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.     

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.     

Offshore Dolly Varden charr (<Emphasis Type="Italic">Salvelinus malma</Emphasis>) in the North Pacific

Many studies have investigated the ecology of charrs in freshwater, however, little is known about charrs in the ocean. This study examined the distribution, seasonal abundance, and some biological features of Dolly Varden (Salvelinus malma) in the Pacific Ocean. An analysis of by-catch data of Japanese offshore salmon monitoring showed that Dolly Varden were distributed across a wide range in the offshore waters of the Pacific Ocean, including the Japan Sea, Bering Sea, and Okhotsk Sea. The catch per unit effort showed a sharp increase from May to August, followed by a sharp decrease in September. Offshore areas served as an important summer habitat for anadromous Dolly Varden.     

Genetic connectivity between sympatric populations of closely related char species,Dolly Varden <Emphasis Type="Italic">Salvelinus malma</Emphasis> and white char <Emphasis Type="Italic">Salvelinus albus</Emphasis>

The closely related chars Salvelinus malma and Salvelinus albus, which sympatrically inhabit the Kamchatka River basin and Kronotsky Lake (Kamchatka), attract the attention of the researchers because of their debated origin and taxonomic status. Previous studies of sympatric populations of these chars revealed small but statistically significant genetic differences between these species at a number of molecular markers, suggesting the presence of the genetic exchange and hybridization. In this study, based on genotypic characterization of nine microsatellite loci, a considerable level of historical and contemporary genetic migration between sympatric populations of these chars was demonstrated. At the individual level, a high degree of hybridization was observed, mainly among the Dolly Varden individuals from the studied populations. The obtained evidences on the genetic connectivity between sympatric S. malma and S. albus do not support the separate species status of S. albus.     

Differentiation of Dolly Varden Char <Emphasis Type="Italic">Salvelinus malma</Emphasis> 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.__________Translated from Genetika, Vol. 41, No. 5, 2005, pp. 626–634.Original Russian Text Copyright © 2005 by Oleinik, Skurikhina, Brykov, Crane, Wenburg.     

Phylogeography of southern Asian Dolly Varden char <Emphasis Type="Italic">Salvelinus malma krascheninnikovi</Emphasis>: Genealogical analysis of mitochondrial DNA

Phylogeography of southern Asian Dolly Varden char was studied using the data on mtDNA variation (regions ND1/ND2, ND5/ND6, and Cytb/D loop) obtained using PCR-RFLP analysis. Analysis of contemporary population genetic structure showed that Salvelinus malma krascheninnikovi throughout the whole species range was characterized by high population differentiation in combination with rather small differences between the populations from remote regions. The genealogy of mtDNA haplotypes was reconstructed and nested clade analysis of geographical distances was performed. Geographical distribution of mtDNA haplotypes of S. m. krascheninnikovi was explained by population genetic processes (restricted gene flow), as well as by historical demographic events (range expansion and fragmentation). It was demonstrated that the main demographic events were associated with cyclic processes of the geological formation of the Sea of Japan and adjacent territories. Furthermore, genealogical tree of S. m. krascheninnikovi contained the traces of secondary contact between isolated phylogeographical lineages.     

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.     

A comparison of different structures and methods for estimating age of northern-form Dolly Varden <Emphasis Type="Italic">Salvelinus malma malma</Emphasis> from the Canadian Arctic

Assessment of anadromous northern-form Dolly Varden Salvelinus malma malma in the Western Canadian Arctic requires reliable methods for estimating ages. Additionally, conservation efforts warrant determining whether fin rays provide a non-lethal alternative to otoliths. Precision and bias of whole and sectioned otoliths, and sectioned pectoral and pelvic fin rays were examined. Two age readers with different levels of experience ageing this species read each structure three times. Coefficient of variation (CV) was calculated to measure precision, and age bias plots were created for each method of preparation/structure within and between readers. The experienced reader demonstrated the highest precision with sectioned otoliths (CV = 1.6 %) followed by whole otoliths (CV = 4.2 %) while pectoral and pelvic fins were the lowest, CV = 7.7 % and 7.5 %, respectively. The age bias plot showed little difference between whole and sectioned otoliths, although greater imprecision/bias was evident for whole otoliths at age ≥9. Compared to otoliths, fin rays produced younger age estimates starting at 5 years; however, pelvic fins were more biased towards younger estimates than pectoral fins. The less experienced reader had greater inconsistencies, tending to overage younger and underage older samples for all methods compared to the more experienced reader, underscoring the importance of experience when estimating age for this species. We conclude that both types of fin rays are a poor non-lethal alternative to otoliths for fish ≥5 years and recommend an experienced ager could use whole otoliths up to age 8 and sectioned otoliths for fish ≥9 years (>500 mm fork length).     

