Population genetic structure of escolar (<Emphasis Type="Italic">Lepidocybium flavobrunneum</Emphasis>)

Escolar (Lepidocybium flavobrunneum) is a large, mesopelagic fish that inhabits tropical and temperate seas throughout the world, and is a common bycatch in pelagic longline fisheries that target tuna and swordfish. Few studies have explored the biology and natural history of escolar, and little is known regarding its population structure. To evaluate the genetic basis of population structure of escolar throughout their range, we surveyed genetic variation over an 806 base pair fragment of the mitochondrial control region. In total, 225 individuals from six geographically distant locations throughout the Atlantic (Gulf of Mexico, Brazil, South Africa) and Pacific (Ecuador, Hawaii, Australia) were analyzed. A neighbor-joining tree of haplotypes based on maximum likelihood distances revealed two highly divergent clades (δ = 4.85%) that were predominantly restricted to the Atlantic and Indo-Pacific ocean basins. All Atlantic clade individuals occurred in the Atlantic Ocean and all but four Pacific clade individuals were found in the Pacific Ocean. The four Atlantic escolar with Pacific clade haplotypes were found in the South Africa collection. The nuclear ITS-1 gene region of these four individuals was subsequently analyzed and compared to the ITS-1 gene region of four individuals from the South Africa collection with Atlantic clade haplotypes as well as four representative individuals each from the Atlantic and Pacific collections. The four South Africa escolar with Pacific mitochondrial control region haplotypes all had ITS-1 gene region sequences that clustered with the Pacific escolar, suggesting that they were recent migrants from the Indo-Pacific. Due to the high divergence and geographic separation of the Atlantic and Pacific clades, as well as reported morphological differences between Atlantic and Indo-Pacific specimens, consideration of the Atlantic and Indo-Pacific populations as separate species or subspecies may be warranted, though further study is necessary.     

Global population divergence of the sea star Hippasteria phrygiana corresponds to the onset of the last glacial period of the Pleistocene

Genetic structure and connectivity of populations of the globally distributed and eurybathic sea star Hippasteria phrygiana (Parelius 1768) were studied in 165 individuals sampled from three oceanic regions: the North Pacific Ocean, the South Pacific Ocean (considered to include the adjacent regions of the Southern Ocean and the southern Indian Ocean) and the North Atlantic Ocean. A nuclear gene region (ATP synthase subunit α intron #5, ATPSα) and a mitochondrial gene region (cytochrome oxidase subunit I, COI) were amplified and sequenced. Significant heterogeneity was detected in an AMOVA analysis among the three sampled oceanic regions for COI, but not for ATPSα. Neither gene showed significant genetic heterogeneity within the North Atlantic, as assessed by ΦST values. Significant heterogeneity was detected for COI (but not ATPSα) in the North Pacific, but the converse was true in the South Pacific. Coalescent simulations suggested that the three regions have been diverging with little or no gene flow for the past 50–75,000 years, a time frame that corresponds to the onset of the last glacial period of the Pleistocene. A possible genetic signature of recent population expansion (or non-neutrality) was detected for each gene in the North Pacific, but not in the other two oceanic regions.     

Genetic stock structure of the swordfish (Xiphias gladius ) inferred by PCR-RFLP analysis of the mitochondrial DNA control region

Restriction fragment length polymorphism (RFLP) analysis was performed on PCR amplified DNA fragments containing the control region of the swordfish (Xiphias gladius Linnaeus, 1758) mitochondrial DNA. A total of 456 individuals comprising 13 local samples (six Pacific, three Atlantic, the Mediterranean Sea, two Indian Ocean and the Cape of Good Hope) were surveyed with four endonucleases (Alu I, Dde I, Hha I and Rsa I), yielding a total of 52 composite genotypes. Within-sample genotypic diversity (H) was high ranging from 0.702 to 0.962 with a value of 0.922 for the pooled sample. Significant geographic variation in the frequencies of genotypes and restriction patterns was revealed. The Mediterranean sample was highly distinct from all other samples. Further, Rsa I digestion revealed high levels of polymorphism in all but the Mediterranean samples, indicating that exogenous swordfishes rarely enter that body of water. Heterogeneity between the North and South Atlantic samples was significant, both of which differed from those of the Pacific. In contrast, the Indian Ocean samples were not significantly different from the samples of South Atlantic and Pacific. Genetic differentiation among the Pacific samples was low. The results indicate that the world-wide swordfish population is genetically structured not only among, but also within ocean basins and suggest that gene flow is restricted despite the absence of geographic barriers. Received: 28 August 1996 / Accepted: 2 October 1996     

