Microsatellite analysis of plaice (Pleuronectes platessa L.) in the NE Atlantic: weak genetic structuring in a milieu of high gene flow

Many species of marine fish are typified by large population sizes, strong migratory behavior, high fecundity, and pelagic eggs and larvae that are passively transported by ocean currents, all features that tend to increase gene flow, and hence reduce genetic partitioning, among localized populations. The plaice, Pleuronectes platessa, is a commercially important demersal species that exhibits all of these characteristics. We analyzed genetic variation at eight microsatellite loci in samples of spawning adults (N = 348) from the coasts of Ireland, Iceland, and, for the first time, from the Baltic Sea. Significant differentiation was observed between Iceland and Irish and Baltic Sea samples. However, there were no genetic differences between Irish and Baltic Sea samples, which contrast with the significant differentiation reported between Baltic Sea and North Sea/Atlantic populations of other flatfish species. To increase the data set, we carried out a cross-calibration exercise, allowing us to perform a joint analysis of data with an earlier study on adult and juvenile plaice (N = 480) collected over a broad geographic range, using six microsatellite loci in common to the two studies. Significant differentiation was observed between fish collected at the northern (Iceland, Faeroes, Norway) and southern (Bay of Biscay) parts of the species range. In contrast, the results showed little evidence of genetic structuring over much of the continental shelf of Europe. We believe that bathymetric and hydrographic barriers are the major factors shaping genetic structure, while lack of structure over much of the European continental shelf may be explained by a combination of past historical events, population structure, and dynamics of the species.     

Genetic variability and level of differentiation in North Sea and Baltic Sea populations of the green alga<Emphasis Type="Italic"> Cladophora rupestris</Emphasis>

Cladophora rupestris is a perennial filamentous macroalga belonging to the Chlorophyta. It is widely distributed on both sides of the northern Atlantic Ocean and penetrates into the brackish Baltic Sea down to ca. 4 psu salinity. In this paper we present evidence for genetic differentiation of a Baltic form of this marine alga. We assessed genetic structure within and among 11 populations ranging along a salinity gradient from the Norwegian coast to the northern Baltic Sea proper. Samples of 328 individuals were studied using starch-gel protein electrophoresis to evaluate genetic variability and interpopulation differentiation based on allozymes. Of 11 loci examined, only one was polymorphic. For this locus, encoding superoxide dismutase (SOD-3), a total of seven alleles were distinguished. We found two genetically differentiated groups of populations of C. rupestris, one Baltic Sea group and one North Sea group, with a distinct border in the southern Kattegat near the entrance to the Baltic Sea. The genetic differentiation for SOD-3, expressed as pairwise F_ST values between the populations, was generally higher within the Baltic Sea group (0.10-0.43) than within the North Sea group (0.05-0.10); in the latter group also fewer pairs of populations differed significantly. Pairs of populations from different groups had the highest F_ST values (0.20-0.60). Hierarchical analysis of variance showed that 29.6% of the total variation in the SOD-3 locus was explained by variation between the two groups, while only 4.2% was explained by variation among the populations within the groups. The remaining variation (66.2%) was found within the populations.     

Comparative study on the population genetics of the red algae <Emphasis Type="Italic">Furcellaria lumbricalis</Emphasis> occupying different salinity conditions

Furcellaria lumbricalis is a red algae occurring in low salinity to fully marine conditions. Here, both putatively neutral and EST-derived microsatellite markers were developed and used to examine the genetic structure of northern European populations inhabiting different salinity conditions ranging from 35 to 3.6 psu. The amount of genetic variation did not differ between ocean and brackish populations, but differences were observed between marker types; EST-derived markers possessed less variation and showed greater differentiation than the putatively neutral microsatellites. No multicopy multilocus genotypes were detected despite expected asexuality in brackish populations. The Bayesian STRUCTURE analysis, when conducted for expressed marker data, indicated the presence of two main clusters, the Atlantic Ocean and Baltic Sea, while no clear structuring was observed based on putatively neutral microsatellites. The moderate level of genetic differentiation at neutral loci is probably due to genetic drift, a feasible explanation considering long distances between many populations, while the high level of differentiation in EST-linked markers reflects selection pressures.     

