View full-text
Alien mollusks within the territory of Ukraine: sources and directions of invasions
Journal Article: Russian Journal of Biological Invasions 01/2010; 1:37-44.
Abstract
The main directions of expansion of alien mollusks in Ukraine are revealed. Expansions of both exotic species and nearby aliens ones are observed. The Danube, Dnieper, and line of water bodies of the Azov–Black Sea region’s seaside are important invasion corridors to the inland waters. The urbanization and reduction of zoogeographical barriers are the basic driving processes in expansion of alien terrestrial snails. Invasions into marine ecosystems are related to transfer of distant exotic species by sea navigation and homogenization of fauna in the East Mediterranean Basin (“pontization” and “mediterranization”)
Comments on this publication
ResearchGate members can add comments. Sign up now and post your comment!
Data provided are for informational purposes only. Although carefully collected, accuracy cannot be guaranteed. The impact factor represents a rough estimation of the journal's impact factor and does not reflect the actual current impact factor. Publisher conditions are provided by RoMEO. Differing provisions from the publisher's actual policy or licence agreement may be applicable.
Page 1
Page 2
Page 3
Page 4
Page 5
Page 6
Page 7
Page 8
Page 1
ISSN 2075�1117, Russian Journal of Biological Invasions, 2010, Vol. 1, No. 1, pp. 37–44. © Pleiades Publishing, Ltd., 2010.
Original Russian Text © M.O. Son, 2009, published in Rossiiskii Zhurnal Biologicheskikh Invasii, 2009, No. 2, pp. 37–48.
37
INTRODUCTION
The expansion of invasive mollusks in Ukraine is
well studied in the faunistic aspect. Available data con�
firm that this territory is a place of crossing of numer�
ous invasion corridors and serves both as a donor and
as a recipient of invaders. The goal of the paper is to
describe the structure of the invasion network of the
territory (a combination of invasion corridors).
MATERIALS AND METHODS
The faunistic data analyzed in this paper are mainly
those published as special reports. Most findings of
alien terrestrial mollusks relate to the process of
urbanization and were made during joint studies of the
urban mollusk faunas in Ukraine [Sverlova and Son,
2006]. The data of the modern faunistic report on the
mollusks of Southeastern Ukraine [Sverlova et al.,
2006] are also used in the present article along with
later publications concerning some specific findings.
The invasions of aquatic mollusks are described in sev�
eral special reports [Son, 2007; Alexandrov et al.,
2007].
RESULTS AND DISCUSSION
Recently, several main directions of expansion of
aquatic mollusks were formed within the territory of
continental Ukraine. Partially, these directions coin�
cide with the largest invasion corridors in Europe.
The range expansion downstream of the Danube
River (southern invasion corridor) resulted in the
appearance within the Ukrainian part of the Danube
basin of such species as Sinanodonta woodiana (Lea,
1834), Corbicula fluminalis (Müller, 1774), C. fluminea
(Müller, 1774), and others [Son, 2007].
The expansion of the Ponto�Caspian mollusks
(Dreissena bugensis Andrusov, 1897, D. polymorpha
(Pallas, 1771), Hypanis spp., etc.) upstream of the
Dnieper River (central corridor) triggered by the lim�
nization of the Dnieper (i.e., turning of the river chan�
nel into a cascade of man�made lakes) opened for
these species the possibility of wide expansion [Alex�
androv et al., 2007]. Creation of large interbasin canals
(the Dnieper–Severskii Donets and the Volga–Don)
resulted in the formation of a hydrographic connec�
tion between the central and northern invasion corri�
dors. In turn, this resulted in the further expansion of
the Ponto�Caspian invaders in the Don and Volga
basins [Panov et al., 2009].
The best studied are the routes of the invasions of
Ponto�Caspian species along the central corridor
through the Dnieper–Bug canal connecting the
Dnieper and Zapadnyi (West) Bug rivers. Recently, the
expansion of D. polymorpha into the Shatskaya group
of lakes (Pulemetskoye, Svityaz’) includes also
another branch of this corridor: the upper Pripyat
River–the Shatsk lakes–the Zapadnyi Bug basin.
In the middle of the 20th century, anthropogenic
destruction of the natural zoogeographic barriers iso�
lating for a long time the aquatic fauna of the Crimea
took place. Before that, these barriers shaped the
fauna of the peninsula in an island manner. The inten�
tional introductions into man�made lakes and con�
struction of the Severo�Krymskii (North Crimean)
Canal had the main impact on the expansion of spe�
cies of aquatic mollusks new to the peninsula. Numer�
Alien Mollusks within the Territory of Ukraine:
Sources and Directions of Invasions
M. O. Son
Kovalevsky Institute of Biology of the Southern Seas (Odessa Branch),
National Academy of Sciences of Ukraine, Odessa, Ukraine
e�mail: michail.son@gmail.com
Received September 7, 2009
Abstract—The main directions of expansion of alien mollusks in Ukraine are revealed. Expansions of both
exotic species and nearby aliens ones are observed. The Danube, Dnieper, and line of water bodies of the
Azov–Black Sea region’s seaside are important invasion corridors to the inland waters. The urbanization and
reduction of zoogeographical barriers are the basic driving processes in expansion of alien terrestrial snails.
Invasions into marine ecosystems are related to transfer of distant exotic species by sea navigation and
homogenization of fauna in the East Mediterranean Basin (“pontization” and “mediterranization”).
DOI: 10.1134/S207511171001008X
Key words: mollusks, Ukraine, alien species, exotic, Ponto�Caspian.
Original Russian Text © M.O. Son, 2009, published in Rossiiskii Zhurnal Biologicheskikh Invasii, 2009, No. 2, pp. 37–48.
37
INTRODUCTION
The expansion of invasive mollusks in Ukraine is
well studied in the faunistic aspect. Available data con�
firm that this territory is a place of crossing of numer�
ous invasion corridors and serves both as a donor and
as a recipient of invaders. The goal of the paper is to
describe the structure of the invasion network of the
territory (a combination of invasion corridors).
MATERIALS AND METHODS
The faunistic data analyzed in this paper are mainly
those published as special reports. Most findings of
alien terrestrial mollusks relate to the process of
urbanization and were made during joint studies of the
urban mollusk faunas in Ukraine [Sverlova and Son,
2006]. The data of the modern faunistic report on the
mollusks of Southeastern Ukraine [Sverlova et al.,
2006] are also used in the present article along with
later publications concerning some specific findings.
The invasions of aquatic mollusks are described in sev�
eral special reports [Son, 2007; Alexandrov et al.,
2007].
RESULTS AND DISCUSSION
Recently, several main directions of expansion of
aquatic mollusks were formed within the territory of
continental Ukraine. Partially, these directions coin�
cide with the largest invasion corridors in Europe.
The range expansion downstream of the Danube
River (southern invasion corridor) resulted in the
appearance within the Ukrainian part of the Danube
basin of such species as Sinanodonta woodiana (Lea,
1834), Corbicula fluminalis (Müller, 1774), C. fluminea
(Müller, 1774), and others [Son, 2007].
The expansion of the Ponto�Caspian mollusks
(Dreissena bugensis Andrusov, 1897, D. polymorpha
(Pallas, 1771), Hypanis spp., etc.) upstream of the
Dnieper River (central corridor) triggered by the lim�
nization of the Dnieper (i.e., turning of the river chan�
nel into a cascade of man�made lakes) opened for
these species the possibility of wide expansion [Alex�
androv et al., 2007]. Creation of large interbasin canals
(the Dnieper–Severskii Donets and the Volga–Don)
resulted in the formation of a hydrographic connec�
tion between the central and northern invasion corri�
dors. In turn, this resulted in the further expansion of
the Ponto�Caspian invaders in the Don and Volga
basins [Panov et al., 2009].
The best studied are the routes of the invasions of
Ponto�Caspian species along the central corridor
through the Dnieper–Bug canal connecting the
Dnieper and Zapadnyi (West) Bug rivers. Recently, the
expansion of D. polymorpha into the Shatskaya group
of lakes (Pulemetskoye, Svityaz’) includes also
another branch of this corridor: the upper Pripyat
River–the Shatsk lakes–the Zapadnyi Bug basin.
In the middle of the 20th century, anthropogenic
destruction of the natural zoogeographic barriers iso�
lating for a long time the aquatic fauna of the Crimea
took place. Before that, these barriers shaped the
fauna of the peninsula in an island manner. The inten�
tional introductions into man�made lakes and con�
struction of the Severo�Krymskii (North Crimean)
Canal had the main impact on the expansion of spe�
cies of aquatic mollusks new to the peninsula. Numer�
Alien Mollusks within the Territory of Ukraine:
Sources and Directions of Invasions
M. O. Son
Kovalevsky Institute of Biology of the Southern Seas (Odessa Branch),
National Academy of Sciences of Ukraine, Odessa, Ukraine
e�mail: michail.son@gmail.com
Received September 7, 2009
Abstract—The main directions of expansion of alien mollusks in Ukraine are revealed. Expansions of both
exotic species and nearby aliens ones are observed. The Danube, Dnieper, and line of water bodies of the
Azov–Black Sea region’s seaside are important invasion corridors to the inland waters. The urbanization and
reduction of zoogeographical barriers are the basic driving processes in expansion of alien terrestrial snails.