Mitochondrial cytochrome b variation in sleeper sharks (Squaliformes: Somniosidae)

Sleeper sharks are a poorly studied group of deep-sea sharks. The subgenus, Somniosus, contains three morphologically similar species: S. microcephalus found in the Arctic and North Atlantic; S. pacificus in the North Pacific; and S. antarcticus in the Southern Ocean. These sharks have been reported mainly in temperate to polar waters and occasionally in subtropical locations. They have not been recorded in tropical waters. This study investigates the relationships among the accepted species of Somniosus through analysis of mitochondrial cytochrome b sequence variation. Seventy-five samples were examined from four sampling locations in the North Pacific, Southern Ocean and North Atlantic. Twenty-one haplotypes were found. A minimum spanning parsimony network separated these haplotypes into two divergent clades, an S. microcephalus and an S. pacificus/antarcticus clade, strongly supporting the distinction of S. microcephalus as a separate species from the Pacific sleeper shark species. Analysis of genetic structure within the S. pacificus/antarcticus clade (analysis of molecular variance, allele frequency comparisons, and a nested clade analysis) showed limited or no differences amongst three populations. Further examination of genetic variation at more variable mtDNA and nuclear markers is needed to examine the species status of S. pacificus and S. antarcticus.     

Mitochondrial control region variability and global population structure in the swordfish,Xiphias gladius

Little is known about what limits genetic exchange in highly vagile, open ocean vertebrate species, such as the swordfish Xiphias gladius L. Reduced abundance of swordfish in some regions, and increased fishing pressure in others, has raised concerns and fueled interest in a more complete evaluation of the resource. In this study, global population structure in swordfish was assessed by sequencing a 300 base pair segment of the 5 end of mitochondrial DNA control region from 159 swordfish collected in three ocean basins: the Mediterranean, Atlantic and Pacific, over the years 1988 to 1994. Among the 159 individuals, 95 polymorphic sites delineated 121 unique haplotypes, indicating a high level of polymorphism on a global scale. A phylogenetic analysis of the unique DNA haplotypes revealed two divergent clades with differing geographic distributions. Phylogeographic concordance of this pattern with that of two other pelagic fish species suggests a biogeographic explanation for this structure. An analysis of molecular variance (AMOVA) revealed significant geographic partitioning of molecular variation among the three ocean basins, indicating that swordfish populations are structured on a global scale. Estimates of genetic exchange among populations within an ocean basin were high, indicating panmixia within ocean basins. Since the haplotypic diversity exhibited by the swordfish control regions is extremely high, much larger sample sizes may be necessary to detect subdivision within ocean basins.     

Global population structure of albacore (Thunnus alalunga) inferred by RFLP analysis of the mitochondrial ATPase gene

The genetic population structure of the highly migratory albacore (Thunnus alalunga) was investigated using restriction fragment length polymorphism (RFLP) analysis of the mitochondrial ATPase gene amplified by the polymerase chain reaction (PCR). 620 individuals comprising 13 geographically distant samples (ten Pacific, two Atlantic and the Cape of Good Hope) were surveyed between 1991 and 1994 with two restriction endonucleases (Mse I and Rsa I), resulting in seven haplotypes. No heterogeneity was observed in the distribution of haplotypes among the ten samples from the North and South Pacific, nor among the samples from North and South Atlantic and Cape of Good Hope. However, highly significant heterogeneity was evident among Atlantic and Pacific samples. Higher haplotypic diversity (h) was observed in the Pacific samples (0.59 to 0.69) than in the Atlantic and Cape samples (0.22 to 0.43). These results suggest greater gene flow between albacore of the northern and southern hemispheres (within oceans) than between the Atlantic and Pacific Oceans.     

