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.     

Roles of sulfate adsorption and base cation supply in controlling the chemical response of streams of western Virginia to reduced acid deposition

Micro-organisms are vital for the functioning of all food webs and are the major drivers of the global biogeochemical cycles. The microbial community compositions and physicochemical conditions of the different water masses in the North Sea, a biologically productive sea on the northwestern European continental shelf, were studied during two summer cruises, in order to provide detailed baseline data for this region and examine its microbial biogeography. For each cruise the stations were clustered according to their physicochemical characteristics and their microbial community composition. The largest cluster, which covered most of the central and northern North Sea, consisted of stations that were characterized by a thermally stratified water column and had low chlorophyll a autofluorescence and generally low microbial abundances. The second main cluster contained stations that were dominated by picoeukaryotes and showed the influence of influxes of North Atlantic water via the English Channel and south of the Shetland Islands. The third main cluster was formed by stations that were dominated by cyanobacteria and nanoeukaryotes in the reduced salinity Norwegian Coastal and Skagerrak waters, while the fourth cluster represented the German Bight, a region with strong riverine input, high nutrient concentrations, and consequently high heterotrophic bacterial and viral abundances. Despite the complex and dynamic hydrographic nature of the North Sea, the consistent distinctions in microbiology between these different hydrographic regions during both cruises illustrate the strong links between the microbial community and its environment, as well as the possibility to use microorganisms for long-term monitoring of environmental change.     

The stock structure and migrations of plaice populations on the west coast of England and Wales

Plaice Pleuronectes platessa populations on the west coast of England and Wales are currently managed as two stocks: in ICES Division VIIa (Irish Sea, Cardigan Bay and St George's Channel), and ICES Divisions VIIf&g (Bristol Channel and Celtic Sea). A total of 13,784 plaice were fitted with Petersen tags and released in these areas during 1979–1980 and 1993–1996. Analysis of the 2788 recaptures received by June 2000 confirmed known spawning and feeding grounds in the region. It showed also that plaice >25 cm L _T tended to undertake extensive spatial movements. At this size, female plaice were likely to be mature or maturing for the first time, whilst males were probably mature. Tag recaptures indicated resident sub-stocks of plaice in the north-east Irish Sea, the south-east Irish Sea, Cardigan Bay and Bristol Channel, a contingent of plaice in all areas that undertook permanent dispersal to other areas, and a contingent which originated in the south-east Irish Sea and migrated to spawn in the Bristol Channel. Plaice originating in the Bristol Channel rarely moved north into ICES Vila. A general hypothesis of plaice population structure in the region is presented and discussed in relation to stock assessment.     

Seasonal changes in frequency of diseases in dab,Limanda limanda,from the southern North Sea

Infestation rates of flatfishLimanda limanda from the southern North Sea (German Bight) were investigated in 1981 and 1982. At three stations, including a dumping area of acid-iron wastes northwest of Helgoland, dabs were examined for the occurrence of lymphocystis and epidermal papillomas. Marked seasonal changes of infestation rates were observed with maxima of diseased fish in March and May (epidermal papillomas 2 % in 1981, 4.1 % in 1982; lymphocystis 7.5 % in 1981, 17.2 % in 1982). Infestation minima were found from June to October. Differences in incidence rates were observed between stations in winter and spring, but not in summer. Both diseases turned out to be size-dependent. Seasonality of the two diseases is discussed in relation to spawning cycle and water quality, including pollution.     

Upper temperature tolerance of North Atlantic and North Pacific geographical isolates ofChondrus species (Rhodophyta)

The upper survival temperature for most isolates ofChondrus crispus from localities ranging from northern Norway and Iceland to Spain, and for an isolate from Nova Scotia, was 28 °C after 2 weeks of exposure to temperatures of 20–31 °C at intervals of 1 °C. An upper survival limit of 29 °C was exhibited by a few European isolates from the English Channel, the North Sea, and one Irish isolate from the upper intertidal. The warm-temperate Japanese speciesC. nipponicus andC. giganteus formaflabellatus survived 30 °C, whereas 29 °C was the upper survival limit for the coldtemperatureC. pinnulatus formapinnulatus from northern Japan. A possible origin ofC. crispus in the north Pacific is discussed.     

