, and ecological differences with regard to salinity tolerance are common. The North Sea–Baltic Sea region provides a unique setting to study evolutionary adaptation during colonization processes at different stages by jointly considering native and non‐native species.     

Timing of downstream migration and food uptake of juvenile North Sea houting stocked in the Lower Rhine and the Lippe (Germany)

In 1992 a stocking programme was established to re‐introduce North Sea houting Coregonus oxyrinchus , which disappeared from the River Rhine in the 1940s, in the Lower Rhine. Juvenile North Sea houting have been stocked since 2001 in a gravel pit lake permanently connected to the Lower Rhine, and in the Lippe, a potamal tributary of the Rhine. Monitoring studies showed that the majority of the small North Sea houting left the gravel pit lake within 4 days after stocking. In the Lippe, some of the juveniles were found in the drift immediately after stocking, generally preferring the middle surface areas of the river. Other North Sea houting waited until dawn before they started their downstream migration. Juveniles immediately started to feed on the zooplankton resources in both waters. Pond studies revealed high growth rates of juvenile North Sea houting, ranging from 0·44 to 0·94 mm day⁻¹. This study suggests that the stocking strategies at both waters of the Lower Rhine comply with the requirements of the migration behaviour of juvenile North Sea houting.     

Zonation of ground beetles (Coleoptera: Carabidae) and spiders (Araneida) in salt marshes at the North and the Baltic Sea and the impact of the predicted sea level increase

The ground beetles and spiders of two salt marshes at the German Northand Baltic Sea coast were investigated by pitfall traps in 1997 and 1998. Whilethe sites at the North Sea coast are tidal salt marshes, the salt marshes at theBaltic Sea are not influenced by tides. Pitfall traps were installed in agradient from 20 to 150 cm above MHT (mean high tide: 157cm + NN, NN: 500 cm above 0 at Amsterdam gauge) atthe North Sea coast or NN at the Baltic Sea coast at six or sevensampling elevations, each with five replicates. Conductivity, water content, organic substance,frequency or duration of floodings, sand content and pH of the soil weredetermined. The flooding regime is the major factor controlling the zonation ofinvertebrates. Two and three invertebrate assemblages at the North and BalticSea, respectively, were distinguished. These corresponded well with thevegetational zones. The border between the two zones was at 60–80cm above MHT at the North Sea. The three zones at the Baltic Seaextended between 20 and 30 cm, 40 to 80 cm and 100 to150 cm above NN. The elevation of the mean abundance of speciesabove MHT or NN was calculated. A tide simulation experiment resulted in ashifting population and in an increasing activity under a tidal regime aspredicted for the global climate change conditions in 2050. From the actualelevation of the mean abundance, the habitat size of salt marsh species wascalculated for a moderate and worse scenario of global climate change. Habitatreduction becomes highest for species of the lower salt marsh zone. Under worseconditions the gradiental length of habitat will only amount to a maximum of 20m at the slopes of the dikes.     

Assessing the role of ontogenetic movement in maintaining population structure in fish using otolith microchemistry

Identifying the mechanisms maintaining population structure in marine fish species with more than a single dispersing life stage is challenging because of the difficulty in tracking all life stages. Here, a two‐stage otolith microchemistry approach to examining life‐stage movement was adopted, tracking a year‐class from the juvenile to adult stage and inferring larval sources from clustering, in order to consider the mechanisms maintaining population structuring in North Sea cod. Clustering of near‐core chemistry identified four clusters, two of which had either a southern or northern affinity and were similar to juvenile edge chemistry. The other two clusters, common to the central North Sea, had intermediate chemical composition and may have reflected either larval mixing in this region or a lack of geographic heterogeneity in the elemental signature. From the comparison of whole juvenile and the corresponding component of adult otoliths, adults from the southern North Sea mostly recruited from adjacent nursery grounds. In contrast, many adults in the northern North Sea had a juvenile chemistry consistent with the Skagerrak and juveniles from the northern Skagerrak site had a near‐core chemistry consistent with the northern North Sea. Similarities in otolith chemistry were consistent with retention of early life stages at a regional level and also juvenile and adult fidelity. The links between the northern North Sea and Skagerrak indicate natal homing, which when considered in the context of genetic evidence is suggestive of philopatry. The approach used here should be useful in exploring the mechanisms underlying population structuring in other species with multiple dispersive life stages and calcified hard parts.     

