Mitochondrial DNA analysis confirms the existence of two glacial races of rainbow smelt Osmerus mordax and their reproductive isolation in the St Lawrence River estuary (Quebec, Canada)

A phylogeographic analysis of mitochondrial DNA variation was performed in order to test the hypothesis that north-eastern North America has been postglacially recolonized by two races of rainbow smelt Osmerus mordax . This was accomplished by documenting the geographical distribution of two major mtDNA phylogenetic clades among 1290 smelt from 49 lacustrine and anadromous populations covering most of the species' native range. The data set was built by combining previously published results with those generated in this study. The two mtDNA clades showed a geographical dichotomy, independent of life-history types, whereby the more eastern populations were either fixed or largely dominated by one clade and western populations for the other. Such geographical pattern implying a phylogenetic discontinuity provided strong evidence for the persistence of smelt in two distinct glacial refugia as well as their differential postglacial dispersal. The most likely refugium for the so-called Atlantic race was the Atlantic coastal plains, whereas that of the so-called Acadian race was the exundated Grand Banks area. Patterns of postglacial dispersal interpreted from palaeogeographic events suggested that the Atlantic race recolonized northern regions about 5000 years prior the Acadian race. Both races came into contact in the St Lawrence River estuary. While gene flow has been possible, the sympatric occurrence in the estuary of anadromous populations alternatively dominated by one mtDNA clade or the other indicated that reproductive isolation mechanisms between the two races developed within this contact zone. This represents the first evidence of secondary intergradation among distinct races of aquatic organisms in an estuarine environment.     

Genetic structure and relationships among anadromous and landlocked populations of rainbow smelt, Osmerus mordax, Mitchill, as revealed by mtDNA restriction analysis

Larvae of anadromous rainbow smelt originating in various spawning tributaries are retained in the St Lawrence estuary. We proposed that these smelt represent one population genetically differentiated from adjacent populations characterized by geographically distinct larval retention areas. We also analysed four landlocked populations to evaluate the phylogenetic basis of dwarf and normal-sized phenotypes and their relation to an adromous smelt. A phylogenetic distinction was revealed between anadromous and landlocked smelt, with only one of the two mtDNA phylogenetic groups of anadromous fish observed among landlocked smelt. Significant geographical heterogeneity in the distribution of mtDNA genotypes was observed among landlocked smelt, suggesting that dwarfism in smelt may be polyphyletic in origin. St Lawrence smelt were genetically identical but distinct from adjacent populations, supporting the proposition that population genetic structure reflects the number of larval retention zones rather than spawning sites.     

Maturation,growth and fecundity of Arctic charr, <Emphasis Type="Italic">Salvelinus alpinus</Emphasis> (L.), life-history variants co-existing in lake systems of Southern Baffin Island,Nunavut, Canada

Two life-history variants of Arctic charr (Salvelinus alpinus), anadromous and lake-resident, have been previously identified in lakes of Southern Baffin Island, Nunavut, Canada. In accordance with classical life-history theory, it is hypothesised that anadromous charr will delay maturation in both size and age, and have increased fecundity (per spawning event) in comparison with lake-resident charr. Sagittal otoliths and biological data were collected for both life-history variants within the three studied lakes: Iqalugaarjuit, Qasigiat and Qinngu. Sagittal otoliths were embedded in epoxy resin, cross-sectioned for age determination, and imaged for back-calculation (size and age). Back-calculated data in each lake were fit to von Bertalanffy growth models for each life-history variant and compared via analysis of residual sums of squares. Anadromous charr had greater mean size at maturity and experienced a delay in mean age at maturity in comparison with lake-residents. The relationship between size and fecundity or egg diameter did not differ between the two life-history variants. Growth models indicate that the overall growth coefficients of lake-resident and anadromous charr were different in all three studied lakes. The Brody growth coefficient for all lake-resident charr populations was greater than anadromous individuals indicating that maximum length was reached at a rapid rate, resulting in a smaller asymptotic length. Indirect evidence suggests that anadromous and lake-resident charr belong to one reproductive population. Future genetic analysis is necessary to further determine the degree of divergence between the life-history variants.     

