Introgression and mitochondrial DNA heteroplasmy in the Baltic populations of mussels Mytilus trossulus and M. edulis

A strong clinal change in salinity occurs between the Baltic Sea and the North Sea, Atlantic Ocean, in the Danish Straits, where hybridization zone between mussels Mytilus edulis and M. trossulus has been reported. Eleven samples of mussels were studied from the Danish Straits and the inner Baltic Sea. Extensive introgression of M. edulis alleles from the North Sea into populations throughout the Baltic was ascertained for mitochondrial DNA (mtDNA) and two nuclear markers (ME15–16 and ITS). In the opposite direction, introgression of M. trossulus alleles into the M. edulis background was observed at the EFbis nuclear marker in populations from Kattegat (Danish Straits). While only M. edulis F (female) mtDNA was present in the Baltic, there were still strong differences in frequencies in the control region length variants between the Danish Straits and the inner Baltic samples, and weaker variation in coding region ND2–COIII haplotype frequencies. In the assays of the two mtDNA regions, various patterns of heteroplasmy were detected in 32% of all the studied individual mussels; this includes the presence of distinct, independently inherited M and F mitochondria in males, as well as the presence of two different distinguishable F genomes. The male-inherited M mtDNA genomes are quite common in the mussels from the Danish Straits, but very rare in males from the inner Baltic. Instead, a recombined control region variant (1r), which seems to have taken over the role of the M genome, was present in a number of specimens in the Baltic. Observations of heteroplasmy for two F genomes in some females and males confirm disruptions of the doubly uniparental inheritance mechanism in the hybrid Baltic Mytilus.     

Enzyme activities and allozyme polymorphism in two genetic types (or sibling species) of the genus Marenzelleria (Polychaeta: Spionidae) in Europe

Two populations of Marenzelleria spp. from the North Sea and the Baltic Sea that had already been characterized by allozyme electrophoresis and found genetically different in 1994 were again examined in 1997. The genetic identity (I = 1.0) examined in 1994 and 1997 shows that the structures of the populations of genetic Type I in the North Sea (Ringkobing Fjord) and genetic Type II in the Baltic Sea (Darss–Zingst Bodden chain) are stable. Three diagnostic gene loci that were determined additionally (citrate synthase, one locus; alanine aminotransferase, two loci) support the hypothesis of the existence of two sibling species of the genus Marenzelleria in Europe. In addition, the activities of ten electrophoretically separated enzyme systems were measured in the worms immediately after removal from their habitat and after they had been kept in the laboratory for 2 months. Under both conditions four enzymes showed significant differences between genetic types. The higher activity of the alanine aminotransferase of the North Sea worms (Type I) is discussed in the context of the different preferred salinities of the two Marenzelleria sibling species. Received: 21 January 1999 / Accepted: 13 July 1999     

Large-scale diversity and biogeography of benthic copepods in European waters

A large-scale database concerning benthic copepods from the Arctic, Baltic Sea, North Sea, British Isles, Adriatic Sea and Crete was compiled to assess species richness, biodiversity, communities, ecological range size and biogeographical patterns. The Adriatic showed the highest evenness and the most species-rich communities. Assemblages from the North Sea, British Isles, Baltic and Crete had a lower evenness. The British Isles were characterised by impoverished communities. The ecological specificity of copepod species showed two diverging trends: higher specificity of species in more diverse assemblages was observed in the Adriatic, North Sea and Baltic. A uniformly high species specificity disregarding sample diversity was found on Crete and in the British Isles. Benthic copepod communities showed distinct patterns that clearly fit the predefined geographical regions. Communities were distinguishable and β-diversity was found to be high around Europe, indicating a high species turnover on the scale of this investigation. The British Isles and the North Sea were found to be faunistic links to the Baltic and the Arctic.     

Phylogeography of Cerastoderma glaucum (Bivalvia: Cardiidae) across Europe: a major break in the Eastern Mediterranean

Molecular variation of the lagoon cockle Cerastoderma glaucum (Poiret, 1789) was examined across the species range along European coasts, from the northern Baltic to the Black Sea and the Caspian Sea. A major phylogeographic break in mitochondrial COI gene sequences (divergence 6.2%) separated a group of Ponto-Caspian and Aegean Sea haplotypes from those to the west of the Peloponnese peninsula in the Mediterranean and in the Atlantic-Baltic sector. A similar subdivision, yet not entirely geographically coincident, was found at 16 allozyme loci (Nei's distance D=0.15). The results imply a long-term isolation of populations in parts of the Eastern Mediterranean or Black Sea basins through the Pleistocene. The subdivision does not concur with previous views on the systematics of C. glaucum complex, but the pattern is notably similar to that described earlier from some fish species. Marked phylogeographic structuring was also found at lower level within the major Mediterranean-Atlantic phylogroup of C. glaucum, which was divided into six regional or local haplotype subgroups. Divergence of these groups may date back one or several major Pleistocene climatic cycles. Local mtDNA diversity was particularly high in a sample from the Ionian Sea, whose mitochondrial identity was of the Mediterranean-Atlantic type, while its nuclear characters were more strongly associated with the Ponto-Caspian type. Patterns of shallower, star-phylogeny type diversity within the Ponto-Caspian phylogroup and in the Baltic Sea area may represent more recent, post-glacial generation of variation.     