Molecular genetic variation in tarpon (<Emphasis Type="Italic">Megalops atlanticus</Emphasis> Valenciennes) in the northern Atlantic Ocean

The tarpon (Megalops atlanticus) is a highly valued game fish and occasional food fish in the eastern and western Atlantic Ocean. Tarpon have a high capacity for dispersal, but some regional biological differences have been reported. In this study we used two molecular genetic techniques—protein electrophoresis of nuclear DNA loci, and restriction fragment length polymorphism analysis of the mitochondrial DNA (mtDNA)—to assess this species population genetic structure in the eastern (coastal waters off Gabon and Sierra Leone, Africa) and western (coastal waters off Florida, Caribbean Sea) Atlantic Ocean north of the equator. Genetic differentiation was observed between tarpon from Africa and tarpon from the western Atlantic Ocean. A unique allele and haplotype, significant differences in allozyme allele and mtDNA haplotype frequencies between the African and western Atlantic samples, and significant F_ST analyses suggest that levels of gene flow between tarpon from these two regions is low. Among the western Atlantic Ocean collections, genetic diversity values and allele and haplotype frequencies were similar. AMOVA analyses also showed a degree of genetic relatedness among most of the western Atlantic Ocean collections: however, some significant population structuring was detected in the allozyme data. A regional jackknifed F_ST analysis indicated the distinction of the Costa Rica population from the other western Atlantic populations and, in pairwise analyses, F_ST values tended to be higher (i.e., genetic relatedness was lower) when the Costa Rican sample was paired with any of the other western Atlantic samples. These data suggest that Costa Rican tarpon could be partially isolated from other western Atlantic tarpon populations. Ultimately, international cooperation will be essential in the management of this species in both the eastern and western Atlantic Ocean.Communicated by P.W. Sammarco, Chauvin     

Genetic diversity of blackspot seabream (<Emphasis Type="Italic">Pagellus bogaraveo</Emphasis>) populations off Spanish Coasts: a preliminary study

Genetic diversity among four natural samples of Blackspot seabream (Pagellus bogaraveo, Brünnich, 1768) from different fishing grounds exploited by Spanish fisheries was analyzed through the use of 12 microsatellite markers. The samples were captured off the Spanish coasts from the Mediterranean Sea to the Cantabrian Sea within the same continental slope. High levels of genetic diversity were revealed for every population and every locus was polymorphic at the 0.95 level. The average number of alleles, average heterozygosity and PIC were found to be 15.75, 0.833 and 0.818, respectively. In general, population differentiation was not detected in these samples. Through AMOVA, a low level of variation between regions (Mediterranean vs. Atlantic samples) was observed, though this was not significant. A larger percentage of total variation was observed inside the ‘within populations’ class. Thus, AMOVA did not reveal any significant population substructure. Moreover, no correlation was found between geographical and F_ST estimates and the observed results did not allow the improvement of a model of isolation by distance. The high homogeneity revealed by means of these markers could indicate the absence of physical frontiers between the geographical areas analyzed in this survey, especially between Atlantic and Mediterranean areas.     

Genetic analysis of tuna populations,<Emphasis Type="Italic"> Thunnus thynnus thynnus</Emphasis> and<Emphasis Type="Italic"> T</Emphasis>.<Emphasis Type="Italic"> alalunga</Emphasis>