Latitudinal variation in growth among Arctic charr in eastern North America: evidence for countergradient variation?

Biological data from 66 populations of Arctic charr, Salvelinus alpinus, from eastern North America were analysed to test the applicability of the countergradient hypothesis as an explanation of differences in seasonally adjusted growth rates. Samples were obtained along a 37° latitudinal gradient and partitioned among anadromous, normal lacustrine, and dwarf lacustrine Arctic charr morphotypes. Models relating length-at-age or age-specific growth rates to latitude were estimated for each morphotype. Length-at-age declined with latitude for anadromous and lacustrine charr. Age-specific growth rates also varied with latitude, particularly for normal lacustrine charr. Results of analyses provide support for the countergradient hypothesis in growth performance of normal lacustrine morphotypes, where northern populations compensate for the shorter growth season with a greater rate of growth than southern populations. Anadromous charr exhibited equivocal evidence of countergradient variation, while results for dwarf lacustrine Arctic charr populations were inconclusive owing to the limited range of ages, and latitudes for which data were available.     

Molecular genetic evidence for reproductive isolation between sympatric populations of smelt Osmerus in Lake Utopia, south-western New Brunswick, Canada

Two morphologically and ecologically distinct forms of smelt, Osmerus, reside sympatrically in Lake Utopia, south-western New Brunswick, Canada. The ‘normal-sized’ form matures at greater than 200 mm standard length, averages about 31–33 gill rakers, and spawns in lake outlets. By contrast, the ‘dwarf-sized’ form matures at less than 150 mm standard length, averages 34–36 gill rakers, and spawns in small streams 3–5 weeks later than the normal form. We tested whether these sympatric forms represented ecological polymorphism within a single population or two reproductively isolated demes by assaying variation within and between forms by mitochondrial DNA (mtDNA) restriction site and nuclear minisatellite DNA analyses. Analysis of smelt mtDNA with twelve restriction enzymes resolved ten composite genotypes (differing by an average 0.27% sequence divergence) which differed markedly in frequency between the forms. Net percentage sequence divergence between the forms was O.l6%. A Wagner parsimony/ bootstrapping analysis of the restriction site presence/absence matrix, however, suggested that there were no significant distinctions between dwarf and normal smelt based on the phylogeny of composite genotypes. Hybridization studies of genomic DNA digests with a minisatellite probe indicated both that nuclear restriction fragment differentiation and the frequency of specific fragments differed significantly between the forms. Significant genetic differentiation between the sympatric forms demonstrates that they are distinct gene pools and reproductively isolated. Our molecular evidence for reproductive isolation between dwarf and normal smelt in Lake Utopia, coupled with the persistent morphological and ecological differentiation between them, argues strongly that they are behaving as distinct species. The Lake Utopia Osmerus populations provide further illustration of the potential for rapid differentiation to the level of biological species in postglacial environments.     

Mitochondrial and Allozyme Genetics of Incipient Speciation in a Landlocked Population of Galaxias Truttaceus (Pisces: Galaxiidae)

