Zoosyst. Evol. 96 (2) 2020, 577-608 | DO! 10.3897/zse.96.52860

Gp Musee TOR
BERLIN

Taxonomic assessment of genetically-delineated species of radicine
snails (Mollusca, Gastropoda, Lymnaeidae)

Maxim V. Vinarski!*, Olga V. Aksenova!?, Ivan N. Bolotov?*

1 Laboratory of Macroecology and Biogeography of Invertebrates, Saint-Petersburg State University, 7/9 Universitetskaya Emb., 199034, Saint-
Petersburg, Russia

2 Omsk State University, 28 Adrianova Str, 644077, Omsk, Russia

3 N. Laverov Federal Center for Integrated Arctic Research, Ural Branch of the Russian Academy of Sciences, 23 Severnaya Dvina Emb.,
163000, Arkhangelsk, Russia

4 Northern (Arctic) Federal University, 17 Severnaya Dvina Emb., 163002, Arkhangelsk, Russia

http://zoobank.org/BSCF 4E 84-1 FEE-46F 3-B275-BAFA6824902B

Corresponding author: Maxim V. Vinarski (radix.vinarski@gmail.com)

Academic editor: Matthias Glaubrecht # Received 2 April 2020 Accepted 4 June 2020 Published | September 2020

Abstract

The article represents an overview of 29 biological species of the radicine snails (genera Ampullaceana Servain, 1882, Bullastra
Bergh, 1901, Racesina Vinarski & Bolotov, 2018, Kamtschaticana Kruglov & Starobogatov, 1984, Myxas G.B. Sowerby I, 1822,
Orientogalba Kruglov & Starobogatov, 1985; Peregriana Servain, 1882, Radix Montfort, 1810, and Tibetoradix Bolotov, Vinarski &
Aksenova, 2018) recovered during our previous molecular taxonomic study (Aksenova et al. 201 8a; Scientific Reports, 8: 11199). For
each species, the following information is provided: scientific name, a (non-exhaustive) list of synonyms, type locality, type materials,
shell and copulative apparatus morphology, distribution, and nomenclatural and taxonomic remarks. The colour images of shell(s) of
each species are also given as well as illustrations of the copulatory apparatuses. We revealed a great conchological variation in the
radicines, both intra- and interspecific, alongside with striking uniformity in the structure of their copulatory apparatuses. The latter
was once thought to be a reliable tool for species delineation and identification in this snail group. The total of 29 species characterised

here represents, probably, only a subset of the global taxonomic richness of the radicine snails, which approaches 50 species.

Key Words

aquatic pulmonates, distribution, Old World, phylogenetics, taxonomy

Introduction

Historically, systematics of freshwater pulmonate gas-
tropods had relied on two basic sources of data — con-
chology and anatomy. The combined use of these sources
allowed researchers to delineate species and higher taxa
and to compile dichotomous keys for identification of
snails (see, for example, Meier-Brook 1983; Taylor 2003;
Kruglov 2005). Since the late 1990s, the novel methods
of molecular phylogenetics and molecular taxonomy
have been widely adopted by malacologists and today
they represent a standard tool for species delimitation in
different families of freshwater pulmonates. Though there
is an almost full agreement that the molecular data are

more exact and ‘objective’ than the morphological ones,
their extensive use has led to some problems in the prac-
tical taxonomy of aquatic Pulmonata. First, many taxa of
snails, established on the basis of morphological informa-
tion, proved non-valid due to the lack of genetic support
(see, for example, Vinarski et al. 2016; Aksenova et al.
2017). On the other hand, some ‘good’ species defined
genetically cannot be identified morphologically as the
ranges of their conchological and anatomical variation
overlap with those of closely-related species (Schniebs et
al. 2011, 2013; Aksenova et al. 2018a). The cryptic spe-
ciation has also been detected within this group (Bargues
et al. 2011; Standley et al. 2013). Lastly, the authors of
some recent molecular works, focused on species diversi-

Copyright Maxim V. Vinarski et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which
permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

578

ty, prefer to deal with impersonal categories such as MO-
TUs (Pfenninger et al. 2006) and clades (von Oheimb et
al. 2011; Clewing et al. 2016), instead of scientific names
(Latin or latinised binomens) that would correspond to
previously-described species of snails. In such cases, the
proper taxonomic work remains unfinished since it is
very difficult to find correspondence between the MOTUs
or impersonal ‘clades’ and biological species, with their
unique names. The use of such nameless entities makes
it almost impossible to treat them in accordance with the
internationally-adopted rules of zoological nomenclature
and impedes the practical issues like conservation plan-
ning or studies of the snail-trematode interactions.
Recently, we published a paper (Aksenova et al. 201 8a)
devoted to the taxonomy, phylogeny and biogeography of
the Old World radicine snails (subfamily Amphipepleinae
Pini, 1877 of the family Lymnaeidae Rafinesque, 1815),
based on an analysis of more than 2600 sequences of two
mitochondrial (COI and 16S rRNA) and one nuclear (28S
tRNA) gene. This analysis allowed us to propose a new
taxonomic structure of the genus Radix Montfort, 1810
s. lato and to recover as many as 35 biological species of

Vinarski, M.V. et al.: A survey of radicine snails (Gastropoda, Lymnaeidae)

radicines, based primarily on the molecular data. Since
2018, one more species of this group, Radix dgebuadzei
Aksenova, Vinarski, Bolotov & Kondakov, 2019, has
been described (Aksenova et al. 2019). A simplified phy-
logenetic tree of the recent Amphipepleinae recovered
during our work is given in Fig. 1.

In that study (Aksenova et al. 2018a), we tried to link
every genetically-defined clade of the genus or species
rank to a certain taxonomic name. In particular, we man-
aged to find proper names for formal designation of al-
most all MOTUs or clades delineated by previous workers
(Pfenninger et al. 2006; von Oheimb et al. 2011; Clewing
et al. 2016). However, in the previous paper, the emphasis
was laid on phylogeny and historical biogeography (Ak-
senova et al. 2018a). Most issues, related to systematics
and nomenclature of the radicines, remained unexplained.

The current study aims at taxononic assessment of
the genetically-defined species of the radicine snails.
We tried to characterise them both morphologically and
geographically and to give the readership some cues on
how to identify these molluscs on the basis of their mor-
phological characters. Additionally, we present here some

Myxas glutinosa
61/0.75 Cerasina luteola
S7/-- Cerasina siamensis
99/1.00 Cerasina oxiana
0.2 Kamtschaticana kamtschatica
33/-- Austropeplea tomentosa
— 29/-- 100/1.00 Austropeplea hispida
49/-- Orientogalba viridis
Orientogalba ollula
1007.00 5570.83 Orientogalba bowelli
Ampullaceana intermedia
53/0.97 99/1.00 Ampullaceana balthica
92/1.00 [7 Ampullaceana ampla
95/1.00 56/0 Ampullaceana sp. Ohrid
81/0.99 Ampullaceana relicta
18/-- : :
Ampullaceana dipkunensis
30/0.71 34/-- Ampullaceana lagotis
59/-- Ampullaceana fontinalis
Peregriana peregra
72/0.79 Peregriana dolgini
88/0.99 ¢ Tibetoradix sp.4
44/0.93 98/1.00 Tibetoradix sp.2
75/0,93 Tibetoradix kozlovi
Tibetoradix hookeri
72/0.98 66/0.65 Tibetoradix sp.3
Tibetoradix sp.1
Radix (Exsertiana) natalensis
100/1.00 Radix (Exsertiana) rufescens
Radix (R.) rubiginosa
95/1.00 97/1.00 Radix (R.) alticola
44 Radix (R.) plicatula
99/1.00 esl Radix (R.) euphratica
28/-- Radix (R.) sp. Trichonis
74/1.00 62/0.67 Radix (R.) brevicauda
Radix (R.) auricularia
90/1.00 Radix (R.) makhrovi

Bullastra cumingiana

Bullastra lessoni

100/0.91

Figure 1. Majority rule consensus phylogenetic tree of the Amphipepleinae recovered from maximum likelihood analysis and
obtained for the complete dataset of mitochondrial and nuclear sequences (three codons of COI + 16S rRNA + 28S rRNA). Black
numbers near nodes are bootstrap support values/Bayesian posterior probabilities. The genus-level clades are highlighted in colour.
The other Lymnaeidae and outgroup taxa are omitted (see Aksenova et al. 2018a for the complete phylogeny, sequence dataset and

methodological details).

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Zoosyst. Evol. 96 (2) 2020, 577-608

taxonomic and nomenclatorial considerations aimed to
substantiate the taxonomic opinions proposed in the pre-
vious article (Aksenova et al. 2018a).

Material and methods

Out of 35 biological species of the radicines delineated
by us (Aksenova et al. 2018a), we were able to study
morphologically 29 taxa. The rest of the species were in-
cluded in our analyses on the basis of sequences available
from GenBank and we had none of our own or museum
materials to examine their morphology. Such species as
Radix sp., an (allegedly) endemic to Lake Trichonis of
Greece (see Aksenova et al. 2018a) and some taxa from
the Tibetan Plateau remain nameless and, though we are
aware of their species status, the full absence of the ma-
terial makes it impossible to form the type series and to
designate the holotype as is required by the international
rules of zoological nomenclature.

The snails for this study were either collected by the
authors from various Old World countries (Russia, China,
Tajikistan, Myanmar, Mongolia and some others) or ex-
amined in the collections of a series of European zoolog-
ical institutions. The full enumeration of these reposito-
ries are given below, in the abbreviations list. In all cases,
when it was possible, we tried to examine the type series
of the studied species and to compare the holotype and
paratypes (or syntypes) with the published descriptions
and our own materials. During our work, we managed
to reveal and study the type series (or possible syntypes)
of 10 valid species. In some cases, high-quality photos
of the type specimens were available for us (for exam-

SH

_SS

A SW

579

ple, those published in Sitnikova et al. 2012). Besides,
we were able to examine the type series of many nominal
species considered below as synonyms. Additionally, we
substantially benefited from the use of some comprehen-
sive taxonomic publications, including high-quality over-
views of the regional faunas (Brandt 1974; Brown 1994;
Gloer 2002; Gloer and PeSi¢ 2012; Gloer and Bossneck
2013) and from studies of the type series (Sitnikova et al.
2012, 2014; Vinarski 201 6a).

The scheme of shell measurements is given in Fig. 2A.
The praeputium:penis sheath ratio has been used as the
characteristics of the proportions of the copulatory appa-
ratus (see Fig. 2). This ratio (also known as the ‘index
of the copulatory organ’, ICA) has been used for spe-
cies delimitation in different genera of the Lymnaeidae
(Falniowski 1980; Jackiewicz 1998; Kruglov 2005; Vi-
narski 2011; Standley et al. 2013). In total, 688 shells of
21 species were measured and 241 specimens of snails
were dissected. All measurements in this paper are given
in millimetres.

The accounts for particular species presented in the
systematic part of this paper include the data on their
original descriptions, type locality, type series and distri-
bution. Additionally, we attempted to give morphological
descriptions of species, with emphasis on those characters
that may help in their identification. We avoided provid-
ing the full synonymies for each species since, in some
cases, (for such widespread and long studied taxa as Ra-
dix auricularia or Peregriana peregra) it would generate
enormous lists of synonyms. In most cases, we included
into synonymy only names with extant type series or such
taxa, whose original descriptions are detailed enough to
warrant sure judgements on their taxonomic identity.

B

Figure 2. The scheme of measurements of the shell (A) and the parts of the copulatory apparatus (B). Abbreviations: SH — shell
height; SW — shell width; SpH — spire height, BWH — body whorl height; AH — aperture height; AW — aperture width; PP — length
of praeputium; PS — penis sheath length. A — after Vinarski (2016b).

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580

List of abbreviations

The acronyms for the zoological repositories:
NHMUK — Museum of Natural History, London, UK;
NMNH — National Museum of Natural History, Paris,
France; ZISP — Zoological Institute of the Russian Acade-
my of Sciences, Saint Petersburg, Russia; ZMB — Natural
History Museum of Berlin/Museum fiir Naturkunde, Ger-
many; ZMUC — Zoological Museum of the Copenhagen
University, Denmark; NHMW — Natural History Museum
of Vienna, Austria, NMG — Natural History Museum of
Gothenburg, Sweden; SMF — Senckenberg Forschungsin-
stitut und Naturmuseum, Frankfurt-am-Main, Germany;
SNSD — Senckenberg Naturhistorische Sammlung Dres-
den, Germany; RMBH — Russian Museum of Biodiversity
Hotspots, Federal Center for Integrated Arctic Research of
the Russian Academy of Sciences, Arkhangelsk, Russia;
LMBI - Laboratory for Macroecology and Biogeography
of Invertebrates, Saint-Petersburg State University, Russia.

The abbreviations for the shell and anatomical
structures and measurements) WN — whorls num-
ber; SH — shell height; SW — shell width; SpH — spire
height; BWH — body whorl height; AH — aperture height;
AW — aperture width; PP — length of praeputium; PS — pe-
nis sheath length; ICA — the index of the copulatory ap-
paratus (= PP:PS).

TL — type locality.

Systematic part
Genus Radix Montfort, 1810

Subgenus Radix s. str.

Radix Montfort 1810: 266.

Gulnaria Turton 1831: 117 (partim).

Cerasina Kobelt 1881: 297.

Auriculariana Servain 1882: 49.

Acuminatiana Bourguignat 1889: 155.
Desertiradix Kruglov and Starobogatov 1989: 23.
Traniradix Kruglov and Starobogatov 1989: 23.
Nipponiradix Kruglov and Starobogatov 1989: 25.
Okhotiradix Kruglov and Starobogatov 1989: 28.
Pamiriradix Kruglov and Starobogatov 1989: 22.
Thermoradix Kruglov and Starobogatov 1989: 19.
Ussuriradix Kruglov and Starobogatov 1989: 24.