Invasions into marine ecosystems are related to transfer of distant exotic species by sea navigation and
homogenization of fauna in the East Mediterranean Basin (“pontization” and “mediterranization”).
DOI: 10.1134/S207511171001008X
Key words: mollusks, Ukraine, alien species, exotic, Ponto�Caspian.
Page 2
38
RUSSIAN JOURNAL OF BIOLOGICAL INVASIONS Vol. 1 No. 1 2010
SON
ous invaders from continental Ukraine were found in
the inland waters of the peninsula: D. bugensis,
D. polymorpha, Viviparus viviparus (Linnaeus, 1758),
Lithoglyphus naticoides (Pfeiffer, 1828), Theodoxus
fluviatilis (Linnaeus, 1758), Fagotia danubialis Bour�
guignat, 1884, Lymnaea stagnalis (Linnaeus, 1758),
L. ovata (Smith, 1889), etc. Previously, these species
were absent from the fauna of the peninsula [Son,
2007].
In addition to the hydrographic network connected
to the North Crimean Canal, a series of other irriga�
tion systems was built on the peninsula which became
the zone of expansion of the Ponto�Caspian invaders
(Zhuravel’ et al., 1968]. This construction resulted in
the creation of interbasin connections, and some small
steppe rivers (Baraboy, Sarata, Kal’mius) were inhab�
ited by many Ponto�Caspian species that were absent
from these waters historically.
Despite the wide distribution of invader mollusks
in the large river basins, the colonization of the river
tributaries up to now shows only a few examples.
Numerous closely situated limans, estuaries, river
deltas, and freshwater and brackish�water lakes often
connected by canals is a characteristic of the territories
close to the northwestern Prichernomor’e and north�
ern Priazov’e (northwestern Black Sea and northern
Sea of Azov coasts). These water bodies form an
important invasion corridor for species able to inhabit
both fresh and brackish waters [Son, 2007]. This route
of invasion typical of Potamopyrgus antipodarum
(Gray, 1843) and Ferrissia fragilis (Tryon, 1863) pro�
vides an exchange of exotic species between down�
stream reaches of the large river systems of the north�
ern Black Sea coast. Originally, this vector of invasions
was named the Black Sea coastal invasion corridor, but
in the classification of the European invasion corri�
dors, the term southern meridional corridor is also in
use [Panov et al., 2009].
Hydroengineering and other anthropogenic
impacts caused changes in the hydrology of the Black
Sea coastal limans. These changes resulted in total
replacement of the faunas in many limans or their sep�
arate parts. The man�made connection of the earlier
isolated limans to the sea or dredging of the historical
mouth zones led to the rise of water salinity in the
water bodies with their consequent “pontization”—
invasion of the diverse Black Sea fauna. This fauna
either partially or totally forced out the Ponto�Cas�
pian fauna or enriched initially poor species compo�
sition [Zambriborshch, 1985]. The Hadjibeyskii,
Tiligul’skii, Dnestrovskii, and many other limans pro�
vide examples of such modifications in the fauna. By
contrast, in the primeval sea limans, the isolation by
dams of some parts or of the whole liman led to the
formation of desalinated zones colonized by freshwa�
ter and Ponto�Caspian fauna (invasion of D. polymor�
pha in the desalinated part of the Dofinovskii liman
and rich fauna of the freshwater and Ponto�Caspian
mollusks in Lake Sasyk). Such modifications of the
ecosystems of limans turned them into zones of mass
naturalization of exotic species of marine (Anadara
sp., Corambe obscura (Verill, 1870), Rapana venosa
(Valenciennes, 1846), Mya arenaria (Linnaeus, 1758),
Mytilopsis leucophaeata (Conrad, 1831)) and freshwa�
ter (S. woodiana, P. antipodarum, Physella spp., etc.)
origin.
Local introduction of aquarium species into natu�
ral and man�made water bodies is common in Ukraine
(Helisoma duryi (Wetherby, 1879), Physella spp., Amp�
ullariidae spp., Melanoides tuberculata (Müller, 1774),
F. fragilis, etc.). Quite often, the water from aquariums
is discarded in open water bodies. In this case, most
thermophilic species as a rule are capable of surviving
either in man�made water bodies subjected to dis�
charge of heated waters or in the southernmost waters
of Ukraine [Son, 2007].
The ecosystems of the Black Sea and the Sea of
Azov are also exposed to invasions of mollusks. The
main direction of the expansion there is the introduc�
tion with ship traffic through the Black Sea straits in
ballast waters (at the stage of pelagic larvae), with bio�
fowling, or in wooden ship hulls (shipworm Teredo
navalis (Linnaeus, 1758)). The majority of mollusk
invaders (Anadara sp., C. obscura, R. venosa,
M. arenaria, Neptunea arthritica (Bernardi, 1857),
Ercolania viridis (A. Costa, 1866), Mytilus edulis (Lin�
naeus, 1758), Mytilus trossulus Gould, 1850, and some
others) penetrated into the Black Sea precisely in this
way [Alexandrov et al., 2007]. The only intentionally
introduced marine species in the Black Sea is mollusk
Crassostrea gigas (Thunberg, 1793) [Zolotarev, 1996].
The drastic change in the salinity of the water of the
Black Sea–Sea of Azov basin resulted in the gradual
expansion of the ranges of Mediterranean fauna, so�
called “mediterranization” [Puzanov, 1965, 1967]. In
addition to the introduction of the Mediterranean spe�
cies (previously absent from the sea except for regions
near the Bosporus Strait) into the Black Sea, the
ranges of these Mediterranean species expanded. This
concerns those species that earlier were present in
some areas of the Black Sea where the water salinity
was the highest. In addition to the naturalization of
these species of the earlier isolated limans and
lagoons, the process of pontization covered large areas
in the Sea of Azov [Anistratenko et al., 2007a, 2008].
A process typical of some desalinated areas of the
Black Sea and Sea of Azov is the introduction of exotic
freshwater species tolerant to high water salinity. These
species come from the rivers (S. woodiana and Corbic�
ula spp. near the Danube River mouth) or with river
ship traffic (introduction of Hypanis glabra (Ostrou�
moff, 1905) to the Taganrog Bay from the Caspian via
the Volga–Don Canal) [Son, 2007; Alexandrov et al.,
2007].
Expansion of terrestrial mollusks is related first of
all to their direct transfer with soil, building materials,
planting stocks, etc. Owing to low mobility, the expan�
sion of these mollusks is often strictly local. After
RUSSIAN JOURNAL OF BIOLOGICAL INVASIONS Vol. 1 No. 1 2010
SON
ous invaders from continental Ukraine were found in
the inland waters of the peninsula: D. bugensis,
D. polymorpha, Viviparus viviparus (Linnaeus, 1758),
Lithoglyphus naticoides (Pfeiffer, 1828), Theodoxus
fluviatilis (Linnaeus, 1758), Fagotia danubialis Bour�
guignat, 1884, Lymnaea stagnalis (Linnaeus, 1758),
L. ovata (Smith, 1889), etc. Previously, these species
were absent from the fauna of the peninsula [Son,
2007].
In addition to the hydrographic network connected
to the North Crimean Canal, a series of other irriga�
tion systems was built on the peninsula which became
the zone of expansion of the Ponto�Caspian invaders
(Zhuravel’ et al., 1968]. This construction resulted in
the creation of interbasin connections, and some small
steppe rivers (Baraboy, Sarata, Kal’mius) were inhab�
ited by many Ponto�Caspian species that were absent
from these waters historically.
Despite the wide distribution of invader mollusks
in the large river basins, the colonization of the river
tributaries up to now shows only a few examples.
Numerous closely situated limans, estuaries, river
deltas, and freshwater and brackish�water lakes often
connected by canals is a characteristic of the territories
close to the northwestern Prichernomor’e and north�
ern Priazov’e (northwestern Black Sea and northern
Sea of Azov coasts). These water bodies form an
important invasion corridor for species able to inhabit
both fresh and brackish waters [Son, 2007]. This route
of invasion typical of Potamopyrgus antipodarum
(Gray, 1843) and Ferrissia fragilis (Tryon, 1863) pro�
vides an exchange of exotic species between down�
stream reaches of the large river systems of the north�
ern Black Sea coast. Originally, this vector of invasions
was named the Black Sea coastal invasion corridor, but
in the classification of the European invasion corri�
dors, the term southern meridional corridor is also in
use [Panov et al., 2009].