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     

Overland movement and migration phenology in relation to breeding of Arctic Terns Sterna paradisaea

It is often implicitly assumed that seabirds migrate using marine environments, but this assumption is increasingly being challenged by electronic tracking data. The arrival and departure routes of Arctic Terns breeding on the North Sea coast of the United Kingdom (UK) are unknown but there has been speculation about the possibility of overland migration. Analysis of light-level geolocator data from birds breeding on the Farne Islands suggests that these birds arrived and left their North Sea colony overland via the Irish Sea, rather than taking coastal routes along the east coast of the UK and through the English Channel. In addition, some departing birds may enter the North Atlantic by crossing Ireland rather than through the Irish Sea. The direction of arrival in spring had a more southerly orientation than the direction of autumn departure. The geolocator data allow migration phenology in relation to breeding to be defined and indicated that the birds arrived around 15 days before the first eggs were laid in the colony. Departure timing may be determined by seasonal progression and not markedly influenced by breeding success. This study supports the idea that overland migration may be a more widespread and consistent strategy for seabirds than has been realized.     

Phylogeography of an estuarine mysid, Neomysis integer (Crustacea, Mysida), along the north-east Atlantic coasts

Aim The brackish water mysid, Neomysis integer, is one of the most common mysid species along the coasts of the north‐east Atlantic. In the present study, the phylogeographical patterns were examined throughout the distribution range of N. integer. In particular, the latitudinal trends in genetic diversity and the distribution of genetic variation were examined in order to elucidate the imprints of the Pleistocene glaciations. Location North‐east Atlantic coasts from the Baltic Sea to the south of Spain. Methods A total of 461 specimens from 11 populations were analysed by means of single‐stranded conformation polymorphism analysis combined with DNA sequencing of a fragment of the mitochondrial cytochrome c oxidase I gene. The genetic structure was examined by using a progression of phylogenetic, demographic and population genetic analyses to elucidate not only the geographical structure, but also the evolutionary history producing that structure. Results The levels of genetic diversity were relatively uniform throughout the distribution range, with the exception of a decline at the northern and southern edges of distribution. A high heterogeneity was observed between the populations analysed (global Φ_ST = 0.787). This is caused by the disparate distribution of the cytochrome oxidase I haplotypes, with several population‐specific haplotypes. A clear genetic break (2.4% sequence divergence) occurred between the southernmost Guadalquivir population and all other populations. Main conclusions The present study corroborates the expectations of the genetic patterns typically observed in an estuarine species. The within‐population variability was low, whereas a significant (moderate to high) divergence was observed between populations. Phylogeographical analysis revealed that northern populations within the English Channel, North Sea and Baltic Sea are characterized by several widespread haplotypes, while the Irish population and all sites south of the Bay of Biscay consist solely of unique haplotypes. This pattern, combined with the relative high levels of genetic diversity, could be indicative for the presence of a glacial refugium in the English Channel region. Under this scenario N. integer must have survived the Last Glacial Maximum in the palaeoriver system present in that region.     

