.31

HARVARD UNIVERSITY

Ernst Mayr Library

of the Museum of

Comparative Zoology

ON THE SPECIES COMPOSITION OF VIVIPARIDS
(Gastropoda, Viviparidae) IN EUROPE AND WESTERN, Agl^Z

by ^W 2 3

mz

E. V. Chernogorenko ^NlVFa^^

m

Zoological Journal, USSR Academy of Sciences, Moscow
1988, Volume 67(5), pp. 645-655, Figs. 1-4

O vidovom sostave viviparid (Gastropoda, Viviparidae)

Evropy i Zapadnoi Azii.

Zoologicheskii Zhurnal, Akademiya Nauk SSSR, Moskva
1988, Volume 67(5), pp. 645-655, Figs. 1-4

Edited and Annotated

by

Kenneth J. Boss

Special Occasional Publication

No. 11

published by the

Department of Mollusks

Museum of Comparative Zoology

Harvard University

Cambridge, Massachusetts 02138

1992

ON THE SPECIES COMPOSITION OF VIVIPARIDS
(GASTROPODA, VIVIPARIDAE) IN EUROPE AND WESTERN ASIAf

by E. V. Chernogorenko
Zoological Institute
Ukrainian Academy of Sciences (Kiev)

Abstract. This is a review of the contemporary species
of Viviparidae of Europe and western Asia. The family
is represented by two genera, Viviparus and
Contectiana. differing in the shape of the shell, the
structure of the radula, and the location of the egg
capsules in the uterus. The genus Viviparus consists
of eight species that fall into three groups. The
first one contains the type-species of the genus, V.
viviparus, as well as V. vistulae. V. ater, and V.
costae. The second one includes V. rossmaessleri (or
V. hunqaricus) . The third one consists of three
species from western Greece (V. hellenicus. V. blanci.
and V. qraecus sp. n.). The genus Contectiana is
represented by eight species falling into two groups:
1) thin-walled - "pond contectiana" - C. contecta and

* Translated through the auspices of the Smithsonian
Institution's Behind-the-Scenes-Volunteer Program and with the
cooperation of Dr. Robert Hershler, Division of Mollusks,
National Museum of Natural History, Smithsonian Institution,
Washington, DC 20560.

C. listeri (V. contectus auct.)» distinguished by their
relative height, and 2) thick-walled - "lake
contectiana" - C. turrita . C. fennica. C. kormosi. C.
ianinensis. C. zebra, and C. ladoqensis sp. n.

It is usually considered that the family Viviparidae is
represented in Europe and western Asia by a single genus of
Viviparus with a few species living today (Zhadin, 1952; Franz,
1932; Zilch, 1955 and, others). The most widely known are the V.
viviparus (L.) and V. contectus (Mill.)/ clearly distinguished by
the shape of their shell, particularly the type of whorls, by the
presence or absence of an umbilicus, and by the thickness of the
shell. Three more species closer to V. viviparus are known to
exist in southern Europe (Zilch, 1955) . Old authors considered
more species in the genus - anywhere from 18 to 37 (Bourguignat,
1880; Westerlund, 1886; Kobelt, 1909).

By identifying only several species of Viviparidae - just
two in the northern half of Europe, the researchers (for example
Zilch, 1955) experienced great difficulty in evaluating the
species to which lake forms with thick shells and bulging whorls
belong. Thus the species mentioned below as Contectiana fennica
is referable to V. viviparus (Zilch, 1955) and to V. contectus
(Kobelt, 1906-1909) . Furthermore in previously cited work,
Bourguignat (1880) identified seven groups within the generic
limits while Kobelt (1909) indicated their specific types.

The oldest descriptions of the species of this family are
very brief, making identification difficult, and which in their
time led to disputes (Watson, 1957) . Meanwhile, a number of
works (Kiister, 1852; Kobelt, 1877, 1906-1909; Zilch, 1955) gave a
more precise picture of the types of samples that are very close
to types based on shells and locality data, thereby enabling us
to utilize these works when making revisions.

