Risto Väinölä

Chloroplast DNA recognizes three refugial sources of European oaks and suggests independent eastern and western immigrations to Finland

Refugial differentiation and routes of postglacial migration are major determinants of the patterns of geographical variation we see in natural populations today. We used patterns of chloroplast DNA (cpDNA) variation to investigate the postglacial colonization history of the European oak species Quercus robur and Q. petraea. By sequencing two cpDNA segments using universal primers, we revealed four polymorphic sites which identify four cytotypes with characteristic geographical distributions. Of these, the principal eastern, central and western cytotypes divide the range into three longitudinal zones, each extending from the south to the north of Europe. This corroborates the idea that the postglacial colonization started from three distinct southerly refugia. The fourth cytotype, restricted to East Anglia, was probably derived from the western type postglacially. As a special problem, we addressed the controversial origin of Q. robur at its northern limits in south-western Finland, where it currently occupies a narrow coastal zone disjunct from the remaining oak range. Using a PCR-RFLP assay that discriminates the eastern cytotype, a contact zone of two cytotypes was identified in the region of the Salpausselkä ridges. This suggests that the marginal northern occurrence was independently colonized both from the east and from the west, across the Baltic Sea.

Global diversity of amphipods (Amphipoda; Crustacea) in freshwater

Amphipods are brooding peracaridan crustaceans whose young undergo direct development, with no independent larval dispersal stage. Most species are epibenthic, benthic, or subterranean. There are some 1,870 amphipod species and subspecies recognized from fresh or inland waters worldwide at the end of 2005. This accounts for 20% of the total known amphipod diversity. The actual diversity may still be several-fold. Amphipods are most abundant in cool and temperate environments; they are particularly diversified in subterranean environments and in running waters (fragmented habitats), and in temperate ancient lakes, but are notably rare in the tropics. Of the described freshwater taxa 70% are Palearctic, 13% Nearctic, 7% Neotropical, 6% Australasian and 3% Afrotropical. Approximately 45% of the taxa are subterranean; subterranean diversity is highest in the karst landscapes of Central and Southern Europe (e.g., Niphargidae), North America (Crangonyctidae), and Australia (Paramelitidae). The majority of Palearctic epigean amphipods are in the superfamily Gammaroidea, whereas talitroid amphipods (Hyalella) account for all Neotropic and much of the Nearctic epigean fauna. Major concentrations of endemic species diversity occur in Southern Europe, Lake Baikal, the Ponto-Caspian basin, Southern Australia (including Tasmania), and the south-eastern USA. Endemic family diversity is similarly centered in the Western Palearctic and Lake Baikal. Freshwater amphipods are greatly polyphyletic, continental invasions have taken place repeatedly in different time frames and regions of the world. In the recent decades, human mediated invasions of Ponto-Caspian amphipods have had great impacts on European fluvial ecosystems.

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.

Diversity and distributions of circumpolar fresh- and brackish-water Mysis (Crustacea: Mysida): descriptions of M. relictaLovén, 1862, M. salemaai n. sp.,M. segerstralei n. sp. and M. diluviana n. sp., based on molecular and morphological characters

Mysid crustaceans of the Mysis relicta species group are widespread throughout the northern Holarctic and play an important role in many fresh- and brackish-water ecosystems. Earlier molecular and morphometric studies already indicated that the conventionally identified Mysis relicta sensu lato comprises several distinct species. Here we present formal taxonomic diagnoses, descriptions and an account of the distributions of Mysis relicta s. str. and three new species split from it, based on comprehensive assessment of both morphological and molecular characters (allozymes, mtDNA). M. relicta Lovén s. str. is the prevalent species in lakes of Northern Europe and peripheral parts of the brackish Baltic Sea. M. salemaai n. sp. inhabits offshore habitats of the Baltic Sea and a range of lakes from the British Isles, southern Scandinavia and Karelia to coastal northern Siberia; at several sites M. relicta and M. salemaai are sympatric. M. segerstralei n. sp. has a circumpolar distribution along the Arctic coasts and islands of Eurasia and North America and also occurs in lakes of these northern regions. M. diluviana n. sp. inhabits continental freshwater lakes of the once-glaciated northern North America. The four species are characterised by unique combinations of alleles at a number of allozyme loci, and most of them by specific mitochondrial DNA lineages diverged by c. 7.5% in the COI gene sequence (cytochrome oxidase subunit I). The most important diagnostic morphological characters include the shape of the posterior emargination of carapace, length of setae on the merus of maxillipede 2, length and shape of spine-setae on maxilla endopod distal segment, number and size of lateral spine-setae on telson, and setation of thoracic endopods. A morphological key to the four species is presented.

Postglacial colonization of Northern Europe by distinct phylogeographic lineages of the bullhead, Cottus gobio.

