Petr Strelkov

Diversity and trans-arctic invasion history of mitochondrial lineages in the North Atlantic Macoma balthica complex (Bivalvia: Tellinidae).

Abstract The history of repeated inter- or transoceanic invasions in bivalve mollusks of the circumpolar Macoma balthica complex was assessed from mtDNA COIII sequences. The data suggest that four independent trans-Arctic invasions, from the Pacific, gave rise to the current lineage diversity in the North Atlantic. Unlike in many other prominent North Atlantic littoral taxa, no evidence for (postinvasion) trans-Atlantic connections was found in the M. balthica complex. The earliest branch of the mtDNA tree is represented by the temperate-boreal North American populations (= Macoma petalum), separated from the M. balthica complex proper in the Early Pliocene at latest. The ensuing trans-Arctic invasions established the North European M. b. rubra, which now prevails on the North Sea and northeast Atlantic coasts, about two million years ago, and the currently northwest Atlantic M. balthica lineage in the Canadian Maritimes, in the Middle Pleistocene. The final reinvasion(s) introduced a lineage that now prevails in a number of North European marginal seas and is still hardly distinguishable from North Pacific mtDNA (M. b. balthica). We used coalescence simulation analyses to assess the age of the latest invasion from the Pacific to the northeast Atlantic. The results refute the hypothesis of recent, human-mediated reintroductions between northeast Pacific and the North European marginal seas in historical times. Yet they also poorly fit the alternative hypotheses of an early postglacial trans-Arctic invasion (< 11 thousand years ago), or an invasion during the previous Eemian interglacial (120 thousand years ago). Divergence time estimates rather fall in the Middle Weichselian before the Last Glacial Maximum, in conflict with the conventional thinking of trans-Arctic biogeographical connections; an early Holocene reinvasion may still be regarded as the most plausible scenario. Today, the most recently invaded Pacific mtDNA lineage is found admixed with the earlier established European Atlantic “rubra” lineage in the Baltic Sea and in Barents Sea populations east of the Varanger peninsula, and it is practically exclusive in the White and Pechora seas. Yet mtDNA does not always constitute an unequivocal taxonomic marker at individual level; the marginal populations represent hybrid swarms of the Atlantic and Pacific lineages in their nuclear genes.

A preliminary revision of the genus Plecotus (Chiroptera, Vespertilionidae) based on genetic and morphological results

The phylogenetic relationships within the genus Plecotus were assessed using molecular as well as morphological methods. With only three species missing, our study is based on an almost comprehensive taxonomic sampling. The genetic analysis comprised 151 individuals from throughout the range. Sequences of two mitochondrial sections, parts of the 16S rRNA gene (16S) and of the control region (CR) were analysed. The morphological analysis of cranial and external characters comprised 697 individuals, including 10 holotypes and one lectotype. Data from 15 craniometric characters of 442 specimens were used in the multivariate analyses. The molecular data identified nine primary clades representing 11 species, 10 of which could be assigned to described taxa, whereas one was described as a new species, Plecotus strelkovi Spitzenberger sp. nov. The tree based on 16S revealed two major lineages, one consisting of only one primary clade restricted to the Mediterranean, the other consisting of eight primary clades representing Eurasian taxa. The morphological analysis revealed five additional species, two of them not described. Together with the recently described P. taivanus, P. sardus and P. balensis, which were not included in our analysis, the genus Plecotus comprises at least 19 more or less cryptic species. Phylogenetic and phenetic analyses resulted in similar but not completely concordant arrangements of the species. The proposed classification relies mainly on the tree based on 16S sequences. The current distribution indicates that 16 species can be linked to arboreal refugia, three to eremial refugia. We assume that speciation within the gleaning, rather slow flying long‐eared bats is due to a multitude of disruption and isolation processes within a formerly continuous range of the broad‐leaved Arcto‐Tertiary forest in which Plecotus probably originated. An exact calibrated molecular dating of the splits is not possible. The Early Oligocene age of the presumed ancestor of the Plecotini and a correlation of the molecular diversifications with palaeogeographic reconstructions suggest that the divergence of the two major lineages may have occurred already during the Middle Miocene, 14.5 Mya.

