A taxonomic revision of two local endemic Radix spp. (Gastropoda: Lymnaeidae) from Khodutka geothermal area, Kamchatka, Russian Far East

ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2014 Magnolia Press

Zootaxa 3869 (5): 585–593

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http://dx.doi.org/10.11646/zootaxa.3869.5.9

http://zoobank.org/urn:lsid:zoobank.org:pub:58477926-D081-4692-8378-53AD8DEBF8FF

A taxonomic revision of two local endemic Radix spp. (Gastropoda: Lymnaeidae)

from Khodutka geothermal area, Kamchatka, Russian Far East

IVAN BOLOTOV1, 3, YULIA BESPALAYA1, OLGA AKSENOVA1, ANDREY AKSENOV1, 2,

NIKITA BOLOTOV1, 2, MIKHAIL GOFAROV1, 2, ALEXANDER KONDAKOV1,

INGA PALTSER1 & ILYA VIKHREV1

1Institute of Ecological Problems of the North, the Ural Branch of Russian Academy of Sciences, Severnaya Dvina Emb. 23, 163000

Arkhangelsk, Russian Federation2Northern (Arctic) Federal University, Severnaya Dvina Emb. 17, 163002 Arkhangelsk, Russian Federation3Corresponding author. E-mail: [email protected]

Introduction

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.

During field expeditions in 2010–2013, lymnaeid snails were collected from eight localities on the Russian Far

East, including the Khodutka geothermal area. Sampled specimens were preserved in 96 % ethanol. Materials are

deposited in the Biological Museum of the Institute of Ecological Problems of the North (INEP), the Ural Branch

of the Russian Academy of Sciences, Arkhangelsk, Russia. Species identification was performed using shell

morphology, soft body anatomy and molecular data. Anatomical and morphological details were photographed

using a stereomicroscope (Leica M165C, Leica Microsystems, Germany). The type specimens of Lymnaea (Radix)

hadutkae Kruglov & Starobogatov, 1989 and L. (R.) thermokamtschatica Kruglov & Starobogatov, 1989 were

studied in the collection of the Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russian

Federation (ZIN).

Molecular analysis was performed in the Institute of Ecological Problems of the North of the Ural Branch of

Russian Academy of Sciences (purification and PCR) and the Engelhardt Institute of Molecular Biology of Russian

Academy of Sciences (sequencing on ABI PRISM 3730). We analyzed Radix spp. samples from Khodutka area, as

Accepted by J. Nekola: 22 Sep. 2014; published: 3 Oct. 2014 585

well as from other Far Eastern localities (Appendix 1). We extracted genomic DNA from individual snails using

the DiatomTM DNA Prep 200 reagents kit (IsoGene Lab., Moscow, Russia). In the present study, we used data

inferred from the mitochondrial cytochrome oxidase subunit I (COI) gene, because DNA barcoding provides an

effective method for species-level identification and is widely accepted for an integrative taxonomy of Radix spp.

(Albrecht et al. 2008; Glöer & Pešić 2008). However, the molecular analyses of the nuclear and mitochondrial

markers are often incongruent (Schniebs et al. 2012) which reveals a limited application of a single mtDNA

marker, which may lead to over-estimation of species diversity. A total 657 bp of the COI gene were amplified and

sequenced using primers LCO1490 and HCO 2198 (Folmer et al. 1994). Additional sequences were obtained from

NCBI’s Genbank (Appendix 2). The alignment of COI sequences was performed using the CrustalW algorithm

implemented in MEGA6 (Tamura et al. 2013). For phylogenetic analyses, each sequence was trimmed, leaving a

588 bp-long COI fragment, according to a minimal length of a sequence in the aligned dataset. The analysis was

performed using 22 unique COI haplotypes, including Galba truncatula and Lymnaea palustris as outgroups. The

sequence dataset was collapsed into haplotypes using an online fasta sequence toolbox (FaBox v1.41, http://users-

birc.au.dk/biopv/php/fabox). The best model of sequence evolution is HKY+G as suggested with respect to the

corrected Akaike Information Criterion of MEGA6 (Tamura et al. 2013). Phylogenetic relationships were

reconstructed based on Bayesian inference implemented in MrBayes version 3.2.2 (Ronquist et al. 2012) using the

following parameters: mcmc ngen = 10,000,000, nchains = 4, samplefreq = 100, temp = 0.1; 25% of the sampled

trees were discarded as burn-in (pre-convergence part). Convergence of the MCMC chains to the stationary

distribution was checked visually based on the plotted posterior estimates using a MCMC trace analysis tool

(Tracer v1.6, http://beast.bio.ed.ac.uk/software/tracer). The effective sample size (ESS) value for each parameter

sampled from the MCMC analysis was always recorded >300. The combined set of trees showed a smooth

frequency plot. We conducted two independent runs of a Bayesian analysis using each outgroup in order to test

their influence on tree topology.