Three major phylogeographic lineages of the cottid fish Cottus gobio (bullhead) were identified in northern Europe from mitochondrial DNA sequences and allozyme data. The largely separate freshwater distributions of the lineages demonstrate distinct postglacial colonization histories. West of the Baltic Sea, Swedish lakes were invaded from the southwest (Germany). Another, eastern lineage has colonized the inland waters northeast and east of the Baltic, from refugia in northwest Russia; this lineage comprises a distinct subgroup found only from Estonia. The third lineage, found south and southeast of the Baltic, probably descended from rivers draining to the Black Sea from the north (e.g. Dnepr). In coastal waters of the Baltic Sea, and in near‐coast inland waters, the lineages are now found intermixed in various combinations. The alternating fresh‐ and saltwater phases of the Baltic basin have variously enabled and disabled the use of coastal waters as colonization routes. Hypotheses on the chronology of dispersal and lineage mixing can be based on the distribution of the marker genes and the palæohydrographical record. The diversity of the Fennoscandian bullhead thus comprises anciently diverged (probably mid‐Pleistocene) refugial lineages that in their freshwater range constitute distinct evolutionarily significant units. The thorough mixing of the various genomic origins in and around the Baltic, however, refutes the controversial view of distinct species status for the western and eastern (‘Cottus koshewnikowi’) bullheads. The postglacial contact of the lineages has created new diversity that cannot be interpreted in a conventional hierarchical framework of taxonomic or conservation units.

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.

Endemic diversification of the monophyletic cottoid fish species flock in Lake Baikal explored with mtDNA sequencing.

In the ancient Lake Baikal in East Siberia, cottoid fishes have diversified into an endemic flock of 33 species. From an ancestral shallow-water, benthic life-style, Baikalian cottoids have shifted to deep-water life in environments even below 1500 m, and also colonized the pelagic habitat. We examined phylogenetic relationships among 22 Baikalian and 10 extra-Baikalian cottoid taxa using a total of 2822 bp of mitochondrial DNA sequence, from complete sequences of ATPase 8 and 6 and cytochrome b genes and the control region. Unlike in earlier studies, we found strong support for a monophyly of the whole endemic Baikalian cottoid diversity. The Baikalian clade, currently assigned to three families and 12 genera, appears to be nested within the Holarctic freshwater genus Cottus. In the molecular phylogeny, all but one of the current Baikalian genera formed well-supported monophyletic groups. However, the topology was inconsistent with the present morphology-based familial subdivision; particularly in positioning the genus Batrachocottus of Cottidae within Abyssocottidae. The branching order of the Baikalian genera could not be resolved completely, however; short basal branches indicate rapid diversification early in the history of the species flock. Using synonymous divergence rates from other fish species for calibration, the diversification of the Baikalian cottoids seems to have started in the Pliocene or early Pleistocene.

Microsatellite variation in ringed seals (Phoca hispida): genetic structure and history of the Baltic Sea population.

Genetic variability and population structure of Baltic ringed seals and an Arctic reference population were assessed using eight microsatellite loci. Ringed seals colonized the Baltic Sea basin soon after deglaciation 11 500 years ago and are supposed to have remained largely isolated from the main Arctic stock since then, ≈1000 generations. In the 1900s the Baltic population declined rapidly, and is now confined to three distinct breeding areas, with N < 6000 seals altogether. Microsatellite heterozygosity in ringed seals was higher than that in the closely related, boreal harbour seal and grey seal, for which the markers were initially developed. This is plausibly attributed to an overall greater population (species) size of ringed seals during the Quaternary. Allele frequency differentiation between the Baltic and Arctic ringed seals, conventionally treated as different subspecies, was weak. Assuming complete isolation, the divergence (FST=0.023) would imply a notably high postglacial effective population size, ≈20 000 for the Baltic population. The isolation assumption however, seems unrealistic in the light of the data: a coalescent-based simulation approach to the likelihood of alternative demographic histories clearly favoured a scenario with recurrent gene flow to the Baltic, over one of complete isolation (drift only). Within the Baltic Sea, no differentiation was found between the Gulf of Finland and the Gulf of Bothnia breeding areas; the recent population decline and split have not yet affected the inbreeding levels of the disjunct breeding stocks.

Phylogeographic analyses of a circumarctic coastal and a boreal lacustrine mysid crustacean, and evidence of fast postglacial mtDNA rates

Phylogeographic structures of two weakly dispersing Mysis sibling species, one with a circumarctic coastal, the other with a boreal lacustrine‐Baltic distribution, were studied from mitochondrial cytochrome c oxidase subunit I (COI) gene sequences. Mysis segerstralei showed high overall diversity and little phylogeographic structure across the Arctic, indicating late‐glacial dispersal among coastal and lake populations from Alaska, Siberia and the north of Europe. A strongly divergent refugial lineage was however identified in Beringia. The boreal ‘glacial relict’Mysis salemaai in turn displayed clear structuring among postglacially isolated Scandinavian lake populations. The inferred pattern of intralake mitochondrial DNA (mtDNA) monophyly in Scandinavia suggested relatively small population sizes and a remarkably fast postglacial mtDNA divergence rate (0.27% per 10 000 years). Nevertheless, the broader phylogeographic pattern did not support distinct eastern and western glacial refugia in Northern Europe, unlike in some other aquatic taxa. In all, the two species comprised three equidistant mitochondrial lineages (approximately 2% divergence), corresponding to M. salemaai, to the bulk of M. segerstralei, and to the Beringian M. segerstralei lineage. The lack of reciprocal monophyly of the two species in respect to their mitochondrial genealogy could indicate postspeciation mitochondrial introgression, also exemplified by an evidently more recent capture of M. segerstralei mitochondria in a Karelian population of M. salemaai. Overall, the data suggest that the continental boreal M. salemaai has a relatively recent ancestry in arctic coastal waters, whereas two other boreal ‘glacial relict’Mysis sibling species in Europe (Mysis relicta) and North America (Mysis diluviana) have colonized inland waters much earlier (approximately 8% COI divergence).

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.