Cryptic taxonomic diversity and high-latitude melanism in the glossiphoniid leech assemblage from the Eurasian Arctic.

The family Glossiphoniidae is a diverse and widespread clade of freshwater leeches, playing a significant role in functioning of aquatic ecosystems. The taxonomy and biogeography of leeches from temperate, subtropical, and tropical regions attracted much attention of zoologists, while their taxonomic richness and distribution in the Arctic are poorly understood. Here, we present an overview of the Eurasian Arctic Glossiphoniidae based on the most comprehensive occurrence and DNA sequence datasets sampled to date. This fauna contains 14 species, belonging to five genera and three subfamilies. One genus and five species are new to science and described here. The world's northernmost occurrences of glossiphoniids are situated on the Taymyr Peninsula at 72° N, although further records at higher latitudes are expected. Most Arctic leeches are characterized by broad ranges crossing several climatic zones (e.g., Glossiphonia balcanica and G. nebulosa), although the distribution of two new species may be confined to the high-latitude areas. The Taymyr Peninsula with the nearby Putorana Plateau represents the most species-rich area (totally 9 species), while the European Arctic, Iceland, Kolyma Highland, and Chukotka Peninsula house depleted faunas (2-4 species per subregion). Finally, we show that the high-latitude melanism is a common phenomenon in glossiphoniid leeches.

A nearly complete database on the records and ecology of the rarest boreal tiger moth from 1840s to 2020.

Global environmental changes may cause dramatic insect declines but over century-long time series of certain species' records are rarely available for scientific research. The Menetries' Tiger Moth (Arctia menetriesii) appears to be the most enigmatic example among boreal insects. Although it occurs throughout the entire Eurasian taiga biome, it is so rare that less than 100 specimens were recorded since its original description in 1846. Here, we present the database, which contains nearly all available information on the species' records collected from 1840s to 2020. The data on A. menetriesii records (N = 78) through geographic regions, environments, and different timeframes are compiled and unified. The database may serve as the basis for a wide array of future research such as the distribution modeling and predictions of range shifts under climate changes. It represents a unique example of a more than century-long dataset of distributional, ecological, and phenological data designed for an exceptionally rare but widespread boreal insect, which primarily occurs in hard-to-reach, uninhabited areas of Eurasia.

Oriental freshwater mussels arose in East Gondwana and arrived to Asia on the Indian Plate and Burma Terrane.

Freshwater mussels cannot spread through oceanic barriers and represent a suitable model to test the continental drift patterns. Here, we reconstruct the diversification of Oriental freshwater mussels (Unionidae) and revise their taxonomy. We show that the Indian Subcontinent harbors a rather taxonomically poor fauna, containing 25 freshwater mussel species from one subfamily (Parreysiinae). This subfamily most likely originated in East Gondwana in the Jurassic and its representatives arrived to Asia on two Gondwanan fragments (Indian Plate and Burma Terrane). We propose that the Burma Terrane was connected with the Indian Plate through the Greater India up to the terminal Cretaceous. Later on, during the entire Paleogene epoch, these blocks have served as isolated evolutionary hotspots for freshwater mussels. The Burma Terrane collided with mainland Asia in the Late Eocene, leading to the origin of the Mekong's Indochinellini radiation. Our findings indicate that the Burma Terrane had played a major role as a Gondwanan "biotic ferry" alongside with the Indian Plate.

Trapped on the Roof of the World: taxonomic diversity and evolutionary patterns of Tibetan Plateau endemic freshwater snails (Gastropoda: Lymnaeidae: Tibetoradix).