Type species. Helix auricularia Linnaeus, 1758

1. Radix (Radix) auricularia (Linnaeus, 1758)
Figs 3A; 4A.

Helix auricularia Linnaeus 1758: 774, 775.

Lymnaea auricularia — Hubendick 1951: 151, figs 96, 97, 99 (partim).

Lymnaea (Radix) auricularia — Kruglov and Starobogatov 1993a: 85,
fig. 1OE; Jackiewicz 1998: 47, figs 64, 65, pl. Il, X.7, XI, 1, 2; Kru-
glov 2005: 250, figs 137(/)}-139; Andreeva et al. 2010: 99, fig. 48.

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Vinarski, M.V. et al.: A survey of radicine snails (Gastropoda, Lymnaeidae)

Lymnaea (Radix) hadutkae Kruglov and Starobogatov 1989: 22,
figs 1(12), 2(7); 1993a: 85, fig. 11D.

Lymnaea (Radix) hakusyensis Kruglov and Starobogatov 1989: 20,
figs 1(8), 2(6); 1993a: 88, fig. 11G.

Lymnaea (Radix) thermobaicalica Kruglov and Starobogatov 1989: 20,
figs 1(10), 2(8); 1993a: 85, fig. 11E.

Lymnaea (Radix) thermokamtschatica Kruglov and Starobogatov 1989:
22, figs 1(9), 2(9); 1993a: 85, fig. 11F.

Radix auricularia — Gloer 2002: 213, fig. 241; Welter-Schultes 2012:
52, textfig; Vinarski and Kantor 2016: 318; Aksenova et al. 2016:
16, figs 1F, G; 2; Gloer 2019: 236, fig. 294.

TL. Europe.

Types. Possibly lost (Vinarski and Kantor 2016).

This species is common in Eurasia, sporadically dis-
tributed in North Africa, introduced into North Ameri-
ca and New Zealand (Hubendick 1951; Charleston and
Climo 1979; Burch 1989; Brown 1994; Kruglov 2005;
Andreeva et al. 2010). It has many times been charac-
terised in literature, both conchologically and anatomi-
cally (Jackiewicz 1998; Gloer 2002; Kruglov 2005). The
typical form of this snail has an ear-shaped shell with
shortened spire and greatly expanded body whorl (see
Fig. 3A). However, R. auricularia exhibits wide variation
is shell shape and proportions (Vinarski 2016b). It may
form local and ecological races, including dwarf races
of geothermal springs, which had been accepted as valid
species (Bolotov et al. 2014; Aksenova et al. 2016, 2017).

The structure of the copulatory apparatus of R. auric-
ularia is quite typical for the genus. The praeputium is
oblong, cylindrical and rather thick; its width is virtual-
ly equal along its whole length (see Fig. 4A). The penis
sheath is much narrower, with a bulbous swelling on its
distal end. The lengths of the praepuptium and the penis
sheath of R. auricularia are nearly equal, though in some
populations, there is a substantial variation in the ICA
values (Vinarski 2011).

A characteristic trait of this species, which distinguish-
es it from the remaining radicines of Europe, is the pres-
ence of freckles on the foot and praeputium (Gloer 2019).

2. Radix (Radix) alticola (Izzatullaev, Kruglov &
Starobogatov, 1983)
Figs 3B, C; 4B; Table 1

Lymnaea (Radix) alticola \zzatullaev et al. 1983: 53, figs 1, 2; Kru-
glov and Starobogatov 1993a: 85, fig. 11C; Kruglov 2005: 261,
figs 146(1)}-148; Sitnikova et al. 2014: 25, fig. 8D.

Radix (Radix) alticola — Vinarski and Kantor 2016: 320.

TL. Tajikistan, a hot spring near the Yashilkul’ Lake (ap-
proximately 37°47'00"N, 72°51'00"E).

Types. ZISP (Vinarski and Kantor 2016). The holo-
type is illustrated by Sitnikova et al. (2014). We studied
both the holotype and the paratypes of R. alticola.

According to our data (Aksenova et al. 2018a), R. al-
ticola is endemic to the High Asia mountains; found in
Tajikistan (from several sites, including the type locality)

Zoosyst. Evol. 96 (2) 2020, 577-608 581

Figure 3. Shells of species in the genus Radix. A. Radix auricularia (28.08.2013, Kazakhstan, Karaganda Region, Suresai River;
LMBI); B. R. alticola (20.06.2012, Tajikistan, a hot spring near Djelandy village; LMBI); C. R. alticola (01.07.2016, Tajikistan, a
warm brook near Dzhaushangoz village; LMBI); D. R. brevicauda (Kashmir, a syntype, NHMUK); E. R. brevicauda (05.08.1948,
Tajikistan, Pamir Mts., Shaimak village, in a warm spring; ZISP); F. R. euphratica (26.06.2016, Tajikistan, Dushanbe, a fountain
near the President’s palace; LMBI); G. R. euphratica (04.07.2016, Tajikistan, a roadside ditch near Kurban-Shakhid village; LMBI);
H. R. euphratica (without date, Iraq, Bagdad; NHMUK); I. R. makhrovi, the holotype (China, Tibet, a roadside ditch west of the
Lhasa River mouth; ZISP); J. R. plicatula, a probable syntype (China, Chusan, NHMUK); K. R. plicatula (26.07.2017, China, Uy-
ghuria, Bagrash-kol’ Lake; LMBI); L. R. plicatula (26.11.2014, China, Bejing, an artificial pond in the former Emperor’s summer
palace; LMBI). Scale bars: 2 mm (B—C, F—L), 5 mm (A, D-E, kK).

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Vinarski, M.V. et al.: A survey of radicine snails (Gastropoda, Lymnaeidae)

Figure 4. Copulatory apparatuses of the species of the genus Radix. A. Radix auricularia (19.07.2007, Russia, Tyumen’ Region,
Vylpos! channel near Labytnangi Town); B. R. alticola (01.07.2016, Tajikistan, a warm brook near Dzhaushangoz village); C. R.
euphratica (04.07.2016, Tajikistan, a roadside ditch near Kurban-Shakhid village); D. R. makhrovi, a paratype (after Aksenova et al.
2018a, modified). E. R. plicatula (26.11.2014, China, Bejing, an artificial pond in the former Emperor’s summer palace). F. R. rubig-
inosa (Malaysia, Kuala-Lumpur); G. R. rufescens (24.11.2016, Myanmar, Yetho River, near the dam). H. R. natalensis (05.08.2018,

Uganda, crater lake Kyamwiga). Scale bars: 2 mm.

and Nepal. In mountain Tajikistan (Pamir), this species
inhabits hot springs and satellite streams with relatively
warm water.

The shell shape of R. alticola is similar to that of R.
auricularia, but the former species is of much smaller
size; its shell height does not exceed 16 mm (see Table 1),
whereas the shells of R. auricularia may reach 35-40 mm
in height. The proportions of the copulatory organ of
R. alticola are very similar to R. auricularia (compare
Fig. 3A and 3B).

3. Radix (Radix) brevicauda (G.B. Sowerby II, 1872)
Fig. 3D, E; Table 1

Limnaea brevicauda G.B. Sowerby II 1872: pl. XV, fig. 105.

Limnaea brevicauda — Hanley and Theobald 1876: 64, pl. 158, fig. 7;
Annandale and Rao 1925: 157, figs 1-6.

Lymnaea (Radix) brevicauda — Subba Rao 1989: 134, figs 302, 303.

Radix brevicauda — Gloer and Bossneck 2013: 153, figs 61-63.

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TL. The type locality was originally stated as ‘Australia’
(Sowerby 1872). It is, however, erroneous. Hanley and
Theobald (1876) gave the proper type locality: Kashmir.
Types. NHMUK. The syntypes were inspected by us.
R. brevicauda inhabits Northern India (Kashmir),
Nepal, China (Western Tibet and Himalaya Range) and,
probably, Tajikistan (Pamir Mts.) [Subba Rao 1989;
Gloer and Boéssneck 2013; Aksenova et al. 2018a]. The
record of this species from Tajikistan is based on numer-
ous shells from ZISP collection (see Fig. 3E), virtually
identical with the syntypes of R. brevicauda. This species
has not been registered from the ex-USSR territory (Vi-
narski and Kantor 2016) and hereby we, for the first time,
include it in the malacofauna of the former Soviet Union.
However, this record is still not corroborated genetically.
The shell of R. brevicauda is ear-shaped, with low spire
and greatly expanded aperture. Generally, in many ways,
it resembles the shell of R. auricularia and Gloer and
Bossneck (2013: 153) note that there are no substantial
differences between the two species, which are probable

583

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Zoosyst. Evol. 96 (2) 2020, 577-608

Ayjeoo] / salzads Xapul / Ja}IeIeYD

‘XIPOY SNUDS SY} UI SoIdads [eIOADS JO sosnyesedde saryejndoo pur s][ays Jo UOTJesLIojovIeYS [eoIsO[OYdIop *] I[QeL

584 Vinarski, M.V. et al.: A survey of radicine snails (Gastropoda, Lymnaeidae)

synonyms. However, Annandale and Rao (1925) reported
the structure of the jaw and radula of R. brevicauda is
distinct from that of R. auricularia. The anatomical struc-
ture of the discussed species, described and illustrated by
Gloer and Bossneck (2013), is typical for the genus Ra-
dix. Our molecular analysis recovered R. brevicauda as a
species sister to R. auricularia (Aksenova et al. 201 8a).

We may indicate some conchological differences be-
tween R. auricularia and R. brevicauda. The latter spe-
cies is of smaller size, the largest syntype shell is 18.1 mm
(our data) and Gloer and Bossneck (2013) mention that R.
brevicauda may reach 20 mm in height. Next, the colu-
mellar depression in shells of R. brevicauda is very prom-
inent, whereas, in R. auricularia, it is typically weakly
developed (compare Fig. 2A and 2D, E). Lastly, the spire
whorls in R. auricularia are usually flattened, while in R.
brevicauda these are visibly convex and rounded.

Nomenclature remark. The name Limnaea brevi-
cauda Sowerby is the oldest available one to designate
a lymnaeid species, sister to R. auricularia, restricted in
its distribution to the Central Asia mountain regions. The
taxonomic identity of R. brevicauda, as well its close af-
finity to R. auricularia, was confirmed by the inspection
of the extant syntypes.

4. Radix (Radix) euphratica (Mousson, 1874)
Figs 3F—-H; 5C. Table 1

Limnaea euphratica Mousson 1874: 40, 41.

Limnaea tenera race euphratica — Annandale and Prashad 1919b: 113,
pl. XUI, figs 3-5.

Limnaea gedrosiana Annandale and Prashad 1919a: 48, pl. VII,
figs 2-4; Annandale and Prashad 1919b: 107.

Limnaea gedrosiana vat. rectilabrum Annandale and Prashad 1919a:
49, pl. VI, figs 1-6

Limnaea iranica Annandale and Prashad 1919a: 43, pl. VII, fig. 1.

Lymnaea gedrosiana — Likharev and Starobogatov 1967: 171, fig. 3.

Lymnaea (Pseudosuccinea) gedrosiana — Annandale and Rao 1925:
172; Subba Rao 1989: 130, fig. 295.

Lymnaea (Pseudosuccinea) gedrosiana f. rectilabrum — Annandale and
Rao 1925: 173.

Lymnaea (Pseudosuccinea) iranica — Annandale and Rao 1925: 172.

Lymnaea (Radix) euphratica — Kruglov and Starobogatov 1993a: 88,
fig. 12F; Kruglov 2005: 273, figs 160, 163.

Lymnaea (Radix) gedrosiana — Kruglov and Starobogatov 1993a: 90,
fig. 14A; Kruglov 2005: 284, figs 164(8), 176.

Lymnaea (Radix) rectilabrum — Kruglov and Starobogatov 1993a: 90,
fig. 13G; Kruglov 2005: 283, figs 164(7), 175.

Radix gedrosiana gedrosiana — Gloer and Pesi¢ 2012: 42.

Radix gedrosiana rectilabrum — Gloer and Pesi¢ 2012: 42.

Radix euphratica — Aksenova et al. 201 8a: 4.

Radix (Radix) euphratica — Vinarski and Kantor 2016: 321.

Radix (Radix) gedrosiana — Vinarski and Kantor 2016: 322.

Radix (Radix) rectilabrum — Vinarski and Kantor 2016: 324.

Radix euphratica — Gloer 2019: 239, fig. 298.

TL. Iraq, vicinity of Es-Samava Town (approximately
31°19'00"N, 45°17'00"E).

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Types. Not traced, but probably in the Zurich Zoolog-
ical Museum (Vinarski and Kantor 2016).

We recorded R. euphratica genetically from such re-
mote countries as Iraq and Turkey in the west and south-
west and Tajikistan in the northwest, also it has been iden-
tified from samples collected in the Krasnodar Region of
Russia and Georgia (Aksenova et al. 2018a; see Fig. 5C).
The records of this species in literature cover also Afghan-
istan, India, Iran, Pakistan, Azerbaijan, Tajikistan, Uz-
bekistan and Turkmenistan (Annandale and Rao 1919a, b;
Likharev and Starobogatov 1967; Subba Rao 1989; Kru-
glov and Starobogatov 1993a; Gloer and PeSi¢ 2012). The
range of R. euphratica seems to be rather wide, stretching
from the Middle East to northern India. The northernmost
localities of this snail are known in the Caucasian region,
in Georgia and European Russia (Aksenova et al. 2018a).