A broad transition zone between an inner Baltic hybrid swarm and a pure North Sea subspecies of Macoma balthica (Mollusca, Bivalvia)

The populations of the bivalve clam Macoma balthica in the low‐salinity Northern Baltic Sea represent an admixture of two strongly diverged genomic origins, the Pacific Macoma balthica balthica (approx. 60% genomic contribution) and Atlantic Macoma balthica rubra (40%). Using allozyme and mtDNA characters, we describe the broad transition from this hybrid swarm to the pure M. b. rubra in the saline North Sea waters, spanning hundreds of kilometre distance. The zone is centred in the strong salinity gradient of the narrow Öresund strait and in the adjacent Western Baltic. Yet the multilocus clines show no simple and smoothly monotonic gradation: they involve local reversals and strong differences between neighbouring populations. The transitions in different characters are not strictly coincident, and the extent of introgression varies among loci. The Atlantic influence extends further into the Baltic in samples from the southern and eastern Baltic coasts than on the western coast, and further in deeper bottoms than at shallow (< 1 m) sites. This fits with the counterclockwise net circulation pattern and with a presumably weaker salinity barrier for invading Atlantic type larvae in saline deeper water, and corresponding facilitation of outwards drift of Baltic larvae in diluted surface waters. Genotypic disequilibria were strong particularly in the shallow‐water samples of the steepest transition zone. This suggests larval mixing from different sources and limited interbreeding in that area, which makes a stark contrast to the evidence of thorough amalgamation of the distinct genomic origins in the inner Baltic hybrid swarm of equilibrium structure.

Sensitivity to stress in the bivalve Macoma balthica from the most northern (Arctic) to the most southern (French) populations: low sensitivity in Arctic populations because of genetic adaptations?

The stress sensitivity, determined in copper exposure experiments and in survival in air tests, and the genetic structure, measured by means of isoenzyme electrophoresis, were assessed in populations of the Baltic clam Macoma balthica (L.) from its southern to its northern distribution limit, in order to test the hypotheses that near the distribution limit the clams would be more stress sensitive and would have a lower genetic variability. The populations in west and north Europe show a strong genetic resemblance. The populations in the sub-Arctic White Sea are genetically slightly different, and show a low stress sensitivity. The populations in the Arctic Pechora Sea are genetically very distant from the other populations, and show the lowest stress sensitivity. Near the southern distribution limit, in agreement with the hypotheses, genetic variability is low and stress sensitivity high. On the other hand, in contrast to expectation, near the northern distribution limit, in the populations of the Pechora Sea, the genetic variability was higher, thus not reduced, and the stress sensitivity was low compared to all other populations. Yet, it remains a question if such is due to gradual physiological acclimatization (and ongoing differential selection) or to genetic adaptation.

Macoma balthica in the White and Barents Seas: properties of a widespread marine hybrid swarm (Mollusca: Bivalvia)

A main molecular subdivision in the circumpolar Macoma balthica complex has been described between Atlantic and Pacific taxa. In NE Europe, the clams of the White and Barents Seas, however, show deviant genetic structures. Using allozyme and mitochondrial DNA data, we explore the hypothesis that these deviations result from hybridization between an Atlantic (M. b. rubra) and an invading Pacific (M. b. balthica) lineage. A practically pure Atlantic Macoma extends from France north to the Varanger Peninsula (NE Norway), whereas populations farther east have genetic compositions intermediate between true Atlantic and true Pacific. Admixture estimates range from 32 to 90% Pacific contribution, with a notable deviation in a nearly pure Atlantic outpost in the Mezen Bay (NE White Sea). The pattern of variation is not one of a simple collinear mixing however. Different characters exhibit different degrees of introgression, and the relative introgression varies regionally. Yet, there are practically no interlocus genotypic disequilibria between the diverged loci, which brings out the White Sea–Barents Sea M. balthica as the best‐documented marine animal hybrid swarms so far, arisen through amalgamation of genomes previously isolated since pre‐Pleistocene times. On top of the main admixture pattern, strong geographical structuring is also seen in characters unrelated to the principal systematic distinction. The persistence of the regional patterns indicates restricted gene flow at the present time, despite the high dispersal potential of the species. The causes of this structuring could be in a complex history of colonization events and features of local hydrography enhancing isolation and divergence of populations.