GEOGRAPHICAL VARIATION OF THE PARASITE, PHYLLOBOTHRIUM DELPHINI (CESTODA), IN BALL'S PORPOISE, PHOCOENOIDES DALLI, IN THE NORTHERN NORTH PACIFIC, BERING SEA, AND SEA OF OKHOTSK

Prevalence of the larval cestode, Phyllobothrium delphini , was estimated from 2,445 Dall's porpoise, Phocoenoides dalli , from the incidental take of the Japanese high seas salmon drift-net fishery in the northwestern North Pacific Ocean, Bering Sea, and a local hand harpoon fishery in the southern Sea of Okhotsk. Prevalence of P. delphini was 22.7% in the northwestern North Pacific Ocean and 1.4% in the Bering Sea. This parasite was not found in the southern Sea of Okhotsk. Geographical differences in the prevalence of P. delphini may be due, at least in part, to regional differences in abundance of elasmobranchs known to feed on marine mammals and suspected as hosts of the parasite. Estimated intensity of infection of individual porpoises by P. delphini was low (estimated mean intensity of 3.5 plerocercoids per animal). This is a low intensity of infection compared to other species of small cetaceans studied and may be due to both differences in regional abundance of elasmobranchs and the comparatively short life span of P. dalli.     

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%.     

The influence of spawning and nursery grounds on the distribution of sole Solea solea (L.) in the Irish Sea, Bristol Channel and adjacent areas

In the coastal waters of west England and Wales, sole, Solea solea (L.), are found in greatest abundance in the north-east Irish Sea and the Bristol Channel. Data from research and charter vessel surveys in these areas showed differences in the spatial distribution of age-groups between the two regions. Juvenile sole (0-group, 1-group and 2-group) were found almost exclusively in the shallow (<20 m deep) parts of the north-east Irish Sea and adults (3-group and older) were also found in the same area and in deeper water. In contrast, while juvenile sole in the Bristol Channel were abundant in and around Carmarthen Bay, Swansea Bay and the Severn estuary, they also occurred frequently in deeper areas of the Bristol Channel down to depths of 40 m. Adult sole in the Bristol Channel were also relatively more abundant at depths >40 m than adults in the Irish Sea. The movements of sole which produce these patterns of distribution in the two areas are also shown by the recaptures of sole tagged in the respective nursery areas. The differences in the distribution of sole in the two areas may be related to the depths of their spawning grounds and the distances that maturing fish must travel to reach them.

In the north-east Irish Sea, sole spawning grounds are generally in waters of <40 m depth and within an extensive area of relatively shallow and gently shelving sediments close to the coastal nursery grounds. The bottom topography of the Bristol Channel is more steeply shelving and the spawning grounds of sole are at a depth of 40–75 m off Trevose Head, ≈100 km from the main nursery areas. The hydrographic conditions in the two areas are described, and it is concluded that one of the more important requirements of a successful spawning ground is the presence of suitable hydrographic conditions to transport eggs and larvae to nursery areas.     

Dynamics of fish diseases in the lower Elbe River

As part of a survey on population dynamics and ecology of fishesin the Elbe River, seasonal and regional fluctuations of external fish diseases were studied from the open North Sea to Hamburg in 1981–1982. Clinical signs of 11 different diseases, several of them not being recorded before, were noted in 22 fish species. Averaged over all samples, the total disease prevalence was below 1 % in 16 species. Highest prevalences were found in smelt (12.7 %), eel (9.2 %), and flounder (5.5 %). The frequency of most diseases increased in larger (older) fish. High prevalences of skeletal abnormalities in cod could be related to abnormal migration habits of diseased fish. Spawning papillomatosis, skin ulceration, and fin rot in smelt occurred predominantly during the spawning season. Most diseases observed occurred at relatively high prevalences in the central Elbe estuary between Cuxhaven and Brunsbüttel. This is most obvious for lymphocystis, fin rot, skin ulceration, and bleaching syndrome in flounder, but such a tendency also seems to occur in cauliflower disease of eel, as well as spawning papillomatosis and pharyngeal granuloma in smelt. This area is less heavily polluted and less frequently affected by oxygen deficiency than the area upstream of Glückstadt, where diseases in general occurred at lower frequencies. Therefore, it is concluded that neither pollution nor lack of oxygen are the main triggers for the outbreak of diseases in Elbe fish. It is supposed that large tidal fluctuations of salinity are a major stress factor for fish in the estuary between Cuxhaven and Brunsbüttel. Flounder from this area usually are in a relatively bad nutritional state. Their condition factor increases significantly towards Hamburg, while their disease prevalence decreases in the same direction.     

