Resource utilization by bull char and cutthroat trout in a mountain stream in Montana,U.S.A.

Resource utilization of sympatric populations of bull char,Salvelinus confluentus, and west-slope cutthroat trout,Oncorhynchus clarki lewisi, were studied by underwater observations of foraging behaviour and microhabitat use, and dietary analysis in a mountain stream of the Flathead River Basin, northwest Montana, U.S.A. Nearly 70% of bull char were categorized as benthic foragers, which moved constantly and captured prey primarily from the streambed, while all cutthroat trout were drift foragers, which held relatively fixed focal points in the midwater layers of pools during foraging. The composition of stomach contents was markedly different between the two species. Bull char fed primarily on baetid mayflies captured from the benthos or drift, whereas cutthroat trout ate primarily terrestrial invertebrates. The species also used different microhabitats. Bull char held positions close to the streambed and rarely strayed far from overhead cover, whereas cutthroat trout held focal points farther above the bed and far from overhead cover. Dietary segregation between these two salmonids appeared to result not only from differences in foraging tactics but also in the foraging microhabitats. Resource partitioning is considered to be one of important mechanisms allowing coexistence of these two stream salmonids.     

Experimentally induced foraging mode shift by sympatric charrs in a Japanese mountain stream

Foraging mode shifts may allow animals flexibility to adaptto a variety of ecological conditions. Theory holds that ectothermssuch as fishes should shift from ambush to active search modeswhen prey density declines, to maintain a minimum encounterrate. Salvelinus malma (Dolly Varden) sympatric withS. leucomaenis(white-spottedchirr) in a northern Japan mountain stream were observed toshift from ambushing drifting invertebrates at fixed focal pointsto picking benthic invertebrates from the substrate along circuitousroutes throughout pools, as drift declined throughout summer.Experimentally reducing drift in two pools during early summerwhen all charr were drift foragers caused subordinate malmato make marked shifts from drift to benthic foraging within1 h, whereas nearly all leucomaenis remained drift foragers.Some charr of both species responded to reduced drift by emigratingfrom pools. Combined results of three replicate experimentsrevealed that malma shifted to benthic foraging when their rateof drift forays was reduced below a specific threshold, therebyupholding predictions of the theory. Adaptive foraging modeshirts may promote coexistence of these congeneric charrs inreaches of Japanese mountain streams     

The Divergence of the Dolly Varden Char Salvelinus malma in the Asian Northern Pacific Populations Inferred from the PCR–RFLP Analysis of Mitochondrial DNA

Genetic differentiation of the Dolly Varden char Salvelinus malmaWalbaum was studied in five populations from the western part of the Northern Pacific. Using restriction analysis (RFLP), we examined polymorphism of three mitochondrial DNA (mtDNA) fragments amplified in polymerase chain reaction (PCR). MtDNA haplotypes were shown to fall into two phylogenetic groups, which probably reflect the existence of two previously described subspecies of Asian Dolly Varden,S. malma malma and S. malma krascheninnikovi. The divergence of mtDNA nucleotide sequences in the Dolly Varden subspecies (about 4%) corresponds to the differences between the valid char species from the genus Salvelinus.     

Effects of forestry on the thermal habitat of Dolly Varden (<Emphasis Type="Italic">Salvelinus malma</Emphasis>)

Forestry activities in riparian areas are known to affect stream communities considerably. Not only do riparian deforestation resulting from agriculture or urbanization developments affect stream communities but extensive commercial plantation and forestry practices can alter stream environments adjacent to remaining, intact or secondary forests. Because forestry often includes the construction of logging roads through the riparian zone, this can directly degrade stream environments. Twelve streams in the Shiretoko Peninsula, Hokkaido were investigated so as to determine the effects of forestry practices on stream temperature, periphyton biomass, grazer (benthic invertebrates) biomass and Dolly Varden (Salvelinus malma Walbaum) biomass. The greater the proportion of planted area in the catchment, the higher the stream temperature. Stream temperature directly affects periphyton biomass and Dolly Varden biomass negatively. Neither stream temperature nor periphyton biomass predicted grazer biomass, whereas a positive correlation was found between grazer biomass and Dolly Varden biomass that forage on invertebrates. The overall results indicated that Dolly Varden in the Shiretoko streams were negatively affected by forestry practices and the resultant stream temperature increases. Without effective future riparian forest management, the complex effects of both riparian disturbance and ongoing global warming could further reduce Dolly Varden populations in the region.     