The high-elevation Tibetan Plateau (western China) is inhabited by a unique, though not particularly species-rich, community of organisms. We explored the species content and evolutionary history of the Tibetan Plateau endemic freshwater snail genus Tibetoradix. Phylogenetic relationships within the genus were reconstructed based on available sequence data. We used a single-rate Poisson Tree Processes approach for species delimitation and compared putative species-level clades with already described taxa. We found that the genus consists of at least 6 species, of which we described 4 as new to science. Shell and soft body morphology was examined and the radula in Tibetoradix was described for the first time. Based on our findings, the diversification of the genus did not result in a prominent morphological differentiation and a number of species can be regarded as morphologically cryptic. Single species found in different drainage areas indicate relatively good passive dispersal abilities of the snails. The allopatric distribution of the species could result from competitive exclusion between them. The absence of Tibetoradix spp. outside the Tibetan Plateau could be explained by a scenario of an "evolutionary trap", where adaptations to high elevation conditions prevented the taxa from a successful colonization of lower elevations.

Phylogeography and ecology of bumble bees on Kolguev Island, a remote European Arctic landmass.

The bumble bee fauna of the Russian Arctic is rather poorly known. Kolguev Island, a remote insular territory in the Barents Sea, is one of the deficiently studied areas. In this study, material on Kolguev's bumble bees is re-examined, phylogeographic data analysed, putative scenarios explaining the origin of the bumble bee fauna on the island discussed, and the biology and phenology of these insular populations described. Five bumble bee species, i.e., Bombusflavidus, B.lapponicus, B.jonellus, B.pyrrhopygus, and B.balteatus, were recorded on this island. All of these species are widespread throughout the Eurasian Arctic. Bumble bee populations on Kolguev Island are characterised by a low level of molecular divergence from mainland populations. Based on paleogeographic reconstructions and phylogeographic patterns, it is hypothesised that the bumble bees appeared on this island in the Early Holocene. The lack of rodents (lemmings and voles) sharply decreases the number of available nesting places for bumble bees on Kolguev Island.

A new freshwater leech species from Asian Swamp Eel stocks in China.

Discoveries of new Hemiclepsis species (Hirudinea: Glossiphoniidae) in East and Southeast Asia were expected. A peculiar freshwater leech was found on the body surface of Asian Swamp Eel Monopterus albus (Synbranchiformes: Synbranchidae) in Jingzhou, Hubei Province, China. Here, we describe this leech species as Hemiclepsis yangtzenensis sp. nov. It could be distinguished from other congeners based on a combination of morphological features such as very small size, translucent body with dense green reticulate markings throughout dorsum, very large posterior sucker with a characteristic reticulate pattern, and two pairs of well-developed drop-like eyespots. It also represents a phylogenetic lineage that is distant from other members of the genus based on the COI and 18S rRNA sequences. This leech species seems to be a specialized fish parasite that could negatively affect farming and wild stocks of Asian Swamp Eel in China.

Is the South African leech Barbronia gwalagwalensis Westergren amp; Siddall, 2004 (Hirudinida: Erpobdelliformes: Salifidae) a Paleotropical species?

The freshwater leech family Salifidae Johansson has a Paleotropical range, with a hotspot of species richness in the Oriental Region, and a few species endemic to Africa, Madagascar, and Reunion. Barbronia gwalagwalensis Westergren Siddall, 2004 was thought to be a characteristic example of the latter group being a lineage endemic to South Africa. However, we found that this species also occurs in Asia (Myanmar and Korea). Our time-calibrated phylogeny based on the mitochondrial cytochrome c oxidase subunit I (COI) gene reveals that the split between the African and Asian populations of this species occurred in the mid-Pleistocene, approximately 1.3 Ma ago (95% HPD 0.7-2.1 Ma). The statistical biogeographic modeling indicates that a B. gwalagwalensis population in South Africa most likely originated due a long-distance dispersal event with a subsequent vicariance (probability = 88.9%). A Late Quaternary range extension towards South Africa is known to occur in some other freshwater taxa (e.g. the freshwater mussel Unio caffer Krauss species group), which agrees with our hypothesis on the ancient origin of the South African B. gwalagwalensis population. Conversely, we can assume that the African population of this species was recently introduced from Asia. If so, the high levels of genetic divergence between African and Asian populations could be a part of a more general phylogeographic pattern historically originated within the Asian subcontinent due to the isolation by orographic or marine barriers. These two alternative hypotheses need further research efforts, i.e. sampling and sequencing of other Barbronia taxa, the ranges of which are situated between South Africa and Southeast Asia, as well as of topotypes of B. weberi (Blanchard, 1897) from Indonesia. Finally, our results highlight that the salifid genus Barbronia Johansson originated in the Oriental Region and that these leeches share both recently and historically high potential for long-distance dispersal events.