The genetic population structures of Atlantic northern bluefin tuna ( Thunnus thynnus thynnus) and albacore ( T. alalunga) were examined using allozyme analysis. A total of 822 Atlantic northern bluefin tuna from 18 different samples (16 Mediterranean, 1 East Atlantic, 1 West Atlantic) and 188 albacore from 5 samples (4 Mediterranean, 1 East Atlantic) were surveyed for genetic variation in 37 loci. Polymorphism and heterozygosity reveal a moderate level of genetic variability, with only two highly polymorphic loci in both Atlantic northern bluefin tuna ( FH* and SOD- 1*) and albacore ( GPI- 3* and XDH*). The level of population differentiation found for Atlantic northern bluefin tuna and albacore fits the pattern that has generally been observed in tunas, with genetic differences on a broad rather than a more local scale. For Atlantic northern bluefin tuna, no spatial or temporal genetic heterogeneity was observed within the Mediterranean Sea or between the East Atlantic and Mediterranean, indicating the existence of a single genetic grouping on the eastern side of the Atlantic Ocean. Very limited genetic differentiation was found between West Atlantic and East Atlantic/Mediterranean northern bluefin tuna, mainly due to an inversion of SOD- 1* allele frequencies. Regarding albacore, no genetic heterogeneity was observed within the Mediterranean Sea or between Mediterranean and Azores samples, suggesting the existence of a single gene pool in this area.     

Evolutionary genetics of the Mytilus edulis complex in the North Atlantic region

Genetic relationships among Mytilus populations throughout the North Atlantic region, including the Mediterranean and the Baltic Sea, were studied using enzyme electrophoresis. Three distinct groups of populations, each of a remarkably wide distribution, can be recognised on the basis of their multilocus allelic composition: (1) M. galloprovincialis L. of the Mediterranean and western Europe; (2) a genetically distinct form of M. edulis Lmk. from both the Baltic Sea and some localities in the Canadian Maritime Provinces (here provisionally termed the trossulus type mussel); and (3) the traditional Atlantic M. edulis populations of northwestern European coasts and most of eastern North America. These groups are regarded as representing three relatively old evolutionary lineages, which all deserve separate and equal systematic status. The main part of the differentiation at most of the loci studied is accounted for by this major systematic pattern, but considerable geographical differentiation within each of the three principal groups was also detected. At single loci, different electromorphs were found to prevail in disjunct populations of M. galloprovincialis (Mediterranean/Britain) and of the trossulus-type mussel (Baltic/eastern Canada). Within the Atlantic M. edulis, a major part of the differentiation is transoceanic. At one locus (Ap), geographic differentiation appeared to be relatively independent of the systematic boundaries; the possible role of interlineage hybridisation in contact areas in regulating the pattern of geographical variation is discussed.     

Genetic structure of the flounders Platichthys flesus and P. stellatus at different geographic scales

The genetic structure of the flounders Platichthys flesus L. and P. stellatus Pallas was investigated on different spatial scales through analysis of allozyme variation at 7 to 24 polymorphic loci in samples collected from different regions (Baltic Sea, North Sea, Brittany, Portugal, western Mediterranean, Adriatic Sea, Aegean Sea and Japan) in 1984 to 1987. No geographic variation was evident within a region. Some pattern of differentiation by distance was inferred within the Atlantic, while the Mediterranean comprised three geographically isolated populations and was itself geographically isolated from the Atlantic (fixed allele differences at up to three loci were found among P. flesus populations from the Atlantic, the western Mediterranean, the Adriatic Sea, the Aegean Sea and also P. stellatus from the coast of Japan). Sea temperature during the reproductive period probably acts as a barrier to gene flow between populations. Genetic distances among European flounder populations (P. flesus) were higher than, or of the same magnitude as, the genetic distance between Pacific (P. stellatus) and European flounder populations, suggesting that P. flesus is paraphyletic and/or there is no phylogenetic basis to recognising P. stellatus as a different species. The divergence between P. flesus and P. stellatus was thus inferred to be more recent than the divergence between the present P. flesus populations from the NE Atlantic and eastern Mediterranean. The eastern Mediterranean populations are thought to originate from the colonisation of the Mediterranean by a proto-P. flesus/P. stellatus ancestor, whereas the present western Mediterranean population has undergone a more recent colonisation event by P. flesus. Patterns of mitochondrial DNA variation, established on a smaller array of P. flesus samples, were in accordance with the geographic patterns inferred from the allozyme survey. In addition, they supported the hypothesis of a two-step colonisation of the western Mediterranean. These results contribute to our understanding of the biogeography of the Mediterranean marine fauna, especially the group of boreal remnants to which P. flesus belongs. Received: 7 February 1997 / Accepted: 26 March 1997     