The diet of whiting Merlangius merlangus in the western Baltic Sea

The diet of whiting Merlangius merlangus in the western Baltic Sea was investigated and compared to the diet in the southern North Sea. Clupeids were important prey in both areas, but especially in the western Baltic Sea where they constituted up to 90% of the diet of larger individuals. Gobies, brown shrimps and polychaetes were the main prey of juveniles in the western Baltic Sea, while a wider range of species were consumed in the North Sea. The shift to piscivory occurred at smaller sizes in the western Baltic Sea and the fish prey consumed was proportionately larger than in the southern North Sea. Estimates of prey abundance and food intake of M. merlangus are required to evaluate its predatory significance in the western Baltic Sea, but its diet suggests that it could be just as significant a fish predator here as in the southern North Sea.     

Female philopatry in coastal basins and male dispersion across the North Atlantic in a highly mobile marine species, the sperm whale (Physeter macrocephalus)

The mechanisms that determine population structure in highly mobile marine species are poorly understood, but useful towards understanding the evolution of diversity, and essential for effective conservation and management. In this study, we compare putative sperm whale populations located in the Gulf of Mexico, western North Atlantic, Mediterranean Sea and North Sea using mtDNA control region sequence data and 16 polymorphic microsatellite loci. The Gulf of Mexico, western North Atlantic and North Sea populations each possessed similar low levels of haplotype and nucleotide diversity at the mtDNA locus, while the Mediterranean Sea population showed no detectable mtDNA diversity. Mitochondrial DNA results showed significant differentiation between all populations, while microsatellites showed significant differentiation only for comparisons with the Mediterranean Sea, and at a much lower level than seen for mtDNA. Samples from either side of the North Atlantic in coastal waters showed no differentiation for mtDNA, while North Atlantic samples from just outside the Gulf of Mexico (the western North Atlantic sample) were highly differentiated from samples within the Gulf at this locus. Our analyses indicate a previously unknown fidelity of females to coastal basins either side of the North Atlantic, and suggest the movement of males among these populations for breeding.     

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.     

Genetic population structure of turbot (Scophthalmus maximus L.) supports the presence of multiple hybrid zones for marine fishes in the transition zone between the Baltic Sea and the North Sea

Genetic population structure of turbot (Scophthalmus maximus L.) in the Northeast Atlantic was investigated using eight highly variable microsatellite loci. In total 706 individuals from eight locations with temporal replicates were assayed, covering an area from the French Bay of Biscay to the Aaland archipelago in the Baltic Sea. In contrast to previous genetic studies of turbot, we found significant genetic differentiation among samples with a maximum pairwise FST of 0.032. Limited or no genetic differentiation was found among samples within the Atlantic/North Sea area and within the Baltic Sea, suggesting high gene flow among populations in these areas. In contrast, there was a sharp cline in genetic differentiation going from the low saline Baltic Sea to the high saline North Sea. The data were explained best by two divergent populations connected by a hybrid zone; however, a mechanical mixing model could not be ruled out. A significant part of the genetic variance could be ascribed to variation among years within locality. Nevertheless, the population structure was relatively stable over time, suggesting that the observed pattern of genetic differentiation is biologically significant. This study suggests that hybrid zones are a common phenomenon for marine fishes in the transition area between the North Sea and the Baltic Sea and highlights the importance of using interspecific comparisons for inferring population structure in high gene flow species such as most marine fishes.     

Genetic Stock Identification of Chum Salmon in the Bering Sea and North Pacific Ocean Using Mitochondrial DNA Microarray

A newly developed DNA microarray was applied to identify mitochondrial (mt) DNA haplotypes of more than 2200 chum salmon in the Bering Sea and North Pacific Ocean in September 2002 and also 2003, when the majority of maturing fish were migrating toward their natal river. The distribution of haplotypes occurring in Asian and North American fish in the surveyed area was similar in the 2 years. A conditional maximum likelihood method for estimation of stock compositions indicated that the Japanese stocks were distributed mainly in the north central Bering Sea, whereas the Russian stocks were mainly in the western Bering Sea. The North American stocks were abundant in the North Pacific Ocean around the Aleutian Islands. These results indicate that the Asian and North American stocks of chum salmon are nonrandomly distributed in the Bering Sea and the North Pacific Ocean, and further the oligonuleotide DNA microarray developed by us has a high potential for identification of stocks among mixed ocean aggregates of high-seas chum salmon.     

