Ilya V. Vikhrev

Diversity, biogeography and conservation of freshwater mussels (Bivalvia: Unionida) in East and Southeast Asia

Recent research efforts have significantly advanced our knowledge on Asian freshwater mussel (Bivalvia: Unionida) diversity and distribution. Here we provide a modern consensus of the diversity, biogeography and conservation of Unionida in the region comprising East and Southeast Asia (excluding Wallacea) and Asian Russia. A data review confirmed the presence of 228 native and 3 non-native Unionida (98% Unionidae, 2% Margaritiferidae), rendering the region a global hotspot of freshwater mussel diversity. Species richness was highest in China (particularly Yangtze basin) in absolute numbers and Cambodia when correcting for country area, and decreased gradually towards the south and steeply towards the north and east. Six of the seven unionid subfamilies are native to the region, with species richness peaking in Southeast Asia for Rectidentinae, Gonideinae, Parreysiinae and Modellnaiinae, China for Anodontinae and Unioninae, and Asian Russia for Margaritiferidae. Conservation status and data collected after 1980 were not available for 61 and 24% of species, respectively. Dams, deforestation and pollution are likely the major threats to mussels in the region, though data in this respect are scarce. The Philippines, Laos, Indonesia, Myanmar and Malaysia are among the countries with the poorest data availability and urgently require research.

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.

Multi-locus fossil-calibrated phylogeny, biogeography and a subgeneric revision of the Margaritiferidae (Mollusca: Bivalvia: Unionoida).

The taxonomy and biogeographic history of the bivalve family Margaritiferidae are controversial because previous molecular studies did not provide a well-resolved phylogenetic framework for these enigmatic freshwater mussels that have extensive but disjunct distribution in North America, Eurasia and North Africa. In this study, we present a new, fossil-calibrated phylogenetic hypothesis based on five molecular markers (∼4kb of total length) for ten species. Our results indicate that all recent margaritiferids are in the single genus, Margaritifera Schumacher, 1816. Additionally, we identified three relatively well-supported phylogenetic clades that are valid subgenera, i.e., Margaritifera s. str. (Holarctic), Margaritanopsis (=Cumberlandia) (southeast North America-southeast Asia disjunct) and Pseudunio (Mediterranean). We suggest that the crown lineage of the Margaritiferidae most likely originated in the Cretaceous (mean age 93Ma, 95% CI 66-126Ma). The combined results of ancestral area reconstructions based on the three different approaches (S-DIVA, DEC and S-DEC) showed that ancient vicariance events could have played an important role in speciation within the family. The rates of mitochondrial evolution of margaritiferids are notably slow, which may be associated with their longevity, long generation time and low metabolic rates. Our findings highlight the complex biogeographic history of the Margaritiferidae as an intermixing of ancient vicariance and dispersal events, which were most likely associated with some inland barriers, continental movements and a sea level dynamic.

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.

Ecology and conservation of the endangered Indochinese freshwater pearl mussel, Margaritifera laosensis (Lea, 1863) in the Nam Pe and Nam Long rivers, Northern Laos

In this paper we present the first ecological data of Indochinese freshwater pearl mussel, Margaritifera laosensis populations. We also provide a comparative study of the ecology of this tropical species with populations of other Margaritiferidae. We conducted surveys in ten tributaries of the River Nam Ou (Middle Mekong Drainage, Northern Laos). Reproductively viable populations were found only in the Nam Long and Nam Pe rivers, which are two of the only three known viable populations of this species in the world. The habitats of M. laosensis include mountainous oligotrophic rivers with circumneutral pH. Optimal mesohabitats are riffles and runs with a median depth of 0.2 m and median current velocity of 0.3 ms−1. Pearl mussels were more common in gravel and fine gravel riverbed substrates. Surveys revealed 252 specimens, but only 78 (31.0%) were alive. The largest mussels observed were 110 mm in length and only 11–12 years of age. The presence of smaller-sized mussels indicates recent recruitment in both populations. The most significant threats to M. laosensis populations are harvest by local people and land development in the River Nam Ou Basin.

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.

Influence of Historical Exploitation and Recovery of Biological Resources on Contemporary Status of Margaritifera margaritifera L. and Salmo salar L. Populations in Northwestern Russia

The influence of the historical exploitiation and recovery of biological resources on populations of freshwater pearl mussel and salmon was studied in the Kozha river (Onega River Basin). The freshwater pearl mussel was characterized by a low population density (0.09 ind./m2, on the average) and number of recruits. The role of the anthropogenic factor in the population is negligible; the effect of the pearl fishing is not observed. The dramatic decrease in the number of Atlantic salmon in the Onega River Basin in the late 20th century is a major reason for the slowed down reproduction of the freshwater pearl mussel. The second factor is the reconstruction of the Onega salmon hatchery in 1984, which led to a decrease in the juvenile salmon population density below the critical level on spawning and nursery grounds. As a result, mollusc reproduction has stopped. An artificial increase in the number of recruits must be developed in order to preserve the salmon population in the Onega River Basin. It is also necessary to forbid salmon fishing, including fishing migratory paths (with the exception of trapping for hatchery needs and traditional harvesting performed by locals in historical settlements).

An integrative approach underscores the taxonomic status of Lamellidens exolescens, a freshwater mussel from the Oriental tropics (Bivalvia: Unionidae)

The Oriental Region harbours the second richest fauna of freshwater bivalves in the world, including many endangered endemic taxa. However, the Oriental fauna of the Unionidae have been very poorly studied using an integrative taxonomic approach, which may provide reasonable revisions of complicated (cryptic) taxa based on morphological, molecular, biogeographic and ecological evidence. Here, we present the first example of an integrative taxonomic revision concerning the status of Unio exolescens Gould (1843), a nominal mussel taxon that was accepted as a valid species within the genus Trapezoideus Simpson (1900). Currently, Trapezoideus exolescens is considered the type of the genus as far as the originally designated type species, U. foliaceus Gould (1843), was considered to be a synonym of T. exolescens. Using nucleotide sequences obtained from mitochondrial (COI and 16S rRNA) and nuclear (28S rDNA) genes, we found that the topotypes of Unio exolescens Gould (1843) cluster together with representatives of another mussel genus, Lamellidens Simpson (1900). Based on these results and on morphological data, we transfer Unio exolescens Gould (1843) from Trapezoideus to Lamellidens and propose Lamellidens exolescens (Gould, 1843) comb. nov. In addition, we revisited the status of Unio foliaceus Gould (1843) as a valid species and the type of the genus Trapezoideus based on the morphological study of the type specimen, although a question concerning the true position of this taxon is still open because its molecular sequences are not available. Our findings highlight that an integrative taxonomic approach is an important tool, particularly when dealing with such species-rich Unionidae fauna as those of the Oriental Realm.

Historical geography of pearl fishing in rivers of the Southern White Sea Region (Arkhangelsk Oblast)

The article provides an overview of literary and archival materials on the historical geography of pearl fishing, as well as a list of rivers of the Southern White Sea Region, where in the 17th–20th centuries pearls were mined. The most significant resources of pearls have been concentrated in the rivers Solza (with the tributary Kazanka) and Kozha (with the tributary Syvtuga) where there was regular and profitable trade of pearls for a long time. At present, pearl mussel populations have been preserved in these rivers, sometimes with a fairly high density. The volumes and frequency of pearl fishing can be indicators of shellfish abundance.