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).     

A new subspecies of the amur grayling Thymallus grubii flavomaculatus ssp. nova (Thymallidae)

A new subspecies of the Amur grayling is described, the yellow-spotted grayling Thymallus grubii flavomaculatus ssp. n. populating the upper reaches of large tributaries of the Lower Amur basin and some rivers flowing to the Tatar Strait, the Sea of Okhotsk, and the Sea of Japan. From nominative subspecies T. g. grubii populating the basin in the upper reaches of the Amur and from other representatives of the genus Thymallus, this subspecies differs in the dorsal fin pattern whose principal trait is a yellow spot on the last two-five interray membranes, in the body coloration, and in some morphological characters. The yellow-spotted grayling in the upper reaches of tributaries sympatrically coexists with the Lower Amur grayling Thymallus sp. No specimens with intermediate characters are found. The sympatric forms spawn at different times, thus being isolated reproductively. Isolation of the yellow-spotted grayling as a subspecies of the Amur grayling T. grubii is confirmed by the results of molecular-genetic investigations.     

New data about distribution of lower Amur grayling <Emphasis Type="Italic">Thymallus tugarinae</Emphasis> and Baikal-Lena grayling <Emphasis Type="Italic">Thymallus</Emphasis> sp. (Thymallidae) in the basins of the upper Amur River and Zeya River

New findings of lower Amur grayling (Lower Amur grayling) Thymallus tugarinae and Baikal-Lena grayling Th. sp. (Th. baicalolenensis) in the basins of the upper Amur River and Zeya River are reported for 2010?2014. Lower Amur grayling has been registered in the tributaries of Argun River and in the middle stream of Ingoda River. Baikal-Lena grayling has been found for the first time in the headstream of Ingoda River. New data about the distribution of this species in the basin of upper stream of Zeya River are reported; particularly, Baikal-Lena grayling has been found in Okonon Lake; this is the first finding of a representative of Thymallidae family in the alpine lakes of the watershed area of Amur River. In the basins of upper Amur River and Zeya River, the summer ranges of upper Amur grayling Th. grubii grubii and lower Amur grayling overlap to some extent; on the other hand, the ranges of Baikal-Lena grayling and lower Amur grayling are disjoint. During the summer period, upper Amur grayling and Baikal-Lena grayling together inhabit the basin of upper stream of Zeya River, but only Baikal-Lena grayling has been found in the headstream of Zeya River. In the basin of upper Amur River (the headstream of Ingoda River), the joint range of these species is still unknown; the sympatric zone is located in Dzhermoltai River (the basin of Onon River).     

A comparative analysis of two Amur grayling species from the <Emphasis Type="Italic">Thymallus</Emphasis> Genus (Salmoniformes; Thymallidae) according to a number of osteological and morphological features in their zone of sympatry

The results of the comparison of the Lower Amur grayling Thymallus tugarinae and the yellowspotted grayling T. flavomaculatus that were collected in the zone of their sympatry in the middle reaches of the Anuy River (the Lower Amur basin) for a number of osteological and morphological features are provided. The work was based on the analysis of X-ray images and a number of plastic features. Considerable differences were detected between the two species in the numbers of the bone elements of the axial skeleton. The differences in the number and topography of certain bone elements correlate with the morphological features of these species. The comparison results for certain features do not correspond with the literature data.     

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.     

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+.     

Graylings of Baikal Lake basin (Thymallus, Thymallidae): Diversity of forms and their taxonomic status

The morphological characters, molecular-genetic features, and patterns of the dorsal fin in different subspecies and forms of Arctic grayling Thymallus arcticus populating Lake Baikal and its tributaries, Irkutsk Reservoir, and Lake Khubsugul have been studied. Three groups are discernable. The first includes the white and black Baikal graylings T. a. baicalensis, as well as the western Siberian T. a. arcticus; the second group is represented by the Kosogol grayling T. a. nigrescens, and the third is composed of T. arcticus ssp. from the Yakchii lakes (the Verkhnyaya Angara basin) with a phenotype close to graylings populating the upper reaches of the Lena River. All of them are distinguished by some morphological characters, elements of the dorsal fin pattern, and by body coloration. The populations of black Baikal graylings are genetically uniform, and their distinctions from white Baikal graylings are insignificant, which agrees with the absence of a considerable divergence of these forms by a complex of meristic characters. It is assumed that in the black and white Baikal graylings the exchange by genetic information has either ceased quite recently, or persists, although, insignificantly. Some genetic remoteness of the west Siberian grayling from Irkutsk Reservoir and Nizhnyaya Tunguska, closely related to the Baikal grayling, is recorded. The formation of the Khubsugul subspecies is possibly a result of the contact of grayling populations during the rearrangements of the river system in the last glaciation period in the upper reaches of Yenisei and Selenga in Mongolia. The habitation in the Baikal system of the Upper Lena graylings indicates a connection between the Lena and Baikal basins in the past. The results of a multivariate analysis of meristic characters and the sequences of mitochondrial DNA confirm the conclusion made by Svetovidov (1931, 1936), concerning the absence of grounds to assign a species status to the Baikal forms.     