New freshwater mussels from two Southeast Asian genera Bineurus and Thaiconcha (Pseudodontini, Gonideinae, Unionidae).

The Mekong and Chao Phraya rivers harbor a species-rich freshwater mussel assemblage containing a large radiation of the Pseudodontini species. Members of the genera Bineurus Simpson 1900 and Thaiconcha Bolotov et al., 2020 primarily inhabit small and medium-sized tributaries of these rivers. Here, we present an integrative taxonomic review of these genus-level clades. We show that Bineurus contains four species: B. mouhotii (Lea, 1863), B. exilis (Morelet, 1866) stat. rev., B. anodontinum (Rochebrune, 1882) stat. rev., and B. loeiensis sp. nov. In its turn, Thaiconcha comprises three species: T. callifera (Martens, 1860), T. munelliptica sp. nov., and T. thaiensis sp. nov. Two species, Pseudodon ovalis Morlet, 1889 and P. thomsoni Morlet, 1884, are considered here as questionable taxa. These findings further highlight that Southeast Asia represents a significant evolutionary hotspot of freshwater mussels, which requires further international collaborative research and conservation efforts.

First freshwater mussel-associated piscicolid leech from East Asia.

Parasites and symbionts of freshwater mussels are poorly understood, although a diverse assemblage of mussel-associated leeches (Glossiphoniidae) was recently described. Here, we report on the discovery of a fish leech (Piscicolidae) in the mantle cavity of the freshwater mussel Cristaria plicata (Unionidae) in the Russian Far East. It is the first member of this leech family being associated with freshwater molluscs. This leech does not match any known genus and species both morphologically and genetically, and is described here as Alexandrobdella makhrovi gen. & sp. nov. It uses mussels as shelter (and probably as a secondary host), while the Amur catfish Silurus asotus (Siluridae) seems to be the primary host. These novel findings indicate that mussel-associated leech assemblage contains at least one piscicolid species. Our fossil-calibrated phylogeny suggests that the crown group of Piscicolidae was originated in the Early Cretaceous. This primarily marine family shares at least five independent colonization events into freshwater environments.

New freshwater mussel taxa discoveries clarify biogeographic division of Southeast Asia.

While a growing body of modern phylogenetic research reveals that the Western Indochina represents a separate biogeographic subregion having a largely endemic freshwater fauna, the boundaries of this subregion are still unclear. We use freshwater mussels (Unionidae) as a model to reconstruct spatial patterns of freshwater biogeographic divides throughout Asia. Here, we present an updated freshwater biogeographic division of mainland Southeast Asia and describe 12 species and 4 genera of freshwater mussels new to science. We show that the Isthmus of Kra represents a significant southern biogeographic barrier between freshwater mussel faunas of the Western Indochina and Sundaland subregions, while the Indian and Western Indochina subregions are separated by the Naga Hills, Chin Hills, and Rakhine Yoma mountain ranges. Our findings highlight that the freshwater bivalve fauna of Southeast Asia primarily originated within three evolutionary hotspots (Western Indochina, Sundaland, and East Asian) supplemented by ancient immigrants from the Indian Subcontinent.

Integrative taxonomy, biogeography and conservation of freshwater mussels (Unionidae) in Russia.