Conchologically, R. euphratica may be distinguished
from the species of Radix described above by its rela-
tively oblong ovate-conical shell, with high spire and
weakly-inflated body whorl. The maximum shell height
is around 20.0 mm (see Table 1). The columellar depres-
sion 1s weakly developed and, in some specimens, may be
not visible. The tangential line of spire is almost straight
or weakly concave that allows us to distinguish this spe-
cies from the three species of Radix discussed above (see
Fig. 3). The morphology of the copulatory apparatus is
typical for the genus (see Fig. 4).

Nomenclature remark. Several nominal species of
radicines, with type localities situated in the Middle East
or the east of Central Asia, were described in the late 19"
— first half of the 20" century. The oldest of them are Lim-
naeus tener Kuster, 1862, Limnaea auricularia var. per-
sica Bourguignat in Issel, 1865 and Limnaea euphratica
Mousson, 1874. The types of L. tener are lost, while the
holotype of L. auricularia var. persica is extant (Sitniko-
va et al. 2012). L. tener and L. auricularia var. persica
share the same shell shape; these are ear-shaped, with low
spire and enlarged aperture (see Fig. 5A, B). Both species
were described from Iran and their conchological simi-
larity, as well as the overlap in geographical distribution,
may indicate these forms are conspecific (or represent the
intraspecific morphs of R. auricularia). Though the type
series of R. euphratica is most probably lost, the shells of
this species collected in Iraq (NHMUK) look like shells
of snails from Tajikistan studied by us both genetically
and morphologically (compare Fig. 3F, G and 3]). It al-
lowed us to select the name Limnaea euphratica Mous-
son, 1874 for designation of this species, since the shell
habitus of both ZL. tener and L. auricularia var. persica 1s
different from that of snails from Iraq.

The absence of shell picture in the original description
of L. euphratica (Mousson, 1874) may be compensated
by the author’s remark that the shell shape of this species
“approaches ... some species of the eastern India [such
as] L. succinea Desh.” (Mousson 1874: 40). Hubendick
(1951) regarded L. succinea as a synonym of Lymnaea lu-
teola, whose habitus indeed resembles that of L. euphrati-
ca, but looks rather different from shells of either L. tener
or L. auricularia var. persica.

Zoosyst. Evol. 96 (2) 2020, 577-608

585

Figure 5. Shells of the Central Asian representatives of the genus Radix. A. Radix tener, the type (after Ktister 1862, slightly mod-
ified); B. R. persica, the holotype (after Sitnikova et al. 2012); C. R. euphratica (30.08.2013, Russia, Krasnodar Region, Yaseni
River, scale bar 2 mm; RMBH),; D. R. tenera race euphratica (Iraq, Samara, after Annandale and Prashad 1919b); E. R. gedrosiana,
a syntype (Pakistan, Baluchistan, Quetta, a pond in the Residency garden; after Annandale and Prashad 1919a); F, G. R. rectilabrum
(Pakistan, Northern Baluchistan, Pishin district, Kushdil Khan reservoir; after Annandale and Prashad 1919a); H. R. iranica, the
holotype (from the “Persian Baluchistan”; after Annandale and Prashad 1919a).

Based on the original descriptions and the study of
specimens from available museum collections (ZISP,
NHMUK, NMNH and NHMW), we consider the three
taxa from the Middle East, described by Annandale and
Prashad (1919), as junior synonyms of R. euphratica.
These are Limnaea gedrosiana, L. gedrosiana vat. recti-
labrum and L. iranica (see Fig. 5D—H). These three spe-
cies are conchologically very similar and their shell traits
correspond well to R. euphratica from Iraq (see Fig. 31).
Specimens of Radix rectilabrum sensu Kruglov and Star-
obogatov 1993a from Tajikistan and Uzbekistan studied
by us (Aksenova et al. 2018a) proved to be genetically
indistinguishable from snails sampled in Iraq.

The taxonomic identity of Limnaea tenera and con-
chologically similar species (Limnaea persica, Limnaea
cor Annandale et Prashad, 1919) needs a further research
by means of an integrative taxonomic analysis of the to-
potypic specimens.

5. Radix (Radix) makhrovi Bolotov, Vinarski &
Aksenova, 2018
Figs 31; 5D.

Radix makhrovi Aksenova et al. 2018a: 9, fig. 7E, G, I.

TL. China, Tibet, a roadside ditch west of the Lhasa Riv-
er mouth, Brahmaputra River basin.
Types. ZISP (holotype, paratypes), RMBH (paratypes).

This species, endemic to Tibet, is fully described in
our previous paper (Aksenova et al. 201 8a), therefore, we
give here only pictures of its shell and copulatory organ.

6. Radix (Radix) plicatula (Benson in Cantor, 1842)
Figs 3J—L; 4E; 5D.

Limnaea plicatula Cantor 1842: 487.

Limnaea swinhoei H. Adams 1866: 319, pl. 33, fig. 13.
Limnaeus plicatulus — Martens 1867: 221.

Limnaea yunnanensis Nevill 1877: 26, 27.

Limnaeus chefouensis Clessin 1878-1886: 391, Taf. 55, fig. 6.
Limnaeus heudi Clessin 1878-1886: 394, Taf. 55, fig. 10.

? Limnaea annamitica Wattebled 1886: 57, pl. IV, fig. 4.
Limnaea (Gulnaria) pettiti Jones and Preston 1904: 142, fig. 3.
Limnaea (Gulnaria) whartoni Jones and Preston 1904: 142, fig. 1.
Limnaea (Gulnaria) lumleyi Jones and Preston 1910: 11, textfig.
Limnaea (Gulnaria) schwilpi Jones and Preston 1910: 10, textfig.
Limnaea (Gulnaria) sinensis Jones and Preston 1910: 11, textfig.
Radix plicatulus — Yen 1939: 65, Taf. 5, fig. 40.

Radix swihhoei — Yen 1939: 66, Taf. 5, fig. 43.

Radix plicatula — Aksenova et al. 2018a: 4

TL. China, Chusan Island.

Types. Probable syntypes are kept in NHMUK (see
Taxonomic remark below).

The actual range of R. plicatula is not ascertained yet.
In our molecular analysis, we studied samples of this spe-

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586 Vinarski, M.V. et al.: A survey of radicine snails (Gastropoda, Lymnaeidae)

cies from Beijing, southern and western China that may
indicate it is distributed throughout the country. The pres-
ence of R. plicatula in adjacent states (Laos, Vietnam) 1s
also very probable, although, as far as we can judge, it
is absent from India (Subba Rao 1989). The records of
R. auricularia swinhoei (Adams) from Thailand (Brandt
1974) may refer to this species, but a special study is
needed to check it.

The shell of R. plicatula is rather oblong, with rela-
tively high spire and moderately inflated body whorl (see
Fig. 3J—L). The copulative apparatus is typical for this
genus (see Fig. 5D). In all individuals dissected in this
study, the penis sheath was slightly longer than the prae-
putium (see Table 1), thus ICA < 1.0.

Taxonomic remark. Radix plicatula is almost indis-
tinguishable from R. euphratica by its shell habitus (com-
pare Fig. 2F and 2L) and the copulatory organ structure,
but molecular analyses have revealed that the two species
are distinct and their ranges are almost non-overlapping
(Aksenova et al. 2018a). We found the shells of this spe-
cies, collected in the type locality, in NHMUK (accession
No. 42.9.30.50.51.487). These possibly once belonged to
Benson’s collection and may be considered the probable
syntypes (see Fig. 3F). Their morphology 1s very similar
to that in R. plicatula snails examined genetically by us.
Having studied the type series of some nominal species
of Radix described from China and South-East Asia (kept
in NHMUK, NMNH and ZMB), we ascertained they are
junior synonyms of R. plicatula (see Vinarski 2016a and
synonymy above).

7. Radix (Radix) rubiginosa (Michelin, 1831)
Figs 4F; 6A-C.

Limnoeus rubiginosus Michelin 1831: 22.

Limnaeus succineus vat. javanicus Mousson 1849: 42.

Limnaeus singaporinus Kuster 1862: 35, pl. 6, fig. 17.

Limnaea crosseana Mabille and Le Mesle 1866: 130, pl. 7, fig. 5.

Limnaeus javanicus — von Martens 1867: 222.

Limnaeus javanicus var. intumescens von Martens 1867: 223.

Limnaeus javanicus var. obesus von Martens 1867: 223.

Limnaeus javanicus var. rubiginosus — von Martens 1867: 224.

Limnaea javanica var. subteres von Martens 1881: 88, figs 6, 7.

Limnaea javanica var. angustior von Martens 1881: 88, pl. 16, fig. 8.

Limnaea javanica vat. porrecta von Martens 1881: 89, figs 9, 10.

Limnaea bongsonensis Bullen 1906: 14, pl. II, figs 5, 6.

Lymnaea auricularia rubiginosa — Hubendick 1951: 154, figs 342, 342;
Brand 1974: 229, pl. 16, fig. 95 (partim).

Lymnaea rubiginosa — van Benthem Jutting 1959: 116.

Radix rubiginosa — Aksenova et al. 2018a: 4

TL. The East-Indies. “The original specimens are said
to come from Bogor in Java” (Brandt 1974: 230).

Types. Probably lost. We failed to find them in
NMNH collection.

This species is endemic to Southeast Asia. Its find-
ings supported by molecular evidence are known from

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a vast territory, 1.e. Thailand, Singapore, Indonesia up
to Lesser Sundas (Flores) and Mascarenes (Réunion)
[Aksenova et al. 2018a]. Is has also been recorded from
Cambodia, Laos and Vietnam (Brandt 1974). However,
the identification of R. rubiginosa, based on shell only,
may be misleading. As Brandt (1974: 230) stated, “this
race has often been confused with L. /uteola Lamarck”.
Therefore, the actual range of R. rubiginosa 1s not satis-
factorily outlined since many historical recordings of it
were shell-based.

A high conical spire, almost straight tangential line
and weakly-inflated body whorl are the most characteris-
tic conchological traits of R. rubiginosa (see Fig. 6A—C)
helping one to distinguish it from the rest of congeners
inhabiting the south of Asia. The variation in the propor-
tions of the copulatory organ of this species has not been
studied. We dissected four specimens collected from dif-
ferent parts of the R. rubiginosa range (Thailand, Malay-
sia, Philippines). In all cases, the penis sheath was shorter
than the praeputium (see Fig. 4F) and the ICA values var-
ied between 1.02 and 1.59.

Taxonomic remark. R. rubiginosa has an extensive
synonymy and a special study is required to clarify the
taxonomic identity of numerous nominal species of Ra-
dix, described from Indonesia and adjacent countries.
Some of these names, such as Radix quadrasi (MoOllen-
dorff, 1898) of Philippines, have been used in recent lit-
erature, but may well be synonyms of R. rubiginosa (see,
for example, Stelbrink et al. 2019).

Subgenus Exsertiana Bourguignat, 1883

Exsertiana Bourguignat 1883: 88.
Raffrayana Bourguignat 1883: 88.
Radix (Exsertiana) — Aksenova et al. 2018a: 6, 9

Type species. This subgenus contains two molecular-
ly-defined species of Radix having an almost entirely
tropical distribution — Radix natalensis (Krauss, 1848)
and R. rufescens (Gray, 1822).

Remark. Bourguignat (1883: 88), in his lengthy ar-
ticle devoted to classification of continental molluscs of
Abyssinia, established two new groups of species with-
in the genus Limnaea — Exsertiana and Raffrayana. He
did not give any diagnoses for the two taxa, only lists of
species included there. In both cases, these species are
currently recognised as junior synonyms of R. natalen-
sis (fide Hubendick 1951; Brown 1994). We selected Ex-
sertiana as a name for designation of a lymnaeid clade
containing R. natalensis. Since Bourguignat (1883) did
not indicate the type species for Exsertiana, we designate
here Limnaeus natalensis var. exsertus Martens, 1866 as
the type taxon for this subgenus. This species 1s usually
regarded as identical to R. natalensis (Hubendick, 1951),
though Mandahl-Barth (1954) accepted it as a valid spe-
cies, closely related to the latter.

Zoosyst. Evol. 96 (2) 2020, 577-608 587

Figure 6. Shells of species of the subgenera Radix s. str. and Exsertiana. A. Radix rubiginosa (Malaysia, Kuala-Lumpur; LMBI); B. R.
rubiginosa (1846, Indonesia, Java Island, “Batavia”; ZMUC); C. R. rubiginosa var. intumescens (Indonesia, Java Island, Surabaya;
ZMB); D. R. natalensis (= Limnaea undussumae) (Congo, Undussuma; ZMB). E. R. natalensis (= Limnaea arabica Smith; the holo-
type; NHMUK); FE. R. natalensis f. gravieri (10.03.1966, Ethiopia, lower Avash valley, swamp between Assaita and Dubte, NHMUK).
G. R. rufescens (24.11.2016, Myanmar, Yetho River, near the dam; LMBI); H. R. rufescens var. patula (India, Ganges River; ZMB);
I. R. rufescens var. chlamys (India, Moradabad; probably syntype; NHMUK). Scale bars: 2 mm (A, B, E, F), 5 mm (C, D, G, I).

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588 Vinarski, M.V. et al.: A survey of radicine snails (Gastropoda, Lymnaeidae)

8. Radix (Exsertiana) natalensis (Krauss, 1848)
Figs 4H; 6D-F; Table 1

Limnaeus natalensis Krauss 1848: 85.

Limnaea hovarum Tristram 1863: 61.

Limnaeus natalensis var. exsertus von Martens 1866: 101, pl. 3, figs 8, 9.

Limnaea electa Smith 1882: 385.

Limnaea caillaudi Bourguignat 1883: 89, figs 100, 101.

Limnaea exserta — Bourguignat 1883: 90.

Limnaea gravieri Bourguignat 1885: 23.

Limnaea nyansae von Martens 1892: 16.

Limnaea arabica Smith 1894a: 142, fig. 3.

