Karyological Differences between Northern Dolly Varden Salvelinus malma malma and White Char Salvelinus albusfrom the Kamchatka River Basin

The karyotypes of northern Dolly Varden and white char, sympatrically inhabiting the Kamchatka River basin, were studied. The karyotype of Dolly Varden was stable: 2n= 78 andNF= 98 + 2, while in white char, polymorphism and mosaicism for the chromosome number were revealed: 2n= 76–79, NF= 98 + 2. Using a routine chromosome staining technique, the karyotype of white char (2n= 78) was shown to be identical to that of Dolly Varden. In both karyotypes, similar sets of marker chromosomes were present: two pairs of submetacentric (SM), one pair of submeta-subtelocentric (SM-ST), one pair of large acrocentric (A), and one pair of large subtelocentric (ST) chromosomes. However, the karyotypes of Dolly Varden and white char differed in the number and location of nucleolus organizer regions (NORs). In Dolly Varden, single NORs located in the telomeric regions of the marker SM-ST chromosomes were observed. In white char, NORs were multiple and located both in the telomeric regions of the marker SM-ST chromosomes and on the short and long arms of large ST chromosomes. The identical marker chromosomes indicate considerable phylogenetic relatedness between Dolly Varden and white char from the Kamchatka River basin. Variation in NORs provides evidence for the reproductive isolation of these chars and their species status.     

Population genetic structure of northern Dolly Varden char Salvelinus malma malma in Asia and North America

The level of genetic differentiation of northern Dolly Varden char Salvelinus malma malma from Asia and North America was evaluated using the data on mtDNA variation (regions ND1/ND2, ND5/ND6, and Cytb/D loop) obtained by means of PCR-RFLP analysis. For S. m. malma, the mean values of haplotype and nucleotide diversity were 0.5261 +/- 0.00388 and 0.001558, respectively. The mean estimate of the population nucleotide divergence constituted 0.055%. It was demonstrated that S. m. malma on the most part of the species range examined (drainages of the Beaufort Sea, Chukotka Sea, Bering Sea, and the Sea of Okhotsk) was characterized by the population genetic structure with the low level of genetic differentiation and divergence. At the same time, populations from the Pacific Ocean Gulf of Alaska demonstrated marked genetic differentiation, supported by the high pairwise phi(ST) values (from 0.4198 to 0.5211) and nucleotide divergence estimates (mean divergence, 0.129%), from Asian and North American populations. Nested analysis of molecular variance (AMOVA) showed that most of the mtDNA variation in S. m. malma fell in the intrapopulation component (72.5%). At the same time, the differences between the populations (21.1%) and between the regions (6.4%) made lower contribution to the total variation.     

Differentiation of Dolly Varden char Salvelinus malma from Asia and North America inferred from PCR-RFLP analysis of mitochondrial DNA

Genetic differentiation of Dolly Varden char Salvelinus malma Walbaum from the Asian and North American Pacific coasts was studied. We examined restriction fragment length polymorphism of three mitochondrial DNA (mtDNA) fragments amplified in polymerase chain reaction, which encoded four NADH dehydrogenase subunits, the cytochrome b gene, and a D-loop segment. The mtDNA haplotypes were shown to form three phylogenetic groups, whose geographic distribution corresponded to three Dolly Varden subspecies: S. malma malma, S. malma krascheninnikovi, and S. malma lordi. The nucleotide sequence divergence between S. malma malma and S. malma krascheninnikovi was 3.8%; between S. malma malma and S. malma lordi, 3.1%; and between S. malma krascheninnikovi and S. malma lordi, 2.5%. The northern Dolly Varden S. malma malma from Asia was shown to be genetically identical to that from North America.     

Biology and fishery of the Dolly Varden charr (Salvelinus malma) on the east coast of Kamchatka

The paper studies populations of the Dolly Varden charr from some rivers in eastern Kamchatka in 2005–2007. Generally small variability is noted in biological parameters (length, weight, age, and sex composition), however, significant variability is revealed in size of individuals within age groups. Long-period cycles are marked in the dynamics of catches. The periodicity of catches of charr in different areas of east Kamchatka in general terms is the same, which gives reason to summarize the nature of its population dynamics in the whole coast.     