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     

Mitochondrial DNA analysis of harbour porpoises (Phocoena phocoena) in the Baltic Sea, the Kattegat–Skagerrak Seas and off the west coast of Norway

Incidental mortality of harbour porpoises, Phocoena phocoena, from captures in fishing gear has been documented consistently as severe in some European waters. Since fishing effort varies greatly among regions, management units must be defined for more effective conservation of this species. In this study, analysis of mitochondrial DNA (mtDNA) restriction fragments was performed to investigate the population structure of harbour porpoises in the Baltic Sea, the joint Kattegat–Skagerrak Seas and off the west coast of Norway. Mitochondrial DNA of 65 harbour porpoises collected from three regions was analysed with nine restriction enzymes. Analysis of the heterogeneity in the frequency distribution of haplotypes among provisional subpopulations revealed significant differences, which supports the recognition of three different subpopulations and thus three separate management units. Furthermore, indices of haplotypic diversity (range: 0.211 to 0.628) and nucleotide diversity (range: 0.070 to 0.168) of these provisional subpopulations were much lower than for western North Atlantic subpopulations, which is consistent with the view that harbour porpoises in European waters, particularly in the Baltic Sea, are depleted. Received: 21 August 1996 / Accepted: 19 September 1996     

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.     

Discrimination among and variation within species of Pseudocalanus based on the GPI locus

Patterns of allozyme variation at the GPI (glucose phosphate isomerase) locus showed that the recently described or clarified Pseudocalanus newmani, P. acuspes, P. moultoni and P. minutus are genetically isolated from each other even though morphologically they are very similar, and in places sympatric. This genetic isolation was inferred from the presence of private alleles in some species, and from differences in their degrees of observed heterozygosity and in the observed frequencies of shared alleles in all the species examined. Temporal variations were not detected in P. newmani, P. acuspes and P. minutus. Populations of P. newmani and P. moultoni from the Atlantic and Pacific coasts of North America have apparently diverged. No such divergence was detected among samples of P. acuspes from Nova Scotia, the Canadian Arctic and the Baltic sea, and of P. minutus from Nova Scotia and Hudson Bay. P. elongatus collected in the English Channel differed from P. acuspes from the Baltic Sea.     

Mytilus trossulus in Northern Europe

From data on allozyme, nuclear DNA and mitochondrial DNA markers, we show that the originally North Pacific/Northwest Atlantic mussel Mytilus trossulus is widespread on North European coasts, earliM. trossuluser thought to be inhabited only by Mytilus edulis. Several local occurrences of , interspersed with a dominant M. edulis, were recorded on the North Sea, Norwegian Sea and Barents Sea coasts of Norway and the Barents and White Sea coasts of Kola Peninsula in Russia. The proportion of M. trossulus genetic background observed at any one site varied from 0 to 95%. These new occurrences are not related to the previously known, introgressed M. trossulus population that occupies the Baltic Sea. The new northern occurrences retain both the F and M M. trossulus mitochondria, which have been lost from the Baltic stock. While hybridization takes place wherever M. trossulus and M. edulis meet, the extent of hybrization varies between the different contact areas. Hybrids are rare, and the hybrid zones are bimodal in the northern areas; more interbreeding has taken place further south in Norway, but even there genotypic disequilibria are higher than those in the steep transition zone between the Baltic mussel and M. edulis: there is no evidence of a collapse toward a hybrid swarm unlike in the Baltic. The Barents and White Sea M. trossulus are genetically slightly closer to the NW Atlantic than NE Pacific populations, while the Baltic mussel has unique features distinguishing it from the others. We postulate that the presence of M. trossulus in Northern Europe is a result of repeated independent inter- or transoceanic cryptic invasions of various ages, up to recent times.     