Population structure of the planktonic copepod Calanus pacificus in the North Pacific Ocean

The pelagic copepod Calanus pacificus ranges nearly continuously across temperate-boreal regions of the North Pacific Ocean and is currently divided into three subspecies—C. pacificus oceanicus, C. pacificus californicus, C. pacificus pacificus—based on subtle morphological differences and geographic location. The relation between geography and genetic differentiation was examined for 398 C. pacificus individuals sampled from six widely distributed locations across the North Pacific, including an open ocean site and coastal sites on both sides of the North Pacific basin. For each individual copepod, the DNA sequence was determined for a 421-bp region of the mitochondrial coxI gene (mtCOI). A total of sixty-three different mtCOI sequences, or haplotypes, were detected, with a sequence divergence between haplotypes of 0.2–3.1%. The number and distribution of haplotypes varied with sampling location; 12 haplotypes were distributed across multiple sampling locations, and 51 occurred at only one location. Five genetically distinct populations were detected based on F _ST values. Haplotype minimum spanning networks, nucleotide divergence and F _ST values indicated that individuals from coastal sites in the North Pacific Ocean were more closely related to each other than to individuals from the open ocean site at Station P. These results provide genetic support for the designation of two subspecies—a coastal subspecies that consists of what is currently referred to as C. p. pacificus and C. p. californicus and an open ocean subspecies C. p. oceanicus. This work also indicates that planktonic copepods with potentially high dispersal capacity can develop genetically structured populations in the absence of obvious geographic barriers between proximate locales within an ocean basin.     

Population structure of the planktonic copepod <Emphasis Type="Italic">Calanus pacificus</Emphasis> in the North Pacific Ocean

The pelagic copepod Calanus pacificus ranges nearly continuously across temperate-boreal regions of the North Pacific Ocean and is currently divided into three subspecies—C. pacificus oceanicus, C. pacificus californicus, C. pacificus pacificus—based on subtle morphological differences and geographic location. The relation between geography and genetic differentiation was examined for 398 C. pacificus individuals sampled from six widely distributed locations across the North Pacific, including an open ocean site and coastal sites on both sides of the North Pacific basin. For each individual copepod, the DNA sequence was determined for a 421-bp region of the mitochondrial coxI gene (mtCOI). A total of sixty-three different mtCOI sequences, or haplotypes, were detected, with a sequence divergence between haplotypes of 0.2–3.1%. The number and distribution of haplotypes varied with sampling location; 12 haplotypes were distributed across multiple sampling locations, and 51 occurred at only one location. Five genetically distinct populations were detected based on F _ST values. Haplotype minimum spanning networks, nucleotide divergence and F _ST values indicated that individuals from coastal sites in the North Pacific Ocean were more closely related to each other than to individuals from the open ocean site at Station P. These results provide genetic support for the designation of two subspecies—a coastal subspecies that consists of what is currently referred to as C. p. pacificus and C. p. californicus and an open ocean subspecies C. p. oceanicus. This work also indicates that planktonic copepods with potentially high dispersal capacity can develop genetically structured populations in the absence of obvious geographic barriers between proximate locales within an ocean basin.     

Colonization of the northwest Atlantic by the blue mussel, <Emphasis Type="Italic">Mytilus trossulus</Emphasis> postdates the last glacial maximum

Blue mussels in the genus Mytilus first arrived in the Atlantic Ocean from the Pacific during the Pliocene, following the opening of the Bering Strait. Repeated periods of glaciation throughout the Pleistocene led to re-isolation of the two ocean basins and the allopatric divergence of Mytilus edulis in the Atlantic and M. trossulus in the Pacific. Mytilus trossulus has subsequently colonized the northwest Atlantic (NW Atlantic) so that the two species are presently sympatric and hybridize throughout much of the Canadian Maritimes and the Gulf of Maine. To estimate when M. trossulus arrived in the NW Atlantic, we have examined sequence variation within a portion of the female mtDNA lineage large untranslated region (F-LUR) for 156 mussels sampled from three Pacific and eleven Atlantic populations of M. trossulus. Although we found no evidence of reciprocal monophyly for Pacific and NW Atlantic M. trossulus, limited gene flow between ocean basins has led to the divergence of unique sequence clades within each ocean basin. In contrast, relative genetic homogeneity indicates high levels of gene flow within each basin. Coalescence-based analysis of the F-LUR sequences suggests that M. trossulus recolonized the NW Atlantic from the northeast Pacific subsequent to a demographic expansion in the Pacific that occurred ~96,000 years before present (ybp). Estimates of timing of divergence for Pacific and NW Atlantic populations and the time since expansion among NW Atlantic sequence clades indicate that M. trossulus arrived in the NW Atlantic more recently, between 20,000 and 46,000 ybp. Given that these estimates overlap with the dates of peak ice in the NW Atlantic during the last glacial maximum (LGM, ~18,000–21,000 ybp), we suggest that colonization of the NW Atlantic by M. trossulus occurred during, but more likely just subsequent to, the LGM and was followed by rapid temporal and spatial expansion in the region.     