Post-pleistocene demographic history of the North Atlantic endemic Irish moss Chondrus crispus: glacial survival, spatial expansion and gene flow

Range expansions and gene flow as micro-evolutionary processes played a leading role in the population demographic history of marine organisms. Herein, we sequenced partial mtDNA Cox1 gene from 26 assigned geographical populations to understand how Irish moss (Chondrus crispus) responded to severe climatic oscillations during the Pleistocene glaciations and contemporary forces such as gene flow. Phylogeographic patterns indicated that haplotype frequency distributions were strongly skewed, with nearly half found only in single samples and thus restricted to a single population. Analysis of molecular variance revealed that most of the variation was within populations with no significant genetic structuring on either side of the Atlantic. Demographic analyses indicated that ISI (Irish Sea and Ireland) and NS (the North Sea) areas experienced a slight trend of increase in population size over time, whereas EC (the English Channel) area experienced expansion beginning approximately 170,000-360,000 BP. The observed complex genetic pattern of C. crispus is consistent with a scenario of multiple unrelated founding events by survival of this species in at least three putative Pleistocene refugia along the European coastline, and subsequent trans-Atlantic dispersal combined with contiguous northward population expansion predating the LGM and geographically gene flow.     

The Distribution and Ecology of Ariosoma Balearicum (Congridae) Leptocephali in the Western North Atlantic

A total of 4589 leptocephali of the congrid eel, Ariosoma balearicum, were examined from 17 cruises in the western North Atlantic Ocean. Myomere counts made on 915 of these indicated there were two ranges of number of myomeres that appear to be associated with separate spawning populations. Those with the higher range (high count: 128–137) were consistently 70–100mm in length in the Sargasso Sea from February to April and 20–80mm in length in the northern Sargasso Sea and Gulf Stream from September to October. Those with the lower range (low count: 120–130) were rare in the northern and eastern Sargasso Sea where they had consistently greater lengths than high count leptocephali and were most abundant in the Florida Current and Providence Channel. The geographic distributions of size and myomere ranges in relation to hydrography provide strong support for the hypothesis that high count eels found along the South Atlantic Bight (SAB) migrate across the Florida Current to spawn in the northwest Sargasso Sea. This migratory pattern is similar to those of Anguilla rostrata and Conger oceanicus, which use the southern Sargasso Sea for development as larvae. However, the distribution of high count leptocephali suggests that they use the entire Sargasso Sea gyre as a development area as larvae before crossing the Florida Current and recruiting to the SAB. The low count eels inhabiting the Bahamas appear to spawn near the banks and their abundance in the Providence Channel and southwest Sargasso Sea suggests most are retained close to the Bahamas. These two distinct styles of spawning, distribution and recruitment of larvae are hypothesized to be related to the different hydrographic regimes of the two juvenile habitats and the resulting constraints on growth and recruitment of larvae. Vertebral and myomere counts reported from other areas suggest there are distinct populations in other regions of the North Atlantic Ocean.     

Multi-torsking: simultaneous measurements of cod behaviour show differences between North Sea and Irish Sea stocks*

