Marine Biology (2000) 137: 827833

Springer-Verlag 2000

T. Wilke E. Rolan G. M. Davis

The mudsnail genus Hydrobia s.s. in the northern Atlantic

and western Mediterranean: a phylogenetic hypothesis

Received: 6 October 1999 / Accepted: 12 September 2000

Abstract The mudsnail taxon Hydrobia s.l. plays a

major role in numerous ecological and parasitological
studies. Yet, its taxonomy is confusing and its biogeography is poorly understood. A principal problem is
that taxa of Hydrobia s.l. belong to morphostatic radiations widely allopatric species radiations with little or
no habitat diversication and a low level of anatomical
change. So far, three morphostatic radiations can be
identied within Hydrobia s.l.: Hydrobia s.s., Peringia,
and Ventrosia. These radiations are identical with the
putative genera of Hydrobia s.l. The purpose of the paper is to infer phylogenetic relationships within one of
these radiations, Hydrobia s.s. in the northern Atlantic
and western Mediterranean. A total of 22 populations,
including topotypical populations of ve commonly
used taxa, were studied genetically using partial sequences of the mitochondrial cytochrome c oxidase
subunit I gene. Based on molecular data and the species
concept we dene, the following species and subspecies
were identied: Hydrobia acuta acuta (Draparnaud,
1805) [=H. minoricensis (Paladilhe, 1875)], H. acuta
neglecta Muus, 1963, and H. glyca (Servain, 1880)
(=H. joossei van Aartsen, Menkhorst and Gittenberger,
1984). One population from Djerba, Tunisia (Hydrobia
sp. A) and one population from Mallorca, Spain (Hydrobia sp. B) are genetically distinct from all other taxa
studied here. Both populations potentially represent new
species. Due to the fact that only one population each
has been studied, and so the degree of variation within
Communicated by J. P. Grassle, New Brunswick
T. Wilke (&) G. M. Davis
The Academy of Natural Sciences,
Department of Malacology,
1900 Benjamin Franklin Parkway,
Philadelphia, PA 19103, USA
e-mail: twilke@acnatsci.org
E. Rolan
Canovas del Castillo 22-5F,
36202 Vigo, Spain

these two taxa cannot be assessed, a nal decision regarding their species level cannot yet be made.

Introduction
Hydrobia s.l. (Hydrobiidae: Rissooidea) is a taxon of
minute, brackish-water to marine snails. The species are
primarily amphi-Atlantic, occurring in coastal estuaries,
salt ponds, and salt marshes of the northern Atlantic,
the Arctic Ocean, and the Baltic Sea, the Mediterranean,
Black, and Caspian Seas.
Hydrobia s.l. is used internationally as a paradigm for
numerous and diverse ecological and parasitological
studies: many species are major hosts for parasites infecting sh, birds, and seals; they usually occur in very
high densities (e.g. 300,000 m)2; Linke 1939) in intertidal settings as major deposit feeders and as a source of
food for birds and other predators. Many Hydrobia spp.
populations are endangered, as they inhabit coastal
habitats that are particularly aected by pollution and
ecological changes. In many cases, the species involved
are unknown to ecologists (and often even to malacologists), making ecological, parasitological, and conservation studies dicult.
A principal problem is that taxa of Hydrobia belong
to morphostatic species radiations, that are, according
to Davis (1993), `` considerable, rapid speciation[s]
with low anatomical diversication there is little or no
habitat diversication, speciation is widely allopatric,
and there are low levels of anatomical change''. According to our anatomical data, there are three distinct
groups within Hydrobia s.l., each with a morphostatic
species radiation: Hydrobia s.s. (sensu Giusti et al.
1998a, b; non sensu Boeters 1984), Ventrosia, and Peringia. These radiations correspond to the putative genera
of Hydrobia s.l. (see Haase 1993; Giusti et al. 1998a).
Whereas dierent radiations are anatomically distinct, taxa within a radiation are almost identical.
Species within a radiation occur in what appear to be
similar habitats, and have until now not been found to