Limnaea elmeteitensis Smith 1894b: 167, fig. 5.

Limnaea humerosa von Martens 1897: 135, pl. 6, fig. 1.

Limnaea undussumae von Martens 1897: 135, pl. I, fig. 18; pl. VI, figs 2, 5.

Limnaea kempi Preston 1912: 190, pl. 32, fig. 1.

Limnaea (Radix) elmeteitensis — Germain 1919: 186.

Limnaea (Radix) gravieri — Germain 1919: 186.

Limnaea (Radix) natalensis — Germain 1919: 185.

Limnaea (Radix) tchadiensis — Germain 1919: 186.

Limnaea (Radix) vignoni — Germain 1919: 186.

Lymnaea exserta — Mandahl-Barth 1954: 71, fig. 32a—c.

Lymnaea natalensis natalensis — Mandahl-Barth 1954: 67, fig. 30a, b.

Lymnaea natalensis caillaudi — Mandahl-Barth 1954: 68, fig. 3la—h.

Lymnaea natalensis nyansae — Mandahl-Barth 1954: 70, fig. 30c—h.

Lymnaea (Radix) natalensis — Brown 1994: 166, figs 76a, b; 79a.

Radix natalensis — Neubert 1998: 354; Lotfy and Lotfy 2015: 29; Gloer
2019 p. 241, fig. 301.

TL. South Africa, Natal.

Types. Probably lost. We failed to find them either in
ZMB collection or in other institutions (NMNH, NHMUK)
known to contain vast collections of African aquatic snails.

R. natalensis is widely distributed in the sub-Saharan
Africa as well as in the Nile Basin of Northern Africa
(van Damme 1984; Brown 1994; Lotfy and Lotfy 2015).
It also occurs in Arabia (Neubert 1998) and Atlantic and
Indian Ocean islands: Anjouan (Comores), Madagascar
(Brown 1994; Stothard et al. 2000) and the Cabo Verde
Islands (Aksenova et al. 201 8a).

The shell of this species is (in its typical form)
ovate-conical, with somewhat oblong and narrow spire
and ovoid body whorl (see Fig. 6D, E). However, some
varieties of R. natalensis possess an almost spheroid shell
with very low spire (see Fig. 6F). The shell height is up to
23 mm (Lotfy and Lotfy 2015).

We studied anatomically a sample of R. natalensis
collected from the crater lake Kyamwiga in Uganda and
found that the soft body anatomy of this snail is general-
ly the same as in the species of Radix discussed above.
Praeputium is relatively narrow and oblong, darkly pig-
mented (grey), whereas the penis sheath is typically light
coloured and much narrower and a little longer than the
praeputium (see Fig. 4H); the ratio of their lengths is
0.96 + 0.07 (see Table 1).

The intraspecific shell variation in R. natalensis is very
prominent (see Fig. 6) that led some authors to split this
species into several ones (Germain 1919; Mandahl-Barth

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1954). Another outcome of this enormous variation is a
plethora of nominal species and variety of names pro-
posed to designate the African Radix snails. The synony-
my given above represents a small part of this abundance
of names (see Germain 1919 for a fuller list of these).

9. Radix (Exsertiana) rufescens (Gray in Sowerby,
1822)
Figs 4, G; 6, G-I; Table 1

Lymnaea acuminata Lamarck 1822: 160 (invalid; a junior homonym of
Lymnaea acuminata Brogniart, 1810).

Limnaea rufescens Sowerby 1822: 44, pl. 178, fig. 2.

Limnaeus chlamys Benson 1836: 744.

Limnaeus amygdalum Troschel 1837: 168.

Limnaeus nucleus Troschel 1837: 171.

Limnaeus patulus Troschel 1837: 167.

Limnaeus prunum Troschel 1837: 170.

Limnaeus sulcatulus Troschel 1837: 167.

Limnaeus amygdalum — Kister 1862: 35, pl. 6, figs 15, 16.

Limnaea acuminata var. rufescens — Nevill 1877: 27.

Limnaea acuminata — Annandale and Prashad 1919c: 140, pl. V, fig. 1;
Germain 1924: 27, figs 23-28.

Limnaea acuminata var. nana Annandale and Prashad 1919c: 141,
pl. IV, fig. 1.

Limnaea chlamys — Annandale and Prashad 1919c: 142, pl. V, fig. 3;
pl. V, fig. 2.

Limnaea (Pseudosuccinea) acuminata — Annandale and Rao 1925: 177.

Lymnaea auricularia rufescens — Hubendick 1951: 157, fig. 344.

Lymnaea (Pseudosuccinea) acuminata — Subba Rao 1989: 126,
figs 254-265, 272.

Radix acuminata — Gloer and Bossneck 2013: 152, figs 52-54, 58-60.

Radix (Exsertiana) rufescens — Aksenova et al. 2018a: 9.

TL. “The East Indies”.

Types. Probably lost.

The range of this species covers the central part of the
tropical Asia; it is known from India, Nepal and Myan-
mar, but absent from Thailand (Brandt 1974; Subba Rao
1989; Gloer and Boéssneck 2013; Aksenova et al, 201 8a).

The shell of R. rufescens is large, its height may ap-
proach 45 mm (see Table 1). Most probably, it is the larg-
est member of the genus. Hubendick (1951: 56) described
its shape in such words: “[shell is] extremely high and
slender. Even the body whorl and the aperture are slen-
der’. However, this description is fully applicable only to
the typical morph of this species. There are varieties with
inflated body whorl and not slender aperture (see Fig. 61).
This high variability allowed some authors to split several
intraspecific morphs or even separate species within the
R. rufescens s. lato (Troschel 1837; Annandale and Pra-
shad 1919c; Subba Rao 1989).

Some shells of R. rufescens look almost indistinguish-
able from shells of certain varieties of R. natalensis (com-
pare Fig. SE and 5H). It is problematic to draw a sharp
boundary between the two species on the basis of their
conchological characters. R. natalensis and R. rubiginosa

Zoosyst. Evol. 96 (2) 2020, 577-608

represent a pair of closely-related vicariant species isolat-
ed by the Middle East, where no representatives of Exser-
tiana occur. Similar to R. natalensis, the relative height of
the spire is the most variable shell trait in R. rufescens.
The next three genera discussed (Ampullaceana, Per-
egriana and Kamtschaticana) represent the parts of the
former subgenus Peregriana s. lato classified within the
genus Lymnaea (Kruglov and Starobogatov 1993b; Kru-
glov 2005) or the genus Radix (Vinarski 2013; Vinarski
and Kantor 2016). From the morphological point of view,
these three genera cannot be distinguished on the basis
of either shell or anatomical characters (Aksenova et al.
2018a) and they share the same Bauplan of the repro-
ductive system. That is why previous authors considered
them as the members of a single subgenus (Peregriana;
see Kruglov and Starobogatov 1993b; Kruglov 2005) or
even lumped them into a single polymorphic species,
Lymnaea peregra s. lato (see Hubendick 1951).

Genus Ampullaceana Servain, 1882

Ampullaceana Servain 1882: 53; Aksenova et al. 2018a: 11.
Biformiana Servain 1882: 47.

Bouchardiana Servain 1882: 53.

Caenisiana Servain 1882: 56.

Effusiana Servain 1882: 49.

Nivalisiana Servain 1882: 54.

Ohridlymnaea Kruglov and Starobogatov 1993b: 166.

Type species. Limnaeus ampullaceus RossmaBler, 1835
=? Ampullaceana balthica (Linnaeus, 1758). According
to Kruglov and Starobogatov (1993b), L. ampullaceus is
a valid species allied to A. balthica.

10. Ampullaceana ampla (W. Hartmann, 1821)
Figs 7A, B; 8A; Table 2

Limneus hartmanni Studer 1820: 93 (nom. nudum).

Limneus auricularia var. ampla Hartmann 1821: 250, Taf. II, fig. 29.

Limnaea ampla — Hartmann 1840-1844: 69, pl. 17.

Gulnaria hartmanni — Hartmann 1840-1844: 71, pl. 18.

Gulnaria monnardi Hartmann 1840-1844: 72, pl. 19.

Lymnaea peregra — Hubendick 1951: 146, figs 1, 9 (partim).

Lymnaea (Peregriana) patula — Kruglov and Starobogatov 1993b:
166, fig. 6D (Lymnaea), non Da Costa 1778; Kruglov 2005: 351,
figs 236(4—5); 241, 242.

Lymnaea tobolica Lazareva 1967: 200, figs 4, 8.

Radix ampla — Gloer 2002: 215, fig. 243; Gloer 2019: 235, fig. 293.

Radix (Peregriana) ampla — Vinarski and Kantor 2016: 327.

Ampullaceana ampla — Aksenova et al. 201 8a: 4.

TL. Germany, Bavaria, Rhein River near Reineck.
Lectotype. Naturmuseum Saint-Gallen, Switzerland
(designated in Vinarski and Gloer 2007).
A. ampla is distributed in Europe (except the north-
ernmost and southernmost latitudes), Western Siberia

589

and the southern part of Eastern Siberia (Gloer 2002,
2019; Vinarski and Gloer 2007; Andreeva et al. 2011;
Welter-Schultes 2012). The shell of this species is rela-
tively large, with low spire and very expanded aperture;
conchologically, A. ampla can sometimes be confused
with R. auricularia, however, the former species has
straight or virtually absent columellar fold, which is not
characteristic for R. auricularia. Shell height may reach
30-32 mm. The lengths of the praeputium and the pe-
nis sheath in this species are roughly equal (see Fig. 8A),
with ICA close to 1.00. In a sample of A. ampla (n = 52),
collected from several water-bodies of Russia, ICA was
equal to 0.88 + 0.13 (min — max = 0.51-1.21).

11. Ampullaceana balthica (Linnaeus, 1758)
Figs 7C, D; 8B; Table 2

Helix balthica Linnaeus 1758: 775.

Turbo patulus Da Costa 1778: 95, pl. V, fig. 17.

Limneus ovatus Draparnaud 1805: 50, pl. U, figs 30, 31, 33.

Limnaeus vulgaris Pfeiffer 1821: 89, Taf. IV, fig. 22.

Limnaea geysericola Beck 1837: 114.

Limnaea ovata vat. inflata Kobelt 1871: 164, Taf. IV, fig. 12.

Lymnaea peregra — Hubendick 1951: 146, figs 1, 9 (partim).

Radix ovata auct.

Lymnaea (Peregriana) balthica — Kruglov and Starobogatov 1983b:
1468, fig. 2(11); 1993b: 165, fig. 4C.

Radix balthica — Gléer 2002: 217, fig. 245; Schniebs et al. 2011: 664,
figs 4-8; Gloer 2019: 237, fig. 296.

Radix (Peregriana) balthica — Vinarski and Kantor 2016: 329.

Ampullaceana balthica — Aksenova et al. 201 8a: 4.

TL. Sweden, the Baltic Sea shore (Linnaeus 1758). The
locality of the neotype is “Sweden, Stockholm” (see Kru-
glov and Starobogatov 1983b for details).

Neotype. ZISP (designated in Kruglov and Starobo-
gatov 1983b).

This is one of the most common and widespread spe-
cies of radicine snails in Europe (Aksenova et al. 2018a).
A single representative of the radicine pond snails was
found in Iceland (Bolotov et al. 2017). The species is
common in various regions of the Northern Palearctic;
distributed in Europe, Siberia and Central Asia (Andree-
va et al. 2010; Schniebs et al. 2011; Vinarski et al. 2017).
A. balthica is characterised by a rather wide variation,
both in conchological and anatomical traits (Summarised
in Schniebs et al. 2011). The taxonomic identity of this
species is based on the neotype (illustrated in Schniebs et
al. 2011, fig. 8).

12. Ampullaceana dipkunensis (Gundrizer &
Starobogatov, 1979)
Figs 7E, F; 8C; Table 2

Lymnaea peregra — Hubendick 1951: 146, figs 1, 9 (partim).
Lymnaea dipkunensis Gundrizer and Starobogatov 1979: 1134, fig. 1(4).

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590 Vinarski, M.V. et al.: A survey of radicine snails (Gastropoda, Lymnaeidae)

Figure 7. Shells of species of the genera Ampullaceana and Peregriana. A. Ampullaceana ampla (Austria, Wallersee Lake; NHMW).
B. A. ampla (Germany, Neckarsteinach; NHMUK). C. A. balthica (14.12.1946, Denmark, Fureso Lake; ZMUC). D. A. balthica (Swe-
den, Stockholm; NMG). E. A. dipkunensis (Russia, Arkhangelsk Region, Nyandomka River; LMBI). F. A. dipkunensis, the holotype
(07.07.1977, Russia, Krasnoyarsk Territory, Gornoye Lake; ZISP). G. Limneus fontinalis, the lectotype (after Forcart 1957). H. A. fon-
tinalis (18.08.2017, Russia, Rostov Region, Veselovskoye Reservoir; LMBI). I. Limneus intermedius (France, Lyon; NHMW). J. A.
lagotis (Russia, Leningrad Region, Lava River near Troitskoye; ZISP). K. A. /agotis (Austria, Burgenland, Bodersdorf; NHMW).
L. Peregriana dolgini (Russia, Republic of Buryatia, Kironskiy Spring; RMBH). M. P. peregra (France, from Draparnaud’s collec-

tion; NHMW). Scale bars: 2 mm (D-F, J, L, M), 5 mm (A-C, H, I, K). G— scale bar absent from the original publication.

Lymnaea (Peregriana) dipkunensis — Kruglov and Starobogatov 1993b:
164, fig. 2B (partim).

Lymnaea (Peregriana) tumida — Kruglov and Starobogatov 1993b: 166,
fig. 6G, non Held 1836 (partim).

Radix (Peregriana) dipkunensis — Vinarski and Kantor 2016: 332 (partim).