Hydroengineering and other anthropogenic
impacts caused changes in the hydrology of the Black
Sea coastal limans. These changes resulted in total
replacement of the faunas in many limans or their sep�
arate parts. The man�made connection of the earlier
isolated limans to the sea or dredging of the historical
mouth zones led to the rise of water salinity in the
water bodies with their consequent “pontization”—
invasion of the diverse Black Sea fauna. This fauna
either partially or totally forced out the Ponto�Cas�
pian fauna or enriched initially poor species compo�
sition [Zambriborshch, 1985]. The Hadjibeyskii,
Tiligul’skii, Dnestrovskii, and many other limans pro�
vide examples of such modifications in the fauna. By
contrast, in the primeval sea limans, the isolation by
dams of some parts or of the whole liman led to the
formation of desalinated zones colonized by freshwa�
ter and Ponto�Caspian fauna (invasion of D. polymor�
pha in the desalinated part of the Dofinovskii liman
and rich fauna of the freshwater and Ponto�Caspian
mollusks in Lake Sasyk). Such modifications of the
ecosystems of limans turned them into zones of mass
naturalization of exotic species of marine (Anadara
sp., Corambe obscura (Verill, 1870), Rapana venosa
(Valenciennes, 1846), Mya arenaria (Linnaeus, 1758),
Mytilopsis leucophaeata (Conrad, 1831)) and freshwa�
ter (S. woodiana, P. antipodarum, Physella spp., etc.)
origin.
Local introduction of aquarium species into natu�
ral and man�made water bodies is common in Ukraine
(Helisoma duryi (Wetherby, 1879), Physella spp., Amp�
ullariidae spp., Melanoides tuberculata (Müller, 1774),
F. fragilis, etc.). Quite often, the water from aquariums
is discarded in open water bodies. In this case, most
thermophilic species as a rule are capable of surviving
either in man�made water bodies subjected to dis�
charge of heated waters or in the southernmost waters
of Ukraine [Son, 2007].
The ecosystems of the Black Sea and the Sea of
Azov are also exposed to invasions of mollusks. The
main direction of the expansion there is the introduc�
tion with ship traffic through the Black Sea straits in
ballast waters (at the stage of pelagic larvae), with bio�
fowling, or in wooden ship hulls (shipworm Teredo
navalis (Linnaeus, 1758)). The majority of mollusk
invaders (Anadara sp., C. obscura, R. venosa,
M. arenaria, Neptunea arthritica (Bernardi, 1857),
Ercolania viridis (A. Costa, 1866), Mytilus edulis (Lin�
naeus, 1758), Mytilus trossulus Gould, 1850, and some
others) penetrated into the Black Sea precisely in this
way [Alexandrov et al., 2007]. The only intentionally
introduced marine species in the Black Sea is mollusk
Crassostrea gigas (Thunberg, 1793) [Zolotarev, 1996].
The drastic change in the salinity of the water of the
Black Sea–Sea of Azov basin resulted in the gradual
expansion of the ranges of Mediterranean fauna, so�
called “mediterranization” [Puzanov, 1965, 1967]. In
addition to the introduction of the Mediterranean spe�
cies (previously absent from the sea except for regions
near the Bosporus Strait) into the Black Sea, the
ranges of these Mediterranean species expanded. This
concerns those species that earlier were present in
some areas of the Black Sea where the water salinity
was the highest. In addition to the naturalization of
these species of the earlier isolated limans and
lagoons, the process of pontization covered large areas
in the Sea of Azov [Anistratenko et al., 2007a, 2008].
A process typical of some desalinated areas of the
Black Sea and Sea of Azov is the introduction of exotic
freshwater species tolerant to high water salinity. These
species come from the rivers (S. woodiana and Corbic�
ula spp. near the Danube River mouth) or with river
ship traffic (introduction of Hypanis glabra (Ostrou�
moff, 1905) to the Taganrog Bay from the Caspian via
the Volga–Don Canal) [Son, 2007; Alexandrov et al.,
2007].
Expansion of terrestrial mollusks is related first of
all to their direct transfer with soil, building materials,
planting stocks, etc. Owing to low mobility, the expan�
sion of these mollusks is often strictly local. After
Page 3
RUSSIAN JOURNAL OF BIOLOGICAL INVASIONS Vol. 1 No. 1 2010
ALIEN MOLLUSKS WITHIN THE TERRITORY OF UKRAINE 39
Table 1. Exotic species of mollusks within the territory of Ukraine
Species Native region Time of intro�duction Distribution in Ukraine References
Freshwater and brackish�water species
Ampullaria sp. Tropical regions 1999 Man�made water bodies of
Southern Ukraine
Son, 2007
Biomphalaria glabrata
Say, 1818
Africa 2000 Man�made water bodies of
Southern Ukraine
Son, 2007
Corbicula fluminalis
(Müller, 1774)
Asia 2000 The Danube basin Son, 2007
C. fluminea (Müller, 1774) Far East 1995 The Danube basin Son, 2007
Ferrissia fragilis North America Beginning of
20th century
The Dnestr, Danube, and
Dnieper deltas, water bodies of
Crimea, cooling ponds in Cen�
tral and Northern Ukraine
Son, 2007; Son, 2007b;
Semenchenko,
Laenko, 2008
Helisoma duryi Africa 2000 Man�made warm water bodies Son, 2007
Hypanis glabra Caspian 2004 Taganrog Bay Son, 2007
Mytilopsis leucophaeata North America 2003 Dnestrovskii Liman Therriault et al. 2004
Melanoides tuberculata Tropical regions 2000 Man�made water bodies in
Southern Ukraine; cooling
pond of Yuzhnoukrainskaya
nuclear power station
Grigorovich et al.
2002; Son, 2007
Set of species Physella
integra (Haldeman,
1841), Ph. heterostropha
(Say, 1817), Ph. acuta
(Draparnaud, 1805)
North America Beginning of
20th century
Predominantly Southern and
Eastern Ukraine
Son, 2007
Potamopyrgus antipodarum New Zealand 1951 Northwestern Black Sea coast Son, 2008
Sinanodonta woodiana Far East 1999 Danube basin; man�made
lake near town of Kotovsk
Son, 2007
Thiara granifera Malaysia 2006 Finding in the Molochnaya
River
Khaliman and Anis�
tratenko, 2006
Sea species
Anadara sp. Attribution of the spe�
cies to A. inaequivalvis
(Bruguiére, 1789) was
disproved (Lutayenko,
2006)
1987 Black Sea and Sea of Azov Zolotarev, 1996;
Lutayenko, 2006
Aporrhais pespelecani
(Linne, 1758)
Mediterranean species
previously found in the
Black Sea only near
the Bosporus
1987 Crimean coast Anistratenko, 1998
Crassostrea gigas Sea of Japan 1900s Crimea, Bay of Odessa Zolotarev 1996, Kovtun
and Zolotarev, 2008
Corambe obscura
(Verrill, 1870)
Atlantic coast of North
America
1989 Black Sea and Sea of Azov Roginskii and
Grintsov, 1990; Zolo�
tarev 1996; Alexan�
drov et al., 2007
Ercolania viridis Caribbean Sea 2001 Bay of Odessa Alexandrov et al., 2007
Mya arenaria Atlantic Ocean 1966 Black Sea and Sea of Azov Zolotarev, 1996
Mytilus edulis and
M. trossulus
Atlantic Ocean 2001 Bay of Odessa Alexandrov et al., 2007
Neptunea arthritica Far Eastern Seas 2000 Kamyshovaya Cove (Crimea) Shadrin et al., 2002
Perna viridis (Linnaeus,
1758)
Pacific region Black Sea (Sevastopol’) Mironov et al., 2002
ALIEN MOLLUSKS WITHIN THE TERRITORY OF UKRAINE 39
Table 1. Exotic species of mollusks within the territory of Ukraine
Species Native region Time of intro�duction Distribution in Ukraine References
Freshwater and brackish�water species
Ampullaria sp. Tropical regions 1999 Man�made water bodies of
Southern Ukraine
Son, 2007
Biomphalaria glabrata
Say, 1818
Africa 2000 Man�made water bodies of
Southern Ukraine
Son, 2007
Corbicula fluminalis
(Müller, 1774)
Asia 2000 The Danube basin Son, 2007
C. fluminea (Müller, 1774) Far East 1995 The Danube basin Son, 2007
Ferrissia fragilis North America Beginning of
20th century
The Dnestr, Danube, and
Dnieper deltas, water bodies of
Crimea, cooling ponds in Cen�
tral and Northern Ukraine
Son, 2007; Son, 2007b;
Semenchenko,
Laenko, 2008
Helisoma duryi Africa 2000 Man�made warm water bodies Son, 2007
Hypanis glabra Caspian 2004 Taganrog Bay Son, 2007
Mytilopsis leucophaeata North America 2003 Dnestrovskii Liman Therriault et al. 2004
Melanoides tuberculata Tropical regions 2000 Man�made water bodies in
Southern Ukraine; cooling
pond of Yuzhnoukrainskaya
nuclear power station
Grigorovich et al.