Fish behaviour can vary considerably in response to environmental and biological conditions. We used electronic Data Storage Tags (DSTs) to investigate the behaviour of cod in different environments; 74 North Sea cod and 20 Irish Sea cod were tagged and released in late-March 1999. So far, 32 tags (>2000 days of data) have been returned from the North Sea and four (>700 days) from the Irish Sea. The maximum recorded depth of a North Sea cod was 84 m, while Irish Sea cod utilised a depth range of 154 m. Active depth changes were of greater magnitude and rate in Irish Sea cod than North Sea cod. In addition, Irish Sea cod were continually active and made proportionally more active depth changes than North Sea cod, which were very active during migration (March–June) but exhibited little vertical movement during the summer months. To our knowledge, this is the first time that the behaviour patterns of cod from different stocks have been measured simultaneously over extended periods at such a fine temporal scale. In addition to describing considerable inter-stock differences in behaviour, our results suggest that while cod in the Irish Sea remain active and highly mobile throughout the year, North Sea cod reduce their movements during the summer feeding period. Our findings will contribute to the development of more biologically realistic management models used for evaluating the efficacy of fisheries technical measures, such as closed areas.     

The occurrence of Calanus finmarchicus (Gunnerus) and Calanus helgolandicus (Claus) in the western Irish Sea

The seasonal abundance of Calanus finmarchicus and Calanus helgolandicusin the North Channel and stratified region of the western IrishSea is reviewed using data collected between 1992 and 1996.Both species occur in the western Irish Sea, but were more abundantin the stratified region during spring. Increased abundanceduring May/June was attributed to an increase in copepoditestages. Calanus helgolandicus dominated in both regions, exhibitingspring and autumn peaks in abundance in the stratified region.It is argued that the presence of ripe females and naupliarstages in the stratified region is evidence of an in situ breedingpopulation, rather than advection of individuals from populationcentres outside the Irish Sea. The lack of geographical separationof the two species in the western Irish sea, and reports thatboth species occur in the Celtic Sea and Malin Shelf, limitthe use of either species as indicators of exchange processesbetween the Irish Sea and neighbouring waters.     

Bioaccumulation of lithium by marine organisms in European, American, and Asian coastal zones: microanalytic study using secondary ion emission

37 species of aquatic mammals, fish, crustacea, annelids, molluscs belonging to cephalopods, gasteropods and lamellibranchs were collected from coastal waters of France: North Sea, English Channel, Atlantic Ocean, Mediterranean Sea, from Greece: Aegian Sea, from North America: Atlantic Ocean and from Japan: Pacific Ocean, Sea of Japan and East China Sea. Microanalyses which were performed on organs and tissues, using secondary ion mass spectrometry, revealed high concentrations of lithium, which is commonly used in human therapy, but is also toxic in low amounts. The retention of this metal by the marine organisms appears as a general phenomenon independent of their biotope and geographical origin; the highest lithium levels were detected in the fish muscles (= edible part).     

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.     

Phylogeography of the common goby, Pomatoschistus microps, with particular emphasis on the colonization of the Mediterranean and the North Sea

The phylogeographical patterns of a small marine fish, the common goby, Pomatoschistus microps, were assessed at 12 sites along the northeastern Atlantic coasts and the western Mediterranean Sea. A combination of two genetic markers was employed: cellulose acetate allozyme electrophoresis (CAGE) and sequence analysis of a 289 bp fragment of the mitochondrial locus cytochrome b. Both markers were congruent in revealing significant differences between samples (global FST = 0.247 for the allozymes and PhiST = 0.437 for the mitochondrial DNA data) and a pattern of isolation-by-distance. Phylogeographical analyses yielded a shallow branching structure with four groups. Three of those were confined to the Atlantic basin and showed a star-like pattern. The fourth group contained a central haplotype occurring at the edges of the species' distribution, accompanied by a few more rare variants, which were restricted to the Mediterranean Sea. A genetic break was observed around the British Isles, with distinct haplotypes dominating at either side of the English Channel. A significantly negative correlation between the degree of genetic diversity and latitude was recorded both for mitochondrial DNA (mtDNA) and allozymes in the Atlantic basin. Gene flow analysis suggested that recolonization of the North Sea and the coasts of western Scotland and Ireland may have taken place from a glacial refugium in the Southern Bight of the North Sea. These results are discussed in the perspective of possible postglacial migration routes of marine fish along the northeastern Atlantic coasts.     

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.