Recently, Sitnikova and Starobogatov (1982) narrowed down
the size of the family, identifying the Bellamyidae and the
Lioplacidae as separate families. At the same time the family in
the modern fauna was represented only in the eastern USA where
for a long time two genera had been identified, and in Europe
(together with eastern Asia) where the aforementioned writers
also numbered two genera - Viviparus and Contectiana . Moreover,
apparently they were guided by the fact that morphological
differences on such a scale as those between V. viviparus and V.
contectus were unconditionally accepted by everyone as being of
generic value in the closely related family of the Bellamyidae.

The purpose of this work is to revise the generic and
specific composition of contemporary representatives of the
families of European and western Asian Viviparidae. A wide-range
of material has been utilized which is kept in the Zoological
Institute of the Ukrainian Academy of Sciences and in the
Zoological Institute of the Russian Academy of Sciences,

including several thousand samples of different ages and sizes
collected primarily in the territory of the [former] USSR. There
were small collections from western Europe that we could use for
comparisons, especially as some of them originated from the type-
localities of nominal species or at least from the same areas.

We used a comparative method (Izzatullayev and Starobogatov,
1984) in analyzing the shells, thereby enabling us to compare the
shells directly on the Raupian parameters (Raup, 1967) . This is
particularly important because usually the corrosion of viviparid
shells causes standard errors in measurements (especially in the
V. viviparus group) and their relationships become meaningless.
The picture obtained from standard measurements is very
distorted, and the changes inside the population from one of the
Raupian parameters - a step along the axis - is very
characteristic of the V. viviparus group.

Anatomical studies were made when we manually opened samples
"fixed" in alcohol and placed them under a stereoscopic
microscope. We also extracted the radulae and prepared
microscopic slides.

In discussing the distribution we used the system of

zoogeographic zoning of continental reservoirs suggested by

Starobogatov (1970, 1983) to whom I am deeply grateful for
assistance and advice in connection with this work.

GENERIC COMPOSITION OF CONTEMPORARY VIVIPARIDAE IN THE OLD WORLD

The study of radulae of Viviparus viviparus and V. contectus
auct. (= listeri Forbes and Hanley) as well as of some other
forms (V. contectus kormosi Kobelt, V. c. turrita Kobelt, V.
acerosus Bourguignat, and Contectiana ladogensis) described
below, has shown that on this basis the Viviparidae patently fall
into two groups. They retain the architaenioglossal type of
radulae (Sitnikova and Starbbogatov, 19 82) that are common to the
Viviparidae with the formula for the transverse row (see diagram")

M, — ,Vfi -

■R

J-M^-M,,

where M]_ and M2 are marginal teeth, J is the initial one [lateral
tooth] , R is the rachidial tooth and the vertical broken lines
represent longitudinal flexions. At the same time they radically
differ in the shape and number of cusps on the teeth. Thus, V.
viviparus and V. acerosus have a large middle tooth with a couple
of cusps on the side. Along the sides of J and H^. ^^® ^^° small
cusps while in R there are three, the medially situated one being
larger. M2 has five small teeth identical in size and shape
(Fig. la). In C. listeri- and C. contecta, on the other hand, as
well as those forms whose shells are close to them, the middle
tooth in R, J and M]_ is always rectangular without cusps on the
edges. Along their sides these teeth have four smaller cusps
smoothly decreasing in size toward the edges of the tooth. M2

has eight identical small teeth (Fig. lb) .

While the Viviparidae have a common structure in a very
simple sexual system, these two groups differ in that the egg
capsules in the uterus of the former are usually in a single row
(Fig. 2a) , whereas in the latter they are in several rows (Fig.
2b) .

All of the aforementioned compels us to agree with the
concept that the Old World Viviparidae should be grouped into two
genera: Viviparus Montfort, 1816 with the type-species Helix
vivipara L. , 1758 based on tautonomy , -'■ and the Contectiana
Bourguignat, 1880 with the type-species Cyclostoma contectum
Millet, 1813, subsequently referred to by Kobelt, 1906-19092.

STRUCTURE OF THE GENUS Viviparus

An analysis of the species structure, as already indicated,
is greatly complicated by corrosion of the shell's upper whorls
and by variations (within the population) of the whorl along-the-
axis. Consequently, situations arise where samples of V.
acerosus recognized by everyone as a separate species (under one
name or another) from the Soviet Black Sea littoral were also
identified a V. viviparus. As a result of a careful comparison
of all samples available to us it was possible to determine that

the species of Viviparus is represented in Europe and the
southern and western coasts of the Black Sea by three subgenera
and seven species.