The diagnoses of two endemics from the Khodutka area are not clear, because all used diagnostic features

show a relatively high interspecific variability within the genus Radix (Glöer & Pešić 2008). The difference

between values of an excess of the last whorl over the shell aperture (Kruglov & Starobogatov 1989; Starobogatov

et al. 2004; Kruglov 2005) is very small and cannot be used as a diagnostic feature because of a high intraspecific

shell variability among R. auricularia populations due to environmental conditions (Stift et al. 2004). Studied

specimens from the Khodutka area belong to R. auricularia based on morphological and anatomical characters

(Table 1 and Figs. 1–3) and molecular data (Appendix 1 and Fig. 5).

Bayesian analysis reveals that R. auricularia is a monophyletic taxon across Eurasia (Bayesian posterior

probabilities (BPP) 1.00), which includes several highly divergent mitochondrial clades (Fig. 5). Two haplotypes

from the Khodutka geothermal area belong to a separate clade, which is well supported (BPP 0.96), and includes

haplotypes from other areas of the Russian Far East, as well as from the Middle East (Tajikistan). The topology of

R. auricularia clade and position of the Khodutka haplotypes remain stable and robust regardless of outgroup. The

mean COI p-distance among haplotypes that belong to this clade, calculated using MEGA6 software (Tamura et al.

2013), is 1.1±0.3 % (0.2–2.0 %).

Taxonomic account

Genus Radix Montfort, 1810

Radix auricularia (Linnaeus, 1758)

(Figs. 1–7, Table 1)

Lymnaea (Radix) hadutkae Kruglov & Starobogatov, 1989 syn. n.: 22, figs. 1.12 & 2.7; Kruglov & Starobogatov, 1993: 85, fig.

11D

Lymnaea (Radix) thermokamtschatica Kruglov & Starobogatov, 1989 syn. n.: 22, figs. 1.9 & 2.9; Kruglov & Starobogatov,

1993: 85, fig. 11E

BOLOTOV ET AL.586 · Zootaxa 3869 (5) © 2014 Magnolia Press

FIGURES 1–7. Radix auricularia (Linnaeus, 1758) from Russian Far East. Figs. 1-4. Shell and proximal part of the female

genitalia. Scale bar 2 mm. Fig. 1. Holotype of Lymnaea (R.) hadutkae Kruglov & Starobogatov, 1989 syn. n. (shell photo: M.V.

Vinarski; genitalia redrawn from the protologue, Fig. 2.7). Fig. 2. Holotype of L. (R.) thermokamtschatica Kruglov &

Starobogatov, 1989 syn. n. (shell photo: M.V. Vinarski; genitalia redrawn from the protologue, Fig. 2.9). Fig. 3. Newly

collected specimen from the Khodutka geothermal area, Kamchatka (photos: O.V. Aksenova). Fig. 4. Specimen from

Malkinskie hot springs, Kamchatka (photos: O.V. Aksenova). Fig. 5. Bayesian phylogram of haplotypes based on

mitochondrial COI gene dataset (Appendixes 1 & 2). The scale bar indicates the branch length. Asterisks: Posterior

probabilities ≥0.95; other significant node support values are mentioned in the figure. Haplotypes exclusively from NCBI’s

Genbank are marked with circumflex accents (˄). The well-supported clade with two haplotypes from the Khodutka

geothermal area is highlighted in blue. Fig. 6. General view of the Khodutka warm lake. Fig. 7. Map of observed localities on

the Russian Far East. The red circles represent species records. Locality codes are given in Appendix 1. The digital elevation

model and other layers of the map were added from Esri Data & Maps 10 dataset (Map: M.Yu. Gofarov).

Zootaxa 3869 (5) © 2014 Magnolia Press · 587RADIX SPP. FROM KHODUTKA GEOTHERMAL AREA

Material examined. Holotype of Lymnaea (Radix) hadutkae (ZIN), Kamchatka: “Khodutkinskiy hot spring,

gradient t° = 15–41°[C], opt. ~ 25–27°[C], 8.ix.1979, N.N. Khmeleva leg.” (no. 510–1982). Holotype of Lymnaea

(Radix) thermokamtschatica (ZIN), Kamchatka: “Khodutkinskiy hot spring, gradient t° = 15–41°[C], opt. ~

25–27°[C], 8.ix.1979, N.N. Khmeleva leg.” (no. 510–1982). Topotypes (INEP), 31 specimens, Kamchatka:

Khodutka geothermal area, warm lake and warm river source, 6th June 2013, Aksenova, Bespalaya & Bolotov leg.