Genetic Divergence of Chars of the Genus <Emphasis Type="Italic">Salvelinus</Emphasis> from Kronotsky Lake (Kamchatka Peninsula)

Chars of the genus Salvelinus, inhabiting lakes and lake-river systems, comprise morphological and ecological forms whose taxonomic status is under dispute. In the present work, we have examined genetic variation and divergence in various chars from the Kronotsky lake basin: the lacustrine chars (white, nose, and longhead) and riverine Dolly Varden Salvelinus malma. The study was conducted using analysis of allozyme and microsatellite loci, myogens, RAPD, and restriction analysis of two mtDNA segments. The estimates of heterozygosity at allozyme and microsatellite loci were similar to the corresponding parameters in populations of northern Dolly Varden and Arctic char. Heterozygote deficit was recorded in both samples of separate forms, and in the total sample of all chars from Kronotsky Lake. For allozyme and microsatellite loci, appreciable genetic differentiation among the samples of different char forms was found, which was comparable to that among the spatially isolated populations of northern Dolly Varden. This result indicates reproductive isolation among the char forms examined. However, this isolation is not complete, because no fixed differences between the forms by any of the genetic systems analyzed were found. The genetic differentiation among different forms of lacustrine chars, which corresponds to the interpopulation rather than interspecific level, is thought to be explained by their comparatively recent divergence.__________Translated from Genetika, Vol. 41, No. 8, 2005, pp. 1096–1107.Original Russian Text Copyright © 2005 by Salmenkova, Omel’chenko, Radchenko, Gordeeva, Rubtsova, Romanov.     

Genetic divergence of mitochondrial DNA in white char <Emphasis Type="Italic">Salvelinus albus</Emphasis> and northern Dolly Varden char <Emphasis Type="Italic">Salvelinus malma malma</Emphasis>

Comparative analysis of mitochondrial DNA variation was performed in white char Salvelinus albus Glubokovsky and in its putative ancestor species, northern Dolly Varden char Salvelinus malma malma Walbaum. Highly statistically significant differentiation of S. albus and S. m. malma in the areas of sympatric (Kamchatka River basin) and allopatric (Kronotskoe Lake and Kronotskaya River) residence was demonstrated. The mtDNA divergence between S. albus and S. m. malma did not exceed the range of intraspecific variation in the populations of northern Dolly Varden char. At the same time, clusterization pattern of the Salvelinus chars provides hypothesis on the common origin of two allopatric populations of white char. Genealogical analysis of haplotypes indicates that S. albus and S. m. malma currently demonstrate incomplete divergence of mitochondrial lineages. The low nucleotide divergence estimates between S. albus and S. m. malma reflect the short time period since the beginning of the divergence of ancestral lineages. These estimates are determined by ancestral polymorphism and haplotype exchange between the diverged phylogenetic groups as a result of introgressive hybridization.     

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.     

Asymmetric competition drives lake use of coexisting salmonids

To what degree are population differences in resource use caused by competition and the occupation of adjacent positions along environmental gradients evidence of competition? Habitat use may be the result of a competitive lottery, or restricted by competition. We tested to what extent population differences in habitat use of two salmonids, cutthroat trout (Oncorhynchus clarki) and Dolly Varden charr (Salvelinus malma) were influenced by interspecific competition. We hypothesized that the depth distribution of Dolly Varden charr would be affected by competition from the more littoral and surface-oriented cutthroat trout, and that the depth distribution of cutthroat trout would be little affected by competition from Dolly Varden charr. Sympatric populations of cutthroat trout and Dolly Varden charr were created by reciprocal transfers of previously allopatric populations in two experimental lakes. We found evidence of asymmetric competition, as Dolly Varden charr were displaced from littoral habitats when sympatric with cutthroat trout, whereas cutthroat trout remained unaffected by the presence of Dolly Varden charr. Evolved differences between the species, and differences between experimental lakes, also contributed to population differences in habitat use, but asymmetric competition remained as the main driver of different depth distributions in sympatry.     

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.     

Karyological Differentiation of Northern Dolly Varden and Sympatric Chars of the Genus <Emphasis Type="Italic">Salvelinus</Emphasis> in Northeastern Russia

In many cases the taxonomic status of sympatric chars (with exception of white-spotted char Salvelinus leucomaenis) is not clear and is actively debated. We karyotyped three pairs of sympatric chars inhabiting the Russian Far East-northern Dolly Varden, S. malma malma, and Lavanidov's char, S. levanidovi, from the Yama River (northern coast of the Sea of Okhotsk, Magadan Region), northern Dolly Varden and Taranetz's char, S. taranetzi, from Lake Achchen (east Chukotka), and northern Dolly Varden and white char, S. albus, from the Kamchatka River (Kamchatka Peninsula). Three of them had similar chromosome numbers. But all chars studied had an individual and discrete set of karyotypic characters, which enabled reliable identification each of them by chromosome number, chromosome arm number, and number and location of active nuclear organizer regions.     

