One Beringian genus less: A re‐assesment of Pacifimyxas Kruglov & Starobogatov, 1985 (Mollusca: Gastropoda: Lymnaeidae) questions the current estimates of Beringian biodiversity

The taxonomic position of three nominal species of lymnaeid snails placed by Kruglov & Starobogatov (1993) into the subgenus Lymnaea (Pacifimyxas) Kruglov & Starobogatov, 1985, has been re‐assessed based on a molecular genetic study of topotypic specimens and an examination of the type series and other materials available. It has been shown that the two species, Lymnaea (Pacifimyxas) magadanensis Kruglov & Starobogatov, 1985 and Lymnaea (Pacifimyxas) streletzkajae Kruglov & Starobogatov, 1985, are identical with the species Kamtschaticana kamtschatica (Middendorff, 1850) and must be treated as its junior synonyms. Hence, Pacifimyxas becomes a junior synonym of Kamtschaticana Kruglov & Starobogatov, 1984. The taxonomic identity of the third species of Pacifimyxas, Lymnaea (Pacifimyxas) perpolita, remains obscure, and this species is considered here as taxon inquirendum. Two other nominal species, Lymnaea aberrans (Westerlund, 1897) and Lymnaea middendorffi (W. Dybowski, 1904), have been synonymized with K. kamtschatica based on morphological and geographical evidence. The lectotype of Limnaea peregra var. middendorffi is designated. The actual level of species richness in the Beringian freshwater malacofauna may be 20–25% lower than it was determined on the basis of the traditional system. Some implications of this outcome for the biogeography of the Beringian freshwater fauna are discussed.     

The species composition and distribution of free-living nematodes (Nematoda) in an area of natural oil and gas seeps in Lake Baikal

The species composition and quantitative parameters of free-living nematode fauna have been studied in an area of natural oil seeps off Cape Gorevoi Utes (Middle Baikal). A total of 34 species of 17 genera, 10 families, and 8 orders have been found, 18 of them endemic to Lake Baikal. The nematode fauna of bitumen volcanoes was found to be represented by three specialized species (Monhystera naphthera Gagarin et Naumova, 2010; Eumonhystera abyssalis Gagarin et Naumova, 2010; and Eutobrilus mirandus Gagarin et Naumova, 2011). Areas of the bottom with scattered oil seeps are dominated by eurybathic species endemic to Lake Baikal. The maximal values of nematode population density and maximum proportion of nematodes in the meiozoobenthos were recorded in areas directly adjacent to oil seeps.     

The endemic Cladophorales (Ulvophyceae) of ancient Lake Baikal represent a monophyletic group of very closely related but morphologically diverse species

Lake Baikal, the oldest lake in the world, is home to spectacular biodiversity and extraordinary levels of endemism. While many of the animal species flocks from Lake Baikal are famous examples of evolutionary radiations, the lake also includes a wide diversity of endemic algae that are not well investigated with regards to molecular‐biological taxonomy and phylogeny. The endemic taxa of the green algal order Cladophorales show a range of divergent morphologies that led to their classification in four genera in two families. We sequenced partial large‐ and small‐subunit rDNA as well as the internal transcribed spacer region of 14 of the 16 described endemic taxa to clarify their phylogenetic relationships. One endemic morphospecies, Cladophora kusnetzowii, was shown to be conspecific with the widespread Aegagropila linnaei. All other endemic morphospecies formed a monophyletic group nested within the genus Rhizoclonium (Cladophoraceae), a very surprising result, in stark contrast to their morphological affinities. The Baikal clade represents a species flock of closely related taxa with very low genetic differentiation. Some of the morphospecies were congruent with lineages recovered in the phylogenies, but due to the low phylogenetic signal in the rDNA sequences the relationships within the Baikal clade were not all well resolved. The Baikal clade appears to represent a recent radiation, based on the low molecular divergence within the group, and it is hypothesized that the large morphological variation results from diversification in sympatry from a common ancestor in Lake Baikal.     