Radix (Peregriana) tumida — Vinarski and Kantor 2016: 336, non Held
1836 (partim).

Ampullaceana cf. dipkunensis — Aksenova et al. 2018a: 4.

TL. Russia, Krasnoyarsk Territory, Gornoye Lake in the

floodplain of the Kureika River upstream of the mouth of
the Dipkun River.

zse.pensoft.net

Holotype. ZISP (see Sitnikova et al. 2014).

A species characterised by a broad shell with well-de-
veloped aperture, relatively-low spire and inflated body
whorl. Its range covers European North of Russia and
Eastern Europe (Aksenova et al. 2018a). Anistratenko et
al. (2018) mentioned it (as Radix tumida) from the West-
ern Ukraine. Specimens, morphogically identified as R.
tumida, are known from the Urals, Western and Eastern
Siberia (Gundrizer 1984; Kruglov 2005; Khokhutkin et
al. 2009; Andreeva et al. 2010).

Conchologically, A. dipkunensis resembles A. ampla,
but differs from the latter by a slightly higher spire and less

Zoosyst. Evol. 96 (2) 2020, 577-608 591

Figure 8. Copulatory apparatuses of the species in the genera Ampullaceana, Peregriana, Kamtschaticana, Tibetoradix, Racesina,
Orientogalba and Bullastra. A. Ampullaceana ampla (11.07.2017, Ukraine, Transcarpathian Region, Irshava River). B. A. balthica
(Germany, Mecklenburg-Vorpommern, lake Torgelower See). C. A. dipkunensis (15.09.2016, Ukraine, Transcarpathian Region,
Tisa River). D. A. fontinalis (20.08.2017, Russia, Rostov Region, Severskiy Donets River). E. A. /agotis (29.06.2016, Tajikistan, a
stream in the Panj River floodplain). F. Peregriana dolgini (13.07.2015, Russia, Tyumen’ Region, a floodplain of the Tol’ ka oxbow).
G. Peregriana peregra (01.07.2006, Russia, Omsk Region, vicinity of Chernoluchye settlement). H. Kamtschaticana kamtschatica
(12.04.2014, Russia, Kamchatka Peninsula, Valley of Geysers). I. Tibetoradix hookeri (09-10.2012, China, Tibet, a brook, 250 km N
of Lhasa). J. 7’ kozlovi (after Aksyonova et al. 2018a, modified). K. Racesina oxiana (04.07.2016, Tajikistan, a roadside ditch near
Kurban-Shakhid village). L. R. siamensis (24.11.2016, Myanmar, a water reservoir on the Yetho River). M. Ampullaceana relicta
relicta (2005, Ohrid Lake). N. Orientogalba ollula (08.07.2010, China, Shaanxi Province, a roadside ditch near Yandi village).
O. Bullastra cumingiana (Philippines, Luzon Island, Lake Sampaloc). Scale bars: 1 mm. B — after Schniebs et al. (2011), modified.

inflated aperture. Besides, Kruglov (2005) reported some 1.7 times longer than the penis sheath. In a sample contain-
differences in the proportions of the copulatory organs of ing 103 specimens of this species, collected from four local-
these two snails. As was stated above, the penis sheath in A. ities in Russia, the mean ICA value was 1.84 + 0.22 (limits
ampla is slightly longer than the praeputium, whereas, in A. of variation: 1.37—2.44) [ Vinarski, unpublished data].

dipkunensis, one may observe an opposite state. According Taxonomic remark. Aksenova et al. (2018a) geneti-
to Kruglov (2005), the praeputium of R. tumida is around cally studied some specimens of radicine snails identified

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A survey of radicine snails (Gastropoda, Lymnaeidae)

Vinarski, M.V. et al.:

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Zoosyst. Evol. 96 (2) 2020, 577-608

as Lymnaea (Peregriana) tumida Held, 1836 sensu Kru-
glov and Starobogatov (1983b, 1993b) and showed that
it is a valid species, distinct from A. ampla. However, to
use this species name 1s hardly acceptable. This taxon is
usually considered as an intraspecific morph of Radix au-
ricularia \iving in large Alpine lakes (Geyer 1927; Gloer
2002). Examination of the topotypes (dried shells) of L.
tumida, kept in different European museums (NHMW,
NMG, ZMUC), allowed Aksenova et al. (2018a) to agree
with this opinion. Therefore, the authors used the next
oldest available name, Lymnaea dipkunensis Gundrizer
et Starobogatov, 1979, to designate this clade. The iden-
tity of this taxon was revealed by means of the inspection
of the type series (see also Sitnikova et al. 2014). The
records of Lymnaea tumida sensu Kruglov and Starobo-
gatov in the Lower Yenissei Basin (Gundrizer 1984), the
type region of L. dipkunensis, give indirect evidence in
favour of their identity. However, this taxonomic hypoth-
esis is in need of future integrative research using the to-
potypes of L. dipkunensis.

13. Ampullaceana fontinalis (Studer, 1820)
Figs 7G, H; 8D; Table 2

Limneus fontinalis Studer 1820: 27.

Lymnaea peregra — Hubendick 1951: 146, figs 1, 9 (partim).

Lymnaea (Peregriana) fontinalis — Kruglov and Starobogatov 1983b:
1469, fig. 2(16), 1993b: 166, fig. 6B; Khokhutkin et al. 2009: 92,
fig. 40; Andreeva et al. 2010: 135, fig. 70.

Ampullaceana fontinalis — Aksenova et al. 2018a: 4.

TL. Switzerland (Studer 1820).

Lectotype. Naturhistorisches Museum der Burgerge-
meinde Bern, Switzerland (see Forcart 1957).

Shell ovate-conical, with prominent, but relative-
ly-low spire and moderately-inflated body whorl. Prae-
putium and penis sheath light-coloured, their lengths
are almost equal, though the praeputium is typically
slightly longer (see Fig. 8D). The mean values of ICA
slightly exceed 1.0 (Kruglov 2005; see also Table 2).
According to genetic data, A. fontinalis is distributed
in Europe, from Switzerland eastward to south of the
European Russia, as well as in Turkey (Aksenova et al.
2018a). The previous recordings of this species from the
Urals and Siberia (Kruglov and Starobogatov 1993b;
Khokhutkin et al. 2009; Andreeva et al. 2010) require
molecular confirmation.

Taxonomic remark. This European species of rad-
icine snails was not listed as valid by most authors
(Hubendick 1951; Gloer 2002; Welter-Schultes 2012),
except by the Russian malacologists (Kruglov and Star-
obogatov 1983b, 1993b; Khokhutkin et al. 2009; Andre-
eva et al. 2010). Aksenova et al. (2018a) have shown
that specimens identified as Lymnaea (Peregriana)
fontinalis sensu Kruglov & Starobogatov, 1993 form a
species-rank clade, sister to A. /agotis. Shells of A. fon-
tinalis sensu Kruglov and Starobogatov (1993b), stud-

593

ied by us, were similar to the lectotype shell illustrated
by Forcart (1957) [see Fig. 7G, H]. The presence of A.
fontinalis in Switzerland (where its type locality hes)
has been confirmed molecularly (Aksenova et al. 2018).
This species is morphologically similar to A. /agotis, but
may be distinguished by a lower spire and more inflated
body whorl.

14. Ampullaceana intermedia (Lamarck, 1822)
Fig. 71; Table 2

Lymnaea intermedia Lamarck 1822: 162.

Limnea intermedia — Michaud 1831: 86, pl. XVI, figs 17, 18.

Limnaeus intermedius — Kuster 1862: 12, Taf. 2, figs 21, 22.

Limnaea intermedia — Sowerby 1872: pl. Ill, fig. 16; Kobelt 1912: 18,
Taf. CCCCLXXXVIII, fig. 2602.

Lymnaea peregra — Hubendick 1951: 146, figs 1, 9 (partim).

Lymnaea (Peregriana) intermedia — Kruglov and Starobogatov 1983b:
1468, fig. 2(9); 1993b: 165, fig. 4B; Khokhutkin et al. 2009: 85,
fig. 36; Andreeva et al. 2010: 125, fig. 64.

Radix (Peregriana) intermedia — Vinarski and Kantor 2016: 331.

Ampullaceana intermedia — Aksenova et al. 2018a: 4.

TL. France, Quercy Plateau.

Types. Most probably lost (Mermod 1952).

Aksenova et al. (2018a) used the binomen Lymnaea
intermedia Lamarck, 1822 to designate a radicine spe-
cies, sister to A. balthica, which is widely distributed in
France and Spain. Since the type specimen, mentioned
by Lamarck (1822), is probably lost, we based our under-
standing of this taxon on examination of both historical
samples of it (see, for example, specimen of L. intermedia
collected in Lyon, France, in the first half of the 19" cen-
tury: Fig. 71) and some old literary sources, dealing with
lymnaeids of Western Europe (Michaud 1831; Kuster
1862; Sowerby 1872; Kobelt 1912).

Conchologically, shells of A. intermedia resemble
those of A. balthica, but can be distinguished from
the latter by higher spire and less inflated body whorl.
Modern European authors do not accept A. intermedia
as a valid species (Gloer 2002; Welter-Schultes 2012),
whereas malacologists of the former USSR still mention
this taxon as a species closely allied to A. balthica (Kru-
glov and Starobogatov 1993b; Khokhutkin et al. 2009;
Andreeva et al. 2010; Vinarski and Kantor 2016). The
concept of this species proposed by Kruglov and Star-
obogatov (1983b, 1993b) coincides with that of old Eu-
ropean authors (Michaud 1831; Kuster 1862; Sowerby
1872). However, we still do not possess any sequence
of A. intermedia from the countries lying east of France
and the actual range of this species remains unknown.
The Russian authors repeatedly recorded this species
from different regions of Russia, including the Urals and
Siberia (Kruglov and Starobogatov 1993b; Khokhutkin
et al. 2009; Andreeva et al. 2010), but all these records
were based solely on morphological data and need to be
confirmed molecularly.

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594

15. Ampullaceana lagotis (Schrank, 1803)
Figs 7J, K; 8E; Table 2

Buccinum lagotis Schrank 1803: 290.

Lymnaea peregra — Hubendick 1951: 146, figs 1, 9 (partim).

Radix lagotis — Gloer 2002: 219, fig. 246; Welter-Schultes 2012: 53,
textfig.; Schniebs et al. 2015: 35, figs 3-5; Gloer 2019: 241, fig. 300.

Lymnaea (Peregriana) lagotis — Kruglov and Starobogatov 1983b:
1469, fig. 2(15); 1993b: 166, fig. 6A; Khokhutkin et al. 2009: 90,
fig. 39; Andreeva et al. 2010: 134, fig. 69.

Radix (Peregriana) lagotis — Vinarski and Kantor 2016: 334.

Ampullaceana lagotis — Aksenova et al. 2018a: 4.

TL. Germany, Bavaria, Danube River.

Types. Lost (Vinarski and Kantor 2016).

This species 1s characterised conchologically by a rela-
tively oblong shell, with high and narrow spire and mod-
erately-inflated body whorl (see Fig. 7J, K). The structure
of the copulatory apparatus in A. /agotis is typical for the
genus. Praeputium is relatively long, its length is about
1.5 times larger than the length of the penis sheath (see
Fig. 8E; Table 2). The range of A. /agotis covers, almost
entirely, Europe, Siberia and some parts of Central Asia,
including Tajikistan and western China (Gloer 2002, 2019;
Andreeva et al. 2010; Welter-Schultes 2012; Vinarski and
Kantor 2016). The molecularly-confirmed records of this
Species were reported from different countries of northern,
central and southern Europe, as well as from Tajikistan
and Siberia (Schniebs et al. 2015; Aksenova et al. 201 8a).

Shell proportions of A. /agotis resemble those of A. in-
termedia; however, A. lagotis typically has a much narrow-
er spire than the latter species (compare Fig. 7I and 7J, K).

16. Ampullaceana relicta (Polinski, 1929)

Note. This species is endemic to the large ancient lakes of
the Balkans (Albrecht et al. 2008; Welter-Schultes 2012:
Gloer 2019). A. relicta is phylogenetically close to A. am-
pla and, most probably, represents a local derivative of
the latter originating as a consequence of its ancient in-
vasion into large lakes. As Aksenova et al. (2018a) have
shown, this species is polytypic and includes two subspe-
cies, which are separately discussed below.

16a. Ampullaceana relicta relicta (Polinski, 1929)
Figs 8M; 9A.

Radix relicta Polinski 1929: 158.

Lymnaea peregra — Hubendick 1951: 146, figs 1, 9 (partim).

Lymnaea (Peregriana) relicta — Kruglov and Starobogatov 1983b:
1469, figs 2(12), 3(12); 1993b: 166, fig. 5C.

Radix relicta — Albrecht et al. 2008: 160, fig. 1; Welter-Schultes 2012:
54, textfig.; Gloer 2019: 244, fig. 305

Ampullaceana relicta relicta — Aksenova et al. 2018a: 4.

TL. Lake Ohrid.

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Vinarski, M.V. et al.: A survey of radicine snails (Gastropoda, Lymnaeidae)

Types. Whereabouts unknown.

This subspecies has been characterised both morpho-
logically and genetically by Albrecht et al. (2008). We
examined the copulatory apparatus of two specimens and
found that its structure 1s typical for the genus. Both prae-
putium and penis sheath are light-coloured; praeuputium is
substantially larger than the penis sheath (see Fig. 8M), the
ratios between their lengths are 1.42 and 1.46, respectively.

A. r. relicta inhabits Ohrid Lake (Albania and Mace-
donia), as well the Drin system, at least upstream to the
dam Globo¢icko, approximately 15 km N of the lake (Al-
brecht et al. 2008).

16b. Ampullaceana relicta pinteri (Schiitt, 1974)
Fig. 9B.