2002; Son, 2007
Set of species Physella
integra (Haldeman,
1841), Ph. heterostropha
(Say, 1817), Ph. acuta
(Draparnaud, 1805)
North America Beginning of
20th century
Predominantly Southern and
Eastern Ukraine
Son, 2007
Potamopyrgus antipodarum New Zealand 1951 Northwestern Black Sea coast Son, 2008
Sinanodonta woodiana Far East 1999 Danube basin; man�made
lake near town of Kotovsk
Son, 2007
Thiara granifera Malaysia 2006 Finding in the Molochnaya
River
Khaliman and Anis�
tratenko, 2006
Sea species
Anadara sp. Attribution of the spe�
cies to A. inaequivalvis
(Bruguiére, 1789) was
disproved (Lutayenko,
2006)
1987 Black Sea and Sea of Azov Zolotarev, 1996;
Lutayenko, 2006
Aporrhais pespelecani
(Linne, 1758)
Mediterranean species
previously found in the
Black Sea only near
the Bosporus
1987 Crimean coast Anistratenko, 1998
Crassostrea gigas Sea of Japan 1900s Crimea, Bay of Odessa Zolotarev 1996, Kovtun
and Zolotarev, 2008
Corambe obscura
(Verrill, 1870)
Atlantic coast of North
America
1989 Black Sea and Sea of Azov Roginskii and
Grintsov, 1990; Zolo�
tarev 1996; Alexan�
drov et al., 2007
Ercolania viridis Caribbean Sea 2001 Bay of Odessa Alexandrov et al., 2007
Mya arenaria Atlantic Ocean 1966 Black Sea and Sea of Azov Zolotarev, 1996
Mytilus edulis and
M. trossulus
Atlantic Ocean 2001 Bay of Odessa Alexandrov et al., 2007
Neptunea arthritica Far Eastern Seas 2000 Kamyshovaya Cove (Crimea) Shadrin et al., 2002
Perna viridis (Linnaeus,
1758)
Pacific region Black Sea (Sevastopol’) Mironov et al., 2002
Page 4
40
RUSSIAN JOURNAL OF BIOLOGICAL INVASIONS Vol. 1 No. 1 2010
SON
introduction to a certain habitat, the species may live
there for decades without moving beyond the borders
of the habitat. Thus, the directions of invasions of ter�
restrial mollusks are determined predominantly by the
routes of transportation and, hence, by trade and eco�
nomic ties between various regions.
The most noticeable process in the invasions of ter�
restrial mollusks is a mass expansion of the Crimean
and Caucasian species towards the steppe zone and
urban territories of other regions in Ukraine. Along
with the aforementioned species, some other mollusks
typical of the steppes also penetrate into urban territo�
ries in Central and Western Ukraine.
Among the species endemic to the Crimea or ear�
lier found in Southern Ukraine only on this peninsula,
the following species were introduced into the steppe
and forest�steppe zones in Ukraine: Mentissa gracili�
costa Rossmässler, 1836, Cecilioides raddei Boettger,
1879, Monacha fruticola (Krynicki, 1833), Phenacoli�
max annularis (Studer, 1820), Brephulopsis bidens
(Krynicky, 1883), Oxychilus diaphanellus (Krynicky,
1833), etc. [Sverlova and Son, 2006]. In turn, Helix
pomatia Linnaeus, 1758 invaded Crimea from conti�
nental Ukraine [Sverlova and Son, 2006].
Such steppe species as Brephulopsis cylindrica
(Mence, 1828), Monacha carthusiana (Müller, 1774),
Xeropicta spp., and Helix albescens Rossmässler, 1839
considerably expanded their ranges owing to introduc�
tion to the urbanized ecosystems in Central, Eastern,
and Western Ukraine [Sverlova et al., 2006; Sverlova
and Son, 2006; Balashov and Vasilyuk, 2007].
Two main directions of the invasion of exotic spe�
cies from the remote regions are clearly distinguish�
able. The first one is the expansion of Balkan and
Mediterranean mollusks (Chondrula microtraga
(Rossmaessler, 1839), Limax flavus (Linnaeus, 1758),
Limax maculatus (Kaleniczenko, 1851), Eobania ver�
miculata (Müller, 1774), Helix lucorum Linnaeus,
1758, Tandonia kusceri (Wagner, 1931), etc.) to the
historical regions of active trade and colonization of
Table 1. (Contd.)
Species Native region Time of intro�duction Distribution in Ukraine References
Rapana venosa Sea of Japan 1954 Southern part of the Sea of
Azov and the Black Sea
Zolotarev, 1996
Teredo navalis Pacific and Atlantic
oceans
750–500 B.C. Black Sea and Sea of Azov Alexandrov et al., 2007
Terrestrial species
Aegopinella nitidula Northwestern Europe End of
20th century
Lviv Sverlova and Son, 2006
Boettgerilla pallens Caucasus 20th century Kyiv, Western Ukraine Sverlova and Son, 2006
Cepaea nemoralis Western Europe Western Ukraine Sverlova and Son, 2006
C. hortensis Central Europe 1990s Western Ukraine Sverlova and Son, 2006
Chondrula microtraga Balkans, Romania,
Asia Minor
1999 Odessa Sverlova and Son, 2006
Cryptomphalus aspersa
(Müller, 1774)
Mediterranean 19th century Several findings in Southern
Ukraine and the Yuzhnyi Bug
River; probably recently
absent from the fauna
Sverlova and Son, 2006
Eobania vermiculata Mediterranean ancient period
or period of
Crimean War
Crimea, Odessa oblast Sverlova and Son, 2006
Helix lucorum Southern Europe, Asia
Minor, Middle East,
Caucasus
19th century Crimea Sverlova, Son, 2006;
Leonov, 2006
Limax maximus Southern and Central
Europe
20th century Western and Central Ukraine,
Crimea
Sverlova and Son, 2006
L. flavus Mediterranean 20th century Odessa, Yalta Sverlova and Son, 2006
Oxychilus draparnaudi Mediterranean,
Western Europe
End of
20th century
Lviv Sverlova and Son, 2006
Stenomphalia ravergieri Caucasus, Transcaucasia 1998 Southeastern Ukraine Sverlova et al., 2006
Tandonia kusceri Balkans, Western
Black Sea coast
Beginning of
20th century
Odessa and Nikolayev oblasts Son, 2004
RUSSIAN JOURNAL OF BIOLOGICAL INVASIONS Vol. 1 No. 1 2010
SON
introduction to a certain habitat, the species may live
there for decades without moving beyond the borders
of the habitat. Thus, the directions of invasions of ter�
restrial mollusks are determined predominantly by the
routes of transportation and, hence, by trade and eco�
nomic ties between various regions.
The most noticeable process in the invasions of ter�
restrial mollusks is a mass expansion of the Crimean
and Caucasian species towards the steppe zone and
urban territories of other regions in Ukraine. Along
with the aforementioned species, some other mollusks
typical of the steppes also penetrate into urban territo�
ries in Central and Western Ukraine.
Among the species endemic to the Crimea or ear�
lier found in Southern Ukraine only on this peninsula,
the following species were introduced into the steppe
and forest�steppe zones in Ukraine: Mentissa gracili�
costa Rossmässler, 1836, Cecilioides raddei Boettger,
1879, Monacha fruticola (Krynicki, 1833), Phenacoli�
max annularis (Studer, 1820), Brephulopsis bidens
(Krynicky, 1883), Oxychilus diaphanellus (Krynicky,
1833), etc. [Sverlova and Son, 2006]. In turn, Helix
pomatia Linnaeus, 1758 invaded Crimea from conti�
nental Ukraine [Sverlova and Son, 2006].
Such steppe species as Brephulopsis cylindrica
(Mence, 1828), Monacha carthusiana (Müller, 1774),
Xeropicta spp., and Helix albescens Rossmässler, 1839
considerably expanded their ranges owing to introduc�
tion to the urbanized ecosystems in Central, Eastern,
and Western Ukraine [Sverlova et al., 2006; Sverlova
and Son, 2006; Balashov and Vasilyuk, 2007].
Two main directions of the invasion of exotic spe�
cies from the remote regions are clearly distinguish�
able. The first one is the expansion of Balkan and
Mediterranean mollusks (Chondrula microtraga
(Rossmaessler, 1839), Limax flavus (Linnaeus, 1758),
Limax maculatus (Kaleniczenko, 1851), Eobania ver�
miculata (Müller, 1774), Helix lucorum Linnaeus,
1758, Tandonia kusceri (Wagner, 1931), etc.) to the
historical regions of active trade and colonization of
Table 1. (Contd.)