Viviparus s. str.
Molluscs belonging to this subgenus have thick-walled shells
without any noticeable sculpture, with moderately protuberant
whorls and a convex tangential line.

V. viviparus (Linnaeus, 1758) (Fig. 3a). More than 3,000
samples are from European USSR. For a long time the use of this
name produced arguments which have now completely ended. In
accordance with the decision of the International Commission on
Zoological Nomenclature (Opinion 573, 1959) under this name we
now accept the form appropriate to the lectotype established by
Watson (1957) . Repeated attempts were made to divide this
species into subspecies. In our opinion, none of them are
successful, although most probably it will be possible to effect
such a division. However, special research is needed for this,
taking into account the changes not in standard measurements but
in Rauperian parameters. This is the most wide-spread species.
Its habitat encompasses all of Europe (except for the far north
and far south) , and the eastern and southern Black Sea littoral.

V. vistulae (Kobelt, 1907) (Fig. 3b) . Several dozen samples

are from the Vistula and Niemen basins. The species is
distinctly characterized in Kobelt's drawings (Kobelt, 1907: 24,
PI. 345, Figs. 214 3-2144) and in lectotype photographs (Zilch,
1955, PI. 3, Fig. 8). It differs from V. viviparus found in the
same locations by a much slower growth of the transverse section
of the shell. If one compares the cross-section of any of the
shell whorls of V. viviparus with those of V. vistulae one sees
that in V. vistulae the preceding turn is wider and the next one
narrower than those of V. viviparus. This signifies that the
shell of V. viviparus is either an oval or an elongated oval,
whereas that of the V. vistulae is a conically-shaped oval. The
difference in shape is caused by the fact that the latter species
tangent line of the upper whorls of the spire (but not the entire
shell) is almost straight while that of the V. viviparus is
distinctly curved or convex. As a rule, they have the same brown
stripes on the shell at almost the same distance from each other.
So far the species is known to exist only in the Vistula and
Niemen basins. Evidently, it is endemic to the Baltic area.

V. ater (Christophori and Jan, 1832) (Fig. 3c).
Several hundred samples are from the northern Black Sea littoral
and central Dnieper River. Nearly all researchers recognize this
as a separate species, but it is more widely known as V. acerosus
(Bgt.). A comparison of the drawing by Kiister (1852: 9, PI. 2,
Figs. 1-5; PI. 4, Fig. 5), recognized by all recent researchers
such as Zilch (1955) for being the most precise representation of

V. ater, with the samples of V. acerosus from Hungary and the
Black Sea coast, revealed that they are identical in accordance
with all the Raupian parameters, thereby compelling us to
classify both forms as a single species. It differs from the V.
viviparus by a more rapid growth of the transverse section of the
shell tube, consequently, the ratio of the subsequent whorls of
V. ater differs from that of V. viviparus just as the latter does
from V. vistulae. This makes the shell of V. ater more oval and
wider even though it has the same number of whorls. Moreover,
the visibly high whorls grow more quickly.

It is located in northern Italy, along the entire Danube
basin, and the eastern Black Sea littoral, at least as far as the
lower Dnieper. It is found sporadically in the central Dnieper
basin. (The Kiev reservoir is close to the mouth of the Teterev,
the Goryn River where it falls into the mouth of the Ust.
Collected Works of V. V. Polischchuk) . A totally unexpected find
of the species near Pyatigorsk was made by E. A. Kazannikov.
They possibly were carried down there, inasmuch as only a few
years before no Viviparidae were found in that region.
Apparently, V. ater is a more oxyphilous species than the V,
viviparus on the Dnieper rapids that existed previously.

V. costae (Mousson, 1863) (Fig. 3d) . Several samples are
from western Georgia. It differs from the V. viviparus by a
slower growth of the transverse section of the shell tube. At

the same time, more than in the previous three species, there is
a noticeable step along the axis which gives a dome-like shape to
the shell and a relatively more noticeable height to the whorls.
It is frequently found together with V. viviparus. Apparently,
it is endemic to the Black Sea littoral areas.

