Olga V. Aksenova

Results of testing the comparatory method: The curvature of the shell valve frontal section is inappropriate as a systematic character for the freshwater pearl mussel of the genus Margaritifera

This paper continues a discussion on the number of pearl mussel species of the genus Margaritifera in northern Europe. A biometric study of 1711 pearl mussel Margaritifera margaritifera shells from 15 rivers in Russia and Latvia (basins of the White and Baltic seas) has been conducted. All the examined samples fall into two groups: the northern group (with the shells more flattened on average, f. margaritifera) and the southern one (with more convex shells, f. elongata); the boundary between these groups is at 63° N. Analysis of intrapopulation variation has shown that the samples contain individuals that correspond to f. margaritifera, f. elongata, and f. borealis. However, any hiatus between these forms is absent in all the samples, and individuals belonging to two intermediate forms are rather frequent. The hypothesis on the species specificity of the shell valve frontal section has not been confirmed based on examination of large shell samples. The pearl mussels inhabiting rivers of Northern Europe belong to a single species, M. margaritifera.

Taxonomy and distribution of freshwater pearl mussels (Unionoida: Margaritiferidae) of the Russian Far East.

The freshwater pearl mussel family Margaritiferidae includes 13 extant species, which are all listed by IUCN as endangered or vulnerable taxa. In this study, an extensive spatial sampling of Margaritifera spp. across the Russian Far East (Amur Basin, Kamchatka Peninsula, Kurile Archipelago and Sakhalin Island) was conducted for a revision of their taxonomy and distribution ranges. Based on their DNA sequences, shell and soft tissue morphology, three valid species were identified: Margaritifera dahurica (Middendorff, 1850), M. laevis (Haas, 1910) and M. middendorffi (Rosén, 1926). M. dahurica ranges across the Amur basin and some of the nearest river systems. M. laevis is distributed in Japan, Sakhalin Island and the Kurile Archipelago. M. middendorffi was previously considered an endemic species of the Kamchatka. However, it is widespread in the rivers of Kamchatka, Sakhalin Island, the Kurile Islands (across the Bussol Strait, which is the most significant biogeographical boundary within the archipelago), and, likely, in Japan. The Japanese species M. togakushiensis Kondo & Kobayashi, 2005 seems to be conspecific with M. middendorffi because of similar morphological patterns, small shell size (<100 mm long) and overlapped ranges, but it is in need of a separate revision. Phylogenetic analysis reveals that two NW Pacific margaritiferid species, M. laevis and M. middendorffi, formed a monophyletic 18S rDNA clade together with the North American species M. marrianae and M. falcata. The patterns that were found in these Margaritifera spp. are similar to those of freshwater fishes, indicating multiple colonizations of Eastern Asia by different mitochondrial lineages, including an ancient Beringian exchange between freshwater faunas across the Pacific.

New taxa of freshwater mussels (Unionidae) from a species-rich but overlooked evolutionary hotspot in Southeast Asia

Southeast Asia harbors a unique and diverse freshwater fauna of Mesozoic origin, which is under severe threat of extinction because of rapid economic development and urbanization. The largest freshwater basins of the region are certainly the primary evolutionary hotspots and they attract the most attention as key biodiversity areas for conservation. In contrast, medium-sized rivers are considered low-importance areas with secondary biodiversity, whose faunas originated via founder events from larger basins during the Pleistocene, although such a scenario has never been tested by using a phylogenetic approach. In this investigation, we used freshwater mussels (Unionidae) as a model to estimate the levels of endemism within the Sittaung, a little-known remote basin in Myanmar, compared with the surrounding larger rivers (Irrawaddy, Salween and Mekong). We discovered that the Sittaung represents an exceptional evolutionary hotspot with numerous endemic taxa of freshwater mussels. On the basis of our extensive dataset, we describe two new tribes, two genera, seven species and a subspecies of Unionidae. Our results highlight that medium-sized basins may represent separate evolutionary hotspots that harbor a number of endemic lineages. These basins should therefore be a focus of special conservation efforts alongside the largest Southeast Asian rivers.