Growth and longevity of Mytilus edulis (L.) from northeast Europe

Abstract The growth of mussels, Mytilus edulis (L.), was studied in most of the northeastern part of their distribution. The growth, longevity and maximal size of molluscs from 24 wild populations and one cultured population located in the White Sea and the southeast part of the Barents Sea were compared. The 25 studied populations were combined in six clusters. The maximal longevity and the size of the mussels varied between 7 and 18 years and 25.5 and 77.7 mm, respectively. The geographical location of the population within the studied region did not affect either maximal longevity or maximal size, or the growth rate of mussels. However, these parameters were influenced by local habitat conditions, primarily connected with the position within intertidal and subtidal zones. Animals inhabiting the intertidal zone were characterized by relatively low growth performance, a short life span and a small size. The longest life span was typical for deep subtidal mussels, whereas the highest growth rate was recorded in the cultured population and in the upper subtidal habitats. Growth patterns of Mytilus edulis in the subarctic White and the Arctic southeast Barents seas are similar to those reported from other parts of the area of distribution. Therefore, growth was mostly determined by local environmental factors, including those related to vertical zonation, rather than by latitude/longitude and related temperature effects.

Phylogeography of amphi-boreal fish: tracing the history of the Pacific herring Clupea pallasii in North-East European seas

BackgroundThe relationships between North Atlantic and North Pacific faunas through times have been controlled by the variation of hydrographic circumstances in the intervening Arctic Ocean and Bering Strait. We address the history of trans-Arctic connections in a clade of amphi-boreal pelagic fishes using genealogical information from mitochondrial DNA sequence data. The Pacific and Atlantic herrings (Clupea pallasii and C. harengus) have basically vicarious distributions in the two oceans since pre-Pleistocene times. However, remote populations of C. pallasii are also present in the border waters of the North-East Atlantic in Europe. These populations show considerable regional and life history differentiation and have been recognized in subspecies classification. The chronology of the inter-oceanic invasions and genetic basis of the phenotypic structuring however remain unclear.ResultsThe Atlantic and Pacific herrings both feature high mtDNA diversities (large long-term population sizes) in their native basins, but an ocean-wide homogeneity of C. harengus is contrasted by deep east-west Pacific subdivision within Pacific C. pallasii. The outpost populations of C. pallasii in NE Europe are identified as members of the western Pacific C. pallasii clade, with some retained inter-oceanic haplotype sharing. They have lost diversity in colonization bottlenecks, but have also thereafter accumulated abundant new variation. The data delineate three phylogeographic groups within the European C. pallasii: herring from the inner White Sea; herring from the Mezen and Chesha Bays; and a strongly bottlenecked peripheral population in Balsfjord of the Norwegian Sea.ConclusionsThe NE European outposts of C. pallasii are judged to be early post-glacial colonists from the NW Pacific. A strong regional substructure has evolved since that time, in contrast to the apparent broad-scale uniformity maintained by herrings in their native basins. The structure only partly matches the previous biological concepts based on seasonal breeding stocks or geographical subspecies designations. The trans-Arctic herring phylogeography is notably similar to those of the amphi-boreal mollusk taxa Macoma and Mytilus, suggesting similar histories of inter-oceanic connections. We also considered the time dependency of molecular rates, critical for interpreting timing of relatively recent biogeographical events, by comparing the estimates from coding and non-coding mitochondrial regions of presumably different mutation dynamics.