Viviparus (Hazayipaludina)
Chernogorenko and Starobogatov, new subgenus

The type, and only, species Viviparus hunaaricus (Hazay,
1881) = V. rossmaessleri Bgt. 1880. The shell is rather thin-
walled with a malleate sculpture, with moderately protuberant
whorls and a convex tangent line.

V. rossmaessleri (Bourguignat, 1880) (Fig. 3e) . Several
dozen samples are from the central Danube basin. It is widely
known in northern Italy under the name V. pyramidalis
Rossmaessler, 1835. However, Zilch (1955) showed that this name
was preoccupied inasmuch as it had been used in 1814 for another
viviparid species. The description and drawing are to be found
in Rossmaessler 's works (1835: 110; 1835a: 19, PI. 7, Fig. 125)
and photographs are in Zilch's work (1955, PI. 4, Fig. 13). In
contradistinction to all preceding species that could be
designated in the V. viviparus group, the shell of V.
rossmaessleri though sturdy, is noticeably more thin-walled and

10

even translucent. Moreover, the malleate sculpture is developed.

Another difference is that its whorls are more concentrated
causing it to resemble three of the species below in the V.
hellenicus group; but unlike them, the tangent-line of V.
rossmaessleri is very convex- This condition is found in shells
of the V. hunaaricus (Hazay) series kept in the Zoological
Institute of Science of the USSR Academy of Sciences, therefore
the latter name should be regarded as synonymous with V.
rossmaessleri and not V. acerosus as is customary in research
works (see, for example. Zilch 1955) . The species is widespread
in northern Italy and the central part of the Danube basin. In
the USSR it may be found in the Trans-Carpathian Region.

Viviparus (Balcanipaludina) Starobogatov, 1986 [1985]

The type-species is Paludina hellenica Clessin, 1879. The
shell is thick-walled without sculpture, with flat whorls and a
straight tangent line.

V. hellenicus (Clessin, 1879) (Fig. 3f ) . Several samples
are from Greece. This species, recognized as being independent
(i.e. Franz, 1932; Zilch, 1955), is distributed in northwestern
Greece and on the Ionian Islands.

11

V. blanci (Bourguignat, 1880) (Fig. 3g) . A few samples are
from Greece. The species is often considered to be an
intraspecif ic form of the preceding one, in a large part due to
contradictions in Bourguignat ' s description of it and the absence
of an original drawing. At the same time, Kobelt's drawing
(1906-1909: 350, Pi. 71, Fig. 34) and Westerlund's description
(1886: 11) unequivocally indicate the existence of the species
that differs from V. hellinicus by a more shapely shell and a
sharper apical angle. It is found in the same areas where the
preceding species is located.

V. qraecus Chernogorenko and Starobogatov, new species (Fig. 3h)

This species is named after the country where it is found.

Material. A holotype and one paratype from Goritsa
(Greece) . They are kept at the Zoological Institute of the
Academy of Sciences of the USSR (Leningrad) .

Description. The shell is almost exactly peg-top-shaped
with nearly flat whorls separated by very small sutures. The
shell's tangent line is almost straight, and the coloring is a
brownish monochrome. The last whorl is very big, somewhat over
0.8 higher than the rest of the shell; its periphery has a
sharply curved angle, somewhat less than a right angle. The

12

following are the measurements (in mm) of the holotype and the
paratype, respectively: shell height, 23.0 and 21.0; shell width,
17.0 and 16.7; the height of the aperture is 13.5 and 12.6; the
width of aperture is 9.2 and 9.0; the height of the spire is 11.7
and 11.0; the height of the last whorl is 19.5 and 18.1; the
width of the last whorl without the orifice is 14.5 and 13.8. As
far as it is possible to determine due to the corrosion, there
are 4.5 and 4.7 whorls, respectively.

Remarks . This species differs from the two preceding which
are closely related by a big apical angle and, therefore, a less
shapely shell. Apparently, this species is located in the same
areas where the two preceding ones are found, namely in
northwestern Greece and on the Ionian Islands.

TABLE FOR DETERMINING CONTEMPORARY
EURO-ASIAN SPECIES OF THE GENUS Viviparus

1. (6) The whorls are greatly flattened or completely flat; the
spire is conical, therefore, the shell's tangent line is straight
or almost straight. subgenus Balcanipaludina .