Galaxias truttaceus is found in coastal rivers and streams in south-eastern Australia. It spawns at the head of estuaries in autumn and the larvae spend 3 months of winter at sea before returning to fresh water. In Tasmania there are landlocked populations of G. truttaceus in a cluster of geologically young lakes on the recently glaciated Central Plateau. These populations have no marine larval stage and spawn in the lakes in spring. Speciation due to land locking is thought to be a frequent occurrence within Galaxias. To investigate the nature of the speciation event which may be occurring within lake populations of G. truttaceus we studied the mitochondrial DNA (mtDNA) and allozyme diversity of both lake and stream populations. Using the presence or absence of restriction sites recognized by 13 six-base restriction endonucleases, we found 58 mtDNA haplotypes among 150 fish collected from 13 Tasmanian and one south-east Australian mainland stream populations. The most parsimonious network relating the haplotypes by site loss or gain was starlike in shape. We argue that this arrangement is best explained by selection upon slightly beneficial mutations within the mitochondrial genome. Gene diversity analysis under Wright's island model showed that the populations in each drainage were not genetically subdivided. Only two of these stream haplotypes were found among the 66 fish analyzed from four lake populations. Despite the extreme lack of mtDNA diversity in lake populations, the observed nuclear DNA heterozygosity of 40 lake fish (0.10355) was only slightly less than that of 82 stream fish (0.11635). In the short time (3000-7000 years) that the lake fish have been landlocked, random genetic drift in a finite, stable-sized population was probably not responsible for the lack of mtDNA diversity in the lake populations. We infer the lake populations have probably experienced at least one, severe, but transitory bottleneck possibly induced by natural selection for life-history characters essential for survival in the lacustrine habitat. If speciation is occurring in the landlocked populations of G. truttaceus, then it may be driven by genetic transilience.     

PHYLOGEOGRAPHIC STRUCTURE IN MITOCHONDRIAL DNA OF THE LAKE WHITEFISH (COREGONUS CLUPEAFORMIS) AND ITS RELATION TO PLEISTOCENE GLACIATIONS

Restriction-fragment length polymorphisms were employed to evaluate the phylogenetic relationships, the genetic diversity and the geographic structure in mitochondrial DNA (mtDNA) lineages of the lake whitefish, Coregonus clupeaformis. Thirteen restriction enzymes that produced 148 restriction fragments were used to assay mtDNAs of 525 specimens collected among 41 populations. The sampling covered the entire range of the species, from Alaska to Labrador. Four distinct phylogeographic assemblages were identified. The Beringian assemblage, confined to Yukon and Alaska, was phylogenetically distinct from other assemblages and exhibited the highest level of nucleotide diversity. The Acadian assemblage was confined to southeastern North America and composed of a unique mtDNA clade. The Atlantic assemblage was confined to southern Québec and the northeastern United States and was also observed among anadromous populations of northern Hudson Bay. This group was highly polymorphic and responsible for most of the mtDNA diversity observed outside Beringia. The Mississippian assemblage occupied most of the actual range of lake whitefish, from the Mackenzie delta to Labrador. Ninety-two percent of all whitefish of this proposed origin belonged to a single mtDNA haplotype. Overall, the diversity, the geographic structure and the times of divergence of mtDNA phylogenetic assemblages correlate with the Pleistocene glaciations classically assumed to have dramatically altered the genetic diversity of northern fishes in recent evolutionary times. Our results emphasize the dominant role of these catastrophic events in shaping the population genetic structure of lake whitefish.     

Evolutionary history of an adaptive radiation in species pairs of threespine sticklebacks (Gasterosteus): insights from mitochondrial DNA

