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Shells of Anodonta, Pseudanodonta and Sinanodonta from Russia. (A) Anodonta anatina, Lake Syrdah, Lena River basin, Eastern Siberia [RMBH biv199_2]. (B) Anodonta anatina, Lake Gusinoye, Yenisei River basin, Eastern Siberia [RMBH biv190_4]. (C) Anodonta anatina, Oka River, Volga River basin, European Russia [RMBH biv573_4]. (D) Anodonta cygnea, Medvezhii Lakes, Moscow, European Russia [RMBH biv194_1]. (E) Pseudanodonta complanata, Khopyor River, Don River basin, European Russia [RMBH 195_1]. (F) Sinanodonta lauta [=S. ovata Bogatov & Starobogatov, 1996, a topotype], Gladkaya River, Russian Far East [RMBH biv225_2]. (G) Sinanodonta schrenkii, Melgunovka River, Khanka Lake basin, Russian Far East [RMBH biv496_3]. (H) Sinanodonta woodiana, non-native population, Yenisei River near Krasnoyarsk, Eastern Siberia [RMBH biv191_3]. Scale bars = 10 mm. (Photos: Ekaterina S. Konopleva [A,B,D–G] and Olga V. Aksenova [C,H]).

Shells of Anodonta, Pseudanodonta and Sinanodonta from Russia. (A) Anodonta anatina, Lake Syrdah, Lena River basin, Eastern Siberia [RMBH biv199_2]. (B) Anodonta anatina, Lake Gusinoye, Yenisei River basin, Eastern Siberia [RMBH biv190_4]. (C) Anodonta anatina, Oka River, Volga River basin, European Russia [RMBH biv573_4]. (D) Anodonta cygnea, Medvezhii Lakes, Moscow, European Russia [RMBH biv194_1]. (E) Pseudanodonta complanata, Khopyor River, Don River basin, European Russia [RMBH 195_1]. (F) Sinanodonta lauta [=S. ovata Bogatov & Starobogatov, 1996, a topotype], Gladkaya River, Russian Far East [RMBH biv225_2]. (G) Sinanodonta schrenkii, Melgunovka River, Khanka Lake basin, Russian Far East [RMBH biv496_3]. (H) Sinanodonta woodiana, non-native population, Yenisei River near Krasnoyarsk, Eastern Siberia [RMBH biv191_3]. Scale bars = 10 mm. (Photos: Ekaterina S. Konopleva [A,B,D–G] and Olga V. Aksenova [C,H]).

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Freshwater mussels are ecosystem engineers and keystone species in aquatic environments. Unfortunately, due to dramatic declines this fauna is among the most threatened globally. Here, we clarify the taxonomy and biogeography of Russian Unionidae species based on the most comprehensive multi-locus dataset sampled to date. We revise the distribution...

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... Haas (1969) considered U. crassus as a single species, but divided it into eight subspecies, which have since been studied to varying degrees (Fig. 1). Recent studies investigating the evolutionary history of the group have led to the elevation of four subspecies (U. c. bruguierianus, U. c. carneus, U. c. ionicus, and U. c. mongolicus) to the species level (Araujo et al., 2018;Klishko et al., 2019;Bolotov et al., 2020;Lyubas et al., 2022). However, U. c. mongolicus was reassigned to a different genus, namely Middendorffinaia (Klishko et al., 2019). ...