Radix pinteri Schutt 1974: 471, pl. 1.

Radix pinteri — Albrecht et al. 2008: 160, fig. 2; Welter-Schultes 2012:
54, textfig.; Gloer 2019: 244, fig. 304.

Ampullaceana relicta pinteri — Aksenova et al. 2018a: 4.

TL. North Macedonia, Lake Prespa near village of Perovo.

Types. Holotype — SMF; paratypes were placed in
NHMW and in a Series of private collections in Germa-
ny and Hungary (Schititt 1974); currently, specimens from
these private collections are kept in the Hungarian Natu-
ral History Museum (Budapest) and in the Natural History
Museum of Bern (Switzerland). We examined the paratypes
kept in NHMW and in the Budapest museum (see Fig. 9B).

This subspecies has been characterised both morpho-
logically and genetically in a series of papers (Schutt
1974; Albrecht et al. 2008; Welter-Schultes 2012). Most
authors regard this taxon as a separate species. For exam-
ple, Albrecht et al. (2008) considered A. 7: relicta and A. r.
pinteri as two sister species from two ‘sister’ lakes, Ohrid
and Prespa (but see Aksenova et al. 2018a). According to
the anatomical data presented in Albrecht et al. (2008), the
morphology of the copulatory apparatus in A. 7 pinteri is
virtually identical with that of the nominative subspecies.

A. r. pinteri is endemic to Prespa Lake (Albania,
Greece and Macedonia).

Genus Peregriana Servain, 1882

Peregriana Servain 1882: 56.

Type species. Buccinum peregrum O.F. Muller, 1774.

17. Peregriana peregra (O.F. Miiller, 1774)
Figs 7M; 8G; Table 2

Buccinum peregrum O.F. Miller 1774: 130.

Lymnaea peregra — Hubendick 1951: 146, figs 1, 9 (partim).

Lymnaea (Peregriana) peregra — Kruglov and Starobogatov 1983b:
1465, fig. 2(2); 1993b: 161, fig. 1B.

Zoosyst. Evol. 96 (2) 2020, 577-608 595

Figure 9. Shells of species of the genera Ampullaceana, Kamtschaticana, Myxas, Tibetoradix, Racesina, Orientogalba and Bullas-
tra. A. Ampullaceana relicta relicta (05.1975, Macedonia, a spring near St. Naum monastery, Ohrid Lake region, NHMW). B. A.
r. pinteri, a paratype (05.06.1973, Macedonia, Prespa Lake near Perovo, MHMW). C, D. Kamtschaticana kamtschatica (Russia,
Kamchatka Peninsula, Valley of Geysers; RMBH). E. Myxas glutinosa (14.09.1963, Kazakhstan, floodplain of the Tobol River near
Kustanay City; ZISP). K, G. Tibetoradix hookeri, two syntypes (NHMUK). H. T kozlovi, the holotype (ZISP). I. Racesina luteola
(India, Bengal, Paksay, Pabna; NHMUK). J. R. oxiana (04.07.2016, Tajikistan, a roadside ditch near Kurban-Shakhid village;
LMBI). K. R. siamensis (“Thailand”; ZMB). L. Orientogalba viridis, a syntype (Guam Island; NMNH). M. O. o/lula (1883, Japan,
Yokohama; NHMUK). N. Limnaea bowelli, a type (after Preston 1909). O. Orientogalba cf. bowelli (30.06.1924, China, Tibet,
Lingka, 14,500 feet a.s.1.; NHMUK). P. Bullastra cumingiana (Philippine, Cebu Island; NHMUK). Scale bars: 2 mm (C, D, F—H,
L, M, O), 5 mm (A, B, E, J-K, N, P).

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596

Radix labiata — Falkner et al. 2002: 94; Gloer 2002: 216, fig. 244; Wel-
ter-Schultes 2012: 53, textfig.; Schniebs et al. 2013: 59, figs 4-8.;
Gloer 2019: 240, fig. 299.

Radix (Peregriana) peregra — Vinarski and Kantor 2016: 326.

Peregriana peregra — Aksenova et al. 201 8a: 4.

TL. Denmark, Copenhagen, Frederiksberg Park, in
swamps (Vinarski and Kantor 2016).

Types. Lost (Nekhaev et al. 2015).

For morphological and molecular characterisation of
this species, see Schniebs et al. (2013) and Vinarski et al.
(2016). P. peregra inhabits Europe (except of the northern
part), the Urals and the southwest part of Western Siberia
(Gloer 2002, 2019; Khokhutkin et al. 2009; Andreeva et
al. 2010; Schniebs et al. 2013; Vinarski et al. 2016). In
many recent publications, this species has been referred
to as Radix labiata (Rossmassler, 1835).

Nomenclatorial note. Falkner et al. (2002) argued that
the lymnaeid species, which had been commonly named
Radix (or Lymnaea) peregra by the European authors,
does not occur in Denmark, the type country of Muller’s
Buccinum peregrum. These authors treated B. peregrum
as a junior synonym of Helix balthica Linnaeus, 1758
and proposed the name Radix labiata for designation of
R. peregra auct. Such authors as Gloer (2002, 2019) and
Welter-Schultes (2012) followed it, but Vinarski (2017)
doubted this decision. According to him, Falkner et al.
(2002) did not present the total evidence for the absence
of R. peregra auct. from Denmark and their assumption
still needs strong confirmation. The type specimens of
Limnaeus pereger var. labiatus (kept in NHMW) were
considered by Vinarski (2017) as juvenile individuals
of R. balthica. Therefore, it was unnecessary to replace
a well-established taxonomic name R. peregra with a
long-forgotten one, R. /abiata.

18. Peregriana dolgini (Gundrizer & Starobogatov,
1979)
Figs 7L; 8F; Table 2

Lymnaea peregra — Hubendick 1951: 146, figs 1, 9 (partim).

Lymnaea dolgini Gundrizer and Starobogatov 1979: 1132, figs 1(2); 2(2).

Lymnaea kurejkae Gundrizer and Starobogatov 1979: 1131, fig. 1(1).

Lymnaea gundrizeri Kruglov and Starobogatov 1983a: 141.

Lymnaea napasica Kruglov and Starobogatov 1983a: 140.

Lymnaea ulaganica Kruglov and Starobogatov 1983a: 141.

Lymnaea (Peregriana) dolgini — Kruglov and Starobogatov 1993b: 170,
fig. 7G.

Lymnaea (Peregriana) gundrizeri — Kruglov and Starobogatov 1993b:
164, fig. 5A.

Lymnaea (Peregriana) kurejkae — Kruglov and Starobogatov 1993b:
170, fig. 7F.

Lymnaea (Peregriana) napasica — Kruglov and Starobogatov 1993b:
169, fig. 7C.

Lymnaea (Peregriana) ulaganica — Kruglov and Starobogatov 1993b:
164, fig. SB.

Radix (Peregriana) dolgini — Vinarski and Kantor 2016: 332 (partim).

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Vinarski, M.V. et al.: A survey of radicine snails (Gastropoda, Lymnaeidae)

Radix (Peregriana) gundrizeri — Vinarski and Kantor 2016: 333 (partim).
Radix (Peregriana) ulaganica — Vinarski and Kantor 2016: 333 (partim).
Radix dolgini — Vinarski et al. 2016: 26, figs 2, 3; Gloer 2019: 239, fig. 297.
Peregriana dolgini — Aksenova et al. 201 8a: 4.

TL. Russia, Krasnoyarsk Territory, a lake in the flood-
plain of the Kureika River, 20 km upstream of its mouth.

Type series. ZISP (see Sitnikova et al. 2014).

Vinarski et al. (2016) presented a full morphological
and molecular account for this species, as well as the data
on its range and a comparison with conchologically simi-
lar radicine taxa. P. dolgini is a species sister to P. peregra
and, probably, represents its vicariant taxon in Northern
Asia (Siberia). The molecularly-confirmed recordings of
P. dolgini are also known from the north-eastern part of
Europe — the Pechora River basin (Aksenova et al. 201 8b).

Genus Kamitschaticana Kruglov & Starobogatov, 1984

Kamtschaticana Kruglov and Starobogatov 1984: 30.

Type species. Limnaeus kamtschaticus Middendorff, 1850.

This taxon was introduced as a ‘section’ of the subge-
nus Peregriana (Kruglov and Starobogatov 1984). Akse-
nova et al. (2018a) elevated it to the genus level.

19. Kamtschaticana kamtschatica (Middendorff, 1850)
Figs 8H; 9C, D; Table 3

Limnaeus kamtschaticus Middendorff 1850: 110; 1851: 295, pl. 30,
figs 11, 12.

Limnaea ovata var. aberrans Westerlund 1897: 125.

Limnaea peregra var. middendorffi W. Dybowski 1904: 52, fig. 7.

Lymnaea peregra — Hubendick 1951: 146, figs 1, 9 (partim).

Lymnaea (Peregriana) kamtschatica — Kruglov and Starobogatov 1984:
30, figs 1(18), 2(7); 1993b: 164, fig. 3A.

Lymnaea (Peregriana) aberrans — Kruglov and Starobogatov 1984: 31,
figs 1(21), 2(3); 1993b: 164, fig. 3E.

Lymnaea (Peregriana) middendorffi — Kruglov and Starobogatov 1984:
31, figs 1(19), 2(5); Kruglov and Starobogatov 1993b: 164, fig. 3D.

Radix (Peregriana) kamtschatica — Vinarski and Kantor 2016: 334; Ak-
senova et al. 2016: 20, fig. II.

Radix (Peregriana) aberrans — Vinarski and Kantor 2016: 327; Akseno-
va et al. 2016: 19, fig. 1H.

Radix (Peregriana) middendorffi — Vinarski and Kantor 2016: 335; Ak-
senova et al. 2016: 21, fig. 1D.

Kamtschaticana kamtschatica — Aksenova et al. 2018a: 4.

TL. Russia, Kamchatka Peninsula, Kamchatka River
(Kruglov and Starobogatov [1984] suggest that the type
specimens were collected from the floodplain of the river).

Lectotype. ZISP (see Kruglov and Starobogatov 1984
for details).

Aksenova et al. (2016) presented a detailed description
of morphology of this snail, accompanied by a molecu-
lar analysis of its phylogenetic and taxonomic affinities.

597

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598 Vinarski, M.V. et al.: A survey of radicine snails (Gastropoda, Lymnaeidae)

Conchologically and anatomically, K. kamtschatica 1s
very similar to other members of the genera Ampullacea-
na and Peregriana and it is virtually impossible to pro-
pose a differential diagnosis for this genus. K. kamtschat-
ica 1s widely distributed throughout north-eastern Asia,
including Kamchatka and Chukotka peninsulas, Trans-
baikalia and the Amur River basin (Kruglov and Starobo-
gatov 1993b; Aksenova et al. 2016). Possibly, this snail
is endemic to Asiatic Russia, though we cannot exclude
that it also inhabits Alaska. In Kamchatka, it forms sta-
ble populations in the geothermal sites (e.g. the Valley
of Geysers) and may live in warm water up to +39.9 °C
(Aksenova et al. 2016).

Genus TJibetoradix Bolotov, Vinarski & Aksenova, 2018

Tibetoradix Aksenova et al. 2018a: 11.

Type species. Lymnaea hookeri Reeve, 1850.

Von Oheimb et al. (2011) have shown that the Tibetan
Plateau is a hotspot of lymnaeid diversity in High Asia,
with several species-level clades of the genus Radix s. lato
restricted to this region. Aksenova et al. (2018a) established
a new radicine genus, 7ibetoradix, to contain six species
endemic to Tibet, four of which remain undescribed.

20. Tibetoradix hookeri (Reeve, 1850)
Figs 81; 9F, G

Lymnaea hookeri Reeve 1850: 49.

Limnaea hookeri — G.B. Sowerby II 1872: pl. XI, fig. 74.

Limnaea hookeri — Annandale and Rao 1925: 167, figs V(7), [X (6).
Lymnaea (Galba) hookeri — Subba Rao 1989: 132.

Tibetoradix hookeri — Aksenova et al. 2018a: 12, figs 6F; 7E, H.

TL. “Thibetan or north side of Sikkim Himalaya, at
18,000 feet elevation” [Reeve, 1850: 49].

Syntypes. NHMUK (examined by us). A syntype
shell was illustrated by Hubendick (1951) and Aksenova
et al. (2018a).

Shell medium-sized (the largest of the two syntypes 1s
16.9 mm high), with high spire and moderately-inflated
body whorl. The shell proportions of 7. hookeri resemble
those of A. /agotis. The copulatory apparatus is typical for
radicines: broad and relatively compact praeputium and
very thin and narrow penis sheath (see Fig. 81); the ratio
of their lengths ranges from 1.25—1.50.

T. hookeri is endemic to China. It occurs in Tibet,
known from the upstream section of the Lhasa River and
a single additional locality (Brahmaputra River basin),
altitude range: 4,540-4,980 m. (Aksenova et al. 201 8a).

Taxonomic remark. Kruglov and Starobogatov
(1993b) classified this species as belonging to the (sub)
genus Orientogalba Kruglov et Starobogatov, 1985 and
treated it as a senior synonym of Lymnaea heptapotami-
ca Lazareva, 1967, described from southern Kazakhstan.

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The examination of the syntypes of L. hookeri has shown
that L. heptapotamica should not be synonymised with
the former species and may well represent a separate tax-
on, not related to the genus Tibetoradix.

21. Tibetoradix kozlovi Vinarski, Bolotov & Aksenova,
2018
Figs 8J; 9H.

Tibetoradix kozlovi Aksenova et al. 2018a: 11, fig. 7A—D.

