A comparative analysis of the Far Eastern grayling species Thymallus tugarinae and Thymallus grubii flavomaculatus based on the data from mitochondrial DNA cytochrome b gene sequencing

Nucleotide sequences of a fragment of mitochondrial DNA cytochrome b gene were obtained from two species of graylings that inhabit the Russian Far East, viz., Thymallus tugarinae and T. grubii flavomaculatus. A phylogenetic analysis of four Thymallus species, whose relationships have been poorly studied, was performed on these new data as well as on several sequences from the NCBI GenBank sequence database. The rate of genetic divergence between the Far Eastern graylings Thymallus tugarinae and T. g. flavomaculatus corresponded to that between European (T. thymallus) and Arctic (T. arcticus) graylings. Moreover, single nucleotide substitutions that result in alterations of the amino-acid structure of protein products (non-synonymic mutations) were revealed between sequences of the cytochrome b gene in the Far Eastern grayling species. The topology of phylogenetic trees, which was composed by means of Bayesian analysis and the maximum parsimony method, showed four independent phylogenetic lineages of graylings. In addition, the phylogenetic relationships between T. tugarinae and T. g. flavomaculatus were supported statistically. The obtained data indicated that Far Eastern graylings T. tugarinae and T. g. flavomaculatus are distinct species and confirmed that they belong to different phylogenetic lineages, as found earlier.     

Biological features of Low Amur grayling Thymallus tugarinae (Salmoniformes: Thymallidae): 2. Growth

We present the results of analysis of the group growth of the Low Amur grayling Thymallus tugarinae from 18 rivers of the Low Amur basin, tributaries of the Bureya and Ussuri rivers. We made use of the published data on the fishes of Sakhalin Island. The dwarf and fast-growing forms are not revealed among the investigated groups of this species. The parameters of growth of the Low Amur grayling in the investigated rivers are different, which is determined by the different conditions of feeding migration connected with the geographical location of the streams, their hydrological features, density of populations, etc. The anthropogenic factor influence is essential. Growth rate of T. tugarinae is lower compared with the other Amur species of graylings.     

New species of grayling <Emphasis Type="Italic">Thymallus tugarinae</Emphasis> sp. nova (Thymallidae) from the Amur River Basin

Data on a new species, low Amur grayling Thymallus tugarinae sp. nova, inhabiting tributaries of the lower and middle current of the Amur River, are presented. This species has been earlier equated with the Amur grayling T. grubii, described by Dybowski (1869) from the rivers Onon and Ingoda (the Upper Amur Basin). The new species differs from other representatives of the genus in the body coloration, the pattern on the upper fin, and certain morphological characters. On most of the Amur Basin, the grayling from the Lower Amur is sympatric with T. grubii and in its tributary Bureya River, also with the Bureya grayling T. burejensis. Individuals with intermediate traits are unknown, pointing to reproductive isolation. These data are supported by the results of molecular-genetic analysis (Froufe et al., 2003, 2005; Knizhin et al., 2004).     

Biological specific features of the Lower Amur grayling Thymallus tugarinae (Salmoniformes: Thymallidae): 1. Age composition

We consider age composition of samples of the Lower Amur grayling Thymallus tugarinae inhabiting water bodies of the lower and middle course of the Amur and several rivers of the northwestern part of Sakhalin Island and the southern coast of the Sea of Okhotsk. Age composition of the Lower Amur grayling from the studied samples is not similar because of several reasons. In rivers of the Amur basin, the species has a maximal age of 6+.     