828

be sympatric (whereas Hydrobia s.l. species belonging to

dierent radiations frequently are sympatric, e.g. Peringia ulvae and Ventrosia ventrosa).
The relative anatomical uniformity and the lack of
clear habitat dierentiation make an objective species
assignment of morphostatic taxa very dicult. Moreover, shell characters and certain anatomical characters
(mainly from the digestive and reproductive system) are
aected by parasitism and environmental factors, such
as substratum and water chemistry. The resulting variation within and among populations may obscure negraded variation among species.
Accordingly, there is much confusion concerning species status. There were, for example, many Hydrobia
species described from the Mediterranean basin in the
nineteenth century. Most of these species are known only
from the type locality; the original shell-based descriptions are vague, and the types, if available, are shells only.
Almost all of these taxa are considered synonyms of one
or another of three rissooidean species: Hydrobia acuta
(Draparnaud, 1805), Ventrosia ventrosa (Montagu, 1803),
and ``Heleobia'' stagnorum (Gmelin, 1791) (Sabelli et al.
1990; Le Renard 1997). The systematics of the few ecologically well-studied Atlantic taxa of Hydrobia s.l. also
are unclear, e.g. the possible conspecicity of the north
European Hydrobia neglecta and the Mediterranean H.
acuta. Although most authors consider them distinct
species, some workers (M. Haase, pers. comm.; Hoeksema 1998) believe, that on the basis of shell and preliminary anatomical data, they are the same. A similar
problem applies for Hydrobia minoricensis (Paladilhe,
Table 1 Hydrobia spp.
Populations of nominal species
studied (*, topotypical
populations). Names of the
regions are underlined (CL,
locality code; NS, number of
specimens sequenced)

1875), described from the island of Menorca (Mediterranean). Almost all workers (e.g. Boeters 1987; Sabelli et al.
1990; Le Renard 1997) consider this species conspecic
either with the North Atlantic Peringia ulvae (Pennant,
1777) or Hydrobia glyca (Servain, 1880), or with the
Mediterranean H. acuta.
Recent studies of Hydrobia s.s. recognize between 1
and 12 species (Nordsieck 1982; Giusti and Pezzoli 1984;
Boeters 1987; Sabelli et al. 1990; Anistratenko and Stadnichenko 1994; Le Renard 1997; Macedo et al. 1999).
These data reect two dierent trends in systematics: (1)
merging taxa that are anatomically similar into a single
taxon, either at or above the species level, using categories like ``morphospecies'' (Giusti and Pezzoli 1984) or
``species complex'' (Giusti and Manganelli 1992) or (2)
assigning species names to slight variation in shell and
anatomical characters (e.g. Nordsieck 1982; Anistratenko and Stadnichenko 1994).
For the northern Atlantic and western Mediterranean
there are currently ve taxa of Hydrobia s.s. ``commonly
in use'' (see Table 1). In the present study we used
partial mitochondrial cytochrome c oxidase subunit I
(COI) gene sequences to infer phylogenetic relationships
within Hydrobia s.s. in the northern Atlantic and western Mediterranean. COI sequences have been shown to
dierentiate reliably among rissooidean taxa from the
superfamily to population level (Davis et al. 1998, 1999;
Hershler et al. 1999; Wilke and Davis 2000; Wilke et al.
in press) and to be congruent with anatomical and/or
allozyme data (Davis et al. 1999; Hershler et al. 1999).
In rissooidean systematics COI data also have been

Locality information

CL

NS

Hydrobia acuta (Draparnaud, 1805)

*Etang du Prevost, Palavas-les-Flots, Herault, France (43.513N; 3.897E)
Patanella, Laguna di Orbetello di Ponente, Grosseto, Italy (42.45N; 11.22E)
Punta del Sabinar (Salinas de Cerrillos), Almer a, Spain (36.699N; 2.673W)
Los Alcazares, Mar Menor, Murcia, Spain (37.73N; 0.85W)
Tunis (Lac de Tunis), Tunis, Tunisia (36.80822N; 10.26293E)
Djerba (Sebkha de Sidi Garous), Medenine, Tunisia (33.82N; 11.06E)

EP
Pa
PS
LA
Tu
Dj

6
4
4
6
4
5

Hydrobia glyca (Servain, 1880)

*San Fernando (San Francisco de As s lagoon), Cadiz, Spain (36.3937N; 6.1375W)
*San Fernando (Salina San Miguel), Cadiz, Spain (36.460N; 6.221W)
Portimao (Salinas Mexilhoeira de Carrigacao), Faro, Portugal (37.141N; 8.529W)
Huelva (near the Casa de Damaso), Huelva, Spain (37.255N; 7.143W)
Faro, Faro, Portugal (37.016N; 7.946W)
O Grove Bay, Pontevedra, Spain (42.474N; 8.855W)
Lostrouc'h, Finistere, France (48.634N; 4.537W) (=Lostrouc'h site in Barnes 1991)
Plouharnel, Morbihan, France (47.583N; 3.126W) (=``site 2'' in Hoeksema 1998)

