Continuous plankton records: Geographical variations in numerical abundance,biomass and production of euphausiids in the North Atlantic Ocean and the North Sea

Geographical variations in the numbers, biomass and production of euphausiids and the contribution of common species to the total are described from samples taken during 1966 and 1967 in the North Atlantic Ocean and the North Sea by the Continuous Plankton Recorder at 10 m depth. Euphausiids were most abundant in the central and western North Atlantic Ocean and the Norwegian Sea. Thysanoessa longicaudata (Krøyer) was numerically dominant. Biomass was greatest in the Norwegian Sea and the north-eastern North Sea where Meganyctiphanes norvegica (M. Sars) accounted for 81 and 59%, respectively, of the total biomass. Production was highest off Nova Scotia and in Iberian coastal waters; the dominant species were T. raschi (M. Sars) in the former area and Nyctiphanes couchi (Bell) in the latter. The mean P:B ratios were correlated with temperature.     

Colonization of Europe by two American genetic types or species of the genus Marenzelleria (Polychaeta: Spionidae). An electrophoretic analysis of allozymes

Allozyme electrophoresis was conducted in an attempt to identify the origin ofMarenzelleria sp. found in the North Sea and Baltic Sea. The analysis covered eight enzymes with ten loti from nine populations found on the North American Atlantic toast, these populations in the North Sea and five populations in the Baltic. The North Sea spionids correspond to the Type IMarenzelleria from North American coastal waters between Barnstable Harbor (Massachusetts) and Cape Henlopen (Delaware). Nei's genetic distance between these North American populations and those from the North Sea wasD = 0.010 to 0.020. TheMarenzelleria sp. found in the Baltic Sea very probably stems from North American populations of Type II found from the region of Chesapeake Bay (Trippe Bay) south to the Currituck Sound (North Carolina). The genetic distance between these North American populations and the Baltic populations isD = 0.000 to 0.001. The invaders appear to have lost little of their genetic variation while colonizing the North and Baltic Seas. Probably, both colonizing events tan be attributed to large numbers of individuals reaching Europe simultaneously on one or more occasions. In addition, aMarenzelleria Type III was found by electrophoresis among specimens from Currituck Sound (North Carolina), rohere it is sympatric withMarenzelleria Type II. Salinity is discussed as an important factor for the establishment ofMarenzelleria Type I in the North Sea and Type II in the Baltic Sea.     

Population dynamics and production of euphausiids. I. Thysanoessa longicaudata in the North Atlantic Ocean

Adults and furciliae of Thysanoessa longicaudata (Krøyer) taken in continuous plankton records (CPR) in 1966 and 1967 were examined and measured. The measurements were converted to biomass using a length/weight equation, and net production values were calculated from these data. Two generations were spawned annually in the eastern North Atlantic Ocean and only one in the colder western waters and in the Norwegian Sea. The timing of reproduction in the spring was correlated with the mean sea-surface temperature. The highest annual production values (up to 12.24 mg dry weight m^-3 year^-1) were recorded in the eastern areas, where the ratio between production and biomass was usually between 4:1 and 8:1 compared with ratios usually between 2:1 and 3:1 in the western waters. The values for daily production per individual ranged from 3.34 g dry weight day^-1 to 17.54 g day^-1. Mortality in the early stages of development was calculated as about 98.2% of the eggs produced.     

Use of seaweeds for monitoring trace elements in coastal waters

Concentrations of a wide range of trace elements: arsenic, cadmium, cobalt, chromium, hafnium, nickel, thorium, uranium, zinc and the rare earth elements, cerium, europium, samarium, terbium and ytterbium were determined by instrumental neutron activation analysis in the brown alga,Fucus vesiculosus from Eckwarder Hörne, North Sea and from Rügen, Baltic Sea. Another brown alga,Sargassum filipendula from Sri Lanka, Indian ocean (representing an unpolluted control station) was similarly investigated. Cobalt, chromium and nickel concentrations were highest inF. vesiculosus from the North Sea while zinc was highest in samples from the Baltic Sea, reflecting high levels of these elements in coastal waters of the North and the Baltic sea. Cadmium, cobalt, nickel and zinc levels were lowest inS. filipendula from Sri Lanka, probably demonstrating lower levels of those elements in coastal waters. Concentration levels of hafnium, thorium, uranium, and the rare earth elements were highest inS. filipendula. Two years later in 1994,S. filipendula along withUlva sp. (green alga) was resampled from the same sampling site, and in addition to the above elements, six other trace elements (Ag, Ba, Br, Rb, Se and Sr) were determined.Sargassium filipendula showed a particular affinity for Ag, As, Br and Sr. For the other elements, marginal concentration differences were observed betweenS. filipendula andUlva sp., probably reflecting the regional background levels. Substantially higher concentrations of Hf, Th, U, and the rare earths were found again in the 1994Sargassum andUlva samples, reflecting the effect of a substrate rich in rare earth elements. The brown algae used in this study may be used to monitor trace elements in coastal waters.     