Identification of reproductively isolated lineages of Amur grayling (Thymallus grubii Dybowski 1869): concordance between phenotypic and genetic variation

We analysed variation at maternally (mitochondrial DNA control region sequences) and bi-parentally (10 microsatellites) inherited genetic markers, as well as across 12 meristic characters in 7 populations of Amur grayling, Thymallus grubii, from eastern Siberia. All three data sets were concordant in supporting the existence of three diagnosable, reciprocally monophyletic, and most probably reproductively isolated, lineages of grayling within the Amur drainage. There was a significant correlation between genetic and phenotypic divergence, both within and among lineages. Two phenotypically distinct forms (with and without an orange spot on the posterior portion of the dorsal fin), found in sympatry in the lower Amur, most likely result from secondary contact, as they demonstrate 4.6% sequence divergence at the mitochondrial DNA control region. This divergence, together with the existence of at least one nearby population of orange spot grayling outside the Amur drainage (0.8% divergence) underscore the palaeo-hydrological complexity of the system, which presumably promoted genetic divergence in a shifting allopatric framework throughout the Pleistocene. Grayling from the upper Amur, corresponding to the type locality for the species, formed a sister group (1.4-1.6% divergent) to the orange spot lineage perhaps diverging in the early Pleistocene (1.4-1.6 Ma).     

Biological and morphological characteristic of the Arctic grayling Thymallus arcticus (Thymallidae) from Alpine Lakes of the basin of the upper reaches of the Angara River

Specific morphological and biological features of the populations of the Arctic grayling Thymallus arcticus from lakes Urunge Nur, Mongosha, Sadaiskoe, and Gusinoe situated in the Upper Sayan Mountains in the sources of the Oka and Kitoi rivers (the basin of the upper reaches of Angara) that have been not studied previously are considered. The results of the multivariate analysis indicated the uniformity of all samples by 12 meristic characters. Graylings of the studied lakes are similar to graylings from the Angara and Nizhnyaya Tunguska rivers and the Irkutsk Reservoir, as well as to the Baikal grayling T. a. baicalensis in body coloration and pattern of the dorsal fin. The samples studied and T. a. baicalensis considerably differ in the above characters from populations inhabiting the basin of the upper reaches of the Ob that belong to the nominative subspecies T. a. arcticus. The differences established between the Angara-Yenisei and the Upper Ob graylings give grounds to doubt the justification of including them into one taxon, T. a. arcticus. The same is evidenced by the results of molecular-genetic studies of graylings from the Palearctic (Froufe et al., 2005). The data obtained indicate the necessity of ascertaining the boundaries of the ranges of different forms of graylings in the Angara-Yenisei and Ob basins and of the revision of their taxonomic status. Populations inhabiting the Angara and Yenisei rivers, except their sources and the section of the lower reaches, should be assigned to the Baikal subspecies, T. a. baicalensis. Graylings from different lakes in the upper reaches of Oka and Kitoi differ in their linear-weight indices. The indices of growth are highest where they occupy a dominant position. Their food in the summer consists mainly of the larval and imaginal stages of amphibiotic and aerial-terrestrial insects. Only in Lake Urunge Nur do graylings of elder age groups use fish of other species for food. The extreme conditions of habitation of the grayling in Lake Gol’tsovoe are the cause of food deficiency, which determines its dwarfism.     

A new species of grayling <Emphasis Type="Italic">Thymallus svetovidovi</Emphasis> sp. nova (Thymallidae) from the Yenisei basin and its position in the genus <Emphasis Type="Italic">Thymallus</Emphasis>

The Upper Yenisei grayling Thymallus svetovidovi sp. nova that inhabits the headwaters of the Yenisei River in Mongolia is described. From the other representatives of the genus Thymallus, the species differs in the elements of body coloration, dorsal fin pattern, some morphometric characters, and genetic characteristics. Besides this species, most of the Yenisei basin is inhabited by Baikal grayling T. baicalensis Dyb., and its low reaches, by the Arctic grayling T. arcticus (Pall.).     