TL. China, Central Tibet, the floodplain of the Requ Qu
River, Yellow River basin, 33°35'20.7"N, 103°05'30.2"E,
alt. 3,470 m.
Types. ZISP (holotype, paratypes), RMBH (paratypes).
This snail, endemic to Tibet, is fully described in our
previous paper (Aksenova et al. 2018a), therefore, we
give here only pictures of its shell and copulatory organ.

Genus Myxas G. B. Sowerby I, 1822

Myxas G.B. Sowerby I 1822: part vii.
Amphipeplea Nilsson 1822: 58.

Type species. Buccinum glutinosum O.F. Muller, 1774.

22. Myxas glutinosa (O.F. Miller, 1774)
Fig. 9E; Table 3.

Buccinum glutinosum O.F. Miller 1774: 129.

Amphipeplea dupuyi Locard 1893: 30.

Amphipeplea mabillei Locard 1893: 30.

Lymnaea glutinosa — Hubendick 1951: 148, fig. 333.

Lymnaea (Myxas) dupuyi — Kruglov and Starobogatov 1985b: 74,
figs 1(3), 2(3); 1993b: 171, fig. 9D (partim).

Lymnaea (Myxas) glutinosa — Kruglov and Starobogatov 1985b: 73,
figs 1(2), 2(2); 1993b: 171, fig. 9B (partim).

Lymnaea (Myxas) mabillei — Kruglov and Starobogatov 1985b: 71,
figs 1(1), 2(1); 1993b: 171, fig. 9A (partim).

Myxas glutinosa — Gloer 2002: 220, fig. 248; Welter-Schultes 2012: 50,
textfig.; Vinarski and Kantor 2016: 316; Gloer 2019: 249, fig. 310.

TL. Not stated in the original description. Most probably,
the type locality should be quoted as Fridrichsdal, a sub-
urb of Copenhagen, Denmark (see Nekhaev et al. 2015
for details).

Types. Lost (Nekhaev et al. 2015).

The taxonomic position and identity of this morpho-
logically-peculiar species have not raised many doubts
and most authors treated it more or less identically (Hu-
bendick 1951; Gloer 2002; Welter-Schultes 2012; but
see Kruglov and Starobogatov 1993b). The results of our
study well correspond to the commonly-accepted con-
cept of M. glutinosa. Shell of this species is very frag-
ile, semi-pellucid and, in a living animal, it is completely

Zoosyst. Evol. 96 (2) 2020, 577-608

covered by the reflected mantle. Shell shape 1s almost
globose, with greatly expanded aperture and very dimin-
ished spire (in some specimens, it is almost invisible).

The structure of the copulatory apparatus of MZ. g/u-
tinosa, aS 1s described by various authors (Hubendick
1951; Kruglov and Starobogatov 1985b; Jackiewicz
1998), is virtually indistinguishable from that of the gen-
era Ampullaceana and Peregriana. However, the length
of the spermathecal duct is different (long in Myxas, very
short or almost absent in Ampullaceana and Peregriana).

The range of M. glutinosa covers Europe (except of
the southern and northern parts), the Urals, Western and
Central Siberia (Khokhutkin et al. 2009; Welter-Schultes
2012; Vinarski et al. 2013).

Genus Racesina Vinarski & Bolotov, 2018

Racesina Vinarski and Bolotov 2018: 332.

Type species. Lymnaea luteola Lamarck, 1822.

Three species included into Racesina by Vinarski and
Bolotov (2018), were previously treated as members
of the (sub-) genus Cerasina Kobelt, 1881 (Subba Rao
1989; Kruglov 2005; Aksenova et al. 2018a). As Vinarski
and Bolotov (2018) have shown, the genus Cerasina sen-
su Kobelt is a junior synonym of Radix and a new generic
name was proposed by these authors for Cerasina sensu
auct. non Kobelt. The type species of the genus inhabits
India and some adjacent countries. Despite some serious
morphological peculiarities of Cerasina sensu Kruglov
and Starobogatov (1993b), the validity of this taxon was
rejected by most authors dealing with the South Asian
malacofauna (Annandale and Rao 1925; Hubendick 1951;
Brandt 1974; Subba Rao 1989). Vinarski (2013; Vinarski
and Kantor 2016) classified Cerasina as a separate genus
within his subfamily Radicinae (see also Zhadin 1952).
The results of the Aksenova et al. (2018a) molecular tax-
onomic study confirmed this opinion. The prostate with
numerous (5-8) internal folds (Hubendick 1951; Kruglov
2005) and a single synapomorphy to sharply distinguish
Racesina from all other genera of radicine snails are dis-
cussed in this paper.

23. Racesina luteola (Lamarck, 1822)
Fig. 91

Lymnaea luteola Lamarck 1822: 160.

Limnaeus cerasum Troschel 1837: 170.

Limnaeus impurus Troschel 1837: 172.

Limnaeus nucleus Troschel 1837: 171.

Limnaeus prunum Troschel 1837: 170.

Limnaea tigrina Dohrn 1858: 134.

Limnaea ovalior Annandale et al. 1921: 572, 573, fig. 13A; pl. VII,
figs 4-6; Annandale and Rao 1925: 186.

Limnaea (Pseudosuccinea) luteola — Annandale and Rao 1925: 183,
fig. [V(1-7), IX.

599

Limnaea (Pseudosuccinea) luteola f. impura — Annandale and Rao
1925: 184, fig. IV(7).

Limnaea (Pseudosuccinea) luteola f. australis — Annandale and Rao
1925: 184, fig. IV(3).

Limnaea (Pseudosuccinea) luteola f. ovalis — Annandale and Rao 1925:
184, fig. [V(4).

Limnaea (Pseudosuccinea) luteola f. succinea — Annandale and Rao
1925: 184, fig. [V(2).

Lymnaea (Radix) luteola — Brandt 1974: 232, pl. 16, fig. 98 (partim).

Lymnaea (Pseudosuccinea) luteola— Subba Rao 1989: 128, figs 275-287.

Lymnaea (Pseudosuccinea) ovalior — Subba Rao 1989: 129, fig. 285.

Lymnaea (Cerasina) luteola — Kruglov and Starobogatov 1993a: 85,
fig. 10C.

Lymnaea (Cerasina) impura — Kruglov and Starobogatov 1993a: 85,
fig. 10D, non Troschel 1837 (partim).

Cerasina luteola — Aksenova et al. 2018a: 4.

Racesina luteola — Vinarski and Bolotov 2018: 331.

TL. India, Bengalia.

Types. Not traced, probably in Muséum d’ Histoire
Naturelle, Geneve, Switzerland.

A highly-variable species, with several intraspecific
‘morphs’ differing from each other by their shell shape
and proportions (Annandale and Rao 1925; Subba Rao
1989). The typical morph (see Fig. 91) has ovate shell
with more or less prominent and wide spire and weak-
ly-inflated body whorl. Aperture is moderately expanded.
Shell surface is smooth and glossy. We had no fixed spec-
imens of R. /uteola for dissection; however, according to
Hubendick (1951) and Kruglov (2005), the genital mor-
phology of this species generally corresponds to that of
the genus Radix, except for the drastically different inter-
nal structure of the prostate. Penis sheath is very narrow
as compared to thick and oblong praeputium, ICA > 1.00
(Hubendick 1951).

R. luteola inhabits India and Nepal (Aksenova et al.
2018a). The recordings of this species from other coun-
tries such as China (south), Sri Lanka and the Andamanes
(Brandt 1974; Subba Rao 1989) need a molecular re-eval-
uation. The findings of “Lymnaea luteola’ in Thailand
and Myanmar (Brandt 1974) belong most probably to R.
siamensis (Sowerby).

24. Racesina oxiana (Boettger, 1889)
Figs 8K; 9J; Table 3

Limnaeus impurus vat. oxiana Boettger 1889: 961, Taf. 27, figs 4, 5.

Cerasina luteola var. oxiana — Zhadin 1952: 177, fig. 79.

Lymnaea (Cerasina) impura — Kruglov and Starobogatov 1993a: 85,
fig. 10D, non Troschel 1837 (partim).

Cerasina impura — Vinarski and Kantor 2016: 338, non Troschel 1837.

Cerasina oxiana — Aksenova et al. 2018a: 4.

Racesina oxiana — Vinarski and Bolotov 2018: 332.

TL. Turkmenistan, middle Amu-Darya River, ‘Tschard-
schui’ (nowadays Turkmenabat).
Types. Not traced.

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600 Vinarski, M.V. et al.: A survey of radicine snails (Gastropoda, Lymnaeidae)

As compared to R. /uteola, this snail is characterised
by a higher spire and ovate-conical shape of shell. In all
other respects, it is conchologically very similar to the
type species of the genus. The copulatory apparatus of R.
oxiana specimens from Tajikistan, dissected by us, was
virtually identical with that of R. /uteola from India stud-
ied and illustrated by Hubendick (1951). Penis sheath 1s
very narrow, its length is almost equal to length of prae-
putium or slightly less (see Fig. 8K).

This species of Racesina is distributed in Central Asia
(within the ex-USSR boundaries) and Nepal (Akseno-
va et al. 2018a). The oldest available name, based on a
type locality situated within this area, is Limnaeus im-
purus var. oxiana Boettger, 1889. Though Kruglov and
Starobogatov (1993b) treated this taxon as identical with
Lymnaea impura, the type locality of the latter species is
in India, where R. oxiana does not occur. L. impura 1s,
most probably, a junior synonym of R. /uteola (Huben-
dick 1951; Brandt 1974).

25. Racesina siamensis (G.B. Sowerby II, 1872)
Figs 8L; 9K; Table 3

Limnaea siamensis G.B. Sowerby II 1872: pl. X, fig. 63.

Limnaea luteola f. siamensis — Annandale and Rao 1925: 185, fig. IV(5, 6).
Lymnaea (Radix) luteola — Brandt 1974: 232, pl. 16, fig. 98 (partim).
Cerasina siamensis — Aksenova et al. 2018a: 4.

Racesina siamensis — Vinarski and Bolotov 2018: 332.

TL. “Siam” (= Thailand).

Types. Not traced.

Aksenova et al. (2018a) used this name as the oldest
one applicable to a species of Racesina which is distrib-
uted in Myanmar and Thailand. Though Brandt (1974)
considered L. siamensis as a junior synonym of Radix
rubiginosa, our data show it is untenable. Despite the
substantial genetic distance separating Racesina siamen-
sis and R. luteola, we were not able to find prominent
morphological differences between them, either in con-
chological or in anatomical characters. It allows us to
treat R. siamensis as a ‘cryptic’ taxon. Note, however,
that we had only a limited number of specimens from
Myanmar and some stable morphological differences be-
tween the two species will probably be found in future
after a thorough study of their intraspecific variation.
There is a strong need for a detailed study of genetic
diversity and phylogeography of the Racesina luteola
species complex.

Morphological note. In a sample of eleven individ-
uals of R. siamensis from Myanmar, dissected by us,
five snails had under-developed copulatory apparatuses
or lacked them altogether. No signs of a heavy parasite
load were seen, thus the hypothesis of the parasitic cas-
tration may be rejected. Such a state (aphally) has been
registered in different families of freshwater pulmonates,
including Physidae, Planorbidae and Lymnaeidae (see Vi-
narski and Palatov 2018 and references therein). For the
radicines, aphally was reported by Arutyunova (1977).

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Aphally in aquatic pulmonates is often viewed as an indi-
rect sign of obligate self-fertilisation (autogamy), which
may facilitate their introduction into new habitats and
rapid establishment of new populations (Beriozkina and
Starobogatov 1991; Jarne et al. 1993). Due to a limited
sample size, it is unclear whether our finding indicates
that R. siamensis 1s a species prone to self-fertilisation.
However, it is a remarkable fact that the aphally was not
found during this research in the rest of the radicine spe-
cies studied anatomically.

Genus Orientogalba Kruglov & Starobogatov, 1985

Orientogalba Kruglov and Starobogatov 1985a: 28.
Viridigalba Kruglov and Starobogatov 1985a: 29.

Type species. Lymnaea heptapotamica Lazareva, 1967.

Kruglov and Starobogatov (1985a) established their
subgenus Lymnaea (Orientogalba) to embrace five nom-
inal lymnaeid species distributed in a vast Pacific Re-
gion, stretching from north-eastern Asia to the Guam and
Hawai. Though, conchologically, these snails resemble
representatives of the genus Galba, their genital mor-
phology is typical for the radicines (Kruglov 2005). The
molecular analysis conducted by Aksenova et al. (2018a)
has shown that there are at least three biological species
within Orientogalba, which are presumably allopatric: O.
viridis (Indonesia and Pacific Islands), O. ollula (Far East
and, probably, some regions of Central Asia) and O. cf.
bowelli (known from Sichuan, China and, probably, also
inhabiting Tibet). The actual range of these species re-
mains unknown since only a limited set of sequences has
been available. The true species content of Orientogalba
is unknown as well, because some nominal species of this
genus, such as Orientogalba heptapotamica (Lazareva)
and O. lenaensis (Kruglov et Starobogatov, 1985) have
not been studied molecularly. The true taxonomic posi-
tion of “Austropeplea viridis’, recently recorded from
Spain as an alien species (Schniebs et al. 2017), must also
be clarified.

In our opinion, the genus Orientogalba is not a junior
synonym of the genus Austropeplea Cotton, 1942, as
was suggested by some authors (Ponder and Waterhouse
1997; Schniebs et al. 2017).

26. Orientogalba viridis (Quoy & Gaimard, 1832)
Fig. 9L

Lymnaea viridis Quoy and Gaimard 1832: 204, pl. 58, figs 16-18.

Lymnaea viridis — Hubendick 1951: 162, fig. 351 (partim).

Lymnaea (Radix) viridis — Brandt 1974: 231, pl. 16, fig. 97.

Lymnaea (Orientogalba) viridis — Kruglov and Starobogatov 1993b:
174, fig. 10C.