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 and assess the conservation status for each species. This fauna comprises 16 native species from 11 genera and 4 tribes: Anodonta, Pseudanodonta (Anodontini); Amuranodonta, Beringiana, Buldowskia, Cristaria, Sinanodonta (Cristariini); Middendorffinaia, Nodularia, Unio (Unionini); and Lanceolaria (Lanceolariini). No country-level endemic species are known in Russia, except for Buldowskia suifunica that may also occur in China. Sinanodonta woodiana, a non-native species, was introduced from China. Russia comprises the northern parts of Western and Eastern Palearctic subregions. The first subregion with six species encompasses a huge area from the western boundary of Russia to the Lena Basin in Siberia. The second subregion with 10 species covers the Amur Basin, rivers east of the Lena Basin, coastal basins of the Japan Sea, and the North Pacific Islands. The fauna of Russia primarily includes widespread generalist species that are here considered Least Concern (LC). However, Buldowskia suifunica and Sinanodonta lauta have restricted distributions and are assessed here as Vulnerable (VU) and Endangered (EN), respectively.

Freshwater mussels (Bivalvia: Unionidae) from the rising sun (Far East Asia): phylogeny, systematics, and distribution.

Freshwater mussels (Bivalvia: Unionidae) is a diverse family with around 700 species being widespread in the Northern Hemisphere and Africa. These animals fulfill key ecological functions and provide important services to humans. Unfortunately, populations have declined dramatically over the last century, rendering Unionidae one of the world's most imperiled taxonomic groups. In Far East Asia (comprising Japan, Korea, and Eastern Russia), conservation actions have been hindered by a lack of basic information on the number, identity, distribution and phylogenetic relationships of species. Available knowledge is restricted to studies on national and sub-national levels. The present study aims to resolve the diversity, biogeography and evolutionary relationships of the Far East Asian Unionidae in a globally comprehensive phylogenetic and systematic context. We reassessed the systematics of all Unionidae species in the region, including newly collected specimens from across Japan, South Korea, and Russia, based on molecular (including molecular species delineation and a COI + 28S phylogeny) and comparative morphological analyses. Biogeographical patterns were then assessed based on available species distribution data from the authors and previous reference works. We revealed that Unionidae species richness in Far East Asia is 30% higher than previously assumed, counting 43 species (41 native + 2 alien) within two Unionidae subfamilies, the Unioninae (32 + 1) and Gonideinae (9 + 1). Four of these species are new to science, i.e. Beringiana gosannensissp. nov., Beringiana fukuharaisp. nov., Buldowskia kamiyaisp. nov., and Koreosolenaia sitgyensisgen. & sp. nov. We also propose a replacement name for Nodularia sinulata, i.e. Nodularia breviconchanom. nov. and describe a new tribe (Middendorffinaiini tribe nov.) within the Unioninae subfamily. Biogeographical patterns indicate that this fauna is related to that from China south to Vietnam until the Mekong River basin. The Japanese islands of Honshu, Shikoku, Kyushu, Hokkaido, and the Korean Peninsula were identified as areas of particularly high conservation value, owing to high rates of endemism, diversity and habitat loss. The genetically unique species within the genera Amuranodonta, Obovalis, Koreosolenaiagen. nov., and Middendorffinaia are of high conservation concern.

Freshwater mussels house a diverse mussel-associated leech assemblage.

Freshwater mussels (Unionida) are one of the most imperiled animal groups worldwide, revealing the fastest rates of extinction. Habitat degradation, river pollution and climate change are the primary causes of global decline. However, biological threats for freshwater mussels are still poorly known. Here, we describe a diverse ecological group of leeches (Hirudinea: Glossiphoniidae) inhabiting the mantle cavity of freshwater mussels. So far, examples of mussel-associated leech species are recorded from East Asia, Southeast Asia, India and Nepal, Africa, and North America. This group comprises a dozen glossiphoniid species with a hidden life style inside the mantle cavity of their hosts largely overlooked by researchers. We show that the association with freshwater mussels evolved independently in three leech clades, i.e. Batracobdelloides, Hemiclepsis, and Placobdella, at least since the Miocene. Seven mussel-associated leech species and two additional free-living taxa are described here as new to science.