SF-a
SF-b
Po
Hu
Fa
OG
Lo
Pl

6
3
6
7
6
6
6
6

Hydrobia joossei van Aartsen, Menkhorst and Gittenberger, 1984

*Laredo (mouth of the Rio Ason), Cantabria, Spain (43.422N; 3.449W)
San Vicente de la Barquera, Cantabria, Spain (43.390N; 4.382W)

La
SV

6
6

Hydrobia minoricensis (Paladilhe, 1875)

*Puerto de Mahon (Colarsega), Menorca, Balearic Islands, Spain (39.896N; 4.261E) PM
Puerto de Pollenca, Mallorca, Balearic Islands, Spain (39.8987N; 3.0820E)
PP

6
5

Hydrobia neglecta Muus, 1963

*Ajstrup Bugt, Mariager Fjord, Jutland, Denmark (56.68N; 10.22E)
Odense Fjord, Fyn Island, Fyn, Denmark (55.50N; 10.53W)
Boiensdorfer Werder, Mecklenburg, Germany (54.0326N; 11.5241E)
Shingle Street (lagoon 6 south), Suolk, Great Britain (52.0390N; 1.4650E)

6
6
3
6

AB
OF
Bo
SS

829

largely congruent with the data from other genes, e.g.

16 S rRNA and 18 S rRNA (Wilke et al. in press). For
these reasons the COI gene appears to be well-suited for
studies of morphostatic radiations in hydrobiid snails.
The present study is not intended to be a formal revision of the genus Hydrobia s.s. More data from the
southern and western Mediterranean, the Black Sea, and
the Caspian Sea basins are needed, and more nominal
species would have to be studied based on morphological comparisons of type material and topotypical populations. Rather, the study establishes a phylogenetic
hypothesis of relationships between morphostatic taxa
that we do consider species based on COI sequencing
data, and the species concept we dene.

Materials and methods

amplify a 658 bp fragment of the COI gene were those of Folmer

et al. (1994). Vent (exo)) DNA polymerase was used for the PCR.
Amplied DNA products were separated by electrophoresis
through a 1% LMP agarose gel. Bands corresponding to fragments
of the correct size were cut out, the slices melted, and the DNA
puried with Wizard PCR preps. COI sequences (forward and
reverse) were determined using the LI-COR DNA sequencer Long
ReadIR 4200 and the Thermo Sequenase uorescent labeled
primer cycle sequencing kit (Amersham) according to the manufacturer's protocols.
A total of 119 individuals were sequenced from 22 populations.
The relatively high number of specimens studied was necessary to:
(a) assess variation within and among populations, (b) to test for
potential sympatry of species from the same radiation, and (c) for
future studies of gene ow.
Sequences for the protein-coding COI gene were aligned by eye
using ESEE 3.0s (Cabot and Beckenbach 1989). The rst 210 bp
beyond each primer were often rich in ambiguities. We therefore
uniformly excluded the rst and last 10 bp of each sequence,
leaving a 638 bp fragment.

Populations studied

Phylogenetic analyses

The study includes ve previously described nominal species of

Hydrobia s.s.: Hydrobia acuta (Draparnaud, 1805); Hydrobia glyca
(Servain, 1880); Hydrobia joossei van Aartsen, Menkhorst and
Gittenberger, 1984; Hydrobia minoricensis (Paladilhe, 1875) and
Hydrobia neglecta Muus, 1963 (Table 1). As there is a considerable
degree of confusion in assigning non-topotypical populations to
described taxa, our study is based on topotypical material. In addition to topotypical populations we studied 17 populations of the
nominal taxa mentioned above throughout the area to access the
degree of infraspecic variation (Table 1; Fig. 1).