Species Native region Time of intro�duction Distribution in Ukraine References
Rapana venosa Sea of Japan 1954 Southern part of the Sea of
Azov and the Black Sea
Zolotarev, 1996
Teredo navalis Pacific and Atlantic
oceans
750–500 B.C. Black Sea and Sea of Azov Alexandrov et al., 2007
Terrestrial species
Aegopinella nitidula Northwestern Europe End of
20th century
Lviv Sverlova and Son, 2006
Boettgerilla pallens Caucasus 20th century Kyiv, Western Ukraine Sverlova and Son, 2006
Cepaea nemoralis Western Europe Western Ukraine Sverlova and Son, 2006
C. hortensis Central Europe 1990s Western Ukraine Sverlova and Son, 2006
Chondrula microtraga Balkans, Romania,
Asia Minor
1999 Odessa Sverlova and Son, 2006
Cryptomphalus aspersa
(Müller, 1774)
Mediterranean 19th century Several findings in Southern
Ukraine and the Yuzhnyi Bug
River; probably recently
absent from the fauna
Sverlova and Son, 2006
Eobania vermiculata Mediterranean ancient period
or period of
Crimean War
Crimea, Odessa oblast Sverlova and Son, 2006
Helix lucorum Southern Europe, Asia
Minor, Middle East,
Caucasus
19th century Crimea Sverlova, Son, 2006;
Leonov, 2006
Limax maximus Southern and Central
Europe
20th century Western and Central Ukraine,
Crimea
Sverlova and Son, 2006
L. flavus Mediterranean 20th century Odessa, Yalta Sverlova and Son, 2006
Oxychilus draparnaudi Mediterranean,
Western Europe
End of
20th century
Lviv Sverlova and Son, 2006
Stenomphalia ravergieri Caucasus, Transcaucasia 1998 Southeastern Ukraine Sverlova et al., 2006
Tandonia kusceri Balkans, Western
Black Sea coast
Beginning of
20th century
Odessa and Nikolayev oblasts Son, 2004
Page 5
RUSSIAN JOURNAL OF BIOLOGICAL INVASIONS Vol. 1 No. 1 2010
ALIEN MOLLUSKS WITHIN THE TERRITORY OF UKRAINE 41
Table 2. Expansion of ranges of terrestrial and freshwater mollusks—neighboring invaders within the territory of Ukraine
Species Change in the range in Ukraine References
Freshwater species
Dreissena bugensis Source range: Bugskii Liman and Dnieper delta. Estab�
lished in all large river basins in Ukraine and in the system of
Severno�Krymskii Canal
Kharchenko, 1995,
Son, 2007a
D. polymorpha Source in Ukraine—the rivers of the Black Sea coast and
limans in continental Ukraine. Expanded range in many
basins, established in many man�made canals and water
bodies all over Ukrainian territory
Kharchenko, 1995;
Son, 2007a
Euxinipyrgula lincta (Milaschev�
itch, 1908)
Relict Ponto�Caspian species. In 2006, found in the
Nizhnednestrovskaya irrigation system (Baraboi River)
Son, 2007
Fagotia danubialis Endemic species of the rivers of Northwestern Black Sea
coast. In 2000, established in the Chernaya River and Cher�
norechenskoe reservoir (Crimea)
Mironov et al., 2002
Hypanis angusticostata (Borcea, 1926) Spread upstream in the Dnieper River cascade of reservoirs Munasypova–Mot�
yash, 2006
H. colorata (Eichwald, 1829) Spread upstream in the Dnieper River cascade of reservoirs.
Spread in series of irrigation systems in different time periods
Munasypova–Mot�
yash, 2006; Son, 2007
H. fragilis (Milaschevitch, 1908) Introduced from the Danube delta to the Dunai–Dnestro�
vskaya irrigation system (Sasyk Lake)
Munasypova–Mot�
yash, 2006
H. pontica (Eichwald, 1838) Spread upstream in the Dnieper River cascade of reservoirs Alexandrov et al., 2007
Lymnaea ovata, L. stagnalis, Lithogly�
phus naticoides, Viviparus viviparus,
Sphaerium corneum (Linnaeus,
1758); S. rivicola (Lamarck, 1818)
The species widely distributed in the freshwaters of conti�
nental Ukraine; introduced into Crimea during hydroengi�
neering in middle of 20th century
Zhuravel’ et al., 1968,
Son, 2007
Theodoxus fluviatilis Widely distributed in the freshwaters of Ukraine. Introductions
into numerous man�made canals and reservoirs of the Dnieper
River, Crimea, Sasyk Lake, etc., noted in different time periods
Son, 2007
Theodoxus donasteri (Lindholm,
1908)
Widely distributed in the downstream reaches of large rivers.
Introduction into Sasyk Lake noted at end of 20th century
Son, 2007
Theodoxus euxinus (Clessin, 1886) Inhabitant of the Black Sea limans. At beginning of 21st
century, introductions into the systems of man�made canals
on the Black Sea coast and in the cooling pond of
Khmel’nitskaya nuclear power station noted
Anistratenko et al.,
2007b; Son, 2007
Terrestrial species
Arion distinctus Species with ambiguous source range. At least part of popu�
lations in the Western and Northern Ukraine introduced
Sverlova, Son, 2006
Bielzia coerulans (Bielz, 1851) Carpathian species; noted in Kyiv in 2002 Sverlova, Son, 2006
Brephulopsis bidens Endemic to Crimea; noted in Odessa oblast at end of 20th century Sverlova, Son, 2006
B. cylindrica Species of the Black Sea coast (presumably, initially
endemic to Crimea); during 20th–21st century, the range
considerably expanded owing to introduction into the urban
ecosystems of Central, Eastern, and Western Ukraine
Sverlova, Son, 2006;
Sverlova et al., 2006
Cecilioides raddei Species lived in Ukraine only within the territory of Crimea;
at end of 20th century noted in the Odessa oblast
Sverlova, Son, 2006
Deroceras caucasicum Species lived in Ukraine only within the territory of Crimea;
at beginning of 21st century noted in city of Donetsk
Sverlova, Son, 2006
Helix albescens Species widely distributed in Southern Ukraine; in 2006
noted within the city of Kyiv
Balashov and Vasi�
lyuk, 2007
H. pomatia Species widely distributed within the territory of continental
Ukraine; introduced into Crimea
Sverlova, Son, 2006
ALIEN MOLLUSKS WITHIN THE TERRITORY OF UKRAINE 41
Table 2. Expansion of ranges of terrestrial and freshwater mollusks—neighboring invaders within the territory of Ukraine
Species Change in the range in Ukraine References
Freshwater species
Dreissena bugensis Source range: Bugskii Liman and Dnieper delta. Estab�
lished in all large river basins in Ukraine and in the system of
Severno�Krymskii Canal
Kharchenko, 1995,
Son, 2007a
D. polymorpha Source in Ukraine—the rivers of the Black Sea coast and
limans in continental Ukraine. Expanded range in many
basins, established in many man�made canals and water
bodies all over Ukrainian territory
Kharchenko, 1995;
Son, 2007a
Euxinipyrgula lincta (Milaschev�
itch, 1908)
Relict Ponto�Caspian species. In 2006, found in the
Nizhnednestrovskaya irrigation system (Baraboi River)
Son, 2007
Fagotia danubialis Endemic species of the rivers of Northwestern Black Sea
coast. In 2000, established in the Chernaya River and Cher�
norechenskoe reservoir (Crimea)
Mironov et al., 2002
Hypanis angusticostata (Borcea, 1926) Spread upstream in the Dnieper River cascade of reservoirs Munasypova–Mot�
yash, 2006
H. colorata (Eichwald, 1829) Spread upstream in the Dnieper River cascade of reservoirs.
Spread in series of irrigation systems in different time periods
Munasypova–Mot�
yash, 2006; Son, 2007
H. fragilis (Milaschevitch, 1908) Introduced from the Danube delta to the Dunai–Dnestro�
vskaya irrigation system (Sasyk Lake)
Munasypova–Mot�
yash, 2006
H. pontica (Eichwald, 1838) Spread upstream in the Dnieper River cascade of reservoirs Alexandrov et al., 2007
Lymnaea ovata, L. stagnalis, Lithogly�
phus naticoides, Viviparus viviparus,
Sphaerium corneum (Linnaeus,
1758); S. rivicola (Lamarck, 1818)
The species widely distributed in the freshwaters of conti�
nental Ukraine; introduced into Crimea during hydroengi�
neering in middle of 20th century
Zhuravel’ et al., 1968,
Son, 2007
Theodoxus fluviatilis Widely distributed in the freshwaters of Ukraine. Introductions
into numerous man�made canals and reservoirs of the Dnieper
River, Crimea, Sasyk Lake, etc., noted in different time periods
Son, 2007
Theodoxus donasteri (Lindholm,
1908)
Widely distributed in the downstream reaches of large rivers.
Introduction into Sasyk Lake noted at end of 20th century
Son, 2007
Theodoxus euxinus (Clessin, 1886) Inhabitant of the Black Sea limans. At beginning of 21st
century, introductions into the systems of man�made canals
on the Black Sea coast and in the cooling pond of
Khmel’nitskaya nuclear power station noted
Anistratenko et al.,
2007b; Son, 2007
Terrestrial species
Arion distinctus Species with ambiguous source range. At least part of popu�
lations in the Western and Northern Ukraine introduced
Sverlova, Son, 2006
Bielzia coerulans (Bielz, 1851) Carpathian species; noted in Kyiv in 2002 Sverlova, Son, 2006
Brephulopsis bidens Endemic to Crimea; noted in Odessa oblast at end of 20th century Sverlova, Son, 2006
B. cylindrica Species of the Black Sea coast (presumably, initially
endemic to Crimea); during 20th–21st century, the range
considerably expanded owing to introduction into the urban
ecosystems of Central, Eastern, and Western Ukraine
Sverlova, Son, 2006;
Sverlova et al., 2006
Cecilioides raddei Species lived in Ukraine only within the territory of Crimea;
at end of 20th century noted in the Odessa oblast
Sverlova, Son, 2006
Deroceras caucasicum Species lived in Ukraine only within the territory of Crimea;
at beginning of 21st century noted in city of Donetsk
Sverlova, Son, 2006
Helix albescens Species widely distributed in Southern Ukraine; in 2006
noted within the city of Kyiv
Balashov and Vasi�
lyuk, 2007
H. pomatia Species widely distributed within the territory of continental
Ukraine; introduced into Crimea
Sverlova, Son, 2006
Page 6
42
RUSSIAN JOURNAL OF BIOLOGICAL INVASIONS Vol. 1 No. 1 2010
SON
the Northwestern Black Sea coast and Crimea (cities
of Odessa, Sevastopol, etc.). Further, the ranges of
these species expand in the southern regions. The sec�
ond direction is the expansion of Central European
species (Arion distinctus Mabille, 1868, Aegopinella
nitidula (Draparnaud, 1805), Oxychilus draparnaudi
(Beck, 1837), Limax maximus Linnaeus, 1758, Cepaea
nemoralis (Linnaeus, 1758), Cepaea hortensis (Müller,
1774), establishing themselves mostly in the forest�
steppe zone [Son, 2004; Leonov, 2006; Sverlova et al.,
2006; Sverlova and Son, 2006].