Palaearctic gastropod gains a foothold in the dominion of endemics: range expansion and morphological change of Lymnaea (Radix) auricularia in Lake Baikal

In and around the endemic-dominated Lake Baikal, palaearctic species are generally restricted to shallow, sheltered bays and in- and out-flowing river floodplains. However, we observed populations of the palaearctic snail Lymnaea (Radix) auricularia on the steep, rocky littoral of Lake Baikal proper. We compared the morphology of 542 shells sampled from this new habitat with potential source populations from conventional habitats. A size-free Discriminant Analysis indicated a strong morphological differentiation of the newly established populations from their likely sources. The new populations had a more compact shell shape with a wide aperture, which may be advantageous in wave-exposed habitats where a firm attachment to the substrate is needed. Shells from the conventional habitats were more elongated, with a narrow aperture, which may be advantageous in habitats that have a dry period where retreating into the mud is required and water loss should be limited. These results may suggest that selection is acting on shell shape in Lake Baikal. The apparent recent arrival of this pandemic gastropod in a habitat previously dominated by endemics constitutes a potential ecological threat and an alert to possible ecological change.     

Ladislavella tumrokensis: The first molecular evidence of a Nearctic clade of lymnaeid snails inhabiting Eurasia

In this study, we provide the first molecular evidence for a possible connection between freshwater mollusc faunas across the Bering Strait via the Beringian Land Bridge using data inferred from gastropods of the family Lymnaeidae. The gastropods collected from geothermal springs in the Tumrok Mountains, West Kamchatka, Russia, share the nuclear internal transcribed spacer 2 (ITS2) and the cytochrome oxidase subunit I gene (COI) haplotypes, thus being as sister to those recorded for lymnaeid snails in the Stagnicola elodes group from Canada and the USA. Two lymnaeid species, Lymnaea (Orientogalba) tumrokensis Kruglov and Starobogatov, 1985 Kruglov, N. D., & Starobogatov, Y. I. (1985). Ob'yom podroda Galba i skhodnykh s nim podrodov roda Lymnaea [The volume of the subgenus Galba and of the other similar subgenera of the genus Lymnaea (Gastropoda, Pulmonata)]. Zoologicheskiy Zhurnal, 64, 24–35.[Web of Science ®] , [Google Scholar] and Lymnaea (Polyrhytis) kurenkovi Kruglov and Starobogatov, 1989 Kruglov, N. D., & Starobogatov, Y. I. (1989). Mollyuski podroda Polyrhytis roda Lymnaea fauny SSSR (Pulmonata, Lymnaeidae) [Molluscs of the subgenus Polyrhytis of the genus Lymnaea of the fauna of the USSR (Pulmonata, Lymnaeidae)]. Zoologicheskiy Zhurnal, 68, 14–20.[Web of Science ®] , [Google Scholar], were described from the Tumrok geothermal locality, but actually they are morphological variations of a single taxon of subspecies rank re-classified here as Ladislavella catascopium tumrokensis. This subspecies is the first discovered representative in the genus, which formed a dwarf race in a geothermal habitat. Our findings highlight the possible exchange between freshwater faunas in Beringia during the Pleistocene and an important role of geothermal ecosystems as possible cryptic refugia for freshwater hydrobionts.     

Concurrent evolution of ancient sister lakes and sister species: the freshwater gastropod genus <Emphasis Type="Italic">Radix</Emphasis> in lakes Ohrid and Prespa

Ancient sister lakes are considered to be ancient lakes lying in close geographic proximity, sharing a related origin and significant time of co-existence, usually having hydrological connection as well as a balanced degree of faunal overlap and distinctness. A paradigm for studying sister lake relationships are the ancient lakes Ohrid and Prespa in the Balkans, which are characterized by high degrees of endemicity. Three general patterns of endemic species can be distinguished for these lakes: (1) taxa that are endemic to either lake, with no close relatives in the respective sister lake, (2) closely related but distinct endemic taxa in both lakes (sister species) and (3) shared endemic taxa occurring in both lakes. In the present paper, two endemic freshwater pulmonate gastropod species, Radix relicta (Lake Ohrid) and R. pinteri (Lake Prespa), are used to study the evolution of presumed sister species based on biogeographical and comparative DNA data from world-wide Radix taxa. Phylogenetic, phylogeographical and parametric bootstrap analyses all suggest a sister group relationship of R. relicta and R. pinteri (pattern 2 of endemic diversity). Sister to these two taxa is the widespread R. ampla, which does not occur in the vicinity of lakes Ohrid and Prespa. The southern feeder spring complexes of Lake Ohrid are inhabited by another lineage (Radix sp. 1), which resembles Radix relicta in morphology/anatomy. For Lake Prespa, the widespread R. auricularia was reported in addition to the endemic R. pinteri. Comparative phylogenetic data favour a western Adriatic zoogeographical affinity of lakes Ohrid and Prespa over an Aegean-Anatolian faunal connection. The status of lakes Ohrid and Prespa as sister lakes is evaluated in the light of current knowledge on gastropod speciation and endemism in these hotspots of biodiversity.     