In preliminary analyses for this study both parsimony and maximum likelihood methods were used to infer phylogenetic relationships within Hydrobia s.s. In all cases, they yielded the same tree
topology (except for some unresolved branches with the parsimony
method). We chose the maximum likelihood method because it
shows one of the best performances of all major phylogenetic
methods (Huelsenbeck 1995). It also incorporates a sophisticated
algorithm for calculating branch lengths in the phylogenetic trees so
that cluster patterns and gaps between the clusters can be inferred.
To nd the optimal transition:transversion ratio for the sequencing analyses in Hydrobia s.s., a series of maximum likelihood
trees was generated using transition:transversion ratios from 1:1 to
10:1; each had the same topology. As the best log likelihood value
was observed in the tree with a ratio of 3:1, this ratio was used for
the analyses.
In order to reduce the extensive data set to a manageable size
for the maximum likelihood analysis, but still be able to demonstrate the degree of variability within and among populations, we
used sequences of the two haplotypes that showed the highest degree of nucleotide diversity within each of the 22 populations.
These 44 sequences were submitted to GenBank (accession nos. AF213344, AF247791AF247807, and AF278798AF278822).
A maximum likelihood tree was constructed using DNAML of
PHYLIP 3.57 (Felsenstein 1993) with the following parameters:
search for best tree, transition:transversion ratio 3.0, empirical base
frequencies (A: 0.22328; C: 0.18153; G: 0.21374; T: 0.38145), one
category of substitution, global rearrangements, and randomized
input order of sequences (25 repetitions). Ventrosia ventrosa and
Peringia ulvae (GenBank accession nos. AF118333 and AF118288)
were used as outgroups.
Bootstrap estimates (200 replicates; no global rearrangement)
were made using program SEQBOOT, in conjunction with
DNAML and CONSENSE of PHYLIP 3.57. For this computationally extensive task, the data set was further reduced to 30 sequences by excluding sequences that diered by only one or two
nucleotides.
The average pairwise number of nucleotide dierences (k), the
nucleotide diversity (p) within groups, and the nucleotide divergence (Dxy) between groups, using the full set of 119 sequences,
were determined with the software program DnaSP 3.0 (Rozas and
Rozas 1999).

DNA isolation and sequencing

The methods of Spolsky et al. (1996) and Davis et al. (1998) were
used for isolating DNA from individual snails. The primers to

Species concept
Fig. 1 Hydrobia spp. Locality information for the populations
studied (Table 1)

The species concept we use to characterize taxa within morphostatic radiations is based on the cohesion species concept of
Templeton (1989): a species is a group of populations of a mono-

830
phyletic lineage which, in sexually reproducing species, have a
species-specic recognition system, the reproductive system. These
populations share the same fundamental niche, i.e. the populations
are potentially interchangeable. Because this study is based on
molecular data, the following criteria were applied to distinguish
morphostatic species: monophyly, cluster patterns in phylogenetic
trees, genetic distances (indicating clear gaps between clusters, a
predetermined distance value cannot be equated with a taxonomic
level), and geographic partitioning of mitochondrial haplotypes.

Results
COI sequencing data
The maximum likelihood tree given in Fig. 2 shows four
main clades within Hydrobia s.s. Clade I comprises all
populations of Hydrobia glyca and H. joossei. There are
three subclades within clade I. The rst subclade consists of some of the specimens from the two topotypical
populations of H. glyca from San Fernando. The second
subclade, which clusters together with the third subclade, comprises the other part of the specimens of topotypical H. glyca, specimens from Huelva, and some
specimens from neighboring Faro. The third subclade
consists of the other specimens from Faro and all
specimens of H. glyca and H. joossei north of Faro. The
second main clade (clade II) consists of the two haplotypes from a population of H. acuta from Djerba (Tunisia). Clade III includes all populations of H. acuta and
H. neglecta as well as topotypical H. minoricensis. The
populations of nominal H. neglecta form a subclade,
distinct from a second subclade consisting of haplotypes
of topotypical H. minoricensis and H. acuta. Clade IV
comprises the second population of H. minoricensis from
Mallorca, Balearic Islands (non-topotypical).
The nucleotide diversity within the clades ranges from
0.00188 (clade II) to 0.00853 (clade I). The nucleotide
divergence between clades varies from 0.03022 (clade III/
clade IV) to 0.05549 (clade II/clade IV) (Table 2).
A bootstrap analysis revealed that two of the main
clades and some of the subclades have relatively low
bootstrap support (Fig. 2). To test the robustness of the
tree, we have analyzed the overall tree structure of 200
pseudotrees generated by nonparametric bootstrap resampling (Felsenstein 1985). These pseudotrees revealed
that the four main clades are distinct, and can be recognized in each of the 200 trees. However, among these
clades ``branch hopping'' is common and partially
causes low bootstrap support. In 56% of the trees, clade IV is basal to clade III that is basal to clade I and
clade II (Fig. 2). In 16% of the trees, clade II is basal to
clade IV that is basal to clade I and clade III. The other
28% of the pseudotrees show nine other combinations
of the four main clades with frequencies of 17%.
Similar eects were found among the subclades. They
are very distinct, but subject to branch hopping, which
decreases bootstrap support.
Problems with branch hopping in rissooidean systematics using sequencing data have been reported before

(Davis et al. 1998; Wilke et al. in press). Apparently, the

problem occurs in taxa that are distinguished by few
synapomorphies (short internodes). Dierent phylogenies are equally likely and/or represented by similar
short trees. Branch hopping is problematic when
studying the course of evolution. Because we are inferring species-level relationships, it has little eect on the
hypotheses in this paper.

