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.

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.

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.

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.

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.

DNA analysis of a non-native lineage of Sinanodonta woodiana species complex (Bivalvia: Unionidae) from Middle Asia supports the Chinese origin of the European invaders.

The Sinanodonta woodiana species complex includes several cryptic species-level phylogenetic lineages, the taxonomic placement of which is unclear. Here, we present the results of molecular analyses of a Sinanodonta sample from Middle Asia (Uzbekistan). The COI haplotype of the Uzbekistan lineage is similar to those of invasive Sinanodonta populations from Europe, Russia and Myanmar. We show that the non-native Sinanodonta flocks in Europe and Middle Asia originated from a single source population in China. Our results reveal that the Sinanodonta woodiana species complex comprises at least six distinct biological species: S. cf. gibba (=temperate invasive lineage), S. cf. woodiana (=tropical invasive lineage), S. schrenkii (=S. amurensis), S. ovata, S. jourdyi, and S. lucida, and two phylogenetic lineages with uncertain taxonomic names.

Discovery of a silicate rock-boring organism and macrobioerosion in fresh water.

Macrobioerosion is a common process in marine ecosystems. Many types of rock-boring organisms break down hard substrates, particularly carbonate rocks and calcareous structures such as dead corals and shells. In paleontology, the presence of rocks with boreholes and fossil macroboring assemblage members is one of the primary diagnostic features of shallow marine paleo-environments. Here we describe a silicate rock-boring organism and an associated community in submerged siltstone rock outcrops in Kaladan River, Myanmar. The rock-boring mussel Lignopholas fluminalis is a close relative of the marine piddocks, and its borings belong to the ichnospecies Gastrochaenolites anauchen. The neotectonic uplift of the area leading to gradual decrease of the sea level with subsequent shift from estuarine to freshwater environment was the most likely driver for the origin of this community. Our findings highlight that rocks with macroborings are not an exclusive indicator of marine paleo-ecosystems, but may also reflect freshwater habitats.

Species Richness, Molecular Taxonomy and Biogeography of the Radicine Pond Snails (Gastropoda: Lymnaeidae) in the Old World.

The radicine pond snails represent a species-rich and widely distributed group, many species of which are key vectors of human and animal trematodoses. Here we clarify the taxonomy, distribution and evolutionary biogeography of the radicine lymnaeids in the Old World based on the most comprehensive multi-locus molecular dataset sampled to date. We show that the subfamily Amphipepleinae is monophyletic and contains at least ten genus-level clades: Radix Montfort, 1810, Ampullaceana Servain, 1881, Peregriana Servain, 1881, Tibetoradix Bolotov, Vinarski & Aksenova gen. nov., Kamtschaticana Kruglov & Starobogatov, 1984, Orientogalba Kruglov & Starobogatov, 1985, Cerasina Kobelt, 1881, Myxas G. B. Sowerby I, 1822, Bullastra Bergh, 1901, and Austropeplea Cotton, 1942. With respect to our phylogeny, species-delimitation model and morphological data, the Old World fauna includes 35 biological species of radicines. Tibet and Eastern Europe harbor the richest faunas, while East Asia and Africa appear to be the most species-poor areas. The radicine clade could have originated near the Cretaceous - Paleocene boundary. The Miocene great lakes in Eurasia seems to be the most important evolutionary hotspots shaping spatial patterns of recent species richness. Finally, we present the first DNA barcode reference library for the reliable molecular identification of species within this group.

Widespread continental mtDNA lineages prevail in the bumblebee fauna of Iceland.

Origins of the fauna in Iceland is controversial, although the majority of modern research supports the postglacial colonization of this island by terrestrial invertebrates rather than their long-term survival in glacial refugia. In this study, we use three bumblebee species as a model to test the hypothesis regarding possible cryptic refugia in Iceland and to evaluate a putative origin of recently introduced taxa. Bombus jonellus is thought to be a possible native Icelandic lineage, whereas B. lucorum and B. hortorum were evidently introduced in the second half of the 20th century. These phylogeographic analyses reveal that the Icelandic Bombus jonellus shares two COI lineages, one of which also occurs in populations on the British Isles and in mainland Europe, but a second lineage (BJ-02) has not been recorded anywhere. These results indicate that this species may have colonized Iceland two times and that the lineage BJ-02 may reflect a more ancient Late Pleistocene or Early Holocene founder event (e.g., from the British Isles). The Icelandic populations of both Bombus lucorum and B. hortorum share the COI lineages that were recorded as widespread throughout Eurasia, from the European countries across Russia to China and Japan. The findings presented here highlight that the bumblebee fauna of Iceland comprises mainly widespread ubiquitous lineages that arrived via natural or human-mediated dispersal events from the British Isles or the mainland.

A new genus and tribe of freshwater mussel (Unionidae) from Southeast Asia.

The freshwater mussel genus Oxynaia Haas, 1911 is thought to be comprised of two geographically disjunct and morphologically variable species groups but the monophyly of this taxon has yet to be tested in any modern cladistic sense. This generic hypothesis has important systematic and biogeographic implications as Oxynaia is the type genus of the currently recognized tribe Oxynaiini (Parreysiinae) and is one of the few genera thought to cross several biogeographically important barriers in Southeast Asia. Morphological and molecular data clearly demonstrate that Oxynaia is not monophyletic, and the type species and its allies (O. jourdyi group) belong to the Unioninae, and more specifically as members of the genus Nodularia Conrad, 1853. Therefore, neither Oxynaia syn. nov. nor Oxynaiini Starobogatov, 1970 are applicable to the Parreysiinae and in the absence of an available name, Indochinella gen. nov. and Indochinellini trib. nov. are described. Several combinations are proposed as follows: Indochinella pugio (Benson, 1862) gen. et comb. nov., Nodularia jourdyi (Morlet, 1886) comb. res., N. gladiator (Ancey, 1881) comb. res., N. diespiter (Mabille, 1887) comb. res. and N. micheloti (Morlet, 1886) comb. res. Finally, we provide an updated freshwater biogeographic division of Southeast Asia.

Climate Warming as a Possible Trigger of Keystone Mussel Population Decline in Oligotrophic Rivers at the Continental Scale.

The effects of climate change on oligotrophic rivers and their communities are almost unknown, albeit these ecosystems are the primary habitat of the critically endangered freshwater pearl mussel and its host fishes, salmonids. The distribution and abundance of pearl mussels have drastically decreased throughout Europe over the last century, particularly within the southern part of the range, but causes of this wide-scale extinction process are unclear. Here we estimate the effects of climate change on pearl mussels based on historical and recent samples from 50 rivers and 6 countries across Europe. We found that the shell convexity may be considered an indicator of the thermal effects on pearl mussel populations under warming climate because it reflects shifts in summer temperatures and is significantly different in viable and declining populations. Spatial and temporal modeling of the relationship between shell convexity and population status show that global climate change could have accelerated the population decline of pearl mussels over the last 100 years through rapidly decreasing suitable distribution areas. Simulation predicts future warming-induced range reduction, particularly in southern regions. These results highlight the importance of large-scale studies of keystone species, which can underscore the hidden effects of climate warming on freshwater ecosystems.