Species pairs of threespine stickleback, Gasterosteus aculeatus, co-exist in several lakes in the Strait of Georgia, southwestern British Columbia. One species, ‘benthics’ is robust-bodied and is morphologically and behaviourally specialized for benthivory. The other species, ‘limnetics’ is specialized for planktivory in open-water habitats of the lakes. We examined mitochondrial DNA restriction site variation in benthic and limnetic sticklebacks as well as in solitary freshwater, anadromous (sea-run), and marine populations to test: (i) if benthic and limnetic pairs have evolved only once or multiple times (parallel evolution) and (ii) if the species have evolved sympatrically, or allopatrically from ‘double invasions’ of lakes by ancestral anadromous/marine sticklebacks. Stickleback mtDNA comprised a single clade with a low (mean = 0.40%) degree of sequence divergence among the 77 haplotypes resolved. Most nucleotide diversity (97%) was found within (rather than among) populations of anadromous/marine sticklebacks whereas most diversity (77%) was found among populations in freshwater sticklebacks. Significant differences in haplotype frequencies were found between benthics and limnetics in three of the four species pair lakes examined, but in all cases the pairs within lakes were characterized by unique assemblages of closely related haplotypes. Hierarchical clustering of divergence estimates suggested that comparable species from different lakes have originated independently in all lakes because in no case did comparable species from different lakes cluster together. Divergent species within lakes tended to be more closely related to one another than to species in other lakes and there were two cases were benthics and limnetics within a particular lake were monophyletic. In two of the four two-species lakes, limnetics were less divergent from putative ancestral anadromous/marine stickleback as predicted by the double invasion hypothesis, but in the two other lakes benthics were less divergent. Our data argue strongly that the species pairs have evolved independently in each lake were they now co-exist. Further, in two lakes our data are consistent with the species having evolved by sympatric divergence, but allopatric divergence followed by introgression of mtDNA that has obscured ancestral relationships cannot be discounted completely. Finally, despite remaining uncertainty about the geography of speciation, the species appear to have evolved in the face of gene flow arguing that natural selection acting on trophic ecology has been a major component of ecological speciation in sticklebacks.     

CORRELATED TROPHIC SPECIALIZATION AND GENETIC DIVERGENCE IN SYMPATRIC LAKE WHITEFISH ECOTYPES (COREGONUS CLUPEAFORMIS): SUPPORT FOR THE ECOLOGICAL SPECIATION HYPOTHESIS

There is ample empirical evidence that phenotypic diversification in an adaptive radiation is the outcome of divergent natural selection related to differential resource use. In contrast, the role of ecological forces in favoring and maintaining reproductive isolation in nature remains poorly understood. If the same forces driving phenotypic divergence are also responsible for speciation, one would predict a correlation between the extent of trophic specialization (reflecting variable intensity of divergent natural selection) and that of reproductive isolation being reached in a given environment. We tested this hypothesis by comparing the extent of morphological and genetic differentiation between sympatric dwarf and normal whitefish ecotypes (Coregonus sp.) from six lakes of the St. John River basin (eastern Canada and northern Maine). Eight meristic variables, 19 morphometric variables, and six microsatellite loci were used to quantify morphological and genetic differentiation, respectively. Dwarf and normal ecotypes in each lake differed primarily by traits related to trophic specialization, but the extent of differentiation varied among lakes. Significant but variable genetic divergence between ecotypes within lakes was also observed. A negative correlation was observed between the extent of gene flow between ecotypes within a lake and that of their morphological differentiation in trophic-related traits. The extent of reproductive isolation reached between dwarf and normal whitefish ecotypes appears to be driven by the potential for occupying distinct trophic niches and, thus, by the same selective forces driving tropic specialization in each lake. These results therefore support the hypothesis of ecological speciation.     

Intraspecific vicariant history and the evolution of adaptive morphological diversity in a fish species (Osmerus mordax)

Vicariant geographic isolation and resource partitioning have long been independently identified as processes contributing to the morphological divergence of closely-related species. However, little is known about the extent to which vicariant history influences the adaptive ecological divergence associated with resource partitioning and trophic specialization within species. The present study thus quantified the contribution of vicariant historical genetic divergence to the adaptive contemporary morphological divergence of intraspecific feeding specialists in the Rainbow smelt (Pisces: Osmerus mordax ). This species is characterized by the polyphyletic origin of two lacustrine feeding specialists originating in two intraspecific lineages associated with independent glacial refuges. The historical genetic segregation was initiated approximately 350 000 years ago, whereas the lacustrine trophic segregation arose within the past 10 000 years. Wild caught lacustrine smelt populations were grouped a priori based on known historical genetic identities (Acadian and Atlantic mitochondrial DNA clades) and contemporary feeding specializations (microphageous and macrophageous morphotypes). The present study demonstrated that independent suites of correlated morphological traits are associated with either vicariant history or contemporary feeding specializations. Second, functionally-similar feeding specialists exhibit distinct morphologies resulting largely from vicariant historical processes. Although, the evolutionary processes producing historical phenotypes remains unknown, the results obtained demonstrate how adaptive radiation associated with ecological resource partitioning and feeding specializations can be strongly influenced by intraspecific phenotypic diversification resulting from relatively recent vicariant histories.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 140–151.     