A taxonomic revision of two local endemic Radix spp. (Gastropoda: Lymnaeidae) from Khodutka geothermal area, Kamchatka, Russian Far East.

Khodutka geothermal area is located near Khodutka and Priemysh volcanoes and is one of the largest geothermal areas of the Kamchatka Peninsula. Vakin (2003) described geological, geochemical and geothermic conditions of this geothermal area in detail. The main thermal water sources have temperatures up to 87°C and a discharge of approximately 150 l×sec. -1 are flows out into the warm lake with dimensions of ca. 250 m length and 80 m width. This warm river is ca. 20 m in width beginning from the lake and flows to the Bolshaya Khodutka River basin. Two local endemic Radix species were described from this geothermal area, especially Lymnaea ( Radix ) hadutkae Kruglov & Starobogatov, 1989 and L. ( R. ) thermokamtschatica Kruglov & Starobogatov, 1989 (Kruglov & Starobogatov 1989, 1993; Kruglov 2005). These species were separated using proportions of shell and reproductive system (Kruglov & Starobogatov 1989, 1993; Starobogatov et al. 2004). According to the diagnosis, L. ( R. ) hadutkae differs in the ear-shape shell, a form of the provaginal duct with cylindrical distal part and conical proximal part, and larger value of the index of the copulatory apparatus (ICA: proportion of the preputium to phallotheca is 1.27) from other species within the section Thermoradix Kruglov & Starobogatov, 1989. The last whorl is large, 0.86–0.89 of the shell height; an excess of the last whorl over upper margin of the aperture is 0.15–0.16 of the aperture height. L. ( R. ) thermokamtschatica has the cylindrical provaginal duct, relatively short bursa duct (1.5X longer than the bursa copulatrix diameter) and very long phallotheca (ICA is 0.77). The last whorl is large, 0.85–0.87 of the shell height; an excess of the last whorl over upper margin of the aperture is 0.2–0.25 of the aperture height. In accordance with an identification key (Starobogatov et al. 2004), the main diagnostic feature is an excess of the last whorl over the shell aperture, which has ≤0.78 and ≥0.80 of the penultimate whorl width in the first and second species, respectively. In the present paper, we revised these taxa using newly collected topotypes and additional Radix spp. specimens from other areas of the Russian Far East.

Radix dolgini: The integrative taxonomic approach supports the species status of a Siberian endemic snail (Mollusca, Gastropoda, Lymnaeidae).

The molecular techniques are the standard tool for the study of the taxonomic position and phylogenetic affinities of the lymnaeid genus Radix Montfort, 1810, and the majority of the European representatives of this taxon have been studied in this respect. However, a plethora of nominal species of Radix described from Northern Asia (Siberia and the Russian Far East) are still characterized only morphologically, raising some doubts concerning their validity. In this paper, we present the triple (morphological, molecular, and zoogeographical) evidence that there is at least one endemic species of Radix, Radix dolgini (Gundrizer and Starobogatov, 1979), widely distributed in Siberia and Western Mongolia. Phylogenetically, it is a sister species to the European R. labiata (Rossmaessler, 1835) [=R. peregra auct.], and their common ancestor most probably lived in the Pliocene, nearly 3.25Myr ago. Our results assume the existence of an extended dispersal barrier for freshwater hydrobionts between Europe and Siberia in the Late Pliocene that may be important for biogeographical explanations. Three other nominal Siberian species of Radix: R. kurejkae (Gundrizer and Starobogatov, 1979), R. gundrizeri (Kruglov and Starobogatov, 1983), and R. ulaganica (Kruglov and Starobogatov, 1983) proved to be the junior synonyms of R. dolgini.