(INEP). Kamchatka: 6 specimens, Paratunskie hot springs, 8th June 2013, Aksenova, Bespalaya & Bolotov leg.; 7

specimens, Malkinskie hot springs, 3rd July 2012, Bespalaya, Bolotov & Vikhrev leg.; 4 specimens, Plotnikova

River (Bolshaya River Basin), 3rd July 2012, Bespalaya, Bolotov & Vikhrev leg. (INEP). Kurile Archipelago: 9

specimens, Kunashir Island, Aliger Lake, 27th July 2011, Bespalaya & Aksenova leg. (INEP). Sakhalin Island: 2

specimens, Tym River, 14th June 2012, Bespalaya, Bolotov & Vikhrev leg. (INEP). Primorye: 3 specimens, a small

pond near Razdolnaya River, 25th June 2012, Bespalaya, Bolotov & Vikhrev leg; 4 specimens, Ussuri River (Amur

River Basin), 20th June 2012, Bespalaya, Bolotov & Vikhrev leg. (INEP). Locality coordinates are given in

Appendix 1.

TABLE 1. The comparison of type specimens of endemic Lymnaea (Radix) species from Khodutka geothermal area with

R. auricularia based on morphological and anatomical distinguishing features (Glöer & Pešić 2008)

Description. Specimens from the Khodutka geothermal area are very small with shell height 7.0–14.6 mm,

width 4.0–9.5 mm. Shell is glossy, dark-yellow or light brownish, elongated, with 4 whorls enlarging slowly (Figs.

1–3). The spire is long, widely conical, in adult shells often destroyed on the top (likely due to the geothermal

water influence). The aperture is ear-shaped, relatively narrow with an angle of 90° or >90° on the top. The inner

lip is folded at the columella, with weakly-sinuated fold. The foot and dorsal part of the preputium bear “freckles”.

The bursa copulatrix is oval or rounded; the bursa duct is thin and very long or long (Figs. 1–3 and Table 1).

According to morphological and anatomical data, specimens from other localities of the Russian Far East are

similar to those found from the Khodutka area (Fig. 4). However, specimens from non-geothermal localities

(Plotnikova River, Aliger Lake, a pond near Razdolnaya River and Ussuri River) are larger with shell height up to

25.7 mm, width up to 20.9 mm, and had a more compact shell shape with a wide aperture.

Remarks. Both of the endemics from the Khodutka geothermal area (Fig. 6) are conspecific to R. auricularia,

based on morphological, anatomical and molecular studies (Table 1 and Figs. 1–5). The same elongated shells with

narrow apertures were reported for several R. auricularia populations from the Baikal Lake area (Stift et al. 2004).

According to our data, R. auricularia is widespread on the Russian Far East, where it was recorded from

Species Region and reference Aperture

ear-shaped

Upper

margin of

aperture

(frontal

view)

Upper

margin of

aperture

(apical

view)

Juveniles

look like

the adults

Bursa duct “Freckles”

on foot and

preputium

R. auricularia Southern Europe (Glöer

& Pešić 2008)

yes 90º or >90º straight no very long yes

R. auricularia Russian Far East,

excluding Khodutka

area (our data, list of

localities see. in

“Material examined”)

yes 90º or >90º straight no very long yes

R. auricularia Khodutka area,

Kamchatka (our data)

yes 90º or >90º straight no very long or

long

yes

Holotype of L. (R.)

hadutkae

Khodutka area,

Kamchatka (Kruglov &

Starobogatov 1989;

Kruglov 2005; our data)

yes 90º straight -- very long no data

Holotype of L. (R.)

thermokamtschatica

Khodutka area,

Kamchatka (Kruglov &

Starobogatov 1989;

Kruglov 2005; our data)

yes >90º straight -- long no data


Kamchatka, Primorye, Sakhalin, and Kuriles (Fig. 7). Zhadin (1952) reported that this species is widespread on the

Pacific Basin, including Kamchatka, Amur Basin and Primorye, which is in agreement with our records. However,

Kruglov & Starobogatov (1993) noted that R. auricularia ranged in Europe, the western part of the Middle East

and Siberia, but not mentioned in the species records from the Far East. Thereafter, Starobogatov et al. (2004)

again reported that species ranged on the Far East, where it was recorded in rivers of the Pacific coast from

Kamchatka to Primorye. Meanwhile, Kruglov & Starobogatov (1989, 1993), Starobogatov et al. (2004) and

Kruglov (2005) listed many local endemic Radix species for different areas of the Russian Far East, which belong

to the sections Ussuriradix Kruglov & Starobogatov, 1989, Nipponiradix Kruglov & Starobogatov, 1989 and

Okhotiradix Kruglov & Starobogatov, 1989. Taxonomic position of these taxa is uncertain and they need in a

current revision based on mitochondrial and nuclear molecular markers.