Genetic relationships among populations of wild resident, and wild and hatchery anadromous brook charr

To determine the genetic relationship of anadromous and resident life-history types within and among drainages, and compare several hatchery strains to their progenitor populations, brook charr Salvelinus fontinalis were examined for allozyme and mitochondrial DNA variation. Greater genetic similarity of sympatric anadromous and resident charr was found compared to similar life-history forms allopatrically, suggesting the two life-history types are not reproductively isolated. Low divergence among the mtDNA haplotypes suggests that the two life-history types are members of the same evolutionary lineage. Population differentiation from mtDNA data exceeded that from estimates based on allozymes. Genetic deviations from expectations suggest that the hatchery strains were derived from few individuals.     

The distribution of divergent mitochondrial DNA lineages of threespine stickleback (Gasterosteus aculeatus) in the northeastern Pacific Basin: post-glacial dispersal and lake accessibility

Aim This study furthers the documentation of the geographical distribution of two divergent (c. 3%) mitochondrial DNA clades in the threespine stickleback (Gasterosteus aculeatus) and tests the hypotheses that the northeastern Pacific distribution has been influenced by post‐glacial colonization and lake elevation and that clade identity is associated with certain morphological attributes such as reduction in body armour. Location Lakes and nearshore marine environments of the eastern Pacific Basin from southcentral Alaska to southeastern British Columbia, (BC) Canada. Methods Restriction enzyme analysis of polymerase chain reaction‐amplified mitochondrial DNA fragments (cytochrome b) from a total of 45 new populations combined with existing data for a further 45 populations. Lake elevation data were collected for 78 localities and tested for an association with mtDNA clade by contingency table analyses. Morphological data were collected on sticklebacks from eight samples representing four lake‐stream systems and tested for differentiation among populations with different mtDNA clade identities using analyses of variance. Results We extend the known distribution of the haplotypes diagnostic of the Trans‐North Pacific Clade (TNPC) southward to mid‐Vancouver Island and, for the first time, on mainland BC, in other island populations far from putative refugia, and in nearby anadromous populations. A morphological analysis indicated that the mainland population with the TNPC was not characterized by reduced spine or lateral plate (‘armour’) traits that characterize some putative relict populations on the Queen Charlotte Islands. We found a significant association between lake elevation and the presence of the TNPC; the TNPC was present more often in lakes located at or lower than 42 m than in higher elevation lakes. Main conclusions Our data support the hypothesis that post‐glacial colonization by TNPC‐bearing marine sticklebacks and aspects of lake ‘accessibility’ were important in determining the distribution of mtDNA clades in the eastern Pacific Ocean basin. More generally, our study demonstrates how processes acting both across immense geographic scales (e.g. pan‐Pacific dispersal) and local scales (lake accessibility contingent on timing and extent of isostatic rebound) may interact to explain biogeographical patterns.     