Flexible seaward migration of Dolly Varden <Emphasis Type="Italic">Salvelinus malma</Emphasis> in the Shiretoko Peninsula,Hokkaido, Japan

The migratory history of Dolly Varden Salvelinus malma was investigated in 30 streams in the Shiretoko Peninsula, Hokkaido, Japan, which is in the southern part of their distribution. Captured Dolly Varden were classified into two categories by their body coloration: silvery fish were thought to be an anadromous form, and non-silvery fish a fluvial form that remains in freshwater throughout its life. Forty-seven of 1,739 Dolly Varden collected in the spring and ten of 2,126 Dolly Varden collected in the autumn were silvery fish. Twenty silvery fish and 120 non-silvery fish were examined by analysis of otolith Sr:Ca ratio. Three silvery fish from autumn and 15 non-silvery fish had migrated to salt water or brackish water. The other specimens of silvery fish were regarded as smolt or pre-smolt before seaward migration. The present study suggests that Dolly Varden in the Shiretoko Peninsula have a flexible migratory history, where some of the non-silvery fish utilize marine environments while some individuals follow an anadromous life history.     

Genetic divergence of populations of the white char <Emphasis Type="Italic">Salvelinus Albus</Emphasis>, northern and southern forms of malma <Emphasis Type="Italic">S. malma</Emphasis> (Salmonidae), by microsatellite loci of DNA

A very small but statistically significant genetic differentiation by allele frequencies (in the Kamchatka basin F _ST = 0.016 and in Kronotskoe Lake F _ST = 0.063) is revealed between sympatric populations of the northern malma Salvelinus malma malma and of the white char S. albus by analysis of nine microsatellite loci. Factor analysis and clusterization of individual multilocal genotypes in the sum sample of sympatric S. malma malma and S. albus also demonstrate their extremely weak genetic differentiation. Divergence between geographically isolated (allopatric) populations of each of these chars exceeds by two-three times the interspecies differentiation in the zones of sympatry. Samples from allopatric populations clusterize by the geographic principle but not by the taxonomic principle. Such a situation may be explained by the absence of reproductive isolation and presence of hybridization between sympatric S. malma malma and S. albus with the simultaneous existence of some limitations of genetic exchange related to the ecological specialization of these chars. Analysis of microsatellite loci revealed highly significant genetic differences between the northern and southern forms of malma S. malma krascheninnikovi by many times exceeding genetic differences between S. malma malma and S. albus. The obtained data do not confirm the independent species status of S. albus but support the opinion of the species level of differences between the northern and southern forms of malma.     

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.     

The trace element contents in tissues of Dolly Varden <Emphasis Type="Italic">Salvelinus malma</Emphasis> Walbaum, 1792 sympatric morphs from Lake Kronotskoe (Kamchatka)

The liver, gonads, gills, and muscles of five trophically distant charr morphs from Lake Kronotskoe, as well as silty sediments of the lake, were tested for trace-element contents. Concentrations of Mn, Fe, Ni, Cu, Zn, Rb, Sr, As, and Se were detectable in the fish tissues, while the xenobiotics Hg, Cd, and Pb were not determined. The current data indicate a strong correlation between trace-element content and the ecology of each morph and no relationship to the fish length. In the majority, the Ni, Cu, and Zn levels were strongly dependent on trophic biology. The contents of Rb, Mn, and Se were dependent on the distribution of the charr morphs along the depth gradient and correlated with the accumulation of these elements in the bottom sediments.     

Morphophysiological patterns of life history strategy adoption in Dolly Varden <Emphasis Type="Italic">Salvelinus malma</Emphasis> in Kamchatka

Growth, age composition, gonad maturity and patterns of life history strategy adoption in the population of Dolly Varden from the Kol River (West Kamchatka) are studied. Major factors controlling differentiation of juveniles and trajectories of ontogenetic developments, growth rate and gametogenesis in generations in particular years. The differentiation in a generation into anadromous and resident groupings is completed mainly at the age of 4 years. The process of formation of resident and migratory life history strategy in Dolly Varden is epigenetic. Bifurcation in the life of one specimen is observed once in life, and that in a generation is observed four times: at the first, the second, the third, or the fourth year. The life cycle of females from the Kol River is invariant until they reach the age of 1 year, and that in males is during the 4 years of the freshwater phase. In Kamchatka, all spawners, including precocious males, may reproduce several times during life.     