Population dynamics and production of euphausiids

Seasonal changes in abundance, size and aspects of the population structure of Meganyctiphanes norvegica (M. Sars) and Nyctiphanes couchi (Bell) are described from samples taken with the Continuous Plankton Recorder at 10 m depth over a 2 yr period (1966 and 1967) in the North Atlantic Ocean and the North Sea. M. norvegica lived for a maximum of just over 2 yr, and adults of both year-classes spawned during a limited breeding season in the spring or summer. N. couchi spawned over a prolonged breeding season, giving rise to a complex of cohorts with overlapping size ranges. It was concluded that 3 or 4 cohorts were spawned in each year and that the maximum life span was probably greater than 1 yr, although maturity may be attained in less than a year. Estimated annual production at 10 m depth for M. norvegica ranged from 0.80 to 18.74 mg m^-3yr^-1 and for N. couchi from 0.67 to 8.23 mg m^-3yr^-1. P:B ratios ranged from 1.3:1 to 6.3:1 for M. norvegica and 4.0:1 to 5.5:1 for N. couchi.     

Dissolved nutrient dynamics along the southwest coastal waters of India during northeast monsoon: a case study

Dissolved nutrients, Chl-a and primary productivity were measured from seven transects along the coastal waters of the southeastern Arabian Sea during northeast monsoon. Ten major estuaries were chosen to study the influence of estuarine discharge on the nutrient dynamics in the coastal waters. The mean water discharge of the estuaries in the north (64.8?±?18?×?10⁵?m³?d^?1) was found to be higher than those in the south (30.6?±?21.4?×?10⁵?m³?d^?1), whereas the nutrient concentrations were found to be higher in the estuaries of the south. The results from the offshore waters were discussed in accordance with the depth contour classification, that is, shelf (depth?≤?30?m) and slope waters (depth?≥?30?m). Our results suggest that the estuarine discharge plays a major role in the nutrient distribution in near shore shelf waters, whereas in shelf and slope waters, it was mainly controlled by in situ biological processes. The inorganic form of N to P ratios were found to be higher than Redfield ratio in slope waters when compared with shelf waters, suggesting that PO₄^3? (<0.15?µmol?L^?1) is a limiting nutrient for primary production. The multivariate statistical analysis revealed that the nutrient dynamics in the coastal waters was controlled by both biological and physical processes.     

Plankton of the fladen ground during FLEX 76 III. Vertical distribution,population dynamics and production of Calanus finmarchicus (Crustacea: Copepoda)

Samples taken in the northern North Sea with the Continuous Plankton Recorder (CPR), the Undulating Oceanographic Recorder (UOR), the Longhurst Hardy Plankton Recorder (LHPR) and by our colleagues from other participating Institutes during the Fladen Ground Experiment (FLEX 76) were used to describe the vertical distribution and population dynamics of Calanus finmarchicus (Gunnerus) and to provide estimates of the production and carbon budget of the population from 19 March to 3 June, 1976. Total production of the 19 March to 3 June, 1976. Total production of the nauplii and copepodite stages (including adults), during the exponential growth phase in May, was estimated to be in the range of 0.49 to 0.91 g C m^-2 d^-1 or 29.0 to 55 g dry wt m^-2 (14.5 to 27.8 g C m^-2) for the three successive 10 d periods in May. Two gross growth efficiencies (K ₁) (20 and 34%), together with the lower value of C. finmarchicus production, were used to calculate the gross ingestion levels of algae as 2.45 and 1.44 g C m^-2 d^-1 (73.5 and 43.2 g C m^-2 over the May period). These ingestion levels, together with the algae ingested by other zooplankton species, are greater than the estimated total phytoplankton production of 45.9 g C m^-2 over the FLEX period. A number of factors are discussed which could explain the discrepancies between the production estimates. One suggestion is that the vertical distribution of the development stages of this herbivorous copepod and their diel and ontogenetic migration patterns enable it to efficiently exploit its food source. Data from the FLEX experiment indicated that the depletion of nutrients limited the size of the spring bloom, but that it was the grazing pressure exerted by C. finmarchicus which was responsible for the control and depletion of the phytoplankton in the spring of 1976 in the northern North Sea.JONSDAP Contribution No. 51     