Multiple species of grayling (Thymallus sp.) found in sympatry in a remote tributary of the Amur River

Large-scale phenotypic and genetic studies of the salmonid genus Thymallus (grayling) in the Palaearctic suggest that most major phylogeographic lineages represent good biological species. Evaluating such a premise in areas involved in palaeo-hydrological dynamics where multiple species are found in sympatry should serve to assess the level of reproductive isolation, the traditional sine qua non of species recognition. Molecular sequence (mtDNA) and microsatellite (nDNA, seven loci) analysis of grayling in the upper Bureya River watershed support the occurrence of three distinctive species of grayling living in sympatry in this large oligotrophic tributary of the Amur River. One of these lineages is primarily found throughout the Lena River basin and is recognized as Baikal-Lena grayling Thymallus baicalolenensis; the second, the upper Amur grayling Thymallus grubii is found over large areas of the Amur catchment including the entire headwater region; and the third, the Bureya grayling Thymallus burejensis is endemic to the study area. A limited number of hybrids were identified, primarily between Baikal-Lena grayling T. baicalolenensis and Bureya grayling T. burejensis with little to no signs of introgression among non-hybrid individuals. Morphological distinctiveness among populations of these species living in sympatry was greater than between populations living in allopatry, suggesting character displacement. Divergence estimates among taxa range up to 6.2 MY, and allopatric origins for all three species’ is suggested. To our knowledge, this is the first data-based confirmation of three species of grayling living in sympatry.     

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.     

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.     

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.     

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.     

Structure of the seismosensory system of <Emphasis Type="Italic">Psettodes erumei</Emphasis> (Psettodidae,Pleuronectiformes)

The structure of the seismosensory system of Psettodes erumei was studied. General and specific features in comparison with the previously studied representatives of the suborder Pleuronectoidei were revealed. Similarity is manifested in the presence of four main head canals, coronal commissure, interrupted supratemporal commissure transformed into a supratemporal canal, and one trunk canal developed on both sides of the body. The seismosensory system of Psettodes erumei is characterized by several plesiomorphous characters: the presence of a sensory segment in lacrimale of the eyed side and supracleithrum, the absence of disruptions in all sensory canals, and a strong branching of cutaneous canaliculi. Specific features of P. erumei involve the presence of a large number of fontanels on angulo-articulare, connection on the eyed side of CIO sites in lacrimale and infraorbitale, and the underdevelopment of the bony vault of the sensory canal in praeoperculum.     

General patterns of sexual dimorphism in graylings (Thymallus), with a comparison to other salmonid species

Among fishes, salmonids (family Salmonidae) have attracted a great deal of research attention focused on sexual dimorphism and associated selective forces. Most of this research has been directed toward anadromous and mostly semelparous salmon and trout (Oncorhynchus, Salmo), and comparatively little is known about intersexual variability in strictly iteroparous freshwater salmonids. We examined a comprehensive data set of 28 linear morphometric characters in 11 of 15 currently recognised species of grayling (Thymallinae, Thymallus), a genus consisting of iteroparous species only, to identify general patterns of intersexual morphological variability. Overall, we found that all grayling species show common sex-specific traits particularly relating to size dimensions of the dorsal, anal, pelvic and pectoral fins. Although the magnitude of sexual dimorphism differed among species, there was no significant phylogenetic signal associated with these differences across the genus. These results are discussed in terms of the assumed selection pressures driving sexual dimorphism in graylings and are compared to existing knowledge in Salmonidae as a whole where similarities and differences with both Salmoninae and Coregoninae exist. The present study provides the first detailed genus-wide comparison of sexually dimorphic phenotypic characters in graylings, and highlights the need for more large-scale comparative studies in multiple salmonid species to better understand general macroevolutionary trends among this important group of freshwater fishes.

     

New species of the genus <Emphasis Type="Italic">Bathycongrus—B. parviporus</Emphasis> (Congridae,Anguilliformes)—from waters of Central Vietnam (Nha Trang and Van Phong Bays)

Bathycongrus parviporus sp. nova is described from specimens collected in the South China Sea, in the coastal waters of central Vietnam. By its characters (short snout, elongate-oval vomerine tooth patch with numerous small teeth of approximately the same size, slender relatively short tail, and residual leptocephalic pigmentation as a series of small melanophores just below lateral line in adults), the species belongs to the group of species of this genus represented by B. bleekeri, B. trimaculatus, and B. unimaculatus in the western part of the Pacific Ocean and B. dubius in the western Atlantic Ocean but differs in small infraorbital pores which are enlarged in all known species of the genus Bathycongrus. By the number of preanal pores, the new species is similar to B. bleekeri from Philippine waters but differs from it in a higher number of vertebrae (120–122), in more numerous rows of teeth on the premaxilloethmoid, in more number of branchiostegal rays (9–10), a shorter head, and in some other proportions of head and body. Topography of canals and pores of the cephalic seismosensory system of B. parviporus sp. nova is described. Morphometric and osteological characters for placing this species in the genus Bathycongrus are noted.     