Echiuropus bekmanae n.sp. (Crustacea, Amphipoda, Carinogammaridae) from Lake Baikal, retrieved by a new deep-water sampling device

Lake Baikal is well known for its exceptional diversity of gammarid amphipods, with 95% of endemics from its 270 described species representing about 20% of the world freshwater fauna. Echiuropus (Asprogammarus) bekmanae n.sp. was collected by an `autonomous trap system', a sampling device thoroughly described here. Although not necrophagous itself, this new species was found in two baited trap samples (at 750 and more than 1600 m) from 1995 and 1996. This rather small (13 mm) deep water species is the eighth species of the subgenus Asprogammarus, of the Baikal endemic genus Echiuropus. Diagnoses of the subgenus and the closer species are given.     

The consortium of the sponge Lubomirskia baicalensis in Lake Baikal,East Siberia

The spatial distribution of the fauna associated with a branched sponge, Lubomirskia baicalensis, endemic of Lake Baikal has been quantitatively studied. The biomass and numbers of three amphipod species which inhabit the sponge correlate (linearly or non-linearly) with the weight of the sponge.     

First report on bacteria of the family <Emphasis Type="Italic">Spirochaetaceae</Emphasis> from digestive tract of endemic gastropods from Lake Baikal

Detection of bacteria of the family Spirochaetaceae in the crystalline style of 11 species of endemic gastropods from Lake Baikal is reported. Investigation by transmission and scanning electron microscopy showed that these spirochetes belonged to the genus Cristispira.     

Heat shock proteins in the mechanisms of stress adaptation in Baikal amphipods and palearctic <Emphasis Type="Italic">Gammarus lacustris</Emphasis> Sars. I. HSP70 family

The participation of HSP70 family proteins in the mechanisms of stress adaptation under thermal and toxic stress in Baikal endemic and palearctic organisms was investigated. The four Lake Baikal endemic species Gmelinoides fasciatus (Stebb.), Eulimnogammarus cyaneus (Dyb.), E. vittatus (Dyb.), Ommatogammarus fl avus (Dyb.) were compared with the palearctic fauna representative Gammarus lacustris Sars. The type of synthesis of HSP70 family heat shock proteins under thermal (exposure at a temperature of 20, 25, and 30°C) and toxic (exposure in CdCl₂ solutions with the concentrations 50, 10, 5, 0.5, and 0.05 mg/l) stress was determined. All species under investigation were shown to have a common trend towards HSP70 content increase; therewith some species-specific characteristics of the type of synthesis of the protein under investigation were observed. The conclusion has been made on HSP70 participation in the mechanisms of thermal and toxic resistance in the studied amphipod species.     

Microsporidian Parasites Found in the Hemolymph of Four Baikalian Endemic Amphipods