2. (3) The apical angle of the shell exceeds 69°. V. qraecus
sp.n.

3. (2) The apical angle of the shell is not greater than 65°.

13

4. (5) The apical angle of the shell does not exceed 56°. V.
blanci Bourg.

5. (4) The apical angle of the shell is not less than 57°. V.
hellenicus Clessin.

6. (1) The whorls are moderately protuberant; the spire is oval,
therefore the shell's tangent line is markedly convex.

7. (8) The shell is relatively thin-walled and translucent with
a clearly malleate sculpture. subgenus Hazayipaludina - V.
rossmaessleri Bourg.

8. (7) The shell has thick non-translucent walls and is without
malleate sculpture. subgenus Viviparus s. str.

9. (10) The height of the penultimate whorl (along the axis) is
not less than half of its width. V. costae Mouss.

10. (9) The height of the penultimate whorl (along the axis) is
less than half of its width.

11.(12) The height of the third- from-the-bottom whorl does not
exceed 0.38 of the height of the penultimate whorl, and in the
most spherical shells it does not exceed 0.30 of the height of
the penultimate whorl. V. ater Christ, and Jan.

14

I

12.(11) The height of the third-from-the bottom whorl is not less
than 0.4 0 of the height of the penultimate whorl, and in the most
spherical shells it is not less than 0.33 of the height of the
penultimate whorl.

13.(14) The tangent line of all upper whorls of the spire except
for the last one is straight or nearly straight. Brownish
stripes are at nearly the same distance from each other. V.
vistulae Kob.

14.(13) The tangent line of all upper whorls of the spire, except
the last one, are distinctly convex. Two brown stripes are
closer together. V. viviparus L.

STRUCTURE OF THE GENUS Contectiana

All forms relegated by us to this genus usually constituted
a single species V. contectus, except for C. ianinensis which was
often considered as the V. pyramidalis form. An analysis of the
extensive material revealed that there are eight independent
species within the structure of the genus Contectiana. The most
indicative was the material from Lake Rytoye (Demidovskiy Rayon
in northern Smolensk Oblast of the western Dvina basin) where
several hundred samples were divided into five species unrelated
to the transitional forms. A comparison of the samples was made

15

easier by the fact that, in contrast to the Viviparus species,
the step-forward along the axis in the Contectiana species is
insignificant. At the same time, the age changes in the shell
proportions are rather marked inasmuch as the tangent line in all
of the species is convex.

Contectiana s. str.

To the subgenus belong molluscs with thin-walled shells and
the whorls of the shell are with a marked shoulder.

C. contecta (Millet, 1813) (Fig. 4a) . Several hundred
samples are from the European part of the USSR. All researchers
(i.e. Kobelt, 1906-1909) take the convex form with a relatively
low spire as a typical form of this species. The species is
found throughout western Europe, except for the Mediterranean,
and in the central part of the Dnieper basin. The most eastern
find has been in the vicinity of Kharkov (collection of I. A.
Krinitskiy in the first half of the 19th century) . It is
impossible to prove the reliability of the find because even if
the species had lived there 150 years ago it could have
completely vanished by now. In general, its habitat may be
considered to be the Baltic, central Dnieper and the central
Danube areas .

16

C. listeri (Forbes and Hanley, 1835) . (Fig. 4b) . Several
hundred samples are from the European part of the USSR. The
species was isolated by Bourguignat (1880) under the name V.
lacustris Beck, however, Beck (according to Kobelt, 1906-1909)
proposed his own appellation - C. contecta instead of Lamarck's
Paludina vivipara. It is also known as V. contectus, V.
russiensis Milach. , and V. rossicus Kob. In native literature it
is referred to as V. contectus. It is distinguished from C.
contecta (also when found together) by a visibly higher spire..
It is the most wide-spread species of the genus. It inhabits all
of Europe except for the far north and the Mediterranean. In the
east it is found as far as the central Ob River basin.

Contectiana ( Kobelt ipaludina)
Chernogorenko and Starobogatov, new subgenus

The type-species of the subgenus is Vivipara contecta
turrita Kobelt (1909). Thick-walled molluscs and those with
moderately rounded whorls belong to this subgenus.