Variation of cytochrome b gene in sympatric chars from Kronotsky Lake (Kamchatka oblast)

In previously identified sympatric char forms (species) from Kronotsky Lake (Kamchatka Peninsula) and a number of other chars from the genus Salvelinus, nucleotide sequences of the mtDNA cytochrome b gene were determined and phylogenetic analysis of these forms was performed. The divergence of the cytochrome b gene in the Kronotsky char group (0.48%) coincided with the interpopulation difference estimates in northern Dolly Varden, which indicates a relatively recent divergence of the chars from Kronotsky Lake. Haplotypes of each form of the Kronotsky Lake chars are divided into two groups, which belonged to two different phylogenetic mtDNA lineages of northern Dolly Varden from Chukotka and Kamchatka. The formation of the Kronotsky char forms proved to be unrelated to the observed variability of the mtDNA cytochrome b gene.     

Genetic divergence of chars of the genus Salvelinus from Kronotsky Lake (Kamchatka Peninsula)

Chars from the genus Salvelinus, inhabiting lakes and lake-river systems, belong to morphologically and ecologically different forms whose taxonomic status is under dispute. In the present work, we have examined genetic variation and divergence in various chars from the Kronotsky lake basin: the lacustrine chars (white, nose, and long-head) and Dolly Varden char Salvelinus malma. The study was conducted using analysis of allozyme and microsatellite loci, myogens, RAPD, and restriction analysis of two mtDNA segments. The estimates of heterozygoisty at allozyme and microsatellite loci were similar to the corresponding parameters in populations of northern Dolly Varden and Arctic char. Heterozygote deficit was recorded in both samples of individual forms, and in the combined sample of all chars from Kronotsky Lake. For both markers, appreciable genetic differentiation among the samples of different char forms was found, which was comparable to that among the spatially isolated populations of northern Dolly Varden. This result indicates reproductive isolation among the char forms examined. However, this isolation is not complete, because no fixed differences between the forms by any of the genetic systems analyzed was found. The genetic differentiation among different forms of lacustrine chars, which corresponds to the interpopulation rather than interspecies level, is thought to be explained by their comparatively recent divergence.     

Microsatellite genetic variation of Asian populations of Dolly Varden char

Genetic variation at eight microsatellite loci was examined in 21 populations of Dolly Varden charrs, Salvelinus malma, representing five geographical regions (Kamchatka Peninsula, Sea of Okhotsk coast, Sea of Japan coast, Sakhalin Island, and Kuril islands). Hierarchical analysis of molecular variance showed that 11% (58% in terms of R-statistics) of the variation was distributed among of northern subspecies of Dolly Varden, Salvelinus malma malma and southern Asian Dolly Varden, S. m. krascheninnikovi while similar values were attributed to the among-regional level within northern Dolly Varden 9% (7%) and southern Asian Dolly Varden 11% (14%). Permutation-based tests indicated a mutational component to genetic differentiation based on allelic size variance and suggested that divergence of the two subspecies had occurred at least 3,000 generations ago. On large spatial scales (within the Asian range of the species), populations clustered according to their geographical location. On smaller scales (within regions and subregions) correlation between genetic and geographic distances was not significant. Northern Dolly Varden has higher allelic diversity and more private alleles than southern subspecies, this probably indicating differences in demographic history.     

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 inter-population heterogeneity of allele frequencies was found for all studied population pairs. For the total population of all islands, the inter-population diversity (GST = 0.188) was comparable to this parameter for the total population from the Kunashir Island (GST = 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 Dolly Varden 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.     

Differentiation of the Dolly Varden Char (Salvelinus malma) and the Taranetz Char (Salvelinus taranetzi) Inferred from the PCR–RFLP Analysis of Mitochondrial DNA

Genetic differentiation of two sympatric char species, the Dolly Varden char (Salvelinus malma Walbaum) and the Taranetz char (S. taranetzi Kaganovski), has been studied. Restriction fragment length polymorphism (RFLP) analysis has been used to compare three mitochondrial DNA (mtDNA) fragments (ND1/ND2, ND5/ND6, and Cytb/D-loop) amplified via polymerase chain reaction (PCR). The divergence between S. malma and S. taranetzi inferred from mtDNA nucleotide sequences is 2.8%; between S. leucomaenis and S. taranetzi, 7.1%; and between S. malma and S. leucomaenis, 7.5%. The absence of common haplotypes and the degree of divergence indicate that the Dolly Varden char and the Taranetz char are genetically isolated and confirm that S. taranetzimay be regarded as a separate species.     

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.     

Is isolation by adaptation driving genetic divergence among proximate Dolly Varden char populations?