Conservation genetics of <Emphasis Type="Italic">Lumnitzera littorea</Emphasis> (Combretaceae), an endangered mangrove,from the Indo-West Pacific

Mangrove forests, with their ecological significance and economic benefits, are vital inter-tidal wetland ecosystems. Lumnitzera littorea (Combreataceae) is a non-viviparous mangrove distributed in tropical Asia and North Australia. Due to natural and human impacts, populations of this species have been isolated, fragmented, and highly disturbed. In China, L. littorea is an endangered species, restricted to small regions of Hainan Island. The genetic composition of five populations of this species from the Indo-West Pacific (South China, Malay Peninsula, Sri Lanka, North Australia) was assessed using inter simple sequence repeat (ISSR) makers. At the species level, expected mean heterozygosity (He) was 0.240 with 75.6% of loci polymorphic (P). However, genetic variation was much lower at the population level (P = 37.1%, He = 0.118). A high coefficient of gene differentiation (Gst = 0.515) and low level of gene flow (Nm = 0.470) indicated significant genetic differentiation among populations. AMOVA also indicated that more than half the total variation (58.4%) was partitioned among populations. The high degree of differentiation observed among populations emphasizes the need for appropriate conservation measures that incorporate additional populations into protected areas, and achieve the restoration of separate, degraded populations.     

Genetic variation between geographic populations of the amphipods Gammarus zaddachi and G. salinus

Spatial and temporal patterns of gene-enzyme variation were estimated in the sibling species Gammarus zaddachi Sexton and G. salinus Spooner by starch gel electrophoresis. Twenty-one G. zaddachi and 18 G. salinus populations from coastal and estuarine areas in the Baltic Sea, North Sea and other localities of north-western Europe were surveyed. Both amphipods display similar electrophoresis patterns of the enzyme systems studied. Considerable interspecific and interpopulational differences were detected in allele frequencies at three highly polymorphic loci, phosphoglucose isomerase (PGI), glutamate oxalacetate transaminase (GOT) and arginine phosphokinase (APK). G. zaddachi exhibits a pronounced genetic heterogeneity in most areas of the sampled range. Populations from northern French and western English coasts differ significantly from the other samples in allele frequencies at the PGI or APK locus, respectively. Baltic populations are widely uniform in their genetic composition but can be distinguished from samples taken at North Sea sites in allele frequencies at the APK locus. The latter reveal a clinal variation, ranging from the Danish to the French coast. In contrast to G. zaddachi, a low degree of genetic differentiation was observed among the G. salinus populations examined. This indicates that migration and interregional mixing may be more important in maintaining the genetic structure than in G. zaddachi which, compared to G. salinus, prefers habitats of lower salinity levels. Evidently, less extensive dispersal capabilities owing to the confinement of G. zaddachi to brackish waters of dilute salt concentrations may account for a diminished gene flow and considerable genetic separation of local populations. This assumption is supported by the genetic homogeneity documented in Baltic G. zaddachi populations. In view of the low and constant salinities in wide areas of this brackish-water sea such barriers do not exist. Survey studies performed with selected populations over a 3-yr period demonstrated a general pattern of temporal constancy in the allozyme variation observed.     

Interspecific variation in thermal denaturation of proteins in the congeneric musselsMytilus trossulus andM. galloprovincialis: evidence from the heat-shock response and protein ubiquitination