Finding a new form of the grayling Thymallus arcticus (thymallidae) in the basin of lake baikal

A dwarf form of the Arctic grayling Thymallus arcticus, inhabiting a group of small lakes at the origin of the Yakchii stream, which flows into the Verkhnyaya Angara River (basin of Lake Baikal), has been found in the northeastern watershed of the Baikal and Lena basins. The form is similar to upper Lena populations in body coloration and the pattern of dorsal fin. Our comparison by meristic characters indicated that the grayling of the Yakchii Lakes is more similar to the fish from Kutima River (basin of the upper courses of the Lena River) than to the black Baikal grayling T. arcticus baicalensis. The presence in Lake Baikal of a population similar to upper Lena graylings may have three causes: (1) possible drainage from Baikal to the pra-Lena via ancient valleys of the Barguzin and the Upper Angara; (2) appearance of transit zones as a result of glacial or tectonic events during the periods of watershed development between the Baikal and Lena basins; (3) the upper Lena grayling could be an endemic of the Baikal basin which was replaced by graylings penetrating from the Yenisei basin and remained in the form of relic populations in the upper courses of certain tributaries of the northern and northeastern parts of Lake Baikal.     

Occupancy of young-of-year Arctic grayling (Thymallus arcticus) in Barrenland streams

Hydrobiologia - Arctic grayling (Thymallus arcticus) is an iconic fish species that is present across the remote subarctic Barrenlands, yet our lack of understanding of their distributional...     

Demography of spawning grayling (Thymallus arcticus) in the Beaverlodge River,Alberta

Changes in a grayling spawning population are described that occurred in relation to both natural and man-induced events. The spawning population was monitored from 1982 to 1987 with a trap on a fishway on the Beaverlodge River, Alberta. Most of the spawning run in each year was tagged with recaptures noted in subsequent years. Male grayling grew slightly faster than females. Growth rates were variable and appeared to be quite low in years of low water flow. Grayling first spawned at 2 to 6 years of age. The majority of tagged grayling would have returned to spawn a second time after a delay of one to two years were it not for the high adult mortality. It is hypothesized that the variable time to first maturity and the delay in the second spawning is a response to highly variable juvenile recruitment. Annual mortality rates increased with age, suggesting an increased vulnerability of larger fish to angling. The population appears highly sensitive to over-exploitation, because of the high mortality rates and the delayed time to spawning of the adults.Deceased.Deceased.     

On the diversity and taxonomic status of graylings (Thymallus Thymallidae) from the Lena River

Two forms of graylings were distinguished in the Lena River, which differ in body coloration, the shape and pattern of the dorsal fin, and biological characteristics. With respect to these traits, the forms are divided between the upper and lower Lena. The first inhabits most of the basin, from the upper courses to the lower reaches, whereas the second inhabits the Lena delta and the adjacent arctic rivers and their tributaries. Comparative analysis of meristic traits revealed their low appropriateness for diagnosing Lena lineages as well as grayling populations inhabiting other northern rivers of Siberia. The body and fin coloration in the upper Lena form significantly differ from that found in forms and subspecies of graylings from the basins of the other rivers in the ranges of Eastern Siberian, Kamchatka, and Alaska. The lower Lena form should be classified with the Eastern Siberian subspecies T. arcticus pallasii and is closer to the populations of graylings from the rivers Anabar and Yana, and the rivers of the Chukot Peninsula and Alaska, and the upper courses of the Missouri. These results are supported by analysis of the mitochondrial DNA of graylings throughout the Paleoarctic. Further studies of these lineages and phylogenetic relationships between Arctic lineages are necessary for the final determination of the taxonomic status of the upper Lena form.     

Component communities of parasites in the grayling Thymallus thymallus (L.) (Salmoniformes, Thymallidae) and the minnow Phoxinus phoxinus (L.) (Cypriniformes, Cyprinidae) from the Pechora River

Component communities of parasites of Thymallus thymallus and Phoxinus phoxinus are described. The communities differ in the number of species and in the number of dominate species, which is one in P. phoxinus and usually two in T. thymallus. In the first case the dominate species is allogenic, while in the second case dominate species are autogenic. Values of the species diversity indexes are also different for these communities. The differences between parasite communities of T. thymallus and P. phoxinus underline positions of these hosts in hydrobiocenose. Species specialists take first place in the parasite communities both of the hosts, that is characteristic for the parasite communities of the fishes from the boreal submountain faunistic complex. Parasite communities in both of the hosts consist of three groups of species discriminated by the ratio of their biomasses, that suggests forming of the groups by a coordination of the species biomasses.     