Seasonal occurrence of the ctenophore Pleurobrachia pileus in the western Dutch Wadden Sea

Due to the short residence time of the water in the western Dutch Wadden Sea the ctenophore Pleurobrachia pileus has to be considered as an allochthonous species, which enters the estuary passively from the North Sea. The seasonal occurrence of this species in the Dutch Wadden Sea is a reflection of its abundance in the coatal zone of the North Sea and is characterized by a pronounced spring peak of up to 17000 individuals per 1000 m³ and a lower autumn peak with numbers up to 400 individuals per 1000 m³. Similar to other areas, predation by Beroë gracilis, another ctenophore species, may be a factor in controlling the P. pileus population in this region, although B. gracilis abundance is low. P. pileus is only important as a predator for a short time, i.e. the month of May, when it reaches maximum biomass values of up to 12 00 mgC per 1000 m³. Both the estimated potential consumption rates in comparison with the available food as well as the observed suboptimal growth rate of the population suggest that in the coastal zone of the North Sea area food conditions may not be optimal for P. pileus.     

Plankton of the fladen ground during FLEX 76 II. Population dynamics and production of Thysanoessa inermis (Crustacea: Euphausiacea)

Samples taken in the northern North Sea with the Continuous Plankton Recorder (CPR), the Undulating Oceanographic Recorder (UOR) and the Longhurst-Hardy Plankton Recorder (LHPR) during the Fladen Ground Experiment in 1976 (FLEX 76) are used to describe the vertical distribution and population dynamics of Thysanoessa inermis (Krøyer) and to provide estimates of the production and carbon budget of the population from 19th March to 3 June 1976. Spawning occurred in late April and early May, in near synchronisation with the start of the spring bloom of phytoplankton. Eggs, nauplii and calyptopes reached maximum abundance in succession, and furciliae were numerous when sampling ceased in early June. Adults increased in length from a mean of 12.1 mm in mid-March to 17.5 mm in early June and the estimated production was 2.40 mg m^-3 over the 74 d period. Total carbon ingested by the population of T. inermis was estimated to be 10 mg C m^-2 d^-1 in the upper 100m which was only 1.5% of the daily primary production of 0.68 gC m^-2 measured over the FLEX period 26 March to 4 June 1976. The grazing by T. inermis on the phytoplankton population was assumed to have little effect on the control and depletion of the spring phytoplankton bloom during FLEX 77.JOSDAP Contribution No. 50     

The population structure of the harbour porpoise,Phocoena phocoena,in Danish waters and part of the North Atlantic

A study of the homogeneity between putative stocks of Phocoena phocoena in the North Sea and inner Danish waters was performed using isozyme electrophoresis. Two polymorphic systems, mannose phosphate isomerase (MPI) and phosphoglucomutase (PGM) were used in the analysis. A two-locus homogeneity test showed an effect of season and locality on the genotypic distribution. This led to a division of the total sample into season and into the two localities, the inner Danish waters (IDW) and the North Sea. The samples were then compared to the Hardy-Weinberg expectations where a deficit in heterozygotes was observed in the North Sea sample in both systems and at the Pgm locus in the total sample, indicating a mixing of sub-populations. A hierarchical contingency table analysis, also based on the genotypic distribution, showed a combined effect of season and locality and only an insignificant effect of sampling period, thus supporting the division into season and locatily. The seasonal effect is further supported by earlier observations of a seasonal migration of harbour porpoises out of the Baltic in winter. A significant difference was observed between the two supposed sub-populations, IDW-summer and North Sea-summer samples, based on the genotypic distribution. This could also be a reflection of the sample sizes. On the basis of samples obtained from Canada, West Greenland and Holland a preliminary study of the population structure on a larger scale, comprising the North East and North West Atlantic, was performed. The results so far accord well with the hitherto accepted sub-populations of P. phocoena in the North Atlantic.     