C. turrita (Kobelt, 1909) (Fig. 4c) . Several dozen samples
are from the Shatskiy Lakes (Volynsk Oblast, USSR) . The
applicability of this name is clearly demonstrated by Kobelt 's
drawing and description (1909: 8, PI. 394, Fig. 2263) and by the
photograph of the lectotype (Zilch, 1955: PI. 3, Fig. 1).

17

This species and all ensuing ones (the C. turrita group-
the lake contectiana) differ from the C. contecta and the C.
lister i (they can be lumped into one group - the C. contecta or
the pond contectiana) by far more evenly rounded whorls without a
seamed shoulder and a fold toward the axis at the lower seam.
Furthermore, it has a much more thick-walled shell, although not
to the same extent as most of Viviparus.

Evidently, it is endemic to the Baltic area; however, within
the USSR it is located only in the Shatskiy Lakes (western Bug
basin) .

C. ianinensis (Mousson, 1859) (Fig. 4d) . Based on research
work it belongs to the genus Contectiana because of the shape of
its shell. (Its anatomy was not studied). The species is
distinguished even from the C. turrita by a higher shell in the
adults and a very curved tangent line. It is located along the
western Balkan shores (northwestern Greece and southern Albania) .

C. fennica (Kobelt, 1909) (Fig. 4e) . Several dozen samples
are from the Shatskiy Lakes. The applicability of this name is
determined by the description and drawing by Kobelt (1909: 9, PI.
394, Figs. 2265-2266) and a photograph of the lectotype by Zilch
(1955: PI. 3, Fig. 9). Unfortunately, in this instance the apex
of the spire is extremely corroded. It differs from C. turrita
that is found together with it by a lower whorl and a large

18

apical angle. It is also located in the Pleshcheyevo Lake
(Yaroslavskaya Oblast) within the borders of the Baltic area.

C. zebra (Kobelt, 1877) (Fig. 4f ) . Several samples are from
the vicinity of Batum. The species is precisely characterized in
the description and drawing of Kobelt (1877: 73, Pi. 138, Fig.
1370) and in a photograph of the lectotype by Zilch (1955: PI. 3,
Fig. 6) . The species differs from C. turrita by a lower spire
and from C. fennica by a smaller apical angle. It comes from
areas near Istanbul. We discovered it for the first time in the
collection at the Zoological Institute from around Batum.
Apparently, the species is endemic to the southern Black Sea
littoral areas.

C. kormosi (Kobelt, 1909) (Fig. 4g) . Several dozen samples
are from the Shatskiy Lakes. The applicability of this name is
determined by the description and drawings of Kobelt (1906-1909:
315, PI. 62, Fig. 12; 1909: 3, PI. 391, Fig. 2242) and the
photograph of the lectotype by Zilch (1955: PI. 3, Fig. 2). It
differs from C. zebra and C. fennica by a large apical angle and
a much lower shell. The species is found throughout the Baltic
area (including the Shatskiy Lakes) , from where all our material
is derived, as well as in the central Danube areas.

19

C. ladoqensis
Chernogorenko and Starobogatov, new species (Fig. 4h)

This species is named after the area where it is found
(Ladoga) .

Material. One holotype and two para types are from the
northern part of Lake Ladoga (collection of the Petrozavodsk
University expedition) . The holotype and one paratype (opened)
are kept in the Zoological Institute, USSR Academy of Sciences
collection.

Description. The shell is almost spherical with greatly
protruding whorls separated by a deep seam. The tangent line of
the whorl is straight. The entire shell is an olive monochrome.
The last whorl is a very large one constituting 0.8 to 0.85 of
the height of the entire shell. Its periphery is uniformly
rounded. The apical angle is around 120°. The umbilicus is wide
open but not rounded. The mouth is oval with a blunt parietal-
palatal angle. The measurements of the holotype shell (in mm) is
as follows: shell height - 29.2, shell width - 25.5, height of
the aperture - 17.7, width of the aperture - 13.2, height of
spire - 13.0, having 5.8 whorls, the height of the last whorl
without the aperture is 22.5. Measurements of the two paratypes
are not given here because the shells are badly damaged.