The genetic population structure of the precominant zooplankter, the copepod Calanus finmarchicus (Gunnerus), was examined to determine whether genetically distinct populations exist in the Gulf of Maine. C. finmarchicus was sampled in three regions of the Gulf of Maine (Great South Channel, spring 1989; northern Gulf of Maine, winter 1990; Great South Channel and Georges Bank, spring 1990). Copepods from seven locations in the Great South Channel, five in the northern Gulf of Maine and four on or near Georges Bank were assayed for allozyme variation and mitochondrial DNA variation of amplified 16S rRNA and cytochrome b genes. Restriction fragment length polymorphism (RFLP) analyses of both mitochondrial DNA genes revealed no variation among any of the individuals assayed. Analysis of five polymorphic allozyme loci revealed that genetic variation among the three geographic regions was low, and genetic identities were high between all locations (I>0.97). Most of the genetic variation was among locations regardless of region. Chi-square tests were used to examine genetic similarity between specific pairs of locations within and between regions. In the northern Gulf of Maine, genetic homogeneity occurred over larger spatial scales (hundreds of km) than in either the Great South Channel or Georges Bank (tens of km). Only copepods from the Bay of Fundy and Nova Scotian Shelf locations were genetically distinct from Wilkinson Basin copepods at two loci. Copepod populations from the northern locations may have been partially isolated or they may represent immigrant populations (e.g., from the Gulf of St. Lawrence). Several pairs of locations were genetically distinct at one or more loci in the two southern regions. Differences between locations in these regions may represent distinct populations advected into the areas at different times or from different sources (e.g., genetic variation may represent a mixture of genetically distinct northern and southern copepod populations). These results suggest extensive gene flow among populations of C. finmarchicus in the Gulf of Maine with some evidence of genetic population subdivision near the Gulf's northeastern and southern boundaries.     

Genetic differentiation in Hediste diversicolor (Polychaeta: Nereididae) for the North Sea and the Baltic Sea

Genetic differentiation between North Sea and Baltic Sea Hediste diversicolor (O.F. Müller, 1776) (Polychaeta: Nereididae) populations was studied by allozyme electrophoresis on starch gel. Thirteen loci were analyzed in eight populations. The level of genetic variation was very low (mean H _o = 0.000 to 0.015). Differentiation between H. diversicolor populations is quite high (F _ST = 0.892) and reflected by three enzyme loci (MDH-I*, MDH-II*, IDH-I*). The reduced gene flow (N _m<1) may be explained by the limited dispersal capacity of the species. Regardless of whether found in the North Sea or Baltic Sea, there appear to be two different genetic types which are parapatric or sympatric in some places. The two types hybridize at three localities, but no signs of hybridization have been found at one (Tallinn). Received: 26 June 1997 / Accepted: 10 September 1997     

Genetic heterogeneity among <Emphasis Type="Italic">Eurytemora affinis</Emphasis> populations in Western Europe

Evolutionary diversification of the broadly distributed copepod sibling species complex Eurytemora affinis has been documented in the northern hemisphere. However, the fine scale geographic distribution, levels of genetic subdivision, evolutionary, and demographic histories of European populations have been less explored. To gain information on genetic subdivision and to evaluate heterogeneity among European populations, we analyzed samples from 8 locations from 58° to 45°N and 0° to 23°E, using 549 base pairs of the mitochondrial cytochrome oxidase subunit I (COI) gene. We discovered three distinct lineages of E. affinis in Western Europe, namely the East Atlantic lineage, the North Sea/English Channel (NSEC) lineage, and the Baltic lineage. These geographically separated lineages showed sequences divergence of 1.7–2.1%, dating back 1.9 million years (CI: 0.9–3.0 My) with no indication of isolation by distance. Genetic divergence in Europe was much lower than among North American lineages. Interestingly, genetic structure varied distinctively among the three lineages: the East Atlantic lineage was divided between the Gironde and the Loire populations, the NSEC lineage comprised one single population unit spanning the Seine, Scheldt and Elbe rivers and the third lineage was restricted to the Baltic Proper (Sweden). We revealed high haplotype diversity in the East Atlantic and the Baltic lineages, whereas in the NSEC lineage haplotype diversity was comparatively low. All three lineages showed signs of at least one demographic expansion event during Pleistocene glaciations that marked their genetic structure. These results provide a preliminary overview of the genetic structure of E. affinis in Europe.     