Deconstructing an assemblage of “turtle” barnacles: species assignments and fickle fidelity in Chelonibia

Barnacles in the genus Chelonibia are commensal with a variety of motile marine animals including sea turtles, crustaceans, and sirenians. We conducted a worldwide molecular phylogenetic survey of Chelonibia collected from nearly all known hosts to assess species relationships, host-fidelity, and phylogeographic structure. Using DNA sequences from a protein-coding mitochondrial gene (COI), a mitochondrial rRNA gene (12S), and one nuclear rRNA gene (28S), we found that of four species, three (C. testudinaria, C. patula, and C. manati) are genetically indistinguishable. In addition, we show each utilizes a rare androdioecious mode of reproduction involving complemental males. In contrast, the fourth species (C. caretta), which is hermaphroditic and specializes on turtles, is genetically distinct—leading to the conclusion that the three former taxa are morphotypes of the same species and should be synonymized under C. testudinaria. Phylogenetic analysis resulted in three geographic clades (Atlantic, Indian Ocean/western Pacific, and eastern Pacific) with haplotype parsimony networks revealing no shared haplotypes among geographic regions. Analysis of molecular variance detected significant differences among sequences by region (p < 0.005); conversely, there were no significant differences among sequences when grouped by host or taxonomic designation. Average pairwise genetic distances were lower between the eastern Pacific and Atlantic clades (0.053 ± 0.006) than between the eastern Pacific and Indian Ocean/western Pacific clades (0.073 ± 0.008), suggesting Atlantic and eastern Pacific populations were connected more recently, perhaps until the rise of the Isthmus of Panama. Host use by Chelonibia morphotypes is discussed along with speculation on possible ancestral hosts and support for a “turtle-first” hypothesis.     

Genetic similarity between samples of the orange roughyHoplostethus atlanticus from the Tasman Sea,South-west Pacific Ocean and North-east Atlantic Ocean

Gel electrophoresis was used to measure genetic variation in the orange roughyHoplostethus atlanticus. Samples were collected on the continental slope in the Tasman Sea, South-west Pacific Ocean and North-east Atlantic Ocean, at various periods from 1982 to 1984. Twenty-two enzymatic loci were resolved in seven samples to give observed heterozygosities between 0.104±0.037 and 0.125±0.044. There was little genetic differentiation between populations separated by a distance of approximately 21 000 km. Tasman-Pacific and Atlantic samples differed significantly in allele frequency at only two loci, while at a third locus a rare allele was found only in the Atlantic Ocean.     

Mitochondrial DNA sequence variation and phylogeography of oceanic insects (Hemiptera: Gerridae: Halobates spp.)

Relatively few insects have invaded the marine environment, and only five species of sea skaters, Halobates Eschscholtz (Hemiptera: Gerridae), have successfully colonized the surface of the open ocean. All five species occur in the Pacific Ocean, H. germanus White also occurs in the Indian Ocean, whereas H. micans Esch- scholtz is the only species found in the Atlantic Ocean. We sequenced a 780 bp long region of the mitochondrial cytochrome oxidase subunit I gene (COI) for a total of 66 specimens of the five oceanic Halobates species. Our purpose was to investigate the genetic variation within species and estimate the amount of gene flow between populations. We defined 27 haplotypes for H. micans and found that haplotype lineages from each of the major oceans occupied by this species are significantly different, having sequences containing five to seven unique base substitutions. We conclude that gene flow between populations of H. micans inhabiting the Atlantic, Pacific, and Indian Ocean is limited and hypothesize that these populations have been separated for 1 to 3 million years. Similarly, there may be limited gene flow between H. germanus populations found in the Pacific and Indian Ocean and between H. sericeus populations inhabiting the northern and southern parts of the Pacific Ocean. Finally, we discuss our findings in relation to recent hypotheses about the influence of oceanic diffusion on the distribution and population structure of oceanic Halobates spp. Received: 29 July 1999 / Accepted: 23 November 1999     