Numerous studies of population genetics in salmonids and other anadromous fishes have revealed that population structure is generally organized into geographic hierarchies (isolation by distance), but significant structure can exist in proximate populations due to varying selective pressures (isolation by adaptation). In Chignik Lakes, Alaska, anadromous Dolly Varden char (Salvelinus malma) spawn in nearly all accessible streams throughout the watershed, including those draining directly to an estuary, Chignik Lagoon, into larger rivers, and into lakes. Collections of Dolly Varden fry from 13 streams throughout the system revealed low levels of population structure among streams emptying into freshwater. However, much stronger genetic differentiation was detected between streams emptying into freshwater and streams flowing directly into estuarine environments. This fine‐scale reproductive isolation without any physical barriers to migration is likely driven by differences in selection pressures across freshwater and estuarine environments. Estuary tributaries had fewer larger, older juveniles, suggesting an alternative life history of smolting and migration to the marine environment at a much smaller size than occurs in the other populations. Therefore, genetic data were consistent with a scenario where isolation by adaptation occurs between populations of Dolly Varden in the study system, and ecological data suggest that this isolation may partially be a result of a novel Dolly Varden life history of seawater tolerance at a smaller size than previously recognized.     

Invading rainbow trout usurp a terrestrial prey subsidy from native charr and reduce their growth and abundance

Movements of prey organisms across ecosystem boundaries often subsidize consumer populations in adjacent habitats. Human disturbances such as habitat degradation or non-native species invasions may alter the characteristics or fate of these prey subsidies, but few studies have measured the direct effects of this disruption on the growth and local abundance of predators in recipient habitats. Here we present evidence, obtained from a combined experimental and comparative study in northern Japan, that an invading stream fish usurped the flux of allochthonous prey to a native fish, consequently altering the diet and reducing the growth and abundance of the native species. A large-scale field experiment showed that excluding terrestrial invertebrates that fell into the stream with a mesh greenhouse reduced terrestrial prey in diets of native Dolly Varden charr (Salvelinus malma) by 46–70%, and reduced their growth by 25% over six weeks. However, when nonnative rainbow trout (Oncorhynchus mykiss) were introduced, they monopolized these prey and caused an even greater reduction of terrestrial prey in charr diets of 82–93%, and reduced charr growth by 31% over the same period. Adding both greenhouse and rainbow trout treatments together produced similar results to adding either alone. Results from a comparative field study of six other stream sites in the region corroborated the experimental findings, showing that at invaded sites rainbow trout usurped the terrestrial prey subsidy, causing a more than 75% decrease in the biomass of terrestrial invertebrates in Dolly Varden diets and forcing them to shift their foraging to insects on the stream bottom. Moreover, at sites with even low densities of rainbow trout, biomass of Dolly Varden was more than 75% lower than at sites without rainbow trout. Disruption of resource fluxes between habitats may be a common, but unidentified, consequence of invasions, and an additional mechanism contributing to the loss of native species Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Evidence for historical introgression along a contact zone between two species of char (Pisces: Salmonidae) in northwestern North America

Phylogeographic analyses can yield valuable insights into the geographic and historical contexts of contact and hybridization between taxa. Two species of char (Salmonidae), Dolly Varden (Salvelinus malma) and bull trout (S. confluentus) have largely parapatric distributions in watersheds of northwestern North America. They are, however, sympatric in several localities and hybridization and some introgression occurs across a broad area of contact. We conducted a comparative phylogenetic analysis of Dolly Varden and bull trout to gain a historical perspective of hybridization between these species and to test for footprints of historical introgression. We resolved two major Dolly Varden mitochondrial DNA (mtDNA) clades (with 1.4-2.2% sequence divergence between haplotypes) that had different geographical distributions. Clade N is distributed across most of the range of Dolly Varden, from southern British Columbia through to the Kuril Islands in Asia. Clade S had a much more limited distribution, from Washington state, at the southern limit of the Dolly Varden range, to the middle of Vancouver Island. The distribution and inferred ages of the mtDNA clades suggested that Dolly Varden survived the Wisconsinan glaciation in a previously unsuspected refuge south of the ice sheet, and that Dolly Varden and bull trout were probably in continuous contact over most of the last 100,000 years. When bull trout were included in the phylogenetic analysis, however, the mtDNA of neither species was monophyletic: Clade S Dolly Varden clustered within the bull trout mtDNA clade. This pattern was discordant with two nuclear phylogenies produced (growth hormone 2 and rRNA internal transcribed sequence 1), in which Dolly Varden and bull trout were reciprocally monophyletic. This discordance between mtDNA- and nDNA-based phylogenies indicates that historical introgression of bull trout mtDNA into Dolly Varden occurred. Percent sequence divergence within these introgressed Dolly Varden (clade S) was 0.2-0.6%, implying that the introgression occurred prior to the most recent glaciation. Our analysis and other evidence of contact between divergent lineages in northwestern North America strongly suggests that the area may be the site of previously unsuspected suture zones of aquatic biotas.     

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.