On the diversity of fishes in mountain lakes of the Amur basin

Results of the study of 11 lakes situated in the mountainous part of the Amur River basin are reported. Six fish species, namely, blunt-snouted lenok Brachymystax tumensis, Baikal-Lena grayling Thymallus baicalolenensis, Siberian stone loach Barbatula toni, Lagowski’s minnow Rhynchocypris lagowskii, Czekanowski’s minnow Rh. czekanowskii, and Amur sculpin Cottus szanaga, were found in five lakes. Four species were found in one lake (Lake Pereval’noe, basin of the Amgun’ River), and two species were found in each of the remaining lakes. Lenok and Amur sculpin were the most common species. The fauna of Lake Okonon (basin of the river Zeya) was the most specific and did not have analogs among the lake faunas; Czekanowski’s minnow and Baikal black grayling were found in this lake. The lenok ecotype that formed in the mountain lakes differed from the river lenok in some regards. The potential pathways of formation and protection of mountain lake ichthyocenoses of the Amur basin are discussed.     

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.     

New data on the distribution of the Upper Lena form of grayling (Thymallidae) in the basin of Lake Baikal and its taxonomic status

The habitation of the Upper Lena form of grayling in the upper reaches of the Tiya (the northwestern tributary of Lake Baikal) and Barguzin rivers, where Baikal grayling Thymallus baicalensis Dyb. also occurs, was established. The listed forms can be diagnosed from the body coloration, dorsal fin pattern, combination of meristic characters, as well as by molecular-genetic methods. The sympatric habitation of the Upper Lena and Baikal graylings in the northern tributaries of Baikal, considerable phenotypic and genetic differences, as well as the absence of individuals with intermediate characters, make it possible to consider them as independent species. The results indicate the need to revise the intraspecies structure of the Arctic grayling T. arcticus (Pall.) and the entire genus Thymallus. The localization of populations of the Upper Lena grayling in the upper reaches of Baikal tributaries testifies to its possible penetration of the Baikal basin by means of temporary river catchments that formed in the past and which are probable at present at watersheds with the Lena River. A wider distribution of this form is apparently hindered by the pressure of aboriginal species.     

Comparative osteology of <Emphasis Type="Italic">Bathylutichthys balushkini</Emphasis> and relationship of the family Bathylutichthyidae (Cottoidei)

The structure of skeleton of Bathylutichthys balushkini was studied and comparative analysis of it with representatives of other families of the suborder Cottoidei was performed. It was found that Bathylutichthyidae are characterized by the presence of a great number of original characters supporting its family status. They include: absence of mesethmoideum; bony canals of seismosensory system on frontale, pteroticum, and parietale and their strong reduction on infraorbitale; and absence of pterosphenoideum, intercalare, pharyngobranchiale 2, and basihyale. At the same time, Bathylutichthyidae retain a considerable number of generalized specific features: location of parietalia laterally to supraoccipitale and absence of their joint between themselves, presence of mesoptergoideum and basibranchialia 2 and 3. Cladistic analysis based on osteological characters and characters of the seismosensory system of 11 families of Cottoidei demonstrates that Bathylutichthyidae are combined into one group with families Rhamphocottidae, Agonidae, Psychrolutidae, Cyclopteridae, and Liparidae occupying an intermediate position between Psychrolutidae and the two last families. The cladogram obtained does not support the point of view of existence in the suborder Cottoidei of an independent superfamily Cyclopteroidea, including families Liparidae and Cyclopteridae.     

Distribution,abundance, and size structure of Amur kaluga <Emphasis Type="Italic">Acipenser dauricus</Emphasis> and Amur sturgeon <Emphasis Type="Italic">A. schrenckii</Emphasis> in the Lower Amur and Amur Estuary

Distribution, population structure, and abundance of Amur kaluga Acipenser dauricus and Amur sturgeon A. schrenckii in the Lower Amur and Amur Estuary have been considered based on the materials of two surveys performed in 2011. Both species occur along almost the entire length of the Lower Amur and in major part of the Amur Estuary. The density of their aggregations has significantly decreased in most regions of the river if compared to the 1960s. In the Amur Estuary, A. dauricus and A. schrenckii are most abundant in freshwater and light salted water bodies of its western part. In 2011, the abundance and biomass of A. dauricus and A. schrenckii in the Lower Amur and Amur Estuary was 345000 specimens and 7110 tons and 289000 specimens and 1946 tons, respectively.