20

Remarks . The species belongs to the group of lake
contectianas but is distinguished from all other species of this
group by an unusually low spire, and from the pond contectianas
by the same characteristics as those found in the lake
contectianas. Below is a table containing the differences among
the contemporary species of the genus Contectiana.

TABLE FOR DISTINGUISHING SPECIES OF THE GENUS Contectiana

1. (4) Shell is thin-walled; whorls are protiiberant and
somewhat uneven. They form a flat shoulder above the seam,
sometimes barely perceptible; their edges toward the lower seam
fit almost perpendicularly. subgenus Contectiana s. str.

2. (3) Width of the penultimate whorl exceeds the distance
(along the axis) from its lower seam to the spire by not less
than 1.6 times. C. contecta (Mill.).

3. (2) Width of the penultimate whorl exceeds the distance
(along the axis) from its lower seam to the spire by not more
than 1.5 times. C. listeri (Forbes and Hanley) .

4. (1) Shell is relatively thick-walled, whorls are evenly
protuberant. Their contour at lower seams strongly bends toward
the axis. sub-genus Kobeltipaludina .

21

5. (6) Height of part of the spire elevated above the last
whorl does not exceed 0.7, which is the elevation of the last
whorl above the mouth. C. ladogensis sp. n.

6. (5) Height of part of the spire elevated above the last
whorl is equal to the elevation of the last whorl above the mouth
(or at least not less than 0.9 of this elevation) or more than
it.

7. (8) Height of an adult shell exceeds the width by not less
than 1.35 times. C. ianinensis (Mousson) .

8. (7) Height of an adult shell exceeds the width by not more
than 1.3 0 times.

9. (12) Shell's apical angle is not less than 110°; height of
the third- from- the-bottom whorl does not exceed 0.42 which is the
height of the penultimate one.

10. (11) Height of the adult shell exceeds its width by not more
than 1.4 times; apical angle is not less than 115°. C. kormosi
(Kob.) .

11. (10) Height of the adult shell exceeds its width by no less
than 1.5 times; apical angle is not more than 114°. C. fennica
(Kob.) .

22

12. (9) Apical angle of the shell does not exceed 108°; height
of third-from-the-bottom whorl is not less than 0.4 3 of the
height of the penultimate one.

13. (14) Width of the penultimate whorl exceeds the distance from
its lower seam (along the axis) to the spire by not less than 1.6
times. C. zebra (Kob.).

14. (13) Width of the penultimate whorl exceeds the distance from
its lower seam (along the axis) to the spire by not more than 1.5
times. C. turrita (Kob.).

BIBLIOGRAPHY

Bourguignat, J. R. 1880. Recensement des Vivipara du systeme
Europeen. Paris, pp. 1-52.

Franz, V. 1932. Viviparus. Morphometrie , Phylogenie und
Geographie der europaischen fossilen und rezenten Paludinen.
Denkschrif ten der Mediz inisch-Naturwissenschaftlen
Gesellschaft zu Jena, 18(1) : 1-160.

Izzatullayev, Z. I. and Ya. I. Starobogatov. 1984. The genus
Melanopsis (Gastropoda, Pect inibranchia ) and its
representatives inhabiting the USSR reservoirs. Zoological

23

Journal, 63(10): 1471-1483.

Kobelt, W. 1877. Iconographie der Land- und Siisswasser-
Mollusken von E. A. Rossmassler, 5. Dresden-Leipzig, pp. 1-
12 7. 19 06-1909. Die Gattung Paludina Lam. Neue
Folge/Martini und Chemnitz Systematisches Conchylien-
Cabinet. Niirnberg, pp. 1-430. 1907. Iconographie der Land-
und Svisswasser-Mollusken von E. A. Rossmassler, Neue Folge
13(3/6) Dresden - Leipzig, pp. 1-65. 1909. Idem, 15(1/4):
1-48.

Kiister, H. C. 1852. Die Gattungen Paludina. Hydrocaena und
Valvata. Martini und Chemnitz Systematisches Conchylien-
Cabinet. Niirnberg, pp. 1-96.