Mitochondrial DNA polymorphisms reveal additional genetic heterogeneity within the Leptasterias hexactis (Echinodermata: Asteroidea) species complex

Sea stars of the Leptasterias hexactis complex are important members of the rocky intertidal community of the temperate and subarctic North American Pacific coasts. Recent attempts to determine the genetic structure and taxonomic relationships of these sea stars have not been completely successful, due to the combination of extreme morphological variability and relative scarcity of taxonomically-informative genetic (allozyme) polymorphisms. The present study used mitochondrial DNA restriction-site polymorphisms for PCR (polymerase chain-reaction)-amplified products to provide additional information about this complex. Five restriction-site haplotypes were identified in southern Alaska and in the Puget Sound region that were allozymically distinct. Because the allozymic differences (Nei's unbiased genetic differences >0.08) are maintained in sympatry, the haplotypes behave evolutionarily as conventional species. Although most haplotypes show deficiencies of heterozygotes for polymorphic allozyme loci, there is no evidence for the existence of cryptic species within any of the haplotypes. Rather than consisting of a few polytypic and cosmopolitan species, the L. hexactis complex is apparently composed of numerous species with more restricted ranges. Determining the exact ranges and taxonomic relationships of the various species will require additional study.     

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.     

Allozyme and mitochondrial DNA variation in orange roughy,Hoplostethus atlanticus (Teleostei: Trachichthyidae): little differentiation between Australian and North Atlantic populations

Allozyme and mitochondrial DNA (mtDNA) genetic variation was compared in orange roughy (Hoplostethus atlanticus Collett) collected from waters off southern Australia and from waters about 22 000 km away in the North Atlantic west of Scotland. Samples were screened for 11 polymorphic allozyme loci and with 9 restriction enzymes. Significant heterogeneity between the two areas was detected for three allozyme loci (ADA ^*, CK ^* and GPI-1 ^*), and the overall G _ST (gene-diversity statistic) value of 1% was small but significant. Significant mtDNA haplotype heterogeneity was observed after ²- of haplotype frequencies but not after a G _ST analysis. Nucleotide sequence-diversity analysis showed very low net divergence (0.0023%) between the two samples. The Australian orange roughy had a lower allozyme heterozygosity and a lower mitochondrial DNA nucleon diversity than the North Atlantic sample. The very limited, although significant, allozyme and mitochondrial DNA heterogeneity between these areas suggests that there is some gene flow between these two populations. The species appears to be widespread, with its presence reported from the southern Pacific, southern Indian, and northern and southern Atlantic Oceans, and it is likely that gene flow between the antipodes is mediated by stepping-stone exchange between adjacent populations rather than by direct migration.     

Allozyme and mitochondrial DNA variation in Cuban populations of the shrimp Farfantepenaeus notialis (Crustacea: Decapoda)

We investigated the genetic diversity among populations of the shrimp Farfantepenaeus notialis, the most abundant penaeid species around Cuba. A total of 25 allozyme loci were analyzed in samples of shrimps from seven localities at the south central platform of the island (Ana María Gulf). Samples from three of these localities and from Batabanó Gulf and Guacanayabo Gulf at the south west and south east platforms of the island, respectively, were also characterized at the mtDNA level through sequence variation of a 2027 bp segment including part of the COI and COIII genes. Of the 25 allozyme loci studied 9 were polymorphic: Akp2, Akp3, AmyB, Est3, Gdh, GP7, and Per1, 2 and 3. In contrast to mtDNA, the pattern of allozyme variation among localities revealed strong population structuring at Ana María Gulf, with significant F _st in all pairwise comparisons. The magnitude of F _st estimates as well as the grouping pattern obtained by a UPGMA analysis based on a distance matrix indicated that the level of differentiation was concordant with the geographical position of the localities and the hydrographic regime. Homogeneity of mtDNA suggested that differentiation of allozyme loci might be due to more recent events rather than historical isolation of the sampled populations. Ana María and Guacanayabo Gulf populations were differentiated by mtDNA from Batabanó Gulf, at the southwestern end of the island. The analysis showed three restriction site differences among them, suggesting genetic isolation of the two regions. The present results also suggest that an artificial introduction of larvae from Tunas de Zaza into Batabanó Gulf, in an effort to repopulate this fishing region, may have been ineffective. Received: 13 December 1999 / Accepted: 2 October 2000     

Mitochondrial DNA lineages in the European populations of mussels (<Emphasis Type="Italic">Mytilus</Emphasis> spp.)

Marine mussels (Mytilus spp.) belong to a group of benthic species crucial to coastal ecosystems in Europe and are important for the cultivation industry. In the present study, the nuclear adhesive protein marker (Me15/16) was used for identification of Mytilus species in coastal areas, on a large geographic scale in Europe. Pure M. edulis populations were found in the White Sea and Iceland. M. edulis, M. trossulus and their hybrids were found in the Baltic Sea and the North Sea (Oosterschelde, The Netherlands). M. galloprovincialis, M. edulis and their hybrids occurred in Ireland. M. galloprovincialis populations were observed in the Sea of Azov (Black Sea), the Mediterranean and Portugal. The mitochondrial (mt) DNA coding-region ND2-COIII was studied by PCR (polymerase chain reaction) and RFLP (restriction fragment length polymorphism) assay methods. The mtDNA control region was studied by PCR. Substantial differentiation in the frequency of female haplotypes among the studied populations in Europe was observed. Despite isolation between the Mytilus taxa on a macro-geographic scale, considerable mitochondrial gene flow occurred between populations, with introgression in hybrid zones on a more local geographic scale in Europe. MtDNA of the Atlantic Iberian (Portugal) population of M. galloprovincialis was more similar to mtDNA in populations of M. galloprovincialis and M. edulis from the Atlantic coasts of the Ireland and M. edulis from the North Sea, than to M. galloprovincialis from the Mediterranean. Lower polymorphism of mtDNA in populations of the Baltic and Azov Sea mussels in comparison with other European populations was observed and can be explained by the recent history of both seas after the Pleistocene glaciation. In the M. galloprovincialis population from the Azov Sea, the presence of the male-inherited (M) genome was demonstrated for the first time by sequencing the control region and was observed at high frequency. Possible influence of mussel culture on geographic distribution of the Mytilus taxa in Europe is discussed.     