Population structure and growth rates at biogeographic extremes: a case study of the common cockle, Cerastoderma edule (L.) in the Barents Sea.

A descriptive study of population structure, growth rates and shell morphometry was conducted on nine intertidal populations of the infaunal bivalve Cerastoderma edule in the Murmansk coast of the Barents Sea. Year-to-year population dynamics was analyzed during 2002-2006 on a tidal flat Dalniy Plaj (eastern Murmansk coast). The region constitutes the northern extremes of C. edule range where populations occupied the middle to low intertidal zone and were characterized by low densities. The distribution of age groups and unstable age structure across years in the cockle populations suggest irregular recruitment. Growth rates and shell morphometry showed little variation across the populations studied, and there were no gradient changes from the west to the east parts of Murmansk coast.

Growth in the bivalve Macoma balthica from its northern to its southern distribution limit: a discontinuity in North Europe because of genetic adaptations in Arctic populations?

The hypothesis was tested that towards a species limit of distribution its performance, such as growth or fitness, decreases. To this end, latitudinal changes in growth, maximum attainable length and genetic constitution were assessed for the Baltic clam, Macoma balthica (L.), at stations ranging from the most southern distribution limit (France) to its most north-eastern limit in the Arctic Pechora Sea (North Russia). Growth was analyzed by means of growth-rings on the shells, the genetic constitution by electrophoretic isoenzyme analysis. Growth patterns and the genetic constitution of populations from West Europe, North Europe and the White Sea were similar, whereas the populations from the Pechora Sea are distinct from the other populations. The performance of clams in the Pechora Sea populations, with a relatively low annual growth but high maximum length, was, in contrast to the hypothesis, not decreased. It is concluded that the Pechora Sea populations form a separate group, genetically different from other European populations and adapted to the Arctic conditions. [KEYWORDS: Arctic; adaptation; distribution limit; genetics; geographic cline; growth; Macoma balthica; shell-length Dutch wadden sea; electrophoretic data; recruitment; gironde; stress]

Genetic, Ecological and Morphological Distinctness of the Blue Mussels Mytilus trossulus Gould and M. edulis L. in the White Sea

Two blue mussel lineages of Pliocene origin, Mytilus edulis (ME) and M. trossulus (MT), co-occur and hybridize in several regions on the shores of the North Atlantic. The two species were distinguished from each other by molecular methods in the 1980s, and a large amount of comparative data on them has been accumulated since that time. However, while ME and MT are now routinely distinguished by various genetic markers, they tend to be overlooked in ecological studies since morphological characters for taxonomic identification have been lacking, and no consistent habitat differences between lineages have been reported. Surveying a recently discovered area of ME and MT co-occurrence in the White Sea and employing a set of allozyme markers for identification, we address the issue whether ME and MT are true biological species with distinct ecological characteristics or just virtual genetic entities with no matching morphological and ecological identities. We find that: (1) in the White Sea, the occurrence of MT is largely concentrated in harbors, in line with observations from other subarctic regions of Europe; (2) mixed populations of ME and MT are always dominated by purebred individuals, animals classified as hybrids constituting only ca. 18%; (3) in terms of shell morphology, 80% of MT bear a distinct uninterrupted dark prismatic strip under the ligament while 97% of ME lack this character; (4) at sites of sympatry MT is more common on algal substrates while ME mostly lives directly on the bottom. This segregation by the substrate may contribute to maintaining reproductive isolation and decreasing competition between taxa. We conclude that while ME and MT are not fully reproductively isolated, they do represent clearly distinguishable biological, ecological and morphological entities in the White Sea. It remains to be documented whether the observed morphological and ecological differences are of a local character, or whether they have simply been overlooked in other contact zones.