Opinion 573. 1959. Determination under the Plenari Powers of a
lectotype for the nominal species Helix vivipara Linnaeus,
1758 and addition to the offical lists of the generic name
Viviparus Montfort, 1810 and the family group name
Viviparidae Gray, 1847 (class Gastropoda) . Bulletin of
Zoological Nomenclature, 17(3-5) : 117-131.

Raup, D. M. 1967. Geometric analysis of shell coiling. Journal
of Paleontology, 41(1): 43-65.

Rossmassler, E. A. 1835. Iconographie der Land- und Susswasser-

24

»

Mollusken. 1. Dresden - Leipzig, pp. 1-132; 1835a. Idem. 2:
pp. 1-26.

Sitnikova, T. Ya. and Ya. I. Starobogatov. 1982. The range and
systematic status of the Architaenioglossa (Gastropoda,
Pectinibranchia) . Zoological Journal, 6^(6): 831-842.

Starobogatov, Ya. I. 1970. Fauna of the Molluscs and the
Zoogeographic Zoning of Continental Reservoirs of the Planet
Earth. Leningrad, Nauka, pp. 1-372.

Starogogatov, Ya. I. 1983. Zoogeographic characteristic of
freshwater malacofauna of the Northern Eurasia. VIII
Malacological Congress, Budapest, p. 13 6.

Watson, H. 1957. VThich of the two common British species of
Viviparus Mont fort named Viviparus viviparus (Linnaeus) (=
Helix vivipara Linnaeus 1758)? Bulletin of Zoological
Nomenclature, 13(2/3): 53-66.

Westerlund, C. A. 1886. Fauna der in der Palaarktischen
Regionen lebenden Binnenconchylien. VI. Fam. Ampullariidae,
Paludinidae, Hydrobiidae, Melaniidae, Valvatidae und
Neritidae. Lund, pp. 1-156.

Zhadin, V. I. 1952. Molluscs of the USSR's Fresh and Brackish

25

Waters. Moscow-Leningrad, USSR Academy of Sciences, pp. 1-
370.

Zilch, A. 19 55. Typen und Typoide des Natur-Museums
Senckenberg, 14: Mollusca Viviparidae. Archiv fiir
Molluskenkunde, 84(1/3): 45-86.

EDITORIAL FOOTNOTES

1. The correct date of Mont fort is 1810, not 1816, and the
correct type-species is, by original designation, Viviparus
fluviorum Montfort 1810 [+ Helix vivipara Linnaeus 1758].

2. The type-species of Contectiana Bourguignat 1880 is, by
original designation and virtual tautonoiay, Vivipara contecta [ =
Cyclostoma contectum Millet de la Turtaudiere 1813].

26

a.

M,

M,

X^jn^J Uo-y ^MZk^

Fig. 1. Structure of the radular teeth.

a) Viviparus viviparus. b) Contectiana listeri

Fig. 2. Female genital system. a) V i v iparus
V i V iparus ■ b) Contectiana turrita. c) albumen
g 1 and. d) egg capsules.

Fig. 3. Viviparus spec ies shel 1 s : a) y_. viv iparus Cwashed up
at Kurshsk-iy Bay, Rybachiy hamlet, Kaliningrad Oblast). b) V_.
vistulae Csame place), c) V_. ater CKiev Reservoir), d) y_. costae
(.Redut Kale, Rosen's col lection) . e) y_. rossmaess 1 er i (syntype) .
fy v. hel lenicus (Corfu Island), g) V_. blanci (Korfu Isl.).
h) V". graecus sp. n. (holotype, Greece, Goritsa). Scale: 10 mm.

Fig. k. Shells of the Contect iana species: a) C. contecta (In
vicinity of Kharkov), b) C. 1 is ter i (Peremut Lake, Volynskaya
Oblast). c) C_. turri ta (near town of Shatska, Volynskaya Oblast).
dl C_. jan inens is (syntype, based on Kobelt, 1909)- e) £. fenn i ca
CLake P leshcheyevo, Yaros lavskaya Oblast). f) C_. zebra (lectotype
based on Zilch, 1955)- g) C. kormos i (Lake Krugloye near Przheval-
skoye, Smolensk Oblast). hT C_- ladogensis sp. n. (holotype).
Scale: 10 mm.

907

I. n

Harvard MCZ Llbran

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