A decline in primary production in the North Sea over 25 years,associated with reductions in zooplankton abundance and fish stock recruitment

Phytoplankton primary production is at the base of the marine food web; changes in primary production have direct or indirect effects on higher trophic levels, from zooplankton organisms to marine mammals and seabirds. Here, we present a new time‐series on gross primary production in the North Sea, from 1988 to 2013, estimated using in situ measurements of chlorophyll and underwater light. This shows that recent decades have seen a significant decline in primary production in the North Sea. Moreover, primary production differs in magnitude between six hydrodynamic regions within the North Sea. Sea surface warming and reduced riverine nutrient inputs are found to be likely contributors to the declining levels of primary production. In turn, significant correlations are found between observed changes in primary production and the dynamics of higher trophic levels including (small) copepods and a standardized index of fish recruitment, averaged over seven stocks of high commercial significance in the North Sea. Given positive (bottom‐up) associations between primary production, zooplankton abundance and fish stock recruitment, this study provides strong evidence that if the decline in primary production continues, knock‐on effects upon the productivity of fisheries are to be expected unless these fisheries are managed effectively and cautiously.     

Francisella noatunensis subsp. noatunensis is the aetiological agent of visceral granulomatosis in wild Atlantic cod Gadus morhua

During the 1980s and 1990s wild-caught cod displaying visceral granulomatosis were sporadically identified from the southern North Sea. Presumptive diagnoses at the time included mycobacterial infection, although mycobacteria were never cultivated or observed histologically from these fish. Farmed cod in Norway displaying gross pathology similar to that identified previously in cod from the southern North Sea were recently discovered to be infected with the bacterium Francisella noatunensis subsp, noatunensis. Archived formalin-fixed paraffin-embedded tissues from the original North Sea cases were investigated for the presence of Mycobacterium spp. and Francisella spp. using real-time polymerase chain reaction, DNA sequencing and immunohistochemistry. Whilst no evidence of mycobacterial infection was found, F. noatunensis subsp. noatunensis was identified in association with pathological changes consistent with Francisella infections described from farmed cod in recent years. This study shows that francisellosis occurred in wild-caught cod in the southern North Sea in the 1980s and 1990s and demonstrates that this disease predates intensive aquaculture of cod.     

Estimation of the load of cyclic organochlorines in North Sea zooplankton

During a North Sea survey in late spring 1986, zooplankton biomass (g dry weight/m²) as well as pollutant concentrations in zooplankton (ng/g dry weight) were determined at 127 stations. On the basis of these data, the load of several cyclic organochlorines in zooplankton (ng/m²) was estimated. It appeared that the highest loads of these pollutants were incorporated in the large zooplankton stocks of the northern and central North Sea, north of 56°N, whereas the highest concentrations were found in zooplankton of the southern North Sea. An explanation for the accumulation of pollutants in sediments as well as in benthos organisms of the central and northern North Sea as found by several authors is discussed in connection with the assumption of a considerably high vertical flux of organic material in these areas. The calculation of total zooplankton content in the North Sea during the late spring survey amounted to about 5 million tons dry weight. This biomass was estimated to contain about 2.6 t PCBs, 19.3 kg p,p′-DDE, 12 kg τ-HCH, 6.4 kg HCB und 5.6 kg α-HCH. These results agree with those on other compartments of the North Sea ecosystem (benthos, fish) found in recent literature. The estimated annual turnover of PCBs in zooplankton (15.2 t) is of the same order of magnitude as the estimated amount transported yearly into the North Sea by rivers and the atmosphere (6–13 t).     

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.     

