ABSTRACT
Freshwater snails
of the family Lymnaeidae are of a great parasitological importance because of
the very numerous helminth species they transmit, mainly trematodiases of large
medical and veterinary impact. The present knowledge on the genetics of lymnaeids
and on their parasite-host inter-relationships is far from being sufficient.
The family is immersed in a systematic-taxonomic confusion. The necessity for
a tool which enables species distinction and population characterization is
evident. This paper aims to review the European Lymnaeidae basing on the second
internal transcribed spacer ITS-2 of the nuclear ribosomal DNA. The ITS-2 sequences
of 66 popu-lations of 13 European and 1 North American lymnaeid species, including
the five generic (or subgeneric) taxa Lymnaea sensu stricto, Stagnicola, Omphiscola,
Radix and Galba, have been obtained. The ITS-2 proves to be a useful marker
for resolving supraspecific, specific and population relationships in Lymnaeidae.
Three different groupings according to their ITS-2 length could be distinguished:
Radix and Galba may be considered the oldest taxa (370-406 bp lengths), and
Lymnaea s. str., European Stagnicola and Omphiscola (468-491 bp lengths) the
most recent, American Stagnicola and Hinkleyia being intermediate (434-450 bp
lengths). This hypothesis agrees with the phylogeny of lymnaeids based on palaeontological
data, chromosome numbers and radular dentition. ITS-2 sequences present a conserved
central region flanked by two variable lateral regions corresponding to the
5..and 3..ends. The number of repeats of two microsatellites found in this conserved
central region allows to differentiate Radix from all other lymnaeids. Phylogenetic
trees showed four clades: (A) Lymnaea s. str., European Stagnicola and Omphiscola;
(B) Radix species; (C) Galba truncatula; and (D) North American stagnicolines.
ITS-2 results suggest that retaining Stagnicola as a subgenus of Lymnaea may
be the most appropriate and that genus status for Omphiscola is justified. Radix
shows a complexity suggesting different evolutionary lines, whereas G. truncatula
appears to be very homogeneous. North American and European stagnicolines do
not belong to the same supraspecific taxon; the genus Hinkleyia may be used
for the American stagnicolines. Genetic distances and sequence differences allowed
us to distinguish the upper limit to be expected within a single species and
to how different sister species may be. S. palustris, S. fuscus and S. corvus
proved to be valid species, but S. turricula may not be considered a species
independent from S. palustris. Marked nucleotide divergences and genetic distances
detected between different S. fuscus populations may be interpreted as a process
of geographic differentiation developping in the present. Among Radix, six valid
species could be distinguished: R. auricularia, R. ampla, R. peregra ( =R. ovata;
=R. balthica), R. labiata, R. lagotis and Radix sp. The information which the
ITS-2 marker furnishes is of applied interest concerning the molluscan host
specificity of the different trematode species. The phylogenetic trees inferred
from the ITS-2 sequences are able to differentiate between lymnaeids transmitting
and those non-transmitting fasciolids, as well as between those transmitting
F. hepatica and those transmitting F. gigantica. The Fasciola specificity is
linked to the two oldest genera which moreover cluster together in the phylogenetic
trees, suggesting an origin of the Fasciola ancestors related to the origin
of this branch. European Trichobilharzia species causing human dermatitis are
transmitted only by lymnaeids of the Radix and Lymnaea s. str.-Stagnicola groups.
Results suggest the convenience of reinvestigating compatibility differences
after accurate lymnaeid species classification by ITS-2 sequencing. Similarly,
ITS-2 sequencing would allow a step forward in the appropriate rearrangement
of the actual systematic confusion among echinostomatids. © 2001 Elsevier Science
B.V. All rights reserved.
ISSN 1567-1348
IF 0
NETHERLANDS