Discussion
Species determined
Hydrobia acuta acuta (=H. minoricensis),
H. acuta neglecta (clade III)
Genetically, populations of nominal H. acuta, H. neglecta, and H. minoricensis cluster together and form a
clade that is distinct from the other taxa studied. Within
this clade, topotypical H. minoricensis cannot be separated from populations of H. acuta. However, one
population from Puerto de Pollensa (Mallorca), which
hitherto was considered to belong to nominal H. minoricensis, formed a separate clade (IV) distinct from the
other taxa in this study.
Within the H. acuta/neglecta/minoricensis clade,
nominal H. neglecta forms a subclade, distinct from
H. acuta. This genetic dierentiation is likely due to
current geographical separation. Nominal H. neglecta
occurs in northwestern Europe, whereas H. acuta can
be found in the Mediterranean. The average geographic
distance between these taxa of 4,000 km of ocean
possibly accounts for the genetic dierentiation of
about 1.4%.
Based on these data, we regard H. neglecta and
H. minoricensis as conspecic with H. acuta. The small
but distinct genetic dierences between nominal H.
acuta and H. neglecta, together with their geographical
separation, may justify considering them subspecies of
the same taxon: H. acuta acuta and H. acuta neglecta.
Hydrobia glyca (=H. joossei) (clade I)
Based on our sequencing data, nominal H. glyca and
H. joossei are conspecic. Within the H. glyca/joossei
clade we have found three genetically distinct groups
(Fig. 2). Two groups were detected in the Bay of Cadiz,
the southern border of the range of H. glyca. A third
group comprises specimens from Faro (Portugal) and all
northern populations of H. glyca, including topotypical
H. joossei. However, we do not presently consider
H. joossei to be a distinct subspecies of H. glyca because
their ranges overlap (e.g. in Faro) and there is an even
higher degree of genetic dierentiation within and between populations from the Bay of Cadiz which cannot
be explained by geographic distance.

831
Fig. 2 Hydrobia spp. Maximum likelihood tree for a
638 bp fragment of the COI
gene. Ventrosia ventrosa and
Peringia ulvae were used as
outgroups. Scale bar indicates
expected numbers of substitutions (0.01 1%). Roman numerals represent the four major
clades discussed. Bootstrap
values (%) are given for major
nodes

Table 2 Hydrobia spp. Average number of nucleotide dierences (k) within (diagonal line) and between (below diagonal line) clades.
Nucleotide diversity (p) and nucleotide divergence (Dxy) in parentheses

Clade
Clade
Clade
Clade

I (n = 58)
II (n = 5)
III (n = 51)
IV (n = 5)

Clade I
H. glyca; H. joossei

Clade II
Hydrobia sp. A

Clade III H. acuta;

H. neglecta; H. minoricensis

Clade IV
Hydrobia sp. B

5.442
30.948
23.652
31.579

1.200 (0.00188)
34.384 (0.05389)
35.400 (0.05549)

4.648 (0.00729)
19.282 (0.03022)

2.400 (0.00376)

(0.00853)
(0.04851)
(0.03707)
(0.04950)

The sample we obtained from Huelva (Spain) contained two sympatric species: H. glyca and H. acuta in a
ratio of about 7:2 (here we present only data for the
specimens of H. glyca, the data for the specimens of
H. acuta will be presented elsewhere). This is the rst
time that we have found two distinct Hydrobia species of
the same radiation in sympatry. We do not know
whether this is a stable, long-term sympatry or if some

specimens were introduced from the Mediterranean

(about 190 km away) to this locality by chance.
Hydrobia sp. A (clade II)
The population from Djerba (Tunisia) is genetically
distinct from all other taxa studied. It clustered apart