At present, approximately 187 genera and over 1300 species of Microsporidia have been described, among which almost half infect aquatic species and approximately 50 genera potentially infect aquatic arthropods. Lake Baikal is the deepest and one of the oldest lakes in the world, and it has a rich endemic fauna with a predominance of arthropods. Among the arthropods living in this lake, amphipods (Crustacea) are the most dominant group and are represented by more than 350 endemic species. Baikalian amphipods inhabit almost all depths and all types of substrates. The age and geographical isolation of this group creates excellent opportunities for studying the diversity, evolution and genetics of host-parasite relationships. However, despite more than 150 years of study, data investigating the microsporidia of Lake Baikal remain incomplete. In this study, we used molecular genetic analyses to detect microsporidia in the hemolymph of several endemic species of amphipods from Lake Baikal. We provide the first evidence that microsporidian species belonging to three genera (Microsporidium, Dictyocoela and Nosema) are present in the hemolymph of Baikalian endemic amphipods. In the hemolymph of Eulimnogammarus verrucosus, we detected SSU rDNA of microsporidia belonging to the genus Nozema. In the hemolymph of Pallasea cancellous, we found the DNA of Microsporidium sp. similar to that in other Baikalian endemic amphipods; Dictyocoela sp. was found in the hemolymph of Eulimnogammarus marituji and Acanthogammarus lappaceus longispinus.     

A transcriptome-wide assessment of differentially expressed genes among two highly divergent,yet sympatric,lineages of the freshwater Atyid shrimp, <Emphasis Type="Italic">Paratya australiensis</Emphasis>

The current study evaluated the possible toxic effects of the water-soluble fraction of crude oil on the general cellular stress-response mechanisms of two dominant representatives of Lake Baikal’s littoral community, the endemic amphipod species Eulimnogammarus verrucosus and E. cyaneus. The acute toxicity effects on the cellular stress-response mechanisms of amphipods were studied in the laboratory by exposing amphipods in water from Lake Baikal to addition of a water-soluble fraction of crude oil at concentrations considered safe for the aquatic environment. The present study found that even short-term exposure to a water-soluble fraction of crude oil at concentration of 50 µg/L, established as the threshold limit for fishery and aquaculture water reservoirs in the Russian Federation, directly affected the general stress-response markers HSP70 and lipid peroxidation and significantly changed the activity of antioxidant enzymes in both studied species. This result confirms the high sensitivity of Baikal endemics to crude oil. Thus, it also indicates that established standards and threshold limit values of oil concentrations estimated for ecological monitoring of general water reservoirs cannot be applied directly to the unique Lake Baikal ecosystem.     

Speciation and chromosomal evolution of the Baikalian endemic chironomids of the genus Sergentia Kief. (Diptera, Chironomidae): Karyotype divergence and chromosomal polymorphism in the populations of deep-water species Sergentia nebulosa Linevitsh et al. and Sergentia assimilis Proviz V. et Proviz L.

Specific karyotype structure and chromosomal polymorphism was investigated in the populations of Sergentia nebulosa Linevitsh et al., 1984 and Sergentia assimilis Proviz V. et Proviz L., 1999, the deep-water endemic chironomid species (Diptera, Chironomidae) from the Baikal Lake. The distinguishing feature of the karyotypes of these species, compared to the other Baikalian Sergentia, is well-developed nucleolus in region 6 of arm C. Both species display the presence of interspecific population polymorphism, determined by the structure of this arm. In some populations, chromosome regions from 4 to 6 contain a homozygous inversion, which is absent in the other populations. The distinguishing karyotype feature of S. assimilis, which shares fluctuating homozygous inversions with the other species, is the presence of two species-specific homozygous inversions. These are the secondary overlapping inversion in arm A, regions 2 to 7, and the inversion in regions 4 to 10 of arm G. Both species of interest contain nucleolus organizer in region 10 of arm G. In populations of S. nebulosa, six heterozygous inversions localized in arms A, B, C, F, and G were discovered. The highest number of heterozygotes for inversions (71%) was observed in the population from Southern Baikal. In arm B of S. assimilis, one heterozygous inversion and heterozygosity for nucleolus organizer in the chromosome region 16 was detected. Chromosomal evolution of Baikalian Sergentia, and the role of inversion polymorphism in the population adaptation is discussed. Original Russian Text ? V.I. Proviz, 2008, published in Genetika, 2008, Vol. 44, No. 12, pp. 1627–1637.     