In recent years, Caucasian and Crimean–Cauca�
sian species (Krynickillus melanocephalus Kalenic�
zenko, 1851, Deroceras caucasicum (Simroth, 1901),
Stenomphalia ravergieri (Ferussac, 1835), Boettgerilla
pallens Simroth, 1912) have actively been introduced
into the various regions of Ukraine [Sverlova et al.,
2006; Sverlova and Son, 2006; Gural’�Sverlova and
Martynov, 2009].
It is not yet possible to affirm with certainty the
allochthonous status of some species that earlier were
classified as endemics of Crimea or the Caucasus (Elia
novorossica (Retowskii, 1888), Helicopsis filimargo
(Krynicky, 1836), Helicopsis retowskii (Clessin,
1883)). These species were found in habitats serving as
refuges for many relict species. In addition, the mol�
lusk fauna in Southeastern Ukraine only relatively
recently began to be studied in detail.
Other directions of invasions of terrestrial mollusks
(from the Carpathians to the plain part of Western
Ukraine, from Western Ukraine to the steppe zone,
etc.) are solitary.
The analysis of certain cases of expansion of terres�
trial alien mollusks shows that, besides weakening of
the zoogeographic barriers, the presence of refuges
plays an important role in their spread. As such ref�
uges, the most important are various urban anthropo�
genic biotopes: greenhouses, parks, etc.
Specific climatic conditions are often formed in
large cities. These conditions may differ from the con�
ditions of relevant physical geographical zones. Con�
sequently, the urban biotopes in general are closer to
each other than to the zonal natural biotopes. This is
why the cities often become places of “accumulation”
of nonendemic species, making possible their further
expansion in a region subject to further anthropogenic
transformation.
CONCLUSIONS
The invasions of mollusks into the Black Sea and
the Sea of Azov, inland waters, and terrestrial ecosys�
tems in fact have no common sources and invasion
vectors and are driven by different mechanisms. The
most uniform in the aspects of invasion vectors and
routes is the situation with alien species in marine eco�
systems. The invasions of mollusks into continental
waters occur predominantly in the estuaries and main
channels of large river basins. This localizes biological
Table 2. (Contd.)
Species Change in the range in Ukraine References
Krynickillus melanocephalus Species lived in Ukraine only within the territory of Crimea;
introduced populations found in cities of Kyiv, Lviv and in
Donetsk oblast
Sverlova, Son, 2006;
Sverlova et al., 2006
Limax maculatus Species lived in Ukraine only within the territory of Crimea;
since end of 20th century widely established in urban eco�
systems of Southern Ukraine
Sverlova, Son, 2006;
Sverlova et al., 2006
Mentissa gracilicosta Endemic to Crimea; in 1994 found in city of Odessa Sverlova, Son, 2006
Monacha carthusiana Species earlier lived only the southern regions of Ukraine;
since end of 20th century, introduced populations appeared
in Western Ukraine
Sverlova, Son, 2006
M. fruticola Species lived in Ukraine only within the territory of Crimea;
established in the northwestern coastal regions of the Black
Sea; noted in Zaporizhzhya oblast
Sverlova, Son, 2006;
Sverlova et al., 2006
Oxychilus diaphanellus Endemic to Crimea; in 2002–2005 introduced into urban
ecosystems of cities of Zaporizhzhya, Donetsk, and Kharkiv
Sverlova et al., 2006
Phenacolimax annularis Species earlier lived in Ukraine only within the territory of
Crimea; at beginning of 21st century noted in city of Odessa
Sverlova, Son, 2006
Xeropicta derbentina (Krynicki, 1836) Species lived in Ukraine only within the territory of Crimea;
established in the northwestern coastal regions of the Black
Sea; noted in Zaporizhzhya oblast
Sverlova, Son, 2006;
Sverlova et al., 2006
X. krynicki (Krynicky, 1836) Species earlier lived in Ukraine only within the territory of
Crimea and in the northwestern coastal regions of the Black
Sea; gradual steady expansion of the range taking place
Sverlova, Son, 2006
RUSSIAN JOURNAL OF BIOLOGICAL INVASIONS Vol. 1 No. 1 2010
SON
the Northwestern Black Sea coast and Crimea (cities
of Odessa, Sevastopol, etc.). Further, the ranges of
these species expand in the southern regions. The sec�
ond direction is the expansion of Central European
species (Arion distinctus Mabille, 1868, Aegopinella
nitidula (Draparnaud, 1805), Oxychilus draparnaudi
(Beck, 1837), Limax maximus Linnaeus, 1758, Cepaea
nemoralis (Linnaeus, 1758), Cepaea hortensis (Müller,
1774), establishing themselves mostly in the forest�
steppe zone [Son, 2004; Leonov, 2006; Sverlova et al.,
2006; Sverlova and Son, 2006].
In recent years, Caucasian and Crimean–Cauca�
sian species (Krynickillus melanocephalus Kalenic�
zenko, 1851, Deroceras caucasicum (Simroth, 1901),
Stenomphalia ravergieri (Ferussac, 1835), Boettgerilla
pallens Simroth, 1912) have actively been introduced
into the various regions of Ukraine [Sverlova et al.,
2006; Sverlova and Son, 2006; Gural’�Sverlova and
Martynov, 2009].
It is not yet possible to affirm with certainty the
allochthonous status of some species that earlier were
classified as endemics of Crimea or the Caucasus (Elia
novorossica (Retowskii, 1888), Helicopsis filimargo
(Krynicky, 1836), Helicopsis retowskii (Clessin,
1883)). These species were found in habitats serving as
refuges for many relict species. In addition, the mol�
lusk fauna in Southeastern Ukraine only relatively
recently began to be studied in detail.
Other directions of invasions of terrestrial mollusks
(from the Carpathians to the plain part of Western
Ukraine, from Western Ukraine to the steppe zone,
etc.) are solitary.
The analysis of certain cases of expansion of terres�
trial alien mollusks shows that, besides weakening of
the zoogeographic barriers, the presence of refuges
plays an important role in their spread. As such ref�
uges, the most important are various urban anthropo�
genic biotopes: greenhouses, parks, etc.
Specific climatic conditions are often formed in
large cities. These conditions may differ from the con�
ditions of relevant physical geographical zones. Con�
sequently, the urban biotopes in general are closer to
each other than to the zonal natural biotopes. This is
why the cities often become places of “accumulation”
of nonendemic species, making possible their further
expansion in a region subject to further anthropogenic
transformation.
CONCLUSIONS
The invasions of mollusks into the Black Sea and
the Sea of Azov, inland waters, and terrestrial ecosys�
tems in fact have no common sources and invasion
vectors and are driven by different mechanisms. The
most uniform in the aspects of invasion vectors and
routes is the situation with alien species in marine eco�
systems. The invasions of mollusks into continental
waters occur predominantly in the estuaries and main
channels of large river basins. This localizes biological
Table 2. (Contd.)
Species Change in the range in Ukraine References
Krynickillus melanocephalus Species lived in Ukraine only within the territory of Crimea;
introduced populations found in cities of Kyiv, Lviv and in
Donetsk oblast
Sverlova, Son, 2006;
Sverlova et al., 2006
Limax maculatus Species lived in Ukraine only within the territory of Crimea;
since end of 20th century widely established in urban eco�
systems of Southern Ukraine
Sverlova, Son, 2006;
Sverlova et al., 2006
Mentissa gracilicosta Endemic to Crimea; in 1994 found in city of Odessa Sverlova, Son, 2006
Monacha carthusiana Species earlier lived only the southern regions of Ukraine;
since end of 20th century, introduced populations appeared
in Western Ukraine
Sverlova, Son, 2006
M. fruticola Species lived in Ukraine only within the territory of Crimea;
established in the northwestern coastal regions of the Black
Sea; noted in Zaporizhzhya oblast
Sverlova, Son, 2006;
Sverlova et al., 2006
Oxychilus diaphanellus Endemic to Crimea; in 2002–2005 introduced into urban
ecosystems of cities of Zaporizhzhya, Donetsk, and Kharkiv
Sverlova et al., 2006
Phenacolimax annularis Species earlier lived in Ukraine only within the territory of
Crimea; at beginning of 21st century noted in city of Odessa
Sverlova, Son, 2006
Xeropicta derbentina (Krynicki, 1836) Species lived in Ukraine only within the territory of Crimea;
established in the northwestern coastal regions of the Black
Sea; noted in Zaporizhzhya oblast
Sverlova, Son, 2006;
Sverlova et al., 2006
X. krynicki (Krynicky, 1836) Species earlier lived in Ukraine only within the territory of
Crimea and in the northwestern coastal regions of the Black
Sea; gradual steady expansion of the range taking place
Sverlova, Son, 2006
Page 7
RUSSIAN JOURNAL OF BIOLOGICAL INVASIONS Vol. 1 No. 1 2010
ALIEN MOLLUSKS WITHIN THE TERRITORY OF UKRAINE 43
invasions within the borders of linear invasion corri�
dors.