Phylogeography of northern Dolly Varden Salvelinus malma malma based on analysis of mitochondrial DNA

The northern Dolly Varden, Salvelinus malma malma, is a typical representative of arctic fauna distributed in northeastern Asia and northwestern North America. Because its spawning habitats were affected by Pleistocene glacial advances over most of its natural range, S. m. malma is among the most interesting objects of phylogeographic and microevolutionary studies. We reconstructed the genealogy of mtDNA haplotypes from 27 Alaskan and Asian populations to study the influence of glacial and geological vicariance events on the contemporary population genetic structure, phylogeographic subdivision and distribution of the northern Dolly Varden. Analysis of restriction site states in three PCR‐amplified mtDNA regions (ND1/ND2, ND5/ND6, Cytb/D‐loop; 47% of the mitochondrial genome) resolved 75 haplotypes in 436 fish. Similar patterns of subspecific variation apparent from hierarchical diversity and nested clade analyses of mtDNA haplotypes identify weak spatial differentiation and low levels of divergence. Our results suggest that (1) demographic history has been influenced by historical range expansions and recent isolation by distance, (2) present populations from Asia and North America were colonized from one main Beringian Refugium, and (3) that this taxon's ancestral population probably experienced a bottleneck in the Beringian Refugium during the late Pleistocene (Wisconsin) glacial period.     

Allozyme Diversity and Genetic Divergence of the Dolly Varden Salvelinus malma Walbaum from the Kuril Islands

Genetic variation was studied in the southern subspecies of the Asian Dolly Varden Salvelinus malma krascheninnikovi from the Kuril Islands. Thirty-six genetic loci controlling 19 enzyme systems were analyzed in 13 Dolly Varden populations from the Shumshu, Paramushir, Onekotan, Rasshua, Simushir, Urup, Iturup, and Kunashir islands. In the studied populations, the proportion of polymorphic loci was 35 to 85% and the mean heterozygosity was 0.104 to 0.173; populations from the Kunashir Island were characterized by maximum heterozygosity. In the island populations examined, significant interpopulation heterogeneity of allele frequencies was found for all studied population pairs. For the all island populations, the interpopulation diversity (G _ST = 0.188) was comparable to this parameter for the populations from the Kunashir Island only (G _ST = 0.170). Genetic distances between populations did not correlate with the corresponding geographical distances, which indicates the lack of a pronounced gene exchange between the island populations. Cluster analysis and multidimensional scaling based on genetic distances did not reveal clear groups among the studied populations but indicated greater similarity within the Iturup–Simushir–Urup–Paramushir group and a greater genetic divergence of the Kunashir, Onekotan, Rasshua, and especially Shumshu populations. In the Shumshu population, allele frequencies indicate the admixture of genes of the northern Dolly Varden. The observed pattern of genetic differentiation was probably caused largely by genetic drift under the conditions of a limited gene flow because of homing (which is typical of the Dolly Varden) and the presence of isolated nonanadromous populations. The population–genetic analysis of the Dolly Varden from the Kuril Islands does not give grounds to distinguish any other isolated char species in this region than S. malma, which is represented by the southern form S. m. krascheninnikovi with an admixture of the northern form S. m. malma in the Shumshu Island.     

Low levels of hybridization between sympatric Arctic char (Salvelinus alpinus) and Dolly Varden char (Salvelinus malma) highlights their genetic distinctiveness and ecological segregation

Understanding the extent of interspecific hybridization and how ecological segregation may influence hybridization requires comprehensively sampling different habitats over a range of life history stages. Arctic char (Salvelinus alpinus) and Dolly Varden (S. malma) are recently diverged salmonid fishes that come into contact in several areas of the North Pacific where they occasionally hybridize. To better quantify the degree of hybridization and ecological segregation between these taxa, we sampled over 700 fish from multiple lake (littoral and profundal) and stream sites in two large, interconnected southwestern Alaskan lakes. Individuals were genotyped at 12 microsatellite markers, and genetic admixture (Q) values generated through Bayesian‐based clustering revealed hybridization levels generally lower than reported in a previous study (<0.6% to 5% of samples classified as late‐generation hybrids). Dolly Varden and Arctic char tended to make different use of stream habitats with the latter apparently abandoning streams for lake habitats after 2–3 years of age. Our results support the distinct biological species status of Dolly Varden and Arctic char and suggest that ecological segregation may be an important factor limiting opportunities for hybridization and/or the ecological performance of hybrid char.     

On morphological diversity of southern Dolly Varden trout Salvelinus malma krascheninnikovi from water bodies of Sakhalin

Morphological and some biological specific features of nine samples of the southern form of Dolly Varden trout Salvelinus malma krascheninnikovi from water bodies of Sakhalin were studied. Comparative analysis was performed of our own estimates and estimates made by other authors of meristic characters of Sakhalin populations of Dolly Varden trout, as well as of samples from water bodies of the Kuril Islands and Western Kamchatka. Sakhalin populations of S. malma krascheninnikovi are characterized by a low level of morphological differentiation and the absence of geographic structure. No clinal latitudinal variation in sample averages was revealed in either of the nine studied meristic characters, including the number of vertebrae for populations of S. malma krascheninnikovi in the range from Bol’shoi Shantar Island to the south of Sakhalin. The cline in vertebral number from Kamchatka, along the Kuril Islands to Hokkaido, is most likely explained by specific features of distribution and introgressive hybridization at the secondary contact of populations of the northern and southern forms of Dolly Varden trout that originated from different glacial refugia and had a different vertebral number rather than by the temperature gradient.     