Acknowledgements

This study was supported by grants from the President of Russia (no. MD-6465.2014.5) and the Ural Branch of

Russian Academy of Sciences (no. 12-M-45-2062 and 12-P-5-1014). We are grateful to Dr. M.V. Vinarski

(Museum of Siberian Aquatic Molluscs, Russia) for providing the high-resolution photos of type specimens of

Lymnaea (Radix) hadutkae and L. (R.) thermokamtschatica. We also are thankful to Dr. P.V. Kijashko and L.L.

Jarochnovich (ZIN, Russia) for their assistance in studies of lymnaeids from the ZIN collection. Special thanks go

to Dr. N. Friberg (National Environmental Research Institute, Oslo, Norway and University of Aarhus, Denmark),

Dr. B.O.L. Demars (The James Hutton Institute, UK), Mr. V.V. Komarov (Volcanoes of Kamchatka Nature Park,

Russia), Mr. O.N. Bespaliy (Arkhangelsk, Russia) and Mr. E.P. Dekin (Elisovo, Russia) for their great help during

our field works. We also thank K. Schniebs (Senckenberg Natural History Collections Dresden, Germany) and an

anonymous reviewer for improvements to the manuscript.

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space. Systematic Biology, 61 (3), 539–542.

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2009 (Mollusca: Gastropoda: Lymnaeidae) in Bulgaria and its taxonomic relationship to other European lymnaeids based

on molecular analysis. North-Western Journal of Zoology, 8 (1), 164–171.

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freshwater invertebrates of Russia and adjacent lands (Vol. 6. Molluscs, Polychaetes, Nemerteans). Nauka, St-Petersburg,

pp. 9–492. [in Russian]

Stift, M., Michel, E., Sitnikova, T.Y., Mamonova, E.Y. & Sherbakov, D.Y. (2004) Palaearctic gastropod gains a foothold in the

dominion of endemics: range expansion and morphological change of Lymnaea (Radix) auricularia in Lake Baikal.

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APPENDIX 1. List of studied Radix auricularia specimens from Russian Far East. Materials are deposited in the

Biological Museum of Institute of Ecological Problems of the North (INEP), the Ural Branch of Russian Academy

of Sciences, Arkhangelsk, Russian Federation.

COI NCBI’s

GenBank acc.

no.

Haplotype

Code

Specimen

Voucher

Locality Habitat type Latitude Longitude

KM067596 PAR1 INEP-378.1 Russia, Kamchatka, Paratunskie

hot springs

Geothermal 52°49'26" 158°09'48"


hot springs

Geothermal 52°49'26" 158°09'48"


hot springs

Geothermal 52°49'26" 158°09'48"


hot springs

Geothermal 52°49'26" 158°09'48"


hot springs

Non-

geothermal

52°49'26" 158°09'48"


hot springs

Geothermal 52°49'26" 158°09'48"

KM067602 KHO1 INEP-375.1 Russia, Kamchatka, Khodutka

geothermal area

Geothermal 52°06'27" 157°39'25"


geothermal area

Geothermal 52°06'27" 157°39'25"


geothermal area

Geothermal 52°06'27" 157°39'25"


geothermal area

Geothermal 52°06'27" 157°39'25"


geothermal area

Geothermal 52°06'27" 157°39'25"


geothermal area

Geothermal 52°06'27" 157°39'25"


geothermal area

Geothermal 52°06'27" 157°39'25"

KM067607 MAL1 INEP-117.1 Russia, Kamchatka, Malkinskie

hot springs

Geothermal 53°19'19" 157°32'14"


hot springs

Geothermal 53°19'19" 157°32'14"


hot springs

Geothermal 53°19'19" 157°32'14"


hot springs

Geothermal 53°19'19" 157°32'14"


hot springs

Geothermal 53°19'19" 157°32'14"

......continued on the next page


http://dx.doi.org/10.1093/molbev/mst197

APPENDIX 2. List of additional COI sequences of lymnaeid species obtained from NCBI’s GenBank including accession number,

taxon, specimen code, locality information and references.