Genetic variation of European grayling (Thymallus thymallus) populations in the Western Balkans

In order to elucidate genetic composition of European grayling (Thymallus thymallus) populations in the Western Balkans, the partial mitochondrial DNA (mtDNA) control region was sequenced and 12 microsatellite loci genotyped in 14 populations originating from tributaries of the Adriatic and Danube drainages. Eleven mtDNA haplotypes were found, one confined to the Adriatic clade, one to the Alpine group and the rest to the ‘Balkan’ grayling phylogenetic clade. Haplotypes from the Balkan clade were confined to the Danube drainage and constituted two groups: northern group with haplotypes found in the Slovenian part of the Danube drainage, and southern group, consisting from Bosnia–Herzegovina and Montenegro. Substantial genetic distance between northern and southern groups of haplotypes (0.75–1.8%) and well supported divisions within the northern group indicate very structured grayling population within the studied Danube basin that most probably did not evolve due to vicariance but rather as a consequence of multiple colonization waves that might have occurred during the Pleistocene. Furthermore, genetic distance of ~4% between Adriatic and Danube populations’ haplotypes, suggest that their separation occurred in mid-Pliocene. These findings imply a complex colonization pattern of the Western Balkans drainages. Microsatellite data also confirm high genetic diversity in Western Balkans populations of grayling (on average 7.5 alleles per microsatellite locus and H _exp 0.58). Limited stocking activities were detected based on microsatellites and mtDNA data. Regarding current knowledge of grayling phylogeography appropriate management strategies were proposed to preserve unique, autochthonous grayling populations in Western Balkan.     

Comparative analysis of oocyte growth in two forms of Baikal grayling <Emphasis Type="Italic">Thymallus baicalensis</Emphasis> (Thymallidae)

Size characteristics of oocytes of the elder generation of the end of previtollogenesis, as well as of the beginning and middle of the phase of cytoplasm vacuolization in two forms of Baikal grayling Thymallus baicalensis were studied. In each form, the dependence of oocyte parameter on the age and size of females is traced. During the studied phases of development—at the termination of the period of previtellogenesis and at the proper beginning of the period of vitellogenesis—oocytes of the white and black graylings have similar sizes; cytoplasm vacuolization in white grayling proceeds less actively. It was established that differences in the diameter of mature ovicells in the black and white Baikal graylings result from dissimilar rate of accumulation of trophic substances in the oocytes of the given forms of this species and are determined by differences in the rate of growth of oocytes during the formation in them of yolk inclusions.     

Geographical variation of allele frequencies at the LDH-A* locus in the arctic grayling Thymallus arcticus Pall. and in the European grayling Thymallus arcticus L

The allele frequencies of LDH-A* locus were studied in the population of Siberian grayling from the Kozhym River (Pechora basin) and in the population of European grayling from Pechora, Mezen', and Vym' rivers (Northern Dvina basin). In samples of both species (n = 134), three LDH-A phenotypes have been identified in total, which proved to be under the control of two alleles: LDH-A*100 and LDH-A*50. The alternative alleles of LDH-A* locus were identified in the populations of Siberian grayling from Kozhym River and in the population of European grayling from the same river and other Pechora tributaries, namely, LDH-A*100 and LDH-A*50 in the Siberian and the European grayling, respectively. However, in the European grayling populations from the Mezen' and Vym' rivers, both alleles occur at the frequencies of the rare LDH-A*100 allele of 0.143 and 0.222, respectively. According to the published data, the frequency of LDH-A*100 allele increases in the European grayling populations of northwestern (Finland) and southern (France) rivers, reaching 0.872 and 1.000 in Rhone and Loire, respectively, i.e., the values characteristic of the Siberian grayling populations.     