Divergence of mitochondrial DNA lineage of the rocky intertidal goby Chaenogobius gulosus around the Japanese Archipelago: reference to multiple Pleistocene isolation events in the Sea of Japan

Geographic variations in the mitochondrial cytochrome b gene sequence (1,062 nucleotides) of the rocky intertidal goby Chaenogobius gulosus were examined and compared against the phylogeographic pattern of its sympatric sister species Chaenogobius annularis, which we studied previously. We collected C. gulosus specimens from 17 localities around the Japanese Archipelago and Korean Peninsula. Thirty-one haplotypes were obtained. The phylogenetic tree showed three geographical lineages, of which one was distributed along the coast of the Sea of Japan, suggesting that a Pleistocene isolation event in the Sea of Japan affected the genetic divergence of this species, similar to that which was observed in C. annularis. The net nucleotide divergence between the Sea of Japan and Pacific Ocean lineages in C. gulosus was much lower than that of C. annularis. Both traditional molecular clock and hierarchical approximate Bayesian computational models suggest that different divergence events affected the genetic divergences of C. gulosus and C. annularis. Our study on the sister species pair strongly supports multiple isolation events in the Sea of Japan, leading to varying levels of genetic divergence between the Pacific Ocean and Sea of Japan lineages in the Japanese coastal marine species.     

55Fe in Pacific Ocean plankton

The specific activity of ⁵⁵Fe in plankton increased from below detectable limits at 20°N to a maximum of 130 nCi/g Fe at 20°S and then decreased to 5 nCi/g Fe at 50°S. Along an east-west track near 15°S a peak of 90 nCi/g Fe occurred at 135°W. It is suggested that this maximum is the result of fallout from French nuclear tests at Mururoa Atoll (22°S; 139°W). The increase in ⁵⁵Fe at high latitudes observed in previous studies in the North Atlantic Ocean and North Pacific Ocean did not occur in the South Pacific Ocean which supports the suggestion that belts of high tropospheric fallout accounted for the increase in the Northern oceans.     

Mitochondrial DNA variation and population structure in the spiny lobster Panulirus argus

Adult spiny lobsters (Panulirus argus) were collected from nine locations throughout the tropical and subtropical northwest Atlantic Ocean and examined for mitochondrial DNA (mtDNA) variation. 187 different mtDNA haplotypes were observed among the 259 lobsters sampled. Haplotype diversity was calulated to be 0.986 and mean nucleotide sequence-diversity was estimated to be 1.44%; both of these values are among the highest reported values for a marine species. Analysis of molecular variance (AMOVA) and phenetic clustering both failed to reveal any evidence of genetic structure within and among populations of P. argus. The present data are consistent with high levels of gene flow among populations of P. argus resulting from an extended planktonic larval stage and strong prevailing ocean currents.     

Song of the humpback whale — Population comparisons

Summary Humpback whale songs recorded on tropical calving grounds exhibit different dialects depending on the oceanic basin. Songs sampled simultaneously from two populations in the North Pacific (Hawaii and Mexico) were essentially identical. These North Pacific songs were clearly different from the song type shared by two populations in the North Atlantic (Cape Verde Islands and West Indies). Songs from the Southern Hemisphere (Tonga) represent a third distinct dialect. Our evidence shows that, despite annual change in song organization, significant differences in humpback song occur between isolated ocean basins, while only subtle differences exist within an oceanic population (Hawaii and Pacific Mexico).     