Pollinators on the polar edge of the Ecumene: taxonomy, phylogeography, and ecology of bumble bees from Novaya Zemlya.

The High Arctic bumble bee fauna is rather poorly known, while a growing body of recent molecular research indicates that several Arctic species may represent endemic lineages with restricted ranges. Such local endemics are in need of special conservation efforts because of the increasing anthropogenic pressure and climate changes. Here, we re-examine the taxonomic and biogeographic affinities of bumble bees from Novaya Zemlya using historical samples and recently collected materials (1895-1925 vs. 2015-2017). Three bumble bee species inhabit the Yuzhny (Southern) Island and the southern edge of Severny (Northern) Island of this archipelago: Bombusglacialis Friese, 1902, B.hyperboreus Schönherr, 1809, and B.pyrrhopygus Friese, 1902. Bombusglacialis shares three unique COI haplotypes that may indicate its long-term (pre-glacial) persistence on Novaya Zemlya. In contrast, Bombushyperboreus and B.pyrrhopygus share a rather low molecular divergence from mainland populations, with the same or closely related haplotypes as those from Arctic Siberia and Norway. A brief re-description of Bombuspyrrhopygus based on the newly collected topotypes is presented. Habitats, foraging plants and life cycles of bumble bees on Novaya Zemlya are characterized, and possible causes of extremely low bumble bee abundance on the archipelago are discussed. The species-poor bumble bee fauna of Novaya Zemlya is compared with those in other areas throughout the Arctic. The mean bumble bee species richness on the Arctic Ocean islands is three times lower than that in the mainland Arctic areas (3.1 vs. 8.6 species per local fauna, respectively). General linear models (GLMs) indicate that this difference can be explained by specific environmental conditions of insular areas. Our findings highlight that the insularity is a significant factor sharply decreasing species richness in bumble bee assemblages on the Arctic Ocean archipelagoes through colder climate (lower summer temperatures), prevalence of harsh Arctic tundra landscapes with poor foraging resources, and in isolation from the mainland.

Eight new freshwater mussels (Unionidae) from tropical Asia.

Freshwater mussels are sensitive to habitat and water quality, revealing the fastest rates of human-mediated global extinction among aquatic animals. These animals are especially diverse in tropical Asia, the faunas of which are characterized by high levels of endemism. Here we describe four new species and four new subspecies of freshwater mussels from Myanmar. Leoparreysia whitteni sp. nov., the smallest representative of this genus, was discovered from the Ayeyarwady and Chindwin rivers. Radiatula myitthanensis sp. nov. and R. chindwinensis sp. nov. were recorded from the Chindwin Basin, and R. mouhoti haungthayawensis ssp. nov. has been discovered from the Haungthayaw River. Indochinella pugio has been revised with a description of three subspecies: I. pugio viridissima ssp. nov. from the Sittaung, Bilin and Bago rivers, I. pugio daweiensis ssp. nov. from the Dawei River, and I. pugio paradoxa ssp. nov. from the Haungthayaw River. Yaukthwa elongatula sp. nov., a peculiar species, conchologically resembling representatives of the genus Solenaia (Gonideinae) with ultra-elongated shell was found in the Chindwin Basin. Our records highlight that tropical Asia harbors numerous, but still overlooked local endemic lineages of freshwater bivalves, which may be on the brink of extinction due to the high anthropogenic and climate change impacts.

Fish hosts, glochidia features and life cycle of the endemic freshwater pearl mussel Margaritifera dahurica from the Amur Basin.