THE GEOGRAPHY OF COEVOLUTION: COMPARATIVE POPULATION STRUCTURES FOR A SNAIL AND ITS TREMATODE PARASITE

Gene flow and the genetic structure of host and parasite populations are critical to the coevolutionary process, including the conditions under which antagonistic coevolution favors sexual reproduction. Here we compare the genetic structures of different populations of a freshwater New Zealand snail (Potamopyrgus antipodarum) with its trematode parasite (Microphallus sp.) using allozyme frequency data. Allozyme variation among snail populations was found to be highly structured among lakes; but for the parasite there was little allozyme structure among lake populations, suggesting much higher levels of parasite gene flow. The overall pattern of variation was confirmed with principal component analysis, which also showed that the organization of genetic differentiation for the snail (but not the parasite) was strongly related to the geographic arrangement of lakes. Some snail populations from different sides of the Alps near mountain passes were more similar to each other than to other snail populations on the same side of the Alps. Furthermore, genetic distances among parasite populations were correlated with the genetic distances among host populations, and genetic distances among both host and parasite populations were correlated with “stepping-stone” distances among lakes. Hence, the host snail and its trematode parasite seem to be dispersing to adjacent lakes in a stepping-stone fashion, although parasite dispersal among lakes is clearly greater. High parasite gene flow should help to continuously reintroduce genetic diversity within local populations where strong selection might otherwise isolate “host races.” Parasite gene flow can thereby facilitate the coevolutionary (Red Queen) dynamics that confer an advantage to sexual reproduction by restoring lost genetic variation.     

Intraspecific Genetic Admixture and the Morphological Diversification of an Estuarine Fish Population Complex

The North-east American Rainbow smelt (Osmerus mordax) is composed of two glacial races first identified through the spatial distribution of two distinct mtDNA lineages. Contemporary breeding populations of smelt in the St. Lawrence estuary comprise contrasting mixtures of both lineages, suggesting that the two races came into secondary contact in this estuary. The overall objective of this study was to assess the role of intraspecific genetic admixture in the morphological diversification of the estuarine rainbow smelt population complex. The morphology of mixed-ancestry populations varied as a function of the relative contribution of the two races to estuarine populations, supporting the hypothesis of genetic admixture. Populations comprising both ancestral mtDNA races did not exhibit intermediate morphologies relative to pure populations but rather exhibited many traits that exceeded the parental trait values, consistent with the hypothesis of transgressive segregation. Evidence for genetic admixture at the level of the nuclear gene pool, however, provided only partial support for this hypothesis. Variation at nuclear AFLP markers revealed clear evidence of the two corresponding mtDNA glacial races. The admixture of the two races at the nuclear level is only pronounced in mixed-ancestry populations dominated by one of the mtDNA lineages, the same populations showing the greatest degree of morphological diversification and population structure. In contrast, mixed-ancestry populations dominated by the alternate mtDNA lineage showed little evidence of introgression of the nuclear genome, little morphological diversification and little contemporary population genetic structure. These results only partially support the hypothesis of transgressive segregation and may be the result of the differential effects of natural selection acting on admixed genomes from different sources.     

Effects of Partially Anadromous Arctic Charr (Salvelinus alpinus) Populations on Ecology of Coastal Arctic Lakes

Little research has been conducted on effects of iteroparous anadromous fishes on Arctic lakes. We investigated trophic ecology, fish growth, and food web structure in six lakes located in Nunavut, Canada; three lakes contained anadromous Arctic charr (Salvelinus alpinus) whereas three lakes did not contain Arctic charr. All lakes contained forage fishes and lake trout (Salvelinus namaycush; top predator). Isotope ratios (δ¹³C, δ¹⁵N) of fishes and invertebrates did not differ between lakes with and without anadromous Arctic charr; if anadromous Arctic charr deliver marine-derived nutrients and/or organic matter to freshwater lakes, these inputs could not be detected with δ¹³C and/or δ¹⁵N. Lake trout carbon (C):nitrogen (N) and condition were significantly higher in lakes with Arctic charr (C:N = 3.42, K = 1.1) than in lakes without Arctic charr (C:N = 3.17, K = 0.99), however, and ninespine stickleback (Pungitius pungitius) condition was significantly lower in lakes with Arctic charr (K = 0.58) than in lakes without Arctic charr (K = 0.64). Isotope data indicated that pre-smolt and resident Arctic charr may be prey for lake trout and compete with ninespine stickleback. Linear distance metrics applied to isotope data showed that food webs were more compact and isotopically redundant in lakes where Arctic charr were present. Despite this, lake trout populations in lakes with Arctic charr occupied a larger isotope space and showed greater inter-individual isotope differences. Anadromous Arctic charr appear to affect ecology and feeding of sympatric freshwater species, but effects are more subtle than those seen for semelparous anadromous species.     