Dispersal of phytoplanktotrophic shipworm larvae (Bivalvia: Teredinidae) over long distances by ocean currents

Shipworms or Teredinidae may be dispersed either as adults in floating wooden objects or as pelagic larvae drifting near the sea surface. Five shipworm species, i.e., half of those having an amphi-Atlantic geographical distribution, are known also to have pelagic phytoplanktotrophic larvae which can be carried by ocean currents. From a series of 742 plankton samples taken from throughout the temperate and tropical North Atlantic Ocean, it can be shown that shipworm larvae are not uncommon in the open sea. Teredinid veligers were found in 19% of all samples taken. One species of larvae, identical in all details to that described by Rancurel (1965), is particularly prevalent and is tentatively identified as Teredora malleolus (Turton). A definitive identification will be possible only after the pelagic larvae of the other Atlantic species are known. The larvae of Teredora malleolus are found throughout the North Atlantic Gyre and the adjacent temperate and tropical seas, and from scattered records in the South Equatorial Current. Larvae of other unidentified Teredinidae species were also found. The distance that larvae may be transported depends upon the length of pelagic larval development and the velocity of the currents. From the known current velocities it can be shown that, even in a few weeks, larvae may be dispersed many hundreds of kilometers. The geographical distribution of shipworm larvae suggests that they are carried along the coasts of continents and even across ocean basins, and that this dispersal must be an important factor in the geographical distribution of the adult forms and in the maintenance of genetic continuity between populations otherwise isolated from one another.Contribution No. 2555 from the Woods Hole Oceanographic Institution, Woods Hole, Massachusetts, USA     

Structure and migration corridors in Pacific populations of the Swordfish Xiphius gladius, as inferred through analyses of mitochondrial DNA

 The swordfish Xiphias gladius is a migratory oceanic species distributed in sub-tropical and temperate waters worldwide. Studies utilizing mitochondrial DNA (mtDNA) have demonstrated genetic subdivision between ocean basins, as well as within the Atlantic basin. However, there has been no support of population subdivision within the Pacific. We sequenced 629 base pairs of the control region for 281 swordfish collected in the Pacific. A rate heterogeneity parameter, alpha, was found to be 0.201, indicating substantial variation in mutation rate within the control region of swordfish. Hierarchical analysis of molecular variance supported significant genetic structuring among Pacific populations. Northern and southern populations in the western Pacific were significantly divergent, while populations in the east appeared to be genetically continuous. Regression analysis supported a correlation of genetic differentiation with geographic distance along a U-shaped corridor of gene flow. These results reveal a pelagic biogeographic pattern heretofore unrecognized in the Pacific, and reject the null hypothesis that Pacific populations of swordfish are unstructured and comprise only a single homogeneous stock. Received: 10 November 1998 / Accepted: 4 February 2000     

Mitochondrial evolution and phylogeography in the hydrozoan<Emphasis Type="Italic"> Obelia geniculata</Emphasis> (Cnidaria)

The distribution and genetic structure of many marine invertebrates in the North Atlantic have been influenced by the Pleistocene glaciation, which caused local extinctions followed by recolonization in warmer periods. Mitochondrial DNA markers are typically used to reconstruct species histories. Here, two mitochondrial markers [16S rDNA and cytochrome c oxidase I (COI)] were used to study the evolution of the widely distributed hydrozoan Obelia geniculata (Linnaeus, 1758) from the North Atlantic and the Pacific and, more specifically, in the context of North Atlantic phylogeography. Samples were collected from six geographic localities between 1998 and 2002. Hydroids from the North Atlantic, North Pacific (Japan), and South Pacific (New Zealand) are reciprocally monophyletic and may represent cryptic species. Using portions of the 16S rDNA and COI genes and the date of the last trans-Arctic interchange (3.1–4.1 million years ago), the first calibrated rate of nucleotide substitutions in hydrozoans is presented. Whereas extremely low substitution rates have been reported in other cnidarians, mainly based on anthozoans, substitution rates in O. geniculata are comparable to other invertebrates. Despite a life history that ostensibly permits substantial dispersal, there is apparently considerable genetic differentiation in O. geniculata. Divergence estimates and the presence of unique haplotypes provide evidence for glacial refugia in Iceland and New Brunswick, Canada. A population in Massachusetts, USA, appears to represent a relatively recent colonization event.Communicated by J.P. Grassle, New Brunswick     

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