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The global decline of freshwater mussels and their crucial ecological services highlight the need to understand their phylogeny, phylogeography and patterns of genetic diversity to guide conservation efforts. Such knowledge is urgently needed for Unio crassus, a highly imperilled species originally widespread throughout Europe and southwest Asia. Recent studies have resurrected several species from synonymy based on mitochondrial data, revealing U. crassus to be a complex of cryptic species. To address long-standing taxonomic uncertainties hindering effective conservation, we integrate morphometric, phylogenetic, and phylogeographic analyses to examine species diversity within the U. crassus complex across its entire range. Phylogenetic analyses were performed using cytochrome c oxidase subunit I (815 specimens from 182 populations) and, for selected specimens, whole mi-togenome sequences and Anchored Hybrid Enrichment (AHE) data on ∼ 600 nuclear loci. Mito-nuclear discor-dance was detected, consistent with mitochondrial DNA gene flow between some species during the Pliocene and Pleistocene. Fossil-calibrated phylogenies based on AHE data support a Mediterranean origin for the U. crassus complex in the Early Miocene. The results of our integrative approach support 12 species in the group: the previously recognised Unio bruguierianus, Unio carneus, Unio crassus, Unio damascensis, Unio ionicus, Unio sesirmensis, and Unio tumidiformis, and the reinstatement of five nominal taxa: Unio desectus stat. rev., Unio gontierii stat. rev., Unio mardinensis stat. rev., Unio nanus stat. rev., and Unio vicarius stat. rev. Morphometric analyses of shell contours reveal important morphospace overlaps among these species, highlighting cryptic, but geographically structured, diversity. The distribution, taxonomy, phylogeography, and conservation of each species are succinctly described.
... Literature on M. margaritifera and conservation reports demonstrate that freshwater mussels are declining globally and 'despite simultaneous efforts to restore mussel habitat over the last 25 years, natural reproduction still does not occur in some countries, such as in the Czech Republic or in Luxemburg' [17]. While conservation priorities over European countries and North America vary widely, ranging from preventing imminent extinction to maintaining living population and habitat reconstruction as well as provision of host fish, human pressure and destruction of habitats continue to constitute the highest risk for extinction of the species. ...
Article
The Freshwater Pearl Mussels M. margaritifera is an endangered Holartic species in the family of Margaritiferadae in the order of Unionoidea and Class of Bivalvia. M. margaritifera is found in North America, Europe and through into Siberia. It has been declining throughout the European part with a negative forecast on survivability of the species [1] IUCN 1. The longevity of M. margaritifera and the fragility of the reproduction conditions as well as habitat selectivity regarding environmental requirements pose difficult niche conditions for the species. Population decline is not only related to species-specific intrinsic characteristics and extrinsic factor like water pollution (IUCN 2), but also caused by the introduction of invasive species such as the less specialised, non-threatened Freshwater Zebra Mussel Dreissena polymorpha (Araujo, 2006). We discuss the vulnerability of the species in the light of the conceptual framework of Modern Synthesis, MS and the Extended Evolutionary Synthesis, EES [2]. In the outlook we review the conservation status and briefly evaluate the opportunities of EES driven approaches for further research on re-establishment of the species [3].
... Consequentially, some systematists have used the term "integrative taxonomy" to name and differentiate this more inclusive approach when addressing taxonomic questions (e.g., Padial & Riva, 2009;Moraes et al., 2021). Integrative taxonomy has been applied to a vast sort of different groups of organisms, including plants, fungi, and invertebrate and vertebrate animals (e.g., Bolotov et al., 2020;Haelewaters et al., 2018;He et al., 2022). However, regarding South American snakes, most studies have extensively focused on the family Dipsadidae (e.g., Abegg et al., 2022;Arredondo et al., 2020;Costa et al., 2022;Trevine et al., 2022;Zaher et al., 2018). ...
Article
The widespread Tantilla melanocephala is a South American snake, which has a convoluted taxonomic history and a poorly addressed systematics. The taxonomic identity and the phylogenetic affinities of their populations are unclear, suggesting the existence of a complex of undescribed species. Here, we preliminarily address the morphological diversity and genetic structure of the T. melanocephala complex through comparative morphological approaches and phylogenetic analyses using maximum likelihood. Our results confirm the presence of hidden diversity within the T. melanocephala complex and strongly indicate that the populations of T. melanocephala from southern South America represent a distinct lineage, which is morphologically diagnosable under a unique combination of character states. Therefore, we formally describe these populations as a new species and present a comparison among all other South American species of Tantilla. We also estimate the divergence times of the main clades within the complex through Bayesian analysis, suggesting a biogeographical scenario of lineage diversification. Finally, our study clarifies aspects of the intricate taxonomy of the T. melanocephala complex, providing a framework for future studies to avoid the description of synonyms.