Induction of sexual reproduction and resting egg production in Brachionus plicatilis by a diet of salt-grown Nannochloris oculata

Rates of population increase in early spring and the sizes of overwintering stocks were calculated for the planktonic copepods Pseudocalanus elongatus and Acartia clausi for a set of areas covering the open waters of the north-east Atlantic Ocean and the North Sea for the period 1948 to 1979. For both species, the rates of population increase were higher in the open ocean than in the North Sea and appear to be related to temperature. The overwintering stocks in the North Sea were larger than those in the open ocean and are probably related to phytoplanton concentration. P. elongatus shows higher overwintering stocks and lower rates of population increase than A. clausi, resulting in different levels of persistence in the stocks of the two species. It is suggested that this difference in persistence is responsible for differences between the two species with respect to geographical distribution in summer and different patterns of year-to-year fluctuations in abundance.     

Calculating carbon biomass ofPhaeocystis sp. from microscopic observations

Conversion factors for calculating carbon biomass ofPhaeocystis sp. colonies and free-living cells were determined from microscopic observations and chemical analysis conducted on cultured and naturalPhaeocystis sp. populations originating from the Southern Bight of the North Sea in 1986 and 1987. They allow calculation, in terms of carbon biomass, of the different forms ofPhaeocystis sp. that succeed each other when the population is growing, on the basis of microscopic observations. The latter include enumerations of free-living cells (flagellated and non-motile) and colonies, as well as colonial biovolume measurement. Specific application to natural populations from Dutch coastal waters during spring 1986 shows that more than 90% ofPhaeocystis sp. carbon biomass is under colonial form, most of it exceeding the grazing characteristics of current zooplankton at this period of the year. Detailed analysis of seasonal changes shows in addition that the size of the colonies greatly increases during the course ofPhaeocystis sp. flowering, reaching sizes as high as 1 mm diameter at the top of the bloom when nutrients are depleted. Physiologically this corresponds to an enhanced synthesis of mucilaginous substances, with the decrease of available nutrients leading to an increasing contribution of the matrix to the total colonial carbon during the course of the bloom. Carbon content ofPhaeocystis sp. colonies therefore greatly varies with their size, ranging from 0.3 to 1430 ngC colony^–1.     

Temporary cyst formation of<Emphasis Type="Italic"> Heterocapsa triquetra</Emphasis> (Dinophyceae) in natural populations

Heterocapsa triquetra (Ehrenberg) Stein is a phototrophic marine dinoflagellate with wide coastal distribution. It is known to be capable of mixotrophy and diel vertical migration. The species was particularly abundant in the Gulf of Finland (the Baltic Sea) during the summers of 1996 and 1998, leading to discolouration of water on the south-west coast of Finland. Large-scale (50 m³) coastal mesocosm experiments in the north-west Gulf of Finland (the Baltic Sea) in the summers of 1996 and 1998 with daily mineral nutrient additions provoked a biomass increase of phytoplankton dominated by H. triquetra. From the first days of the experiment temporary cysts of H. triquetra were found in the bottom sediment water of the mesocosms. Maximum temporary cyst production rates reached values up to 20×10⁶ cysts m^–2 day^–1, accounting for <1% of the depth-integrated motile population size. The environmental features favouring temporary cyst production remain uncertain; zooplankton grazing and nutrient stress are potential factors. Temporary cysts of H. triquetra were observed in a unialgal culture (f/2 medium) isolated in summer 1999 from Eel Pond (Woods Hole, Mass., USA).Communicated by M. Kühl, Helsingør     

Strong variability in bacterioplankton abundance and production in central and western Bay of Bengal

With large influx of freshwater that decreases sea-surface salinities, weak wind forcing of <10 m s⁻¹ and almost always warm (>28°C) sea-surface temperature that stratifies and shallows the mixed layer leading to low or no nutrient injections into the surface, primary production in Bay of Bengal is reportedly low. As a consequence, the Bay of Bengal is considered as a region of low biological productivity. Along with many biological parameters, bacterioplankton abundance and production were measured in the Bay of Bengal during post monsoon (September–October 2002) along an open ocean transect, in the central Bay (CB, 88°E) and the other transect in the western Bay (WB). The latter representing the coastal influenced shelf/slope waters. Bacterioplankton abundances (<2 × 10⁹cells l⁻¹) were similar to those reported from the HNLC equatorial Pacific and the highly productive northern Arabian Sea. Yet, the thymidine uptake rates along CB (average of 1.46 pM h⁻¹) and WB (average of 1.40 pM h⁻¹) were less than those from the northwestern Indian Ocean. These abundances and uptake rates were higher than those in the oligotrophic northwestern Sargasso Sea (<7 × 10⁸ cells l⁻¹; av 1.0 pM h⁻¹). Concentrations of chlorophyll a (chl a), primary production rates and total organic carbon (TOC) were also measured for a comparison of heterotrophic and autotrophic production. In the WB, bacterioplankton carbon biomass equaled ∼ 95% of chl a carbon than just 31% in the CB. Average bacterial:primary production (BP:PP) ratios accounted for 29% in the CB and 31% in the WB. This is mainly due to lower primary productivity (PP) in the WB (281 mg C m⁻² d⁻¹) than in the CB (306 mg C m⁻² day⁻¹). This study indicates that bacteria–phytoplankton relationship differs in the open (CB) and coastal waters (WB). Higher abundance and contrastingly low bacterial production (BP) in WB may be because of the riverine bacteria, brought in through discharges, becoming dormant and unable to reproduce in salinities of 28 or more psu. Heterotrophic bacteria appear to utilize in situ DOC rather rapidly and their carbon demand is ∼50% of daily primary production. It is also apparent that allochthonous organic matter, in particular in the western Bay, is important for meeting their carbon demand.     