*—These species were used as outgroups for Bayesian analysis.

**—Data source: (1) Bargues, M.D., Artigas, P., Khoubbane, M., Ortiz, P., Naquira, C. and Mas-Coma, S. (2011) Human and

animal fascioliasis vectors in Cajamarca, Peru: DNA assessment of Galba truncatula, Galba neotropica and Lymnaea

schirazensis and analysis of vector overlap implications (unpubl.); (2) International Barcode of Life (iBOL), Biodiversity

Institute of Ontario, University of Guelph, Canada (unpubl.); (3) Oheimb, P.V.v., Albrecht, C., Riedel, F., Du, L., Yang, J. et al.

(2011) Freshwater biogeography and limnological evolution of the Tibetan Plateau—insights from a plateau-wide distributed

gastropod taxon (Radix spp.). PLoS ONE, 6(10), e26307. doi:10.1371/journal.pone.0026307; (4) Vinarski, M. V., Schniebs, K.,

Glöer, P., & Hundsdoerfer, A. K. (2011) The taxonomic status and phylogenetic relationships of the genus Aenigmomphiscola

Kruglov and Starobogatov, 1981 (Gastropoda: Pulmonata: Lymnaeidae). Journal of Natural History, 45(33–34), 2049–2068.

doi: 10.1080/00222933.2011.574800; (5) Quintela, M., Johansson, M.P., Kristjansson, B.K., Barreiro, R., Laurila, A. (2014)

AFLPs and mitochondrial haplotypes reveal local adaptation to extreme thermal environments in a freshwater gastropod. PLoS

ONE, 9(7), e101821. doi:10.1371/journal.pone.0101821; (6) Albrecht, C., Wolff, C., Glöer, P., & Wilke, T. (2008) Concurrent

evolution of ancient sister lakes and sister species: the freshwater gastropod genus Radix in lakes Ohrid and Prespa.

Hydrobiologia, 615, 157–167. doi: 10.1007/978-1-4020-9582-5_11

APPENDIX 1. (continued)

COI NCBI’s

GenBank acc.

no.

Haplotype

Code

Specimen

Voucher

Locality Habitat type Latitude Longitude

KM067612 PLO1 INEP-231.1 Russia, Kamchatka, Plotnikova

River (Bolshaya Basin)

Non-

geothermal

53°06'14" 157°45'16"

KM067613 PLO1 INEP-231.3 Russia, Kamchatka, Plotnikova

River (Bolshaya Basin)

Non-

geothermal

53°06'14" 157°45'16"

KM067614 ALI1 INEP-241 Russia, Kunashir Island, Aliger

Lake

Non-

geothermal

44°02'50" 145°44'38"

KM067615 TYM1 INEP-498 Russia, Sakhalin Island, Tym

River

Non-

geothermal

50°54'45" 142°39'48"

KM067618 RAZ1 INEP-500 Russia, Primorye, small pond near

Razdolnaya River

Non-

geothermal

43°46'47" 131°57'38"

KM067616 USS1 INEP-499.1 Russia, Primorye, Ussuri River

(Amur Basin)

Non-

geothermal

44°04'52" 133°50'43"

KM067617 USS2 INEP-499.3 Russia, Primorye, Ussuri River

(Amur Basin)

Non-

geothermal

44°04'52" 133°50'43"

Haplotype

code

COI NCBI’s

GenBank acc. no.

Species Specimen

Voucher

Locality Data

source**

TRU1 HE610435 Galba truncatula* No data Peru, Cajamarca (1)

PAL1 GU680909 Lymnaea stagnalis* 09PROBE-

02278

Canada, Manitoba (2)

TAJ1 JN794513 Radix auricularia UGSB 7532 Tajikistan, Gorno-Badakhshan (3)

TIB1 JN794432 R. cf. auricularia (clade 1) UGSB 7400 China, Tibet (3)

GER1 FR797876 R. auricularia Moll S1332 Germany (4)



GER4 FR797879 R. auricularia Moll 53070 Germany (4)

BAL1 JN624792 R. balthica No data Iceland, Lake Mývatn (5)

REL1 EU818822 R. relicta No data Albania, Pole, Lake Ohrid (6)

AMP1 EU818837 R. ampla No data Austria, Salzburger Land, Lake

Wallersee

(6)

LAB1 EU818832 R. labiata No data Albania, Dibre, Cidhne, gorge of

Pr. i Setes

(6)


A taxonomic revision of two local endemic Radix spp. (Gastropoda: Lymnaeidae) from Khodutka geothermal area, Kamchatka, Russian Far East

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