Icy heritage: ecological evolution of the postglacial Baltic Sea reflected in the allozymes of a living fossil,the priapulid Halicryptus spinulosus

Genetic variation of 16 allozyme loci in 397 Halicryptus spinulosus (Priapulida) revealed overall polymorphism of P=0.438 and Hardy-Weinberg expectations for heterozygosity of H _e=0.060 for Baltic Sea stocks, H _e=0.143 for the White Sea and H _e=0.121 for Iceland. Maximal unbiased standard distances of D=0.0693 separated Baltic and White Sea populations. Nordic and Baltic populations could be distinguished by allozymes, but Baltic subsamples proved cohesive. Gene flow amounted to effective exchange values per generation of N _m=2.94 over 650 km of continuous habitat, N _m=10.65 over 175 km, and N _m=13.85 over 20 km. Gene flow started to decrease with geographic distance beyond a dispersal threshold of 20 km, but hierarchical G _ST-statistics indicated light isolation by distance beyond a minimum of 8 km. Gene flow is high for a benthic worm assumed to lack dispersal by pelagic larva, a paradox which cannot be resolved now. Baltic populations are characterized by lower heterozygosity than Nordic stocks. In the Baltic Basin, temporally continuous brackish-water conditions have only existed for the past 7000 years. The two possible colonization routes of H. spinulosus to the geologically young Baltic Sea imply genetic drift, whether by founder effect (sweepstake colonization from Iceland) or by refugial bottlenecking during the Ancylus phase of the Baltic Basin after a direct connection to the White Sea had been sequestered. Continued genetic drift is emphasized by lower heterozygosity in the ecologically unstable Belt Sea compared to the central Baltic. Allozymes falsify the reduced-mutability hypothesis to explain bradytelic evolution of Priapulida. Regional genetic homogeneity, ample polymorphism, and preference for anoxic black mud qualify H. spinulosus populations as indicators of microevolutionary responses to water circulation regimes or pollution in the Baltic Sea.     

Are there two species of the polychaete genus Marenzelleria in Europe?

North Sea and Baltic Sea populations of the introduced polychaete Marenzelleria viridis (Verrill, 1873) reproduce at different times (spring and autumn, respectively). Enzyme separation by starch gel electrophoresis revealed major differences between specimens from the Baltic Sea and those from the North Sea (collected in 1992 and 1993) but a high degree of homogeneity among populations from the same sea. Three enzyme loci, glucose-6-phosphate isomerase (GPI-A, GPI-B) and malate dehydrogenase (MDH), were fixed to 100% by different alleles in the North and Baltic Sea populations, respectively. Different alleles are dominant for mitochondrial aspartate aminotransferase (mAAT) with allele frequencies of ca. 0.97 in all sampled populations from the North Sea and Baltic Sea, respectively, but heterozygotes were found in all populations. These genetic differences could be due to environmentally induced selection or genetically different origins of the populations, suggesting that populations of the genus Marenzelleria in the North and Baltic Seas may be two different species.     

Microsatellite variation and significant population genetic structure of endangered finless porpoises (Neophocaena phocaenoides) in Chinese coastal waters and the Yangtze River

The finless porpoise (Neophocaena phocaenoides) inhabits a wide range of tropical and temperate waters of the Indo-Pacific region. Genetic structure of finless porpoises in Chinese waters in three regions (Yangtze River, Yellow Sea, and South China Sea) was analyzed, including the Yangtze finless porpoise which is widely known because of its highly endangered status and unusual adaptation to freshwater. To assist in conservation and management of this species, ten microsatellite loci were used to genotype 125 individuals from the three regions. Contrary to the low genetic diversity revealed in previous mtDNA control region sequence analyses, relatively high levels of genetic variation in microsatellite profiles (H _E = 0.732–0.795) were found. Bayesian clustering analysis suggested that finless porpoises in Chinese waters could be described as three distinct genetic groups, which corresponded well to population “units” (populations, subspecies, or species) delimited in earlier studies, based on morphological variation, distribution, and genetic analyses. Genetic differentiation between regions was significant, with F _ST values ranging from 0.07 to 0.137. Immigration rates estimated using a Bayesian method and population ancestry analyses suggested no or very limited gene flow among regional types, even in the area of overlap between types. These results strongly support the classification of porpoises in these regions into distinct evolutionarily significant units, including at least two separate species, and therefore they should be treated as different management units in the design and implementation of conservation programmes.     