832

from H. acuta and H. glyca, although it is apparently

more closely related to H. glyca than to H. acuta. The
average genetic distance between these taxa (Hydrobia
sp. A vs. H. glyca 5.02%; Hydrobia sp. A vs. H. acuta 5.57%) is much higher than we have found within
species of Hydrobia s.l. Thus, Ventrosia ventrosa shows
an average infraspecic variation of 0.28% and Peringia
ulvae 1.21% (Wilke and Davis 2000). Based on our COI
sequencing data, the population from Djerba possibly
belongs to a new species. However, data from more
populations would be necessary to assess the degree of
intra- and interpopulation variation and to establish a
well-founded, species-level hypothesis.
Hydrobia sp. B (clade IV)
The COI sequencing data show that one population of
``H. minoricensis'' from Mallorca is genetically distinct
from topotypical H. minoricensis from neighboring
Menorca (about 100 km of ocean away). In fact, in the
maximum likelihood tree it forms its own clade, distinct
from all other taxa (Fig. 2). The genetic distance to
other Hydrobia spp. ranges from 3.02% (H. acuta) to
5.55% (Hydrobia sp. A) (Table 2). These numbers are
well above typical infraspecic variations within
hydrobiid taxa. The faunas of Mallorca and Menorca
are generally very similar. Considering the relatively
high capacity of passive dispersal in Hydrobia species
(e.g. oating, transport by water birds; see Wilke and
Davis 2000 for details) compared with the relatively
small geographic distance between Mallorca and Menorca, founder eects as a possible explanation are unlikely. Moreover, the relatively high genetic diversity
within the population from Mallorca (0.63%) would not
support the hypothesis of founder eects. Data from
more populations from the area would be necessary to
solve the problem.
Biogeographical implications
We propose the following distribution patterns for
Hydrobia spp. in the northern Atlantic and western
Mediterranean (Fig. 3). The distribution of H. acuta is
divided into two segments. H. acuta acuta occurs in the
central and western Mediterranean, including Spain,
France, Italy, and northern Tunisia. It likely occurs
also on the Mediterranean coasts of Algeria and Morocco although records from there still need conrmation. H. acuta neglecta occurs in the eastern part of the
English Channel, the North Sea, the Baltic Sea, and the
Norwegian Sea. H. glyca occurs in the area between the
southern and northern ranges of H. acuta. The northern border of its range is not exactly known, but based
on the general distribution patterns of benthic fauna in
the English Channel (Cabioch et al. 1977), it may extend from southwest of Manche (France) along the
Atlantic coasts of France, northern Spain, Portugal,

Fig. 3 Hydrobia spp. Hypothesized distributions in the northern

Atlantic and western Mediterranean. Question marks indicate areas
where records still have to be conrmed

and southern Spain to the Strait of Gibraltar. Here

there is a sharp boundary just northeast of Tarifa,
where H. acuta occurs (based on unpublished 16 S
rRNA sequencing data). Hydrobia sp. A is known, thus
far, from the island of Djerba (Tunisia). It does not
seem to occur in northern Tunisia or other parts of the
western Mediterranean, but it likely occurs in other
parts of the Gulf of Gabes and perhaps eastwards as
far as Libya or even Egypt. Hydrobia sp. B also has a
very restricted range, being reported only on the island
of Mallorca. It does not occur on the neighboring island of Menorca.
Acknowledgements This study was in part supported by a grant
of the German Academic Exchange Service (DAAD) to T. Wilke,
in part by N.I.H. grants I P50 AI3946 (the Tropical Medical
Research Center, Shanghai, China) and AI 11373 (G.M. Davis),
and in part by grant DGES PB85-0235 (``Proyecto FAUNA'', E.
Rolan). We thank Prof. F. Giusti (University of Siena, Italy), Dr.
A. Falniowski (Jagiellonian University, Krakow, Poland), and
three anonymous reviewers for their critical and helpful comments
on an earlier version of the manuscript. The study could not have
been done without the kind support of many ecologists and
malacologists who provided samples or assistance in collecting,
including Dr. R. Barnes (University of Cambridge, United
Kingdom), Dr. L. Cabioch (Station Biologique de Rosco,
France), Mr. J. Quintana Cardona (Menorca, Spain), Dr. C.
Casagranda (University of Marseille, France), Dr. P. Drake (Instituto de Ciencias Marinas de Andaluc a, Spain), Dr. T. Jensen
(University of Aarhus, Denmark), Dr. G. Manganelli (University
of Siena, Italy), Mr. A. Penas Mediavilla (Vilanova, Spain), Mr.
L. Murillo (Cartagena, Spain), and Dr. U. Walter (Biological
Station Boiensdorf, Germany).

833

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