Mesozooplankton communities in the Aegean and Black Seas: a comparative study

An attempt was made for a comparative study of the mesozooplankton communities between the Aegean and Black Seas. These areas are in connected through the Sea of Marmara, the Bosporus and the Dardanelles straits. The comparison was mainly based on data collected both in coastal and offshore waters, by the Hellenic Centre for Marine Research (Greece), the Shirshov Institute of Oceanology (Russia), the Institute of Biology of Southern Seas (Ukraine) and the Institute of Oceanology (Bulgaria). Our data revealed important differentiation in terms of species composition and interannual fluctuations of zooplankton quantity. Among the common copepod species, those presenting an eurytherm and/or euryhaline character (e.g. Acartia clausi, Paracalanus parvus, Penilia avirostris, Podon polyphemoides) were found dominant in the Black Sea and common in the Aegean Sea. Decreased dissimilarity of species composition seems to exist between the North Aegean Sea and the Black Sea, as well as between the coastal areas of the Black Sea and those affected by pollution in the Aegean Sea. A common feature of both seas is the existence of spatial differentiation in terms of the quantity of zooplankters due to anthropogenic and/or natural influences. The pattern of this differentiation seems to vary interannually in the Black Sea, depending either on the abundance of mesozooplankton predators (indigenous or invasive species) or on global climatic changes. Although the Black Sea is generally richer in zooplankton than the Aegean Sea, the significant decrease in the quantity of zooplankton from the early 1980s to the mid-1990s, due to the above factors, resulted in quasi-similar values in both areas. Black Sea mesozooplankton has revealed strong interannual variability both quantitatively and qualitatively, whereas no significant variability was detected in the Aegean Sea. This difference could be attributed both to variant anthropogenic forcing on these ecosystems and to the sensitive character of the semi-enclosed Black Sea.Communicated by M.E. Vinogradov, Moscow     

Polymorphism of mannose phosphate isomerase in North Sea and Baltic Sea populations of the amphipods Gammarus zaddachi and G. salinus

Genetic variation at the mannosephosphate isomerase (MPI) locus was examined in the euryhaline sibling species Gammarus zaddachi Sexton and G. salinus Spooner. Both crustacean amphipods share identical enzyme mobilities, following electrophoresis on vertical starch gels. The MPI locus turned out to be highly polymorphic; it is encoded by 6 alleles in G. zaddachi and 7 alleles in G. salinus. Geographic variation of allelic diversity was studied in samples from 9 G. zaddachi and 10 G. salinus populations, primarily obtained from Baltic Sea and North Sea sites. Patterns of inter- and intraspecific heterogeneity are described. Differences in allelic composition exist between Baltic and North Sea samples of G. salinus. In G. zaddachi, levels of polymorphism are higher in North Sea populations than in those from Baltic Sea areas. The significance of these findings is discussed in the light of previous biochemical genetic investigations on the population structure of the two amphipods considered.     

Are there physiological races of the sea anemone Diadumene luciae?

The chemical activators of feeding behaviour for coelenterates have been generally regarded as species specific. Recently, however, and English (North Sea) population of the sea anemone Diadumene (Haliplanella) luciae (Verrill) and an American (Californian) population of the same species were found to respond to different chemical activators. The results of further examination of this apparent anomaly show that a third population (from the English Channel, selected because of differences in colour variety and type of habitat from the North Sea population) shared the same activators as the other English population. There was no correlation between the identities of the feeding-behaviour activators and either the type of habitat or coloration of the three populations. Since the only physical factor shared by anemones with similar activators (i.e., the two English populations) was geographical, it is tentatively concluded that physiological races of D. luciae may be evolving, independently of morphological or differently coloured races, in different parts of the world.     