Growth and reproductive characteristics of rarely observed resident female Dolly Varden (Salvelinus malma malma) in North America

The replacement of native species by invasive species is one of the most critical threats to the biodiversity of aquatic systems today. However, little is known about potential effects of species invasions on the genetic diversity of indigenous species in cases where the latter coexist with the invader. Here we present an example of the indigenous amphipod Gammarus roeselii, which has been partly replaced by the invasive Dikerogammarus villosus in Lake Constance (C-Europe) and now mostly exists in small, isolated populations. We compared the genetic diversity, population structure, indicators for bottlenecks and migrations rates of G. roeselii before and after the invasion event from samples collected between 1999 and 2013. We expected a genetic impoverishment in the reduced and segregated populations of G. roeselii. However, no genetic measure on G. roeselii differed temporally or spatially over the investigated period, which indicates that D. villosus has not yet had an impact on G. roeselii at the genetic scale. Hence, even though a decline in population size of G. roeselii was found in Lake Constance, our results on the genetic scale contribute to recent findings that the overall impact of D. villosus on native species is not as strong as often discussed.

     

Isolated Fluvial Population of Dolly Varden Salvelinus malma on Bering Island (Commander Islands)

Journal of Ichthyology - For the first time on Bering Island (Commander Islands), an isolated population of northern Dolly Varden Salvelinus malma has been found. It inhabits only the Vodopadnaya...     

Genetic analysis of sympatric char populations in western Alaska: Arctic char (Salvelinus alpinus) and Dolly Varden (Salvelinus malma) are not two sides of the same coin

The North Pacific Ocean has been of great significance to understanding biogeography and speciation in temperate faunas, including for two species of char (Salmonidae: Salvelinus) whose evolutionary relationship has been controversial. We examined the morphology and genetics (microsatellite and mitochondrial DNA) of Arctic char (Salvelinus alpinus) and Dolly Varden char (Salvelinus malma) in lake systems in western Alaska, the eastern and western Arctic, and south of the Alaskan Peninsula. Morphologically, each lake system contained two forms: one (Arctic char) largely confined to lake habitats and characterized by greater numbers of pyloric caeca, gill rakers, and shallower bodies, and another (Dolly Varden) predominated in adjacent stream habitats and was characterized by fewer pyloric caeca, gill rakers, and deeper bodies. MtDNA partial (550 bp) d-loop sequences of both taxa were interspersed with each other within a single 'Bering' clade and demographic inferences suggested historical gene flow from Dolly Varden to Arctic char had occurred. By contrast, the taxa were strongly differentiated in sympatry across nine microsatellite loci in both lakes. Our data show that the two taxa are highly genetically distinct in sympatry, supporting their status as valid biological species, despite occasional hybridization. The interaction between these species highlights the importance of the North Pacific, and Beringia in particular, as an evolutionary wellspring of biodiversity.     

Development of Artificial Hybrid between Two Deepwater Sympatric Forms of Dolly Varden Salvelinus malma complex (Salmonidae) from Lake Kronotskoe (Eastern Kamchatka)

Journal of Ichthyology - The development of embryos and larvae of the hybrid (smallmouth charr females × bigmouth charr males) between unique endemic deepwater forms of northern Dolly Varden...     

Sympatric Diversification of Dolly Varden Salvelinus malma (Salmonidae) in an Extremely Small Ecosystem

Journal of Ichthyology - A planktivorous form of Salvelinus malma characterized by lacustrine winter spawning was discovered for the first time in north-east Asia in an isolated water system with...     