Orientogalba viridis — Aksenova et al. 201 8a: 4; Gloer 2019: 246, fig. 307.

TL. The Pacific, Marian Archipelago, Guam Island.
Syntypes. MNHN (examined by us).

Zoosyst. Evol. 96 (2) 2020, 577-608

Our concept of this species is based on the type series
of O. viridis (see Fig. 9L). Shell is almost globose, with
relatively wide and short spire and heavily-inflated body
whorl. Aksenova et al. (2018a) recorded O. viridis from
Indonesia. Numerous recordings of this species from oth-
er areas, including China, Mongolia, Thailand, Siberia,
Spain and some others (Hubendick 1951; Brandt 1974;
Kruglov and Starobogatov 1985a, 1993b; Schniebs et al.
2017; Vinarski et al. 2017), were based chiefly on mor-
phological evidence and, therefore, the actual range of O.
viridis sensu Aksenova et al. (201 8a) is unclear.

27. Orientogalba ollula (Gould, 1859)
Figs 8M; 9N; Table 3

Limnaea ollula Gould 1859: 40.

Limnaeus pervius von Martens 1867: 221.

Lymnaea viridis — Hubendick 1951: 162, fig. 351 (partim).

Galba pervia — Zhadin 1952: 176, fig. 77.

Lymnaea (Orientogalba) ollula — Kruglov and Starobogatov 1993b:
175, fig. 10C.

Orientogalba ollula — Vinarski and Kantor 2016: 357; Aksenova et al.
2018a: 4.

TL. China, streams and marshes of Hong Kong Island.

Lectotype. National Museum of Natural History, Smith-
sonian Institution, Washington, USA (Johnson 1964).

Though the type material of L. ollula Gould is extant,
we were unable to study it. Instead, we examined a small
series of shells of this species collected in Yokohama, Japan
(NHMUK). Shell is ovoid, with shortened spire and strong-
ly-inflated body whorls. Generally, it much resembles the
shell of O. viridis and we could not delimit these two taxa
by means of conchology. Like most species of radicines
discussed above, O. ollula is characterised by the dispro-
portion in sizes between the penis sheath and praeputium,
the latter being much larger and broader (see Fig. 8M).

Taxonomic remark. This species, described from
eastern China, has usually been considered as a junior
synonym of O. viridis (Hubendick 1951; Brandt 1974);
however, Kruglov and Starobogatov (1993b) regarded it
as a Separate taxon. The results of our study allowed us to
accept their opinion and use the name O. ollula to label
a species of Orientogalba, widely distributed in China
(found also in South Korea and Nepal). Perhaps, all re-
cordings of “Lymnaea viridis” from Siberia, Russian Far
East and Mongolia (Kruglov and Starobogatov 1985a,
1993b; Vinarski and Kantor 2016; Vinarski et al. 2017)
should also be referred to as O. ollula.

28. Orientogalba cf. bowelli (Preston, 1909)
Fig. 9N, O

Limnaea bowelli Preston 1909: 115, fig. 1.

Limnaea bowelli — Annandale and Rao 1925: 169, fig. V(6).
Lymnaea (Galba) bowelli — Subba Rao 1989: 131, figs 298, 299.
Orientogalba cf. bowelli — Aksenova et al. 2018a: 4.

601

TL. Tibet, “Te-ring Gompa, in a small hill stream arising
from a spring, 14,000 feet: also from Mangtsa, 14,500
feet; High Hill, Gompa, Gyantse valley in a small hill
stream, among moss and stones, 14,500 feet; and Gyan-
ise=-f3s120sfeete

Types. Zoological Survey of India, Kolkata (fide Sub-
ba Rao 1989).

This poorly-studied species inhabits China: Tibet, Si-
chuan Province (Subba Rao 1989; Aksenova et al. 2018a)
and possibly lives in other parts of the High Asia (Kruglov
and Starobogatov 1985a; Kruglov 2005). Conchologically,
it represents a typical Orientogalba and the traits allowing
one to distinguish it surely from the two above-mentioned
species of this genus are unknown. A special study of in-
traspecific variation in O. cf. bowelli and allied species is
needed to delineate them on the basis of morphological
data. The data on anatomy of O. cf. bowelli are scarce.
Though Kruglov (2005) gave a rather detailed description
of its genital morphology, the picture provided by this au-
thor (Kruglov 2005: fig. 90) illustrates an animal belong-
ing to the genus Galba, not Orientogalba.

Taxonomic remark. The application of the binomen
Limnaea bowelli Preston is difficult. Recent authors tend
to consider it as amember of the (sub)genus Galba, whose
distribution is restricted to High Asia (Subba Rao 1989;
Kruglov 2005; Vinarski and Kantor 2016). On the other
hand, Hubendick (1951) hypothesised that Limnaea bow-
elli is identical with Radix auricularia, whereas Brandt
(1974) synonymised L. bowelli with L. viridis. The type
series was unavailable for Aksenova et al. (201 8a) and the
authors conditionally attached the Preston’s taxon name
to a species of Orientogalba found in Sichuan Province
of China. Having examined several small samples of this
species kept in NHMUK and NHMN, we found that these
shells may belong to Orientogalba; some of them are very
similar to shells of O. viridis, whereas others were more
or less like Preston’s original drawing (see Fig. 9N, O).

Genus Bullastra Bergh, 1901

Bullastra Bergh 1901: 254.

Type species. Bullastra velutinoides Bergh, 1901.

A genus with South Asian — Australasian distribution.
It includes several nominal species living in Philippines,
Indonesia and mainland Australia (Kruglov and Star-
obogatov 1993a; Puslednik et al. 2009). Aksenova et al.
(2018a) have studied one of these species.

28. Bullastra cumingiana (L. Pfeiffer, 1845)
Figs 8N; 9P; Table 3

Amphipeplea cumingiana Pfeiffer 1845: 68.

Amphipeplea cumingi Pfeiffer 1854-1860: 5, pl. U, figs 3, 4.
Bullastra velutinoides Bergh 1901: 254, pl. 20, figs 22-34.
Lymnaea cumingiana — Hubendick 1951: 162, fig. 355.
Bullastra cumingiana — Aksenova et al. 201 8a: 4.

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602 Vinarski, M.V. et al.: A survey of radicine snails (Gastropoda, Lymnaeidae)

TL. Philippines, island of Luzon, Naga, province of
South Camerines.

Probable syntypes. ZMB (see Vinarski 2016a
for details).

Hubendick (1951) provided a description of shell, rad-
ula and genital morphology of this peculiar species. By
its almost globose shell, with extremely reduced spire and
extended and reflected mantle border, B. cumingiana su-
perficially resembles M. glutinosa, however, the alleged
phylogenetic affinity between the two snails has been re-
jected by Hubendick (1951) and Kruglov (2005).

Two specimens of B. cumingiana originated from
Lake Sampaloc (Philippines, Luzon Island; LMBI collec-
tion) were dissected during this study. The structure of the
copulatory apparatus is typical for radicines. Praeputium
is enlarged and widened, its length is around twice that
of the penis sheath length (see Fig. 8N). Penis sheath is
short and narrow. The ICA values of the two dissected in-
dividuals were 2.11 and 2.13. Both parts of the copulatory
apparatus are light-coloured.

B. cumingiana is endemic to the Philippine Islands
(Hubendick 1951).

Discussion

In this article, we attempted to present the taxonomic ac-
counts for all species of the Old World radicines geneti-
cally delineated up to now. The 35 biological species of
radicine snails recovered during our molecular taxonom-
ic study (Aksenova et al. 201 8a) by no means exhaust the
overall diversity of extant species in this group. There
are morphologically-distinct species of the radicines still
not studied genetically and their actual taxonomic status
and phylogenetic affinities remain unclear. Such species
of Central Asia as Radix obliquata (Martens, 1864) and
R. cucunorica (Mollendorff, 1902) may be mentioned
here as two examples. In our opinion, the actual glob-

al taxonomic richness of the radicines may approach 50
valid species.

These molluscs exhibit an impressive variation in
their shell characters, including shell size, shape, num-
ber of whorls, aperture proportions and so on (see Figs
2, 4). Strikingly, this wide repertoire of shell forms is
not accompanied by any substantial anatomical diver-
sity. As we tried to show, the structure of the copulato-
ry apparatus remains virtually the same within the Old
World radicines (see Figs 3, 7). Although the species
slightly differ from each other by the ICA values, the
overall morphological scheme of this organ 1s identical
within the group. Though Kruglov (2005) used the ICA
as a tool for delineation and identification of radicine
species, the usefulness of this ratio for lymnaeid sys-
tematics is sometimes questioned. Some authors consid-
ered its intraspecific variation too high to provide sig-
nificant and reproducible results (Schniebs et al. 2011,
2013). Our results have shown that there are no hiatuses
in the ICA values of sister species (see Tables 1—3) and,
thus, this ratio is almost useless for species diagnostics.
Though the species may differ by some qualitative char-
acters, such as pigmentation of praeputium (compare
colouration of this organ in two sister species, A. fontin-
alis and A. lagotis: Fig. 8D, E), we do not have informa-
tion how widely this trait varies amongst populations of
the same species.

A comparison of conchological, anatomical and karyo-
logical traits of the radicine genera discussed in Aksenova
et al. (201 8a) and this paper is given in Table 4. The struc-
ture of the copulatory organ within radicines is so uniform
that it led some researchers to a drastic reduction of the
number of valid species they agreed to accept (see, for ex-
ample, Hubendick 1951; Jackiewicz 1998). On the other
hand, there were malacologists with a strong inclination to
species splitting. For example, Kruglov and Starobogatov
(1993a, b) delineated more than 90 nominal radicine spe-
cies within the Northern Eurasia alone. The recent works,

Table 4. A comparison of conchological, anatomical and caryological features of the genera and subgenera of the Old World radicines*.

Genus Shell height Shell shape Spermathecal Mantle Haploid Number of
(Subgenus) duct border chromosome prostate
number folds
Racesina Up to 25 mm}| Ovoid to ovate-conical Smooth | High and relatively ?Reflected ? 5-8
wide
Myxas Globose Flattened, very low Reflected 1
Radix s. str. Up to 35 mm] Ear-shaped to ovate- | Sculptured Low and acute Not reflected
conical

Exsertiana Up to 30 mm Ovate-conical
Ampullaceana |Up to 25 mm) Ear-shaped to conical Low to relatively

high and obtuse
Peregriana Up to 25 mm | Ovate-conical to conical Medium-sized and

obtuse

Kamtschaticana | Up to 20 mm | Ovate-conical to almost Low and obtuse

globose
Tibetoradix Up to 15 mm Ovate-conical High to medium-

sized, narrow and
obtuse

Orientogalba Ovate-conical to almost High to low, obtuse

globose
Bullastra Up to 30 mm Globose Smooth | Very low, reduced, Reflected

* After Aksenova et al. (201 8a), modified.

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obtuse

Zoosyst. Evol. 96 (2) 2020, 577-608

based on the integrative taxonomic approach, have shown
that many of these nominal taxa lack molecular support
and should be synonymised (Aksenova et al. 2016, 2017;
Vinarski et al. 2016). The ratio between valid and non-val-
id species in this group may be around 1:4 (Vinarski et al.
2016).

Unfortunately, due to overlap of morphological traits, tn-
cluding both shell and soft body characters, between close-
ly allied species of radicines (Schniebs et al. 2011, 2013),
to develop a dichotomous key for identification of snails
is next to impossible. The molecular methods are most re-
liable for species determination in this group (Schniebs et
al. 2011, 2013; Vinarski et al. 2016), though there are mor-
phologically-distinct taxa (such as Myxas glutinosa), whose
correct identification is possible on the basis of shell charac-
ters alone. The use of the reproductive anatomy characters,
such as the ICA, is very limited due to the relative unifor-
mity of the copulatory organ structure within the radicines.

Though the evolutionary processes in this group have
rarely been studied and discussed, the high species rich-
ness and endemism of Radicinae in the mountain part of
Central Asia is remarkable. This region forms an obvi-
ous hotspot of diversity of this group, with at least one
endemic genus (7ibetoradix), which includes not less
than six narrow-range species. In addition, at least four
species of the genus Radix (R. alticola, R. brevicauda, R.
makhrovi, Radix dgebuadzei) are restricted in their dis-
tribution to High Asia. This highlights the significance
of the region for evolution of the current diversity of
the subfamily Amphipepleinae and the probable role of
mountain refugia for speciation in lymnaeid snails (see
Aksenova et al. 2018a for details).

Acknowledgements

We thank museum curators, whose help greatly facilitat-
ed our work with collections: Dr. Pavel V. Kiyashko and
Mrs. Lidiya L. Yarokhnovich (ZISP), Dr. Anita Eschner
(NHMW), Dr. Jonathan Abblett (HNMUK), Dr. Ted von
Proschwitz (NMG) and Dr. Katrin Schniebs (SNSD).
We thank also Peter Gloer (Hetlingen, Germany) and
Katrin Schniebs for sending us some samples of lymnae-
ids (Bullastra cumingiana, Peregriana peregra) used in
this study. The four referees (Peter Gloer, Zoltan Fehér,
Thomas Wilke and an anonymous one) are acknowledged
for their critical comments on the earlier versions of the
text. The writing of this paper was supported by the Rus-
sian Scientific Fund (project No. 19-14-00066). Some
molecular analyses and some fieldwork were made with
financial support of the Russian Ministry of Science and
Higher Education (projects Nos. MK-1720.2019.4 and
0409-2019-0042 (AAAA-A17-117033010132-2). The
Russian Foundation for Basic Research (project Nos. 19-
04-00270_a and 20-04-00361_a) supported the examina-
tion of museum collections. The work in Hungarian Nat-
ural History Museum was made possible by a grant from
the Synthesys+ program (project No. HU-TAF 2697)

603

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