... It should be noted that freshwater mussels are characterized by more or less expressed regional endemism, especially in tropical areas Konopleva et al., 2019). The situation in which the native range of one species covers two different biogeographical subregions very rarely occurs and requires direct connections between remote freshwater systems, for example, the unique trans-Beringian range of Beringiana beringiana (Unionidae) (Bolotov et al., 2020;Lopes-Lima et al., 2020). Moreover, the species composition of the majority of subregional unionid faunas in the world has already been identified . ...
... Many species of freshwater mussels may be distinguished based on conchological features, but there are complexes of cryptic species that do not share clear diagnostic characters and cannot be divided using the morphological characters alone. The genera Sinanodonta, Lamellidens, Lens and Yaukthwa are good examples of such taxonomically complicated clades (Bolotov et al., 2020;Bolotov, Konopleva, et al., 2022;Konopleva et al., 2019;Lopes-Lima et al., 2020;Pfeiffer et al., 2021). The best way to identify freshwater mussel species from these groups is the DNA-based approach supplemented with morphological data and biogeographical evidence . ...
Article
1. Freshwater mussels (order Unionida) are a popular group for a wide array of non-taxonomic research driven by questions related to their functions in ecosystems, to relationships among species, to biogeochemical and morphometric patterns of certain species and others. Valid taxonomy and correct species identification are key requirements for all of these aspects. However, species-level identification of representatives from this group is a rather complicated task and should be based on an integrative approach, combining DNA sequences, morphological and anatomical investigation and biogeographical estimates. 2. This article reconsiders a selection of non-taxonomic scientific works (N = 25), containing misidentified occurrences of freshwater mussel species. The dataset contains records of the endangered Margaritifera margaritifera (endemic to eastern North America and Europe) from the Philippines and West Africa, as well as occurrences of the Nearctic Gonidea angulata from West Africa and the Middle East. Several Palearctic and Nearctic unionid species were erroneously reported from the Indus River, Pakistan. Subfossil shells of the native Simpsonella sp. from a prehistoric site in the Philippines were misidentified as the invasive Sinanodonta woodiana that was introduced to the islands in the 20th century. Samples of the tropical lineage of S. woodiana from Indonesia were mistaken for the native Pilsbryoconcha exilis, and vice versa. Salinity tolerance and morphometric characteristics of the estuarine clam Geloina sp. (Cyrenidae) from Sumatra were examined, but these data were published as belonging to the strictly freshwater S. woodiana. 3. It is clear that this information, being reused by researchers, conservationists and stakeholders, will lead to incorrect conclusions on the range, status, biogeochemistry, morphometry and ecological tolerance of certain freshwater mussel taxa, including invasive and endangered species (the so-called ‘error cascades’ in biological sciences caused by ‘bad taxonomy’). 4. To reduce the growing body of literature containing misidentifications of the Unionida taxa, practical recommendations are proposed for researchers, who include freshwater mussels in non-taxonomic surveys, as well as for journal editors dealing with articles that focus on these animals.
... Details of the shell structure were studied using Canon EOS 7D DSLR camera (Canon Inc., Japan). Collected specimens of freshwater mussels were compared with descriptions of freshwater bivalve species of the studied region published by Bolotov et al. (2020) and with the voucher specimens deposited in RMBH. The results of the morphological identification was verified based on nucleotide sequences of the specimens of bivalves using the Basic Local Alignment Search Tool, BLAST (Johnson et al., 2008). ...