Margaritiferidae is a small freshwater bivalve family with 16 species. In spite of a small number of taxa and long-term history of research, several gaps in our knowledge on the freshwater pearl mussels still exist. Here we present the discovery of host fishes for Margaritifera dahurica, i.e. Lower Amur grayling, sharp-snouted lenok, and blunt-snouted lenok. The host fishes were studied in rivers of the Ussuri Basin. The identification of glochidia and fish hosts was confirmed by DNA analysis. The life cycle of M. dahurica and its glochidia are described for the first time. The SEM study of glochidia revealed that the rounded, unhooked Margaritifera dahurica larvae are similar to those of the other Margaritiferidae. Margaritifera dahurica is a tachytictic breeder, the larvae of which attach to fish gills during the Late August - September and finish the metamorphosis in June. Ancestral host reconstruction and a review of the salmonid - pearl mussel coevolution suggest that the ancestral host of the Margaritiferidae was a non-salmonid fish, while that of the genus Margaritifera most likely was an early salmonid species or their stem lineage. The overfishing of lenoks and graylings appears to be the most significant threat for this rare mussel species.

A new genus and two new species of freshwater mussels (Unionidae) from western Indochina.

The systematics of Oriental freshwater mussels (Bivalvia: Unionidae) is poorly known. Here, we present an integrative revision of the genus Trapezoideus Simpson, 1900 to further understanding of freshwater mussel diversity in the region. We demonstrate that Trapezoideus as currently circumscribed is non-monophyletic, with its former species belonging to six other genera, one of which is new to science and described here. We recognize Trapezoideus as a monotypic genus, comprised of the type species, T. foliaceus. Trapezoideus comptus, T. misellus, T. pallegoixi, and T. peninsularis are transferred to the genus Contradens, T. subclathratus is moved to Indonaia, and T. theca is transferred to Lamellidens. Trapezoideus prashadi is found to be a junior synonym of Arcidopsis footei. Trapezoideus dallianus, T. nesemanni, T. panhai, T. peguensis, and two species new to science are placed in Yaukthwa gen. nov. This genus appears to be endemic of the Western Indochina Subregion. The two new species, Yaukthwa paiensis sp. nov. and Y. inlenensis sp. nov., are both endemic to the Salween River basin. Our results highlight that Southeast Asia is a species-rich freshwater mussel diversity hotspot with numerous local endemic species, which are in need of special conservation efforts.

Discovery of Novaculina myanmarensis sp. nov. (Bivalvia: Pharidae: Pharellinae) closes the freshwater razor clams range disjunction in Southeast Asia.

The razor clam genus Novaculina represents an example of a marine-derived, secondary freshwater group. It was thought to comprise three species: N. gangetica (Ganges and smaller basins in Bangladesh and northwestern Myanmar), N. siamensis (Bang Pakong and Pasak rivers in Thailand and Mekong River in Vietnam), and N. chinensis (lower Yangtze River, China). Here we describe Novaculina myanmarensis sp. nov., an additional species from the Ayeyarwady and Salween basins representing a divergent lineage that appears to be sister to N. gangetica. This new record closes a Novaculina range disjunction between northwestern Myanmar and Thailand. The populations of this novel species share a shallow molecular divergence from each other indicating potential dispersal events between the two distant freshwater basins during the Late Pleistocene. Our ancestral area modeling suggests that the MRCA of Novaculina crown group was a salt-tolerant freshwater species. The recent Novaculina species most likely originated via allopatric speciation. Our findings highlight that generalist estuarine species could have played the role as a source for bivalve expansions into freshwater and that western Indochina is a separate biogeographic subregion, which is clearly distinct from India. A new synonymy is proposed as follows: Pharellinae Stoliczka, 1870 = Novaculininae Ghosh, 1920 syn. nov.

An example of a possible leech-bryozoan association in freshwater.

Associations of various invertebrate species with bryozoans and sponges are a well-known marine phenomenon but such epizooic communities are far less diverse in freshwater environments. Here an occurrence of numerous leeches Alboglossiphoniacf.papillosa (Braun, 1805), in interstitial spaces between zooids of a colony of the freshwater bryozoan species Plumatellaaff.fungosa (Pallas, 1768) in Eastern Siberia is described. To the best of our knowledge, this record appears to be the first known example of a leech-bryozoan association, although such relationships deserve further research.