REFERENCES
Aleksandrov B., Boltachev A., Kharchenko T., Lyash�
enko A., Son M., Tsarenko P., Zhukinsky V. Trends of
Aquatic Alien Species Invasion in Ukraine, Aquat.
Invasions 2007, vol. 2, pp. 215–242.
Anistratenko, V., Anistratenko, O., and Khaliman, I.,
Gastropods of the Sea of Azov: Zoogeographical Com�
position and Characteristics of Biology as a Function of
Salinity, Ruthenica, 2008, vol. 18, pp. 9–16.
Anistratenko, V.V., Anistratenko, O.Yu., and Khali�
man, I.A., Composition of the Molluscan Fauna of the
Sea of Azov as a Function of Its Salinity, Dopov. NAN
Ukr., 2007a, vol. 4, pp. 161–166.
Anistratenko, V.V., Identification Guide to Gastropod
Mollusks of the Order Pectinibranchia (Gastropoda,
Pectinibranchia) of the Fauna of Ukraine. Part 1:
Marine and Brackish�Water Mollusks, Vestn. Zool.,
1998, issue 8, pp. 3–65.
Anistratenko, V.V., Protasov, A.A., and Babariga, S.P.,
The First Discovery of the Gastropod Mollusk Theo�
doxus euxinus (Gastropoda, Neritidae) in the Water
Reservoir�Cooler of Khmel’nitskaya Atomic Power
Plant, Vestn. Zool., 2007b, vol. 41, p. 168.
Balashov, Z.O. and Vasilyuk, O.V., Finding of a Colony
of Helix albescens (Gastropoda, Geophila, Helicidae)
in Kyiv, Nauk. Zap. Derzhav. Prirodoznav. Muz., 2007,
vol. 23, pp. 227–228.
Grigorovich, I.A., MacIsaac, H.J., Shadrin, N.V., and
Mills, E.L., Patterns and Mechanisms of Aquatic
Invertebrate Introductions in the Ponto�Caspian
Region, Can. J. Fish. Aquat. Sci., 2002, vol. 59,
pp. 1189–1208.
Gural'�Sverlova, N.V. and Martynov, V.V., The First
Finding of Terrestrial Mollusks of the Genus Elia
(Clausiliidae) in Ukraine, Ruthenica, 2009, vol. 19,
pp. 29–30.
Khaliman, I.A. and Anistratenko, V.V., Melanoides
granifera, One More Invader Molluscan Species in the
Fauna of Ukraine, Vestn. Zool., 2006, vol. 40, p. 320.
Kharchenko, T.A., Dreissena: Range, Ecology, and
Biological Hindrances Gidrobiol. Zh., 1995, vol. 31,
no. 3, pp. 3–21.
Kovtun, O.A. and Zolotarev, V.N., The First Finding of
the Pacific Giant Oyster Crassostrea gigas (Bivalvia,
Ostreidae) in the Gulf of Odessa (Black Sea), Vestn.
Zool., 2008, vol. 42, p. 262.
Leonov, S.V., Historical Aspects of the Use of Terres�
trial Snails by Humans in the Crimea, in Fauna,
ekologiya i vnutrividovaya izmenchivost’ nazemnykh
mollyuskov v urbanizirovannoi srede (Fauna, Ecology,
and Intraspecific Variability of Terrestrial Mollusks in
Urban Environment), Lvov: Gos. Prirodoved. Muzei,
2006, pp. 180–188.
Lutaenko, K.A., The Fauna of Bivalve Mollusks of the
Subfamily Anadarinae (Arcidae) in Southern India,
Byul. Dal’nevost. Malakol. Obshch., 2006, vol. 10,
pp. 102–121.
Mironov, S.S., Shadrin, N.V., and Grintsov, V.A., New
Molluscan Species in Sea and Continental Waters of
the Crimea, Ekol. Morya, 2002, vol. 61, p. 43.
Munasypova�Motyash, I.A., Modern Fauna of Bivalve
Mollusks of the SubfamilyLimnocardiinae (Bivalvia,
Cardiidae) of the Northwestern Black Sea Region,
Vestn. Zool., 2006, vol. 40, pp. 41–48.
Panov, V.E., Alexandrov, B., Arbaciauskas, K., Binime�
lis, R., Copp, G.H., Grabowski, M., Lucy, F., Leuven,
R.S.E.W., Nehring, S., Paunovic, M., Semenchenko, V.,
and Son, M.O., Assessing the Risks of Aquatic Species
Invasions via European Inland Waterways: From Con�
cepts to Environmental Indicators, Integr. Environ.
Assess. Manag., 2009, vol. 5, pp. 110–126.
Puzanov, I.I., Mediterraneanization of the Black Sea
and the Prospects of Its Enhancement, Zool. Zh., 1967,
vol. 46, pp. 1287–1297.
Puzanov, I.I., Successive Stages of Mediterraneaniza�
tion of the Black Sea Fauna: New Data, Gidrobiol. Zh.,
1965, vol. 1, pp. 54–56.
Roginskaya, I.S. and Grintsov, V.A., Sea Slugh
Doridella obscura Verill: A New Invader Species in the
Black Sea, Okeanologiya, 1990, vol. 30, pp. 855–857.
Semenchenko, V. and Laenko, T., First Record of the
Invasive North American Gastropod Ferrissia fragilis
(Tryon, 1863) from the Pripyat River Basin, Belarus,
Aquat. Invasions, 2008, vol. 3, pp. 80–82.
Shadrin, N.V., Mironov, S.S., and Golikov, A.N., The
Finding in the Black Sea of Living Samples of Neptunea
arthritica (Bernardi, 1857) (Gastropoda, Buccinidae),
Ekol. Morya, 2002, vol. 62, p. 29.
Son, M.O., Mollyuski�Vselentsy V Presnykh I Solono�
vatykh Vodakh Severnogo Prichernomor’ya (Mollusks�
Invaders in Fresh and Brackish Waters of Northern
Black Sea Region), Odessa: Druk, 2007.
Son, M.O., Native Range of the Zebra Mussel and
Quagga Mussel and New Data on Their Invasions
within the Ponto�Caspian Region, Aquat. Invasions,
2007a, vol. 2, pp. 174–184.
Son, M.O., New Findings of Tandonia kusceri (Pulmo�
nata, Milacidae) in Ukraine, Vestn. Zool., 2004, vol. 38,
p. 76.
Son, M.O., North American Freshwater Limpet Ferris�
sia fragilis (Tryon, 1863)—the Cryptic Invader in the
Northern Black Sea Region, Aquat. Invasions, 2007b,
vol. 2, pp. 55–58.
Son, M.O., Rapid Expansion of the New Zealand Mud
Snail Potamopyrgus antipodarum (Gray, 1843) in the
Azov–Black Sea Region, Aquat. Invasions, 2008, vol. 3,
pp. 335–340.
Sverlova, N.V. and Son, M.O., Mollusks�Invaders and
Their Place in Urban Malacocenoses, in Fauna,
ekologiya i vnutrividovaya izmenchivost' nazemnykh
mollyuskov v urbanizirovannoi srede (Fauna, Ecology,
and Intraspecific Variability of Terrestrial Mollusks in
Urban Environment), Lvov: Gos. Prirodoved. Muzei,
2006, pp. 42–59.
Sverlova, N.V., Martinov, V.V., and Martinov, O.V.,
Study of Terrestrial Malacofauna (Gastropoda: Pulmo�
nata) of Southeastern Ukraine, Nauk. Zap. Derzhav.
Prirod. Muz., 2006, vol. 22, pp. 35–46.
ALIEN MOLLUSKS WITHIN THE TERRITORY OF UKRAINE 43
invasions within the borders of linear invasion corri�
dors.
REFERENCES
Aleksandrov B., Boltachev A., Kharchenko T., Lyash�
enko A., Son M., Tsarenko P., Zhukinsky V. Trends of
Aquatic Alien Species Invasion in Ukraine, Aquat.
Invasions 2007, vol. 2, pp. 215–242.
Anistratenko, V., Anistratenko, O., and Khaliman, I.,
Gastropods of the Sea of Azov: Zoogeographical Com�
position and Characteristics of Biology as a Function of
Salinity, Ruthenica, 2008, vol. 18, pp. 9–16.
Anistratenko, V.V., Anistratenko, O.Yu., and Khali�
man, I.A., Composition of the Molluscan Fauna of the
Sea of Azov as a Function of Its Salinity, Dopov. NAN
Ukr., 2007a, vol. 4, pp. 161–166.
Anistratenko, V.V., Identification Guide to Gastropod
Mollusks of the Order Pectinibranchia (Gastropoda,
Pectinibranchia) of the Fauna of Ukraine. Part 1:
Marine and Brackish�Water Mollusks, Vestn. Zool.,
1998, issue 8, pp. 3–65.