Population genetic structure of the endangered and endemic medicinal plant <Emphasis Type="Italic">Commiphora</Emphasis><Emphasis Type="Italic">wightii</Emphasis>

Commiphora wightii is a medicinally important endangered species endemic to the Thar Desert of Rajasthan, India and adjoining areas of Pakistan. The populations of this species are declining sharply because of its extensive use as a natural herb. Random amplified polymorphic DNA analysis was conducted to find the genetic variation among 7 populations of C. wightii. Of the 100 random primers screened, 44 primers yielded 220 loci. Statistical analysis indicated low genetic diversity (H _pop = 0.0958; I = 0.1498; mean polymorphic loci = 14.28%), and high genetic differentiation among the populations (G _ST = 0.3990; AMOVA Φ _ST of 0.3390; Bayesian θ ^(II) = 0.3002). The low genetic diversity may be due to geographic isolation and restricted gene flow (N _m = 0.7533) between the fragmented populations. Unsustainable utilization of the plant has fragmented the population continuum which served the purpose of genetic exchange between populations. Mantel’s test was performed which revealed a highly significant positive correlation between genetic and geographic distance (r ² = 0.614, P = 0.023) among the populations studied. Low variation can also be attributed to poor seed setting and the slow growth pattern of the species, which is also an apomict. In UPGMA dendrogram the Commiphora wightii samples were divided into two major and one minor cluster. These findings can serve as a guide to preserving the genetic resources of this medicinal plant species.     

<Emphasis Type="Italic">Diplazium</Emphasis> hybrids involving <Emphasis Type="Italic">D. plantaginifolium</Emphasis> and <Emphasis Type="Italic">D</Emphasis>. <Emphasis Type="Italic">ternatum</Emphasis> from Mexico and Central America

Here we evaluate the origins and relationships of Mexican and Central American Diplazium hybrids derived from crosses involving either D. plantaginifolium or D. ternatum. Based on study of live plants and herbarium specimens, we distinguish D. ×verapax from the similar D. riedelianum and present evidence that the former is a sterile hybrid derived from a cross between D. plantaginifolium and D. werckleanum. We also describe new hybrids, D. ×torresianum and D. ×subternatum from Mexico and northern Central America. Both involve D. ternatum as one parent. Diplazium. cristatum is the other putative parent of D. ×torresianum, and D. plantaginifolium is the second parent of D. ×subternatum. We also designate lectotypes for D. cordovense and D. dissimile.     

<Emphasis Type="Italic">Ty</Emphasis>1<Emphasis Type="Italic">/copia</Emphasis>- and <Emphasis Type="Italic">Ty</Emphasis>3<Emphasis Type="Italic">/gypsy</Emphasis>-like DNA sequences in <Emphasis Type="Italic">Helianthus </Emphasis>species

Two repeated DNA sequences isolated from a partial genomic DNA library of Helianthus annuus, p HaS13 and p HaS211, were shown to represent portions of the int gene of a Ty3 /gypsy retroelement and of the RNase-Hgene of a Ty1 /copia retroelement, respectively. Southern blotting patterns obtained by hybridizing the two probes to BglII- or DraI-digested genomic DNA from different Helianthus species showed p HaS13 and p HaS211 were parts of dispersed repeats at least 8 and 7 kb in length, respectively, that were conserved in all species studied. Comparable hybridization patterns were obtained in all species with p HaS13. By contrast, the patterns obtained by hybridizing p HaS211 clearly differentiated annual species from perennials. The frequencies of p HaS13- and p HaS211-related sequences in different species were 4.3x10(4)-1.3x10(5) copies and 9.9x10(2)-8.1x10(3) copies per picogram of DNA, respectively. The frequency of p HaS13-related sequences varied widely within annual species, while no significant difference was observed among perennial species. Conversely, the frequency variation of p HaS211-related sequences was as large within annual species as within perennials. Sequences of both families were found to be dispersed along the length of all chromosomes in all species studied. However, Ty3 /gypsy-like sequences were localized preferentially at the centromeric regions, whereas Ty1/ copia-like sequences were less represented or absent around the centromeres and plentiful at the chromosome ends. These findings suggest that the two sequence families played a role in Helianthusgenome evolution and species divergence, evolved independently in the same genomic backgrounds and in annual or perennial species, and acquired different possible functions in the host genomes.     