Zoogeographical implications of variation in mitochondrial DNA of Arctic grayling (Thymallus arcticus)

Most of the northern half of North America's freshwater fish survived the last glacial period in unfrozen refugia peripheral to the ice sheets. In our study, the question of which refugia Arctic grayling ( Thymallus arcticus ) inhabited during the Wisconsinan Ice Age, and how they subsequently dispersed to their present geographical range, was examined using mitochondrial DNA (mtDNA) analysis. mtDNA from 12 T. arcticus populations was analysed by direct sequencing and restriction fragment length polymorphism analysis (RFLP). Our data support the hypotheses that T. arcticus had a large refugial population in the Bering Refuge (shown by high mtDNA diversity in extant Alaskan populations) and that British Columbia was colonized from the Bering Refuge (shown by mtDNA haplotype similarities). Our data also show that a disjunct southern set of populations in Montana is significantly different from the northern grayling, in terms of restriction haplotype frequency and distinguishing sequence characteristics. Sequence results yielded an estimated divergence time of 370 000 years between the northern and Montana grayling haplotypes. We conclude that T. arcticus survived the Wisconsinan glaciation in at least two refugia: (i) the Bering Refuge north of the ice sheets; and (ii) either the Upper Missouri or the southwest Alberta Refuge, south of the ice sheet.     

Distribution and Biological Features of Grayling Thymallus thymallus (Thymallidae) in the European North-East of Russia

Journal of Ichthyology - The distribution and some ecological features of the European grayling Thymallus thymallus are studied in the extreme European Northeast of Russia. In many water systems of...     

Familiarity with a partner facilitates the movement of drift foraging juvenile grayling (Thymallus thymallus) into a new habitat area

Preferring one social partner over another can enhance fitness. This paper reports that juvenile grayling were significantly more likely to enter and forage in new, upstream habitats when paired with familiar versus unfamiliar social partners. Fish paired with unfamiliar partners or when alone were more reluctant to enter the new area. The entry times for both fish in a familiar pair were significantly correlated, but uncorrelated for unfamiliar fish. These differences between familiars and unfamiliars were consistent over a 2-week period. Fish with familiar partners spent more time within three body lengths of each other than did those with unfamiliars. The results are discussed in relation to optimality models of drift foraging, which do not included sociality. It is suggested that the social dimension creates a more dynamic foraging response to variable environmental conditions and could have consequences for growth.     

The susceptibility of grayling (Thymallus thymallus) to experimental infections with the monogenean Gyrodactylus salaris

The pathogenic monogenean Gyrodactylus salaris infecting Atlantic salmon (Salmo salar) is found to attach and reproduce under laboratory conditions on several species in the subfamily Salmoninae other than the Atlantic salmon. The gyrodactylid species Gyrodactylus thymalli infecting grayling (Thymallus thymallus) in another subfamily, Thymallinae, is previously said to be very similar to G. salaris based on morphometry and genetical analysis which prompted the present laboratory experiments to test the susceptibility and resistance of grayling to G. salaris. All 0+ and 1+ grayling became infected with G. salaris during the experimental infection procedure. However, both innate resistant and susceptible grayling were found. In susceptible individually isolated fish, parasite reproduction lasted for more than 35 days. Parasite reproduction also occurred among grouped grayling as judged from the duration of infection of more than 50 days. However, grayling susceptibility as judged from G. salaris reproduction, was very limited. Hence, the results indicate significant biological differences between the function of Atlantic salmon and grayling as host for G. salaris. The grayling is interpreted as unable to sustain G. salaris in nature which implies that G. thymalli is not conspecific with G. salaris. However, G. salaris dispersal by grayling cannot be excluded.