Genetic evidence for cryptic speciation in the marine bryozoan Alcyonidium gelatinosum

The common subtidal marine bryozoan Alcyonidium gelatinosum (L.) is one of the most confusing species of a taxonomically very difficult genus. It shows massive variation for almost all morphological characters. A number of sympatric colonies collected off the coast of Lundy (Bristol Channel) could be clearly divided on morphological grounds into two quite distinct types. Using starch gel electrophoresis the products of a range of genetic enzyme loci were compared between the two morphotypes and also between these and other Bristol Channel populations of A. gelatinosum. Minimal differences were observed between one Lundy morphotype and the other allopatric populations, which it was consequently concluded were conspecific. The second morphotype showed considerable genetic differentiation [genetic identity (Nei, 1972) =0.417] from the other Lundy population, with no common alleles at some loci. The probability of obtaining the observed results by chance in a single population is significantly low (P<10^–14). It is therefore concluded that the two morphotypes of A. gelatinosum from Lundy are not conspecific.     

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.     

The influence of temperature on the development of Baltic Sea sprat (Sprattus sprattus) eggs and yolk sac larvae

In spring 2004 and 2005 we performed two sets of experiments with Baltic sprat (Sprattus sprattus balticus Schneider) eggs and larvae from the Bornholm Basin simulating ten different temperature scenarios. The goal of the present study was to analyse and parameterise temperature effects on the duration of developmental stages, on the timing of important ontogenetic transitions, growth during the yolk sac phase as well as on the survival success of eggs and early larval stages. Egg development and hatching showed exponential temperature dependence. No hatching was observed above 14.7°C and hatching success was significantly reduced below 3.4°C. Time to eye pigmentation, as a proxy for mouth gape opening, decreased with increasing temperatures from 17 days post hatch at 3.4°C to 7 days at 13°C whereas the larval yolk sac phase was shortened from 20 to 10 days at 3.8 and 10°C respectively. Maximum survival duration of non-fed larvae was 25 days at 6.8°C. Comparing the experimental results of Baltic sprat with existing information on sprat from the English Channel and North Sea differences were detected in egg development rate, thermal adaptation and in yolk sac depletion rate (YSDR). Sprat eggs from the English Channel showed significantly faster development and the potential to develop at temperatures higher than 14.7°C. North Sea sprat larvae were found to have a lower YSDR compared to larvae from the Baltic Sea. In light of the predictions for global warming, Baltic sprat stocks could experience improved conditions for egg development and survival.     

Genetic structure and allozyme variation of sea bass (Dicentrarchus labrax and D. punctatus) in the Mediterranean Sea

This paper reports data on 28 allozyme loci in wild and artificially reared sea bass (Dicentrarchus labrax) samples, originating from either coastal lagoon or marine sites in the Mediterranean Sea. F _ST analysis (θ estimator) indicated strong genetic structuring among populations; around 34% of the overall genetic variation is due to interpopulation variation. Pairwise θ estimates showed that, on average, the degree of genetic structuring was much higher between marine populations than between samples from lagoons. Six polymorphic loci showed differences in allele frequencies between marine and lagoon samples. Multivariate analyses of individual allozymic profiles and of allele frequencies suggested that different arrays of genotypes prevail in lagoons compared to marine samples, particularly at those loci that, on the basis of previous acclimation experiments, had been implicated in adaptation to freshwater. On the other hand, variation at “neutral” allozyme loci reflects to a greater extent the geographic location of populations. Allozyme differentiation was also studied in a D. labrax population from the Portuguese coast. Average genetic distance between this population and the Mediterranean populations was quite high (Nei's D = 0.236) and calls into question the taxonomic status of the Portuguese population. Finally, genetic relationships between D. labrax and D. punctatus were evaluated. Average Nei's D was 0.648, revealing high genetic differentiation between the two species, even for two sympatric populations of these species in Egypt; thus gene flow was not indicated between species. Received: 24 October 1996 / Accepted: 27 November 1996