<Emphasis Type="Italic">Cantharellus</Emphasis> sect. <Emphasis Type="Italic">Amethystini</Emphasis> in Asia

In this contribution on the genus Cantharellus in Asia, C. subvaginatus is described from the Republic of Korea as a close relative to the Chinese C. vaginatus, which is here reported for the first time from India. Both species are here placed in Cantharellus subg. Cantharellus sect. Amethystini, together with the Indian C. pseudoformosus (syn.: C. umbonatus) and the Malayan C. subamethysteus. As such, Asia has suddenly become the continent with the highest diversity for Amethystini. Species delimitation in sect. Amethystini is molecularly supported by a combined phylogenetic analysis of rDNA sequences obtained for LSU and ITS and additionally suggests the existence of a still undescribed species in North America. Character variability is discussed for all known members of Amethystini, including atypical specimens of the North American C. lewisii that are morphologically more reminiscent of the South Korean C. subvaginatus.     

Genetic relationships of Chukchi charr <Emphasis Type="Italic">Salvelinus andriashevi</Emphasis> and Taranetz charr <Emphasis Type="Italic">Salvelinus taranetzi</Emphasis>

On the basis of sequence analysis of the mitochondrial DNA (mtDNA) control region (CR), cytochrome b (Cytb), and cytochrome oxidase-1 (CoI) genes, the relationships of endemic species Salvelinus andriashevi Berg, 1948, represented by the only population from Lake Estikhed (Chukotka), were estimated. The data on the genealogical analysis of mtDNA haplotypes supported phylogenetic closeness of S. andriashevi and S. taranetzi. It was also demonstrated that the specimens of Chukchi charr, along with Salvelinus sp. 4 (Lake Nachikinskoe), S. krogiusae (Lake Dal’nee), S. boganidae and S. elgyticus (Lake El’gygytgyn), and S. a. erythrinus from Canada’s Northwest Territories (NWT) belonged to the Arctic group of Taranetz charr. The problem of coincidence of taxonomic differentiation of charrs of the genus Salvelinus based on morphological and genetic analyses is discussed.     

Systematics and biology of the East Siberian char <Emphasis Type="Italic">Salvelinus leucomaenis</Emphasis>

A review that considers in detail for the first time the taxonomic position, phenetic diversity, and principal features of the endemic species East Siberian char Salvelinus leucomaenis in the range from the northeast of Asia to Japan based on own and published data is provided. In comparison with other species of chars, S. leucomaenis is characterized by a smaller morphological and ecological diversity. A migratory form feeding on fish is dominant in the range. At the same time, parallel to large migratory forms, other life forms—resident riverine and lacustrine-riverine forms represented by males and females and riverine and dwarf males were found in S. leucomaenis. Individuals with various color variations occur everywhere. Resident forms are rare in the north of the range; they are dominant in the south of the range, in Japan.     

Stream-dwelling Dolly Varden (<Emphasis Type="Italic">Salvelinus malma</Emphasis>) density and habitat characteristics in stream sections installed with low-head dams in the Shiretoko Peninsula,Hokkaido, Japan

Numerous low-head dams have been constructed for erosion control and installed in almost all mountainous streams in Japan. Shiretoko Peninsula, Hokkaido, is considered as one of the most intact areas in Japan, however, more than 331 low-head dams have been installed in 39 streams of the peninsula since the 1960s. In this study, the density of stream-dwelling Dolly Varden (Salvelinus malma) and the habitat characteristics in dam-installed sections were investigated in Shiretoko streams and compared with unaltered sections in which no dams have been installed as a reference. Fish density in dam-installed sections was significantly lower than reference sections. Canopy cover, maximum temperature, wetted width, frequencies of pebble, bedrock and boulder, and areas of rapid, glide and riffle was significantly differed between dam-installed and reference sections. High stream temperature, occurrence of glides, and loss of boulder in dam-installed sections may reduce the suitability of habitats for fish and minimize the population size. We suggest that negative effects of dam installations on Dolly Varden populations have occurred in Shiretoko streams.