Seasonal population structure,vertical distribution and migration of the chaetognath Sagitta elegans in the Celtic Sea

The biology of the chaetognath Sagitta elegans Verrill has been much researched, but detailed studies of population structure have generally been conducted in coastal water where dynamic tidal conditions may cause difficulty in interpretation of data. The resolution of sampling examining vertical distribution and diurnal migration has also been rather coarse. During a series of eight cruises to a seasonally thermally stratified sampling site in the Celtic Sea in 1978 and 1979, detailed vertical zooplankton profiles were taken to study the seasonal population structure, vertical distribution and migration of this species. The overwintering stock of S. elegans (22 to 52 individuals m^-2, 0 to 90 m) had a wide range of lengths (5 to 20 mm) and matured in 1978 from early March, spawning several times before dying out by late July. Young produced by the overwintering stock started to mature in July and population numbers reached their highest in August (2483 m^-2, 132.8 mg C m^-2) when sea temperature peaked (17.1°C). By October, the population of S. elegans declined (284 m^-2), which was thought to be due to a combination of lower sea-water temperature, competition for and availability of food, and predation. Because of the length range of the overwintering population (5 to 20 mm), it is assumed that reproduction continued at a low level over the winter, although eggs were not found in January and February, the coldest months of the year. In summer, the smallest S. elegans (2 to 6 mm) were found in the near-surface waters and did not migrate, but as their lengths increased they occupied deeper depth ranges and a portion of the population started to migrate diurnally. Individuals which did not migrate and stayed in the warmer surface waters, or those which migrated into it, matured faster than those remaining in the colder water below the thermocline. Migration to surface waters by mature individuals seemed to be stopped by high surface temperatures (17°C) and a sharp thermocline (3 C°). As sea temperature increased during the year from the winter minimum of 7.7°C, S. elegans matured at a progressively shorter length (14 mm in March 1978 to 10 mm in August). There are probably only three generations of S. elegans a year in the Celtic Sea.     

Hermaphroditism in brown shrimp: lessons from field data and modelling

This study was undertaken to further clarify whether the brown shrimp, Crangon crangon (Linnaeus 1758), is a gonochorist, a facultative or an obligate hermaphrodite. Juvenile shrimps were sampled from intertidal habitats along the German Wadden Sea coast with a push net and from a power plant water inlet to quantify the share of primary females. Length-based sex ratios were determined for about 27,000 individuals using external characteristics. Observed sex ratios were mainly female-biased, and also large males occurred regularly in the catch. This indicates that sex at hatch is not male as would be characteristic for an obligate protandric hermaphrodite and that not all male shrimps change sex. A cohort-based computer simulation, including sex-specific growth rates, mortality and seasonally varying recruitment, generated sex ratios comparable to the field. The observed decline in the proportion of males with increasing size can be explained solely by faster growth of females without involving hermaphroditism. Based on temperature-dependent growth and moult rates as well as length-specific numbers of eggs per female, the potential egg production of primary and secondary females was modelled, yielding contributions of secondary females of <1%. Sex change in C. crangon has previously been observed and may be interpreted as an evolutionary relict of this species having evolved in a habitat characterized by lower population densities, lower predation levels and increased longevity compared to today’s living conditions in North Sea coastal waters.     

Population genetic analysis of Ensis directus unveils high genetic variation in the introduced range and reveals a new species from the NW Atlantic

We report current genetic variation of populations of the razor shell Ensis directus (Conrad 1843) (Mollusca: Bivalvia: Pharidae) in native (North American) and introduced (European) ranges using nuclear and mitochondrial sequence-based markers. We expected less variation within the introduced range, especially considering the frequent mass mortality events observed in Europe since the species was recorded for the first time in 1978. However, we found higher variation in Europe. The possible significance of temporal fluctuations of genetic variation, limited effect of random genetic drift, and multiple introductions are discussed. Interestingly, the multiple-introduction hypothesis contrasts with the gradual colonisation of European coastal waters but is supported by trained clustering analysis and by the intensity of transatlantic shipping. Genetic and morphometric evidence strongly supports that examined individuals from a supposed E. directus population from Newfoundland (Canada) belong to a separate species. This new Ensis is formally described here and named E. terranovensis n.sp.