Native range of the zebra mussel and quagga mussel and new data on their invasions within the Ponto-Caspian Region

Aquatic Invasions (2007) Volume 2, Issue 3: 174-184
URL: http://www.aquaticinvasions.ru
© 2007 European Research Network on Aquatic Invasive Species

174
Research article
Native range of the zebra mussel and quagga mussel and new data on their
invasions within the Ponto-Caspian Region
Mikhail O. Son
Odessa Branch Institute of Biology of the Southern Seas, National Academy of Sciences of Ukraine,
Odessa, Ukraine
E-mail: michail.son@gmail.com

Received 19 July 2007; accepted in revised form 15 August 2007

Abstract
This is a special review on the native ranges of the zebra mussel (Dreissena polymorpha polymorpha)
and quagga mussel (Dreissena bugensis). Revision of museum collections showed that the Dniepr Delta
is a native locality of the quagga mussel, for example in the Lower Southern Bug and that Dreissena
presbensis occurs in the Aegean Sea Basin in Greece. The native range of the zebra mussel includes
Danube, Dniestr, Berezan, Southern Bug, Dniepr, Molochnaya, Don, Kuban, Kamchia, and Veleca river
basins; isolated and semi-isolated relic estuarian reservoirs along the Bulgarian, Romanian, Ukrainian,
and Russian Black and Azov seas coasts; and