The genetic population structure of lacustrine sockeye salmon, <Emphasis Type="Italic">Oncorhynchus nerka,</Emphasis> in Japan as the endangered species

Lacustrine sockeye salmon (Oncorhynchus nerka) are listed as an endangered species in Japan despite little genetic information on their population structure. In order to clarify the genetic diversity and structure of Japanese populations for evaluating on the bottleneck effect and an endangered species, we analyzed the ND5 region of mitochondrial DNA (mtDNA) and 45 single nucleotide polymorphisms (SNPs) in 640 lacustrine sockeye salmon in Japan and 80 anadromous sockeye salmon in Iliamna Lake of Alaska. The genetic diversity of the Japanese population in both mtDNA and SNPs was significantly less than that of the Iliamna Lake population. Moreover, all Japanese populations had SNP loci deviating from the HWE. In spite of low genetic diversity, the SNP analyses resulted that the Japanese population was significantly divided into three groups. These suggest that Japanese sockeye salmon populations should be protected as an endangered species and genetically disturbed by the hatchery program and transplantations.     

On the origin of Solovetskaya vendace <Emphasis Type="Italic">Coregonus albula</Emphasis> and of the syamozero smelt <Emphasis Type="Italic">Osmerus eperlanus</Emphasis>

The genetic analysis is made on the population of the European smelt Osmerus eperlanus occasionally introduced to Lake Syamozero (Karelia) and of the vendace Coregous albula supposedly acclimatized in Solovetskie Islands as a result of fish cultural activities of the Solovetskii Monastery. They are compared with several natural populations—possible donors for both introductions. Genetic variation in a sample of smelt was estimated by means of restrictase analysis of mtDNA (fragment ND1/ND2) in samples of vendace—by means of allozyme analysis of six isoenzyme systems. The probable population for the Syamozero smelt is the population of Onega Lake in spite of the previously noted greater morphological similarity of colonizers with the smelt of Ladoga Lake. A high level of genetic variation in the Syamozero smelt in comparison with native populations indicates that, beside the introduction from Onega Lake, there were repeated introductions from neighboring water bodies. The genetic analysis of the Solovetskaya vendace does not prove that the vendace appeared on islands due to acclimatization. Frequencies of alleles of allozyme loci in the Solovetskaya population significantly differ from frequencies in continental populations. Still, it is compared with some populations of the Arkhangelsk oblast. Estimations of genetic diversity of the Solovetskaya vendace turned out to be comparable with those in native populations.     

Extant and extinct forms of arctic charr Salvelinus alpinus (L.) complex from the Leprindo lake system (Transbaikalia): Differentiation in life history, morphology, and genetics

Interconnected lakes Bol’shoe Leprindo and Maloe Leprindo in Transbaikalia hosted large (extinct) and dwarf charr forms. Rarely “small” individuals intermediate in size between these forms are caught. In order to assess morphological, ecological, and genetic differentiation of sympatric charr forms and parapatric charr populations we studied their meristic and morphometric characters, feeding, breeding, and growth; we also investigated variation at 8 microsatellite loci using DNA isolated both from contemporary and historic samples. Profound differences were found between large and dwarf charr in growth rate, feeding (piscivores and highly specialized zooplanktivores, respectively), spawning time, and morphology. Dwarf charr from the two lakes demonstrate minor differences in morphology and growth rate. “Small” individuals are morphologically similar with dwarf charr and spawn together with them, they are recruited from dwarf form in late ontogeny as the result of transition to piscivorous feeding and growth acceleration. Microsatellite analysis showed that: (1) large and dwarf charr forms display high degree of genetic differentiation and reproductive isolation; (2) dwarf charr from interconnected lakes belong to different isolted populations; (3) “small charr” are genetically identical with dwarfs. The degree of ecomorphological and genetic differentiation between large and dwarf forms places charr from Leprindo Lakes among the most strongly differentiated Arctic charr forms’ flocks known at the vast range of S. alpinus complex.     