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Concentrations of the chemical elements were analyzed in the shells of a bivalve mollusk species (Beringiana beringiana), water, and bottom sediments from seven lakes located on the Kamchatka Peninsula, the Kurile Islands, Sakhalin Island, and Primorsky Krai (Northeast Asia). A principal component analysis allowed to determine three factors those were related to environments in the waterbodies. We revealed two groups of samples corresponding to large geographical regions using the determined factors. Statistically significant differences were found between geographical groups of samples, and higher values of element distribution coefficients were determined for samples from lakes on the Kamchatka Peninsula. The highest concentrations of lithophilic elements were measured in the shells from Lake Kurazhechnoye (Kamchatka Peninsula). The highest concentrations of Al and Mg were detected in shells from Lake Peschanoye (Kunashir Island). In Lake Chernoye (Sakhalin Island), the highest concentrations of Sr and Sb in the shells were detected. Zn, Fe, Pb, and rare earth elements were present in large concentrations in the shells from Lake Vaskovskoye, Primorsky Krai. The shells of the Beringian freshwater mussel show large phenotypic plasticity, and their shape demonstrates significant relationships with various environmental parameters, that were assessed based on the geochemical indicators.
... Комиссаровка L. maacki и L. chankensis были сведены в синонимы к L. grayii (J. E. Gray, 1833) (Bolotov et al. 2020;Lopez-Lima et al. 2020), имеющему обширный ареал от бассейна Янцзы до Амура. Очевидно, что для подобных выводов необходимы, как минимум, дополнительные исследования, поскольку на наличие двух российских видов ланцеолярий указывают морфологические различия как взрослых (Богатов 2022), так и личиночных (Саенко 2008) раковин, а также разные экологические предпочтения видов с указанием на то, что в бассейне Ханки они не встречаются совместно (Прозорова 2021). ...
Article
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Information on the distribution of two endemic species of large bivalves in the Primorsky Krai has been clarifi ed. Both species are listed in the Red Data Book of the Russian Federation: Lanceolaria maacki Moskvicheva, 1973 and L. chankensis Moskvicheva, 1973. Currently known populations of these species are listed together with an assessment of their condition. A reduction in the number and area of distribution of the species is demonstrated. The main threats and ways for the Lanceolaria species conservation in Lake Khanka and the Ussuri River are discussed.
... After removing the soft parts, the shells of adult mussels were washed, dried and labeled in the laboratory, and photographed using a digital camera. Identification was made according to the most recent taxonomic revisions 23,24 and confirmed by DNA data of mollusks collected in the abovementioned localities 23 . ...
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Freshwater mussels of the genus Buldowskia (Bivalvia, Unionidae) are distributed from the Amur River basin in Russia and China southward to the Korean Peninsula and some Japanese islands. This work is an integrative morphological study of Buldowskia suifunica glochidia from locations in the Primorsky Territory, the Russian Far East. Glochidia of B. suifunica, taken from the same gill have asynchronous development. The external and internal morphology of its shell has been characterized. The morphology of its sensory system, within three stages of larval development (immature, intermediate and mature glochidia), consists of hair cells as well as nonhair cells. Their muscle system is composed of massive adductor and minor muscle bundles. The FMRFamid-ergic nervous system turned out to be a complex system includes basal cells (neurons), their neurites and anterior neurons. FMRFamide and tubulin was found in all neurons. Glochidia of B. suifunica have only four 5-HT-lir neurons. We concluded that B. suifunica glochidial nervous system differs from those of the larval systems of planktotrophic marine mollusks.
... To test this hypothesis, we compared trace element composition in marine and freshwater mussel shells on the basis of several environmental indicators, considering two contrasting sites across Northern Eurasia, one in the European Subartic and another one in the Kamchatka Peninsula. The selected regions are well-studied in the course of fresh-and brackish-water mollusks and there are several well-known localities of widely distributed freshwater (Unio sp., Anodonta sp., and Beringiana sp.) and brackish-water (Mytilus sp.) species of mussels [24,25]; those were used in the presented study as representative sampling sites. ...
... The primary identification of collected bivalve mollusks was carried out based on standard and special keys [24,25]. The comparative analysis of the shell morphology was carried out, taking into account the structure of the pseudocardinal and lateral teeth, muscle attachment scars, shell shape, and umbo position [26,27]. ...
... Analyses of seven COI gene sequences for marine mussels with length of 663 bp demonstrated that the mollusk specimens belong to species M. edulis and M. trossulus. Analyses of five nucleotide sequences of COI gene for freshwater mussels with length of 660 bp yielded B. beringiana, A. anatina, U. tumidus, and U. pictorum specimens at the species level, in accordance with conchological features [25]. ...