Genetic Polymorphism of a Pelagic Fish Species,Little Baikal Oilfish <Emphasis Type="Italic">Comephorus dybowski</Emphasis>, Deduced from Analysis of Microsatellite Loci

Intraspecific genetic polymorphism of a Baikal Lake endemic, little Baikal oilfish (Comephorus dybowski Korotneff, 1905), was evaluated based on microsatellite analysis. Six microsatellite loci designed for the European sculpin, Cottus gobio, were used. Each locus was tested using 25 to 32 individuals from each of the Baikal basins (southern, middle, and northern). Analysis of genetic differentiation (F _ST and R _ST) revealed no statistical significant differences between the samples. The data showed that the target species was represented by a single panmictic stable population.__________Translated from Genetika, Vol. 41, No. 7, 2005, pp. 919–924.Original Russian Text Copyright © 2005 by Teterina, Sukhanova, Bogdanov, Anoshko, Kirilchik.     

Speciation of sponges in Baikal-Tuva region: an outline

Lake Baikal is known for its high percentage of endemic fauna and flora. The most abundant sessile animal taxa in the littoral zone of Baikal are the photosymbiotic sponges. These endemic sponges are grouped to the family Lubomirskiidae and are separated, based on molecular data, from the cosmopolitan family Spongillidae Gray 1867. In the present review, recent data on the potential driving forces of the rapid speciation in Lake Baikal have been unified. Current data suggest that the genetic repertoire of the sponges was sufficiently large to cope with the major cold events, occurring 2.8–2.5 and 1.8–1.5 Ma. It is proposed that during those periods of climatic incisions founder populations were separated from the parental cosmopolitan sponge population and developed subsequently to the array of endemic species. To clarify whether the endemic sponge fauna is indeed restricted to Lake Baikal only or whether there exist related taxa in other lakes, a collection of sponges in the 750 km distant Lake Chagytai was performed. This lake harbours unexpectedly large populations of the endemic species Baikalospongia dzhegatajensis . As the habitat of the Lake Chagytai (algal habitat) differs strongly from that of Lake Baikal (rocky habitat), it is proposed that after the formation of the initial founder population in response to the climatic shift, subsequent speciation was driven by habitat differences.     

Expression patterns and organization of the hsp70 genes correlate with thermotolerance in two congener endemic amphipod species (Eulimnogammarus cyaneus and E. verrucosus) from Lake Baikal

We studied various aspects of heat‐shock response with special emphasis on the expression of heat‐shock protein 70 (hsp70) genes at various levels in two congener species of littoral endemic amphipods (Eulimnogammarus cyaneus and E. verrucosus) from Lake Baikal which show striking differences in their vertical distribution and thermal tolerance. Although both the species studied demonstrate high constitutive levels of Hsp70, the thermotolerant E. cyaneus exhibited a 5‐fold higher basal level of Hsp70 proteins under normal physiological conditions (7 °C) and significantly lower induction of Hsp70 after temperature elevation compared with the more thermosensitive E. verrucosus. We isolated the hsp70 genes from both species and analysed their sequences. Two isoforms of the cytosolic Hsp70/Hsc70 proteins were detected in both species under normal physiological conditions and encoded by two distinct hsp/hsc70 family members. While both Hsp70 isoforms were synthesized without heat shock, only one of them was induced by temperature elevation. The observed differences in the Hsp70 expression patterns, including the dynamics of Hsp70 synthesis and threshold of induction, suggest that the increased thermotolerance in E. cyaneus (compared with E. verrucosus) is associated with a complex structural and functional rearrangement of the hsp70 gene family and favoured the involvement of Hsp70 in adaptation to fluctuating thermal conditions. This study provides insights into the molecular mechanisms underlying the thermal adaptation of Baikal amphipods and represents the first report describing the structure and function of the hsp70 genes of endemic Baikal species dwelling in thermally contrasting habitats.     

Materials on the flora of diatoms (Centrophyceae) from lakes of the Baikal Region and Transbaikalia

The species composition of Centrophyceae (Bacillariophyta) in the phytoplankton of nine lakes in the Baikal Region and Transbaikalia was studied using electron microscopy (SEM) methods. Nineteen species and varieties of Centrophyceae were recorded; Aulacoseira distans var. septentrionalis and Cyclotella arctica were registered in this region for the first time. Aulacoseira baicalensis, Cyclotella minuta, and Stephanodiscus inconspicuus were also found in the studied lakes and attributed to being endemic to Lake Baikal.