Aquatic food‐web dynamics following incorporation of nutrients derived from Atlantic anadromous fishes

Changes in the isotopic composition (δ¹³C and δ¹⁵N) in biofilm, macro‐invertebrates and resident salmonids were used to characterize temporal dynamics of marine derived nutrients (MDNs) incorporation between stream reaches with and without MDN inputs. Five Atlantic rivers were chosen to represent contrasting MDN subsidies: four rivers with considerable numbers of anadromous fishes; one river with little MDN input. Rainbow smelt Osmerus mordax, alewife Alosa pseudoharengus, sea lamprey Petromyzon marinus and Atlantic salmon Salmo salar, were the primary anadromous species for the sampled rivers. Regardless of the spatial resolution or the pathway of incorporation, annual nutrient pulses from spawning anadromous fishes had a positive effect on isotopic enrichment at all trophic levels (biofilm, 1·2–5·4‰; macro‐invertebrates, 0·0–6·8‰; fish, 1·2–2·6‰). Community‐wide niche space shifted toward the marine‐nutrient source, but the total ecological niche space did not always increase with MDN inputs. The time‐integrated marine‐nutrient resource contribution to the diet of S. salar parr and brook trout Salvelinus fontinalis ranged between 16·3 and 36·0% during anadromous fish‐spawning periods. The high degree of spatio‐temporal heterogeneity in marine‐nutrient subsidies from anadromous fishes lead to both direct and indirect pathways of MDN incorporation into stream food webs. This suggests that organisms at many trophic levels derive a substantial proportion of their energy from marine resources when present. The current trend of declining anadromous fish populations means fewer nutrient‐rich marine subsidies being delivered to rivers, diminishing the ability to sustain elevated riverine productivity.     

Contrasting patterns of mitochondrial DNA and microsatellite introgressive hybridization between lineages of lake whitefish (Coregonus clupeaformis); relevance for speciation

We performed a combined analysis of mitochondrial DNA (mtDNA) and microsatellite loci among lake whitefish (Coregonus clupeaformis) populations in order to assess the levels of congruence between both types of markers in defining patterns of genetic structuring, introgressive hybridization and inferring population origins in the hybrid zone of the St. John River basin. A second objective was to test the hypothesis that secondary contact between glacial lineages always resulted in the occurrence of sympatric dwarf and normal whitefish ecotypes. Fish were sampled from 35 populations and polymorphism was screened at mtDNA and six microsatellite loci for a total of 688 and 763 whitefish, respectively. Four lakes harbouring a single whitefish population of normal ecotype admixed with mtDNA haplotypes of different lineages were found. This confirmed that secondary contact between whitefish evolutionary lineages did not always result in the persistence of reproductively isolated ecotypes. Microsatellites further supported the definition of distinct glacial lineages by identifying lineage-specific allelic size groups. They also further supported the hypothesis that ecotypes originated from either a single founding lineage (sympatric divergence) or following secondary contacts between lineages (allopatric divergence), depending on the lake. In general, however, the pattern of population differentiation and introgressive hybridization observed at microsatellites was in sharp contrast with that depicted by mtDNA variation. Both factorial correspondence analysis and analysis of admixture proportion revealed a much more pronounced pattern of introgressive hybridization than depicted by mtDNA analyses. Variable levels of introgression indicated that environmental differences may be as important as the historical contingency of secondary contact in explaining the persistence of sympatric ecotypes and the differential pattern of introgressive hybridization among lakes. Whitefish populations from the St. John River basin hybrid zone represent a rare illustration of a continuum of both morphological and genetic differentiation within a given taxon, spanning from complete introgression to possibly complete reproductive isolation, depending on lakes. Thus, each lake may be viewed as a different temporal snapshot taken throughout the gradual process of speciation.