Variability of microsatellite DNA and genetic differentiation of populations in the resident form of Dolly Varden Salvelinus malma krascheninnikovi from Sakhalin

Variability in 11 microsatellite loci is followed for nine populations of the resident southern form of Dolly Varden Salvelinus malma krascheninnikovi from Sakhalin. In all studied populations, the level of genetic differentiation is significant with global values of F _st and R _st 0.182 and 0.329, respectively. On the NJ tree, two clusters are separated significantly: the former is represented by the populations from southwestern Sakhalin (the Novoselka River basin), and the latter includes all populations from southern and eastern Sakhalin. Abrupt size variations (bottleneck events) are suggested for several studied populations, and 50% reductions of allelic richness and average heterozygosity are observed in two lacustrine populations of the Novoselka River basin in comparison to the riverine populations of this basin. The results suggest an occurrence of a comparatively recent gene flow between the populations of resident Dolly Varden from Sakhalin and absence of reproductive isolation between the anadromous and resident forms. The degradation of the majority of Dolly Varden populations can be expected in the near future, and conservation projects should be developed.     

Introgressive hybridization between Dolly Varden Salvelinus malma and white‐spotted charr Salvelinus leucomaenis on Hokkaido Island, Japan

Intra‐ and interspecific phylogenetic analysis of Dolly Varden Salvelinus malma and white‐spotted charr Salvelinus leucomaenis throughout Hokkaido Island was conducted using nucleotide sequences of the mitochondrial DNA control region and three nuclear DNA markers [ribosomal DNA (ITS‐1), microsatellite ( u ‐85), SINE ( Fok ‐223)]. A total of 19 mtDNA haplotypes from 271 individuals of Dolly Varden, nine haplotypes from 134 white‐spotted charr were defined, and identified two well‐supported monophyletic clades for each species. Two haplotypes of Dolly Varden, however, were clustered together in the white‐spotted charr clade. Those haplotypes were found only from several rivers in the Shiretoko Peninsula, and a neighbouring river of that region. Analyses of ribosomal DNA and Fok ‐223 loci revealed that both species are characterized by having completely species‐specific diagnostic sequence and fragment patterns. Dolly Varden of the Shiretoko populations are typically allopatric in distribution where white‐spotted charr do not currently occur. In addition, incongruence in genetic relationships between mtDNA and nuclear DNA markers give strong evidence of historical mtDNA introgression between Dolly Varden and white‐spotted charr. Some white‐spotted charr diagnostic alleles in a nuclear microsatellite locus ( u ‐85) were found in some Shiretoko Dolly Varden populations, suggesting that introgressive hybridization might have also occurred in relatively recent contact and potentially ongoing evolutionary event. The present study presents an example of historical hybridization and introgression at the southernmost distribution limits of Dolly Varden.     

Basic description and some notes on the evolution of seven sympatric morphs of Dolly Varden Salvelinus malma from the Lake Kronotskoe Basin

The study examines the basic morphological and ecological features of Dolly Varden from Lake Kronotskoe (Russia, Kamchatka). Seven valid morphs different in head proportions, feeding, timing, and place of spawning have been determined in this ecosystem. The basic morphometric characteristics clearly separate Lake Kronotskoe morphs from each other, as well as from its potential ancestor (Dolly Varden). According to CVA analysis, the most notable morphological characteristics determining the mouth position are the length of a lower jaw and rostrum. Furthermore, five of seven morphs inhabit different depth zones of the lake and feed on different food resources. Our data suggest that reproductive isolation may be maintained by temporal/spatial isolation for two morphs with lacustrine spawning, and by spatial isolation only for the rest of the morphs with riverine spawning. The sympatric diversity of the Lake Kronotskoe charrs is exceptionally wide, and there are no other examples for seven sympatric morphs of genus Salvelinus to coexist within a single ecosystem. This study puts forward a three‐step hypothetical model of charr divergence in Lake Kronotskoe as a potential ground for future studies.     

Biology of resident Dolly Varden trout Salvelinus malma (Salmonidae) from a river with increased natural concentration of toxicants and suspension (Eastern volcanic belt of Kamchatka)

Specific biological features of the population of resident Dolly Varden trout Salvelinus malma inhabiting upper reaches of the Falshivaya River whose water in mean water period has natural increased concentration of copper (0.006–0.018 mg/L), zinc (0.061–0.072 mg/L), and suspension (6–8 mg/L) are described. Its spawning was recorded in mouths of tributaries where (unlike the main river) water is not toxic, its mineralization is two times lower, and turbidity does not exceed 1 mg/L. Among juveniles that left the ground, individuals with scoliosis (11%), undeveloped fins (16%), and exophthalmia occur. The numbers of the population are maintained due to early maturation and spawning before the accumulation in the body of a lethal dose of toxicants and critical amount of functional disturbances. In prespawning fish (3+), multiple pathologies of the histological structure of the gills, liver, and spleen up to necroses were recorded. On average, 10% of maturing oocytes in the ovaries are at different stages of apoptosis. The frequency of pathologies correlates with the concentration in the liver of copper, zinc, and sulphur whose concentration significantly increases with an increase in body weight.