<i>Ladislavella tumrokensis</i>: The first molecular evidence of a Nearctic clade of lymnaeid snails inhabiting Eurasia

<p>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 <i>Stagnicola elodes</i> group from Canada and the USA. Two lymnaeid species, <i>Lymnaea</i> (<i>Orientogalba</i>) <i>tumrokensis</i> Kruglov and Starobogatov, <a href="#cit0033" target="_blank">1985</a> and <i>Lymnaea</i> (<i>Polyrhytis</i>) <i>kurenkovi</i> Kruglov and Starobogatov, <a href="#cit0034" target="_blank">1989</a>, were described from the Tumrok geothermal locality, but actually they are morphological variations of a single taxon of subspecies rank re-classified here as <i>Ladislavella catascopium tumrokensis</i>. 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.</p

Range map of <i>Margaritifera middendorffi</i> (Rosén, 1926) and <i>Margaritifera togakushiensis</i> (Kondo and Kobayashi, 2005).

<p>Circles—<i>M</i>. <i>middendorffi</i> locations, squares—<i>M</i>. <i>togakushiensis</i> locations. Green circles and squares are representing recent viable populations (observed since 2000), white circles—old records (until 2001), yellow squares—records without exact dates. Grey areas indicate approximate modern species ranges (showing only the large river systems). Species locality numbers on the map correspond to numbers in <a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0122408#pone.0122408.s003" target="_blank">S3 Table</a>.</p

Map of location of the field study areas.

<p>1—Kamchatka Peninsula (2012, I. Bolotov, Y. Bespalaya, I. Vikhrev, M. Gofarov), 2—Central Sakhalin (2012, same team), 3—southern Sakhalin (2011–2012, same team & Y. Kolosova, O. Aksenova), 4—Kunashir Island (2011, same team & Y. Kolosova, O. Aksenova), 5—Primorye (2012, same team), 6—Transbaikalia (2004–2011, O.K. Klishko). Data on the studied river sites are presented in <a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0122408#pone.0122408.s001" target="_blank">S1</a>–<a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0122408#pone.0122408.s004" target="_blank">S4</a> Tables.</p

Range map of <i>Margaritifera dahurica</i> (Middendorff, 1850).

<p>Green circles are representing recent viable populations (observed since 2000), white circles—old records (until 2000). Question mark is indicated an uncertain record from the Langry River [<a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0122408#pone.0122408.ref069" target="_blank">69</a>], [<a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0122408#pone.0122408.ref026" target="_blank">26</a>]. Grey areas are indicated an approximate modern species range (it is shown only for the large river systems). Species locality numbers on the map correspond to numbers in <a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0122408#pone.0122408.s002" target="_blank">S2 Table</a>.</p

Shells of <i>Margaritifera dahurica</i> (Middendorff, 1850).

<p>A—Lectotype (ZISP: no. 7a). B—specimen from Komarovka River, Razdolnaya River basin, Primorye. Photos by I.V. Vikhrev. Scale bar—2 cm.</p

Shells of <i>Margaritifera togakushiensis</i> (Kondo and Kobayashi, 2005) and <i>Margaritifera middendorffi</i> (Rosén, 1926).

<p>A—Holotype of <i>M</i>. <i>togakushiensis</i> [<a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0122408#pone.0122408.ref018" target="_blank">18</a>]: 137, figs <a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0122408#pone.0122408.g005" target="_blank">5</a>–<a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0122408#pone.0122408.g008" target="_blank">8</a>. B—Lectotype of <i>M</i>. <i>middendorffi</i> (ZISP: no. 6). Photo by I. V. Vikhrev. Scale bar—2 cm.</p

Shells of <i>Margaritifera laevis</i> (Haas, 1910).

<p>A—Sennaya River, Kunashir Island. B—Tym’ River, Sakhalin Island. Photos by I. V. Vikhrev. Scale bar—2 cm.</p

General plan of soft tissue of three Far Eastern margaritiferid species.

<p>A—<i>Margaritifera dahurica</i> (Middendorff, 1850). B—<i>M</i>. <i>laevis</i> (Haas, 1910). C—<i>M</i>. <i>middendorffi</i> (Rosén, 1926). Scale bar—1 cm.</p

Typical habitats of the Eastern Asian margaritiferid species.

<p>A—<i>Margaritifera dahurica</i> (Middendorff, 1850): the Ilystaya River, Primorsky Kray. B—<i>M</i>. <i>middendorffi</i> (Rosén, 1926): the Nachilova River, Kamchatka. C—<i>M</i>. <i>middendorffi</i> (Rosén, 1926) & <i>M</i>. <i>laevis</i> (Haas, 1910): the Golovnina River, Kunashir Island. Photos by Y.V. Bespalaya, Y.S. Kolosova & I.V. Vikhrev.</p

Bayesian phylogeny of <i>Margaritifera</i> spp. haplotypes.

<p>The scale bar indicates the branch length. Asterisks: Posterior probabilities ≥0.95; other significant support node values are mentioned in the figure. For detailed locality and specimen data for analyzed haplotypes, see the supplementary materials (<a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0122408#pone.0122408.s001" target="_blank">S1</a> and <a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0122408#pone.0122408.s002" target="_blank">S2</a> Tables). A—The 18S rDNA gene dataset. B—The COI gene dataset.</p