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The accumulation of trace metals in the shells of bivalves allows quantitative assessments of environmental pollution and helps to reconstruct paleo aquatic environments. However, the understanding on how marine and freshwater mollusks control the level of trace elements in their shells remains very limited. Here, we compared the trace element composition of marine and freshwater bivalves from boreal and subarctic habitats, using examples of widely distributed species of marine (Mytilus edulis, M. trossulus) and freshwater (Anodonta anatina, Unio sp., Beringiana beringiana) mussels. Sizable differences in several trace element concentrations were detected between different species, depending on their environmental niches. A multiparametric statistical treatment of the shell’s elemental composition allowed to distinguish the impact of external factors (water and sediment chemical composition) from active metabolic (biological) control. In particular, the obtained results demonstrated that Ba:Ca and Pb:Ca ratios in mussels’ shells are closely related to the primary productivity of aquatic ecosystems. The Mn:Ca ratio allowed to constrain the environmental conditions of mussels’ species depending on the trophic state of inhabited waterbody. Overall, the marine mussels exhibited stronger biological control of trace element accumulation, whereas trace element pattern in shells of freshwater mussels was chiefly controlled by environmental factors. The obtained results might help to use the trace element composition of bivalves in distinguishing marine and freshwater habitats of mollusks in paleo environments.
... Recently, laser ablation multi-collector inductively coupled plasma mass spectrometry (LA-MC-ICP-MS) instrumentation has been widely used to determine the in situ major trace element and isotopic compositions of titanites [1,14,[16][17][18][19][20][21][22][23][24][25][26][27][28]. However, the lack of matrix matching standard materials is a significant barrier to the widespread adoption of this technology [29]. ...
... The Mogok titanite samples can meet the requirement of younger standard samples. trace element and isotopic compositions of titanites [1,14,[16][17][18][19][20][21][22][23][24][25][26][27][28]. However, the lack of matrix matching standard materials is a significant barrier to the widespread adoption of this technology [29]. ...
... Natural dispersal of freshwater mussels mostly occurs at the larval stage together with their fish hosts, and usually requires direct connections between freshwater basins, because they are unable to cross oceanic barriers 2,12-15 . Hence, freshwater mussels are considered to be among the best model organisms for biogeographic and paleogeographic reconstructions [16][17][18][19][20][21] . ...
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... The published distribution maps of B. beringiana depict a continuous range in the Russian Far East [13]. However, the list of its findings (Figure 2) suggests that it occurs as scattered enclaves, mostly associated with the lower reaches of large rivers (Indigirka, Kolyma, Anadyr, Kamchatka, and their tributaries). ...
Article
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The origin of the fauna of Beringia is a notable biogeographical puzzle. Large mussels of the genus Beringiana inhabit both Northeast Asia and the northwestern part of North America and thus provide an important model to investigate the paleobiogeography of Beringia and the past and current intercontinental species exchanges. Data on Beringiana distribution, morphology, genetics, and taxonomy are fragmentary or questionable. In this study, we summarized the data on its distribution in Northeast Asia, performed genetic analysis (cox1), and studied the variation in the shell morphology in samples from four isolated populations, including the putative sympatric species. Over ten large enclaves of Beringiana are currently known in Northeast Asia (east to the Verkhoyansk Range), mostly in the lower reaches of large rivers in northeastern Yakutia, Kamchatka, Magadan Oblast, and Khabarovsk Krai. These enclaves are far apart (several hundreds of kilometers) because the mussel is associated with muddy sand or pebbly sand bottom substrates and spreads through its glochidia, which are parasitic on the fish. Shell morphology can be highly variable even in a single population, overlapping the diagnoses of several Beringiana species (which are currently not recognized as valid). Our analysis of the cox1 sequence in four populations identified all individuals as B. beringiana. We evaluated the possible current and probable Late Pleistocene–Early Holocene freshwater and marine pathways of B. beringiana dispersal through the area of former Beringia, including the current intercontinental migration of glochidia on fish.