Ruthenica, 2013, vol. 23, No. 1: 41-58.
Published online April 5, 2013
© Ruthenica, 2013
http: www.ruthenica.com
ABSTRACT. The problem of generic classification of
the basommatophoran family Lymnaeidae Rafinesque,
1815 is reviewed as well as recent theoretical approach-
es to genus delineation in the zoological systematics.
Two main versions of the generic system of lymnaeid
snails are: 1) bigeneric approach where all diversity of
lymnaeid species is split between two genera; and 2)
polygeneric approach suggesting that there are several
(about twenty) genera in the family. The first version is
presented in systems developed by Kruglov and Staro-
bogatov [1993] and Jackiewicz [1993, 1998]. The sec-
ond one is most commonly accepted in current West-
ern European and Northern American literature [Burch,
1989; Falkner et al., 2001; Glöer, 2002]. However, there
are no grounds to delimit lymnaeid genera objectively
as the solution critically depends on what taxonomic
methodology (cladistic or ‘evolutionary’ taxonomy) is
followed by a particular author. The ‘evolutionary’ tax-
onomic methodology (sensu Mayr) is favorable to the
bigeneric approach, whereas the cladistic (Hennigian)
methodology leads to the separation of a series of taxa
of generic rank within Lymnaeidae. It is impossible to
prefer one approach to another ultimately since the
problem of acceptability of paraphyletic taxa is still not
resolved. The co-existence of two different generic sys-
tems of the same family is therefore inevitable. Different
criteria of generic rank were critically discussed in per-
spective of their applicability to lymnaeid taxonomy.
The morphological and ecological criteria as well as the
criterion of hybridizability proved to be controversial
and their use gives no key to select one of these ap-
proaches. The fourth criterion, that of monophyly, is
more usable now, when the data of molecular phyloge-
netics are available. The recent advances in molecular
taxonomy of pond snails have been reviewed briefly. It
seems very reliable that the family consists of two large
monophyletic clades of deep origin that differ from
each other by chromosome number albeit there are no
morphological characters to distinguish surely between
representatives of these clades. It is impracticable to
assign the generic rank for these clades due to their
huge internal heterogeneity (morphological and eco-
logical). The most reliable cladistic solution is to regard
the two deep lymnaeid clades as separate subfamilies
each containing a set of genera that are internally ho-
mogeneous enough to comply with most of criteria of
the genus rank. The scheme of lymnaeid classification
proposed here includes the nominotypical subfamily
Lymnaeinae (type genus Lymnaea Lamarck, 1799) with
haploid chromosome number equal to 18 (rarely 19),
and the new one Radicinae subfam.n. (type genus
Radix Montfort, 1810). The latter taxon embraces gen-
era and species of Lymnaeidae characterized by 16 or
(most often) 17 chromosome pairs. Radicinae is, most
probably, derived clade as compared to Lymnaeinae,
however, there are no morphological synapomorphies
to support it. The bigeneric system is, however, still
acceptable for those who uses generic criteria pro-
posed by “evolutionary systematics” such as “princi-
ple of the same degree of difference” [Golikov, Staro-
bogatov, 1988] and others.
Introduction
The basommatophoran family Lymnaeidae
Rafinesque, 1815 is a very diverse and almost glo-
bally distributed group of freshwater snails. The
global species diversity of this family is estimated
from nearly 100 [Strong et al., 2008] to not less
than 250 [Kruglov, 2005] extant species. There is,
however, no agreement among today’s systemati-
cists on how many species and higher taxa (genera,
subgenera) should be recognized within the family,
and several alternative systems were proposed in
the second half of the 20th century, i.e. before the
ongoing molecular revolution in the zoological sys-
tematics. The systems are those of Hubendick
[1951], Kruglov and Starobogatov [1993a, b; Kru-
glov, 2005], and Jackiewicz [1993, 1998]. In many
aspects these systems do not correspond to each
other and even are contradictory in some regards.
In particular, the question of how many lymnaeid
genera should be accepted has not been answered
unambiguously. Some researchers accept only two
genera and a plethora of subgenera within a large
cosmopolitan genus Lymnaea Lamarck, 1799 [Jack-
iewicz, 1998; Kruglov, Starobogatov, 1993a, b; Kru-
glov, 2005]. Other workers prefer to treat these
subgenera (including Lymnaea s.str.) as distinct
genera [Ponder, Waterhouse, 1997; Falkner et al.,
2001; Glöer, 2002]. As a result, we have as many as
three different systems on the generic level even for
the very thoroughly studied lymnaeids of Europe
One, two, or several? How many lymnaeid genera are there?
M.V. VINARSKI
Museum of Siberian Aquatic Molluscs, Omsk State Pedagogical University. 14 Tukhachevskogo
Emb., Omsk, Russian Federation, 644099. e-mail: radix.vinarski@gmail.com
urn:lsid:zoobank.org:pub:79A89EB9-8698-4B68-9136-6F59FE4AEB05