Mollusks in the Zoobenthos of Relict Lakes with Abnormally High Biological Production in the Eastern European Subarctic

Species diversity and density of mollusks have been studied in Vashutkiny Lakes (Bol’shezemel’skaya Tundra, northeastern Europe). These lakes are comparable to northern boreal lakes in mollusk species diversity and density levels. Two hypotheses proposed by Zvereva are analyzed: on the relict origin of the ecosystem of these lakes and on the leading role of intrazonal factors in the formation of their abnormally high level of production. Both hypotheses are confirmed by recent data. The production of these lakes is mainly determined by their flowing regime, which offers intense convective heat exchange between water masses and suprapermafrost taliks under the lakes and leads to the accumulation of summer heat in friable Quaternary sediments, which are widespread in lake depressions. Intense water exchange together with strong wind-induced mixing hinders the stratification of water masses.

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 and Lymnaea (Polyrhytis) kurenkovi Kruglov and Starobogatov, 1989, 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.

Reproduction of Pisidium casertanum (Poli, 1791) in Arctic lake

Freshwater invertebrates are able to develop specific ecological adaptations that enable them to successfully inhabit an extreme environment. We investigated the brooding bivalve of Pisidium casertanum in Talatinskoe Lake, Vaigach Island, Arctic Russia. Here, quantitative surveys were conducted, with the collection and dissections of 765 molluscs, on the basis of which analyses on the brood sacs length (marsupia) and the number and size of embryos, were performed. In this study, the number of brooded embryos was positively correlated with the parents shell length. The number of extramarsupial embryos was much lower than the number of intramarsupial embryos. Our research also showed that the brood sac length and embryos within one individual can vary significantly. Thus, we detected that P. casertanum has a specific brooding mechanism, accompanied by asynchronous development and embryos release by the parent. We suggest that such a mode could result in the coin-flipping effect that, presumably, increases the population breeding success in the harsh environment of the Arctic lake.

Occurrence of a Sphaerium species (Bivalvia: Sphaeriidae) of Nearctic origin in European Arctic Russia (Vaigach Island) indicates an ancient exchange between freshwater faunas across the Arctic

Although a recent series of studies has examined the phylogeography and diversity of Arctic invertebrates related to the glacial history on a pan-Arctic scale, large gaps remain in the knowledge on the origin, dispersal and biogeography of freshwater mollusks in the Russian Arctic. In samples from two lakes on Vaigach Island, Arctic European Russia, we found five specimens of an undescribed Sphaerium species (Bivalvia: Sphaeriidae), which is most closely related to Nearctic species (particularly Sphaerium rhomboideum), with the majority of known occurrences in the Great Lakes region of North America. This finding suggests a Nearctic rather than Palearctic origin of this lineage. Genetic evidence demonstrated the existence of freshwater refugia in the European Arctic during the last glacial maximum, and a Nearctic origin of the Vaigach lineage would suggest the existence of a phylogeographic element distinct from other bivalve molluscs of Northern and Alpine Europe origin inhabiting the European Arctic.

Two Pisidium species inhabit freshwater lakes of Novaya Zemlya Archipelago: the first molecular evidence

The information regarding freshwater mollusks of water bodies of Novaya Zemlya Archipelago is extremely limited. In 2015, the study of the lakes of the Southern Island of Novaya Zemlya Archipelago was undertaken within the framework of the research expedition of the “Arctic Floating University”. In general, two bivalve species were found in the lakes, Pisidium cf. conventus (Clessin, 1877) and P. waldeni (Kuiper, 1975). It is a northernmost record for Pisidium in the Palearctic Region. The present study provides the first occurrence of P. waldeni on the Novaya Zemlya Archipelago. The results of molecular analyses show that P. cf. conventus from Svyatoe Lake clusters together with the clade Odhneripisidium Kuiper, 1962. Our data are in accordance with the tabula rasa hypothesis, which states that the contemporary freshwater invertebrate fauna of the Novaya Zemlya originates through recent immigration processes after the Last Glacial Maximum.