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.     

New indicator species in the Baltic zooplankton in 1972

The occurrence of zooplankton species, indicating a salinewater influx from the North Sea into the Baltic Sea in 1972, is described. Particular attention is paid to 8 species so far unknown in this region (Ectopleura dumortieri, Rosacea plicata, Tomopteris kefersteini, Aetideus armatus, Centropages typicus, Eucalanus elongatus, Metridia lucens, Electra pilosa). The appearance of these zooplankters in the Southern Baltic Sea is discussed in relation to the hhdrographic changes taking place in this area in 1972. Based on these data, assessments are made regarding the intensity of dynamic exchange processes in the Baltic and North Atlantic waters in 1972.     

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.     

Repeated trans-Arctic invasions in littoral bivalves: molecular zoogeography of the<Emphasis Type="Italic"> Macoma balthica</Emphasis> complex

From a geographical survey of allozyme variation, a history of repeated trans-Arctic invasions since the Plio-Pleistocene is suggested for circumboreal bivalves of the Macoma balthica complex. A principal genetic subdivision, involving several nearly diagnostic loci and Nei's distance D=0.6, distinguishes the clams of the NE Pacific from those of the NE Atlantic. The Pacific taxon is however also present in Europe, in disjunct isolates in the Baltic Sea and White Sea basins. Nevertheless, these populations have marked Atlantic introgressive elements in their gene pools (ca. 30%). Two further population types are recognized, one in the St. Lawrence estuary, Quebec, the other in Varangerfjorden, NE Norway; the latter appears a mixture of Pacific and Atlantic components in almost equal proportions, in local genetic equilibrium (a hybrid swarm). Populations in temperate North America fall outside the circumboreal M. balthica complex discussed here (D=1.0), and are referred to M. petalum. In a scenario of the history and evolution of the M. balthica complex and the similarly subdivided Mytilus edulis complex, the divergence between Pacific and Atlantic taxa started after an initial introduction of Pacific ancestors to the Atlantic basin, enabled by the Pliocene opening of the Bering Strait. During the Pleistocene and Holocene, the ocean basins were, for the most part, effectively isolated, but occasional re-invasions have taken place, causing secondary contacts of the diverged bivalve types on the Atlantic coasts. The recently re-invaded Pacific taxa in northern Europe now seem to thrive only in the extreme marginal environments. Exact dating of the re-invasions is not possible from current data. Apart from the divergence through isolation, hybridization and introgression have significantly molded the present affinities within the M. balthica complex. A formal taxonomic treatment of reticulate and hybridizing lineages is problematic; yet to recognize the evolutionary and systematic diversity within the M. balthica complex, a subspecies distinction between the NE Atlantic clams and those from the Pacific, Baltic and White Sea basins is suggested.Communicated by L. Hagerman, Helsingør     

Abundance,distribution and prey composition of scyphomedusae in the southern North Sea

The annual cycle of abundance and distribution of the scyphozoan medusae Aurelia aurita, Cyanea lamarckii, C. capillata and Chrysaora hysoscella were studied in the southern North Sea in 2004 and 2005. Three different patterns of seasonal occurrence of medusae were distinguished: (1) the early occurring C. lamarckii (February–August), (2) C. capillata and A. aurita (April–August) and (3) the late appearing C. hysoscella (July/August–September). Cyanea lamarckii was the most frequently encountered species in this study; its highest mean abundance was 1.8 ± 2.7 ind. 100 m⁻³. The prey spectra of C. lamarckii, C. capillata and C. hysoscella contained several copepod and other crustacean species and thus make them potential competitors with fish larvae. Medusae in this study also consumed fish eggs and larvae, including clupeids, in all months analysed. Although peak spawning of sprat (Sprattus sprattus) coincides with the maximum abundance of medusae (May–June) the relative low abundance of all medusae species in this study makes jellyfish predation unlikely to be a factor controlling sprat recruitment in the time frame investigated.