Anistratenko, V.V., Protasov, A.A., and Babariga, S.P.,
The First Discovery of the Gastropod Mollusk Theo�
doxus euxinus (Gastropoda, Neritidae) in the Water
Reservoir�Cooler of Khmel’nitskaya Atomic Power
Plant, Vestn. Zool., 2007b, vol. 41, p. 168.
Balashov, Z.O. and Vasilyuk, O.V., Finding of a Colony
of Helix albescens (Gastropoda, Geophila, Helicidae)
in Kyiv, Nauk. Zap. Derzhav. Prirodoznav. Muz., 2007,
vol. 23, pp. 227–228.
Grigorovich, I.A., MacIsaac, H.J., Shadrin, N.V., and
Mills, E.L., Patterns and Mechanisms of Aquatic
Invertebrate Introductions in the Ponto�Caspian
Region, Can. J. Fish. Aquat. Sci., 2002, vol. 59,
pp. 1189–1208.
Gural'�Sverlova, N.V. and Martynov, V.V., The First
Finding of Terrestrial Mollusks of the Genus Elia
(Clausiliidae) in Ukraine, Ruthenica, 2009, vol. 19,
pp. 29–30.
Khaliman, I.A. and Anistratenko, V.V., Melanoides
granifera, One More Invader Molluscan Species in the
Fauna of Ukraine, Vestn. Zool., 2006, vol. 40, p. 320.
Kharchenko, T.A., Dreissena: Range, Ecology, and
Biological Hindrances Gidrobiol. Zh., 1995, vol. 31,
no. 3, pp. 3–21.
Kovtun, O.A. and Zolotarev, V.N., The First Finding of
the Pacific Giant Oyster Crassostrea gigas (Bivalvia,
Ostreidae) in the Gulf of Odessa (Black Sea), Vestn.
Zool., 2008, vol. 42, p. 262.
Leonov, S.V., Historical Aspects of the Use of Terres�
trial Snails by Humans in the Crimea, in Fauna,
ekologiya i vnutrividovaya izmenchivost’ nazemnykh
mollyuskov v urbanizirovannoi srede (Fauna, Ecology,
and Intraspecific Variability of Terrestrial Mollusks in
Urban Environment), Lvov: Gos. Prirodoved. Muzei,
2006, pp. 180–188.
Lutaenko, K.A., The Fauna of Bivalve Mollusks of the
Subfamily Anadarinae (Arcidae) in Southern India,
Byul. Dal’nevost. Malakol. Obshch., 2006, vol. 10,
pp. 102–121.
Mironov, S.S., Shadrin, N.V., and Grintsov, V.A., New
Molluscan Species in Sea and Continental Waters of
the Crimea, Ekol. Morya, 2002, vol. 61, p. 43.
Munasypova�Motyash, I.A., Modern Fauna of Bivalve
Mollusks of the SubfamilyLimnocardiinae (Bivalvia,
Cardiidae) of the Northwestern Black Sea Region,
Vestn. Zool., 2006, vol. 40, pp. 41–48.
Panov, V.E., Alexandrov, B., Arbaciauskas, K., Binime�
lis, R., Copp, G.H., Grabowski, M., Lucy, F., Leuven,
R.S.E.W., Nehring, S., Paunovic, M., Semenchenko, V.,
and Son, M.O., Assessing the Risks of Aquatic Species
Invasions via European Inland Waterways: From Con�
cepts to Environmental Indicators, Integr. Environ.
Assess. Manag., 2009, vol. 5, pp. 110–126.
Puzanov, I.I., Mediterraneanization of the Black Sea
and the Prospects of Its Enhancement, Zool. Zh., 1967,
vol. 46, pp. 1287–1297.
Puzanov, I.I., Successive Stages of Mediterraneaniza�
tion of the Black Sea Fauna: New Data, Gidrobiol. Zh.,
1965, vol. 1, pp. 54–56.
Roginskaya, I.S. and Grintsov, V.A., Sea Slugh
Doridella obscura Verill: A New Invader Species in the
Black Sea, Okeanologiya, 1990, vol. 30, pp. 855–857.
Semenchenko, V. and Laenko, T., First Record of the
Invasive North American Gastropod Ferrissia fragilis
(Tryon, 1863) from the Pripyat River Basin, Belarus,
Aquat. Invasions, 2008, vol. 3, pp. 80–82.
Shadrin, N.V., Mironov, S.S., and Golikov, A.N., The
Finding in the Black Sea of Living Samples of Neptunea
arthritica (Bernardi, 1857) (Gastropoda, Buccinidae),
Ekol. Morya, 2002, vol. 62, p. 29.
Son, M.O., Mollyuski�Vselentsy V Presnykh I Solono�
vatykh Vodakh Severnogo Prichernomor’ya (Mollusks�
Invaders in Fresh and Brackish Waters of Northern
Black Sea Region), Odessa: Druk, 2007.
Son, M.O., Native Range of the Zebra Mussel and
Quagga Mussel and New Data on Their Invasions
within the Ponto�Caspian Region, Aquat. Invasions,
2007a, vol. 2, pp. 174–184.
Son, M.O., New Findings of Tandonia kusceri (Pulmo�
nata, Milacidae) in Ukraine, Vestn. Zool., 2004, vol. 38,
p. 76.
Son, M.O., North American Freshwater Limpet Ferris�
sia fragilis (Tryon, 1863)—the Cryptic Invader in the
Northern Black Sea Region, Aquat. Invasions, 2007b,
vol. 2, pp. 55–58.
Son, M.O., Rapid Expansion of the New Zealand Mud
Snail Potamopyrgus antipodarum (Gray, 1843) in the
Azov–Black Sea Region, Aquat. Invasions, 2008, vol. 3,
pp. 335–340.
Sverlova, N.V. and Son, M.O., Mollusks�Invaders and
Their Place in Urban Malacocenoses, in Fauna,
ekologiya i vnutrividovaya izmenchivost' nazemnykh
mollyuskov v urbanizirovannoi srede (Fauna, Ecology,
and Intraspecific Variability of Terrestrial Mollusks in
Urban Environment), Lvov: Gos. Prirodoved. Muzei,
2006, pp. 42–59.
Sverlova, N.V., Martinov, V.V., and Martinov, O.V.,
Study of Terrestrial Malacofauna (Gastropoda: Pulmo�
nata) of Southeastern Ukraine, Nauk. Zap. Derzhav.
Prirod. Muz., 2006, vol. 22, pp. 35–46.
Page 8
44
RUSSIAN JOURNAL OF BIOLOGICAL INVASIONS Vol. 1 No. 1 2010
SON
Therriault, T.W., Docker, M.F., Orlova, M.I., Heath,
D.D., and MacIsaac, H.J., Molecular Resolution of
the Family Dreissenidae (Mollusca: Bivalvia) with
Emphasis on Ponto�Caspian Species, Including First
Report of Mytilopsis leucophaeata in the Black Sea
Basin, Mol. Phylogenet. Evol., 2004, vol. 30, pp. 479–
489.
Zambriborshch, F.S., Modern Tendencies in Changes
of Ichthyocenoses in the Black Sea, Vopr. Ikhtiol., 1985,
no. 4, pp. 688–690.
Zhuravel', P.A., Bogolyubova, M.M., and Zagubi�
zhenko, N.I., On the Radiation of Mollusks in Water
Bodies of Ukraine and Crimea through Channels, Irri�
gation Systems, and Pipelines, in Mollyuski i ikh rol' v
ekosistemakh (Mollusks and Their Role in Ecosys�
tems), Leningrad: Nauka, 1968, pp. 29–30.
Zolotarev, V., The Black Sea Ecosystem Changes
Related to the Introduction of New Mollusk Species,
Mar. Ecol., 1996, vol. 17, pp. 227–236.
RUSSIAN JOURNAL OF BIOLOGICAL INVASIONS Vol. 1 No. 1 2010
SON
Therriault, T.W., Docker, M.F., Orlova, M.I., Heath,
D.D., and MacIsaac, H.J., Molecular Resolution of
the Family Dreissenidae (Mollusca: Bivalvia) with
Emphasis on Ponto�Caspian Species, Including First
Report of Mytilopsis leucophaeata in the Black Sea
Basin, Mol. Phylogenet. Evol., 2004, vol. 30, pp. 479–
489.
Zambriborshch, F.S., Modern Tendencies in Changes
of Ichthyocenoses in the Black Sea, Vopr. Ikhtiol., 1985,
no. 4, pp. 688–690.
Zhuravel', P.A., Bogolyubova, M.M., and Zagubi�
zhenko, N.I., On the Radiation of Mollusks in Water
Bodies of Ukraine and Crimea through Channels, Irri�
gation Systems, and Pipelines, in Mollyuski i ikh rol' v
ekosistemakh (Mollusks and Their Role in Ecosys�
tems), Leningrad: Nauka, 1968, pp. 29–30.
Zolotarev, V., The Black Sea Ecosystem Changes
Related to the Introduction of New Mollusk Species,
Mar. Ecol., 1996, vol. 17, pp. 227–236.
Loading ...
Resources
-
155.72 KB · Uploaded Jan 26, 2012 by Mikhail Son