Population genetic structure of North Atlantic, Mediterranean Sea and Sea of Cortez fin whales, Balaenoptera physalus (Linnaeus 1758): analysis of mitochondrial and nuclear loci

Samples were collected from 407 fin whales, Balaenoptera physalus , at four North Atlantic and one Mediterranean Sea summer feeding area as well as the Sea of Cortez in the Pacific Ocean. For each sample, the sex, the sequence of the first 288 nucleotides of the mitochondrial (mt) control region and the genotype at six microsatellite loci were determined. A significant degree of divergence was detected at all nuclear and mt loci between North Atlantic/Mediterranean Sea and the Sea of Cortez. However, the divergence time estimated from the mt sequences was substantially lower than the time elapsed since the rise of the Panama Isthmus, suggesting occasional gene flow between the North Pacific and North Atlantic ocean after the separation of the two oceans. Within the North Atlantic and Mediterranean Sea, significant levels of heterogeneity were observed in the mtDNA between the Mediterranean Sea, the eastern (Spain) and the western (the Gulf of Maine and the Gulf of St Lawrence) North Atlantic. Samples collected off West Greenland and Iceland could not be unequivocally assigned to either of the two areas. The homogeneity tests performed using the nuclear data revealed significant levels of divergence only between the Mediterranean Sea and the Gulf of St Lawrence or West Greenland. In conclusion, our results suggest the existence of several recently diverged populations in the North Atlantic and Mediterranean Sea, possibly with some limited gene flow between adjacent populations, a population structure which is consistent with earlier population models proposed by Kellogg, Ingebrigtsen, and Sergeant.     

Cryptic species in marine polychaete and their independent introduction from North America to Europe

The vast body of ballast water carried across oceans by freight ships represents a major source for the introduction of foreign species into marine ecosystems. The worm Marenzelleria viridis, originally found only in North America, appeared in estuaries of the North Sea in 1979 and 6 years later also in the Baltic, where it has developed into a major faunal element. Two competing hypotheses are discussed here: either both populations owe their presence to a single introductory event in the North Sea, or each population originated from a separate introduction. Our phylogeographic analysis of Baltic, North Sea and American Marenzelleria, based on mitochondrial 16S rDNA sequences (326- bp segment) of 98 individuals from 17 localities on the North American, North Sea, and Baltic coasts not only favors the two-event hypothesis, but also separates the locations of origin for the introductions. Eighteen mitochondrial genotypes were identified altogether. In agreement with allozyme data, three lineages were identified: genotypes assigned to the same lineage differed from each other by up to 5 point mutations, and those assigned to different lineages differed by up to 17. The existence of three morphologically indistinguishable, and thus cryptic, species is therefore suggested. The individuals from the Baltic Sea probably originated from the Atlantic coast of the United States between Chesapeake Bay and Georgia, and the North Sea populations may stem from the U.S. coast region north of Chesapeake Bay to Nova Scotia. Despite their similar morphologies, the two European Marenzelleria species may differ ecologically with respect to their preference for habitat salinity. Assuming that transport via ballast water occurs quite frequently, we hypothesize that both European cryptic species of Marenzelleria may originally have been introduced to both the North Sea and the Baltic Sea but that neither of them was able to proliferate in both water bodies owing to their differential physiological performances at high and low salinities.      

Introduced marine species of the North Sea coasts

About 80 non-indigenous species are assumed to have been introduced into the North Sea by transoceanic shipping and aquaculture. The number is certainly underestimated as most small organisms received insufficient attention at the species level. Also, the seafaring tradition of the North Sea countries is much longer than our biological surveys are. Most exotic invertebrates originate from the western Atlantic and were introduced by shipping, while most algae stem from the Pacific and came with the introduced oysters. A peak of newcomers was observed in the 1970s. Most of the arrivals became established in brackish environments, at harbor sites and in the vicinity of oyster farms, fouling on hard substrates or living as epibionts. A few live in sediments, are holoplanktonic or are parasites. At the open coast, approximately 6% of the macrobenthic species are exotics, while in estuaries their share is up to 20%. Most exotics have been encountered in the southern North Sea first, and many did not spread further north. About 25% of the established non-natives are widespread and attain locally high abundances. As a consequence, some inshore habitats are entirely dominated by exotics. The overall effect on the ecosystem seems to be more additive than one of displacement. This suggests that the coastal biota of the North Sea are quite capable of accommodating newcomers. However, this is no guarantee that the next introduced species may not cause severe ecological change or economic harm. There is a need to minimize the risk of unintentional introductions by ballast water treatment and by adhering to quarantine procedures in aquaculture. Current research on exotics in the North Sea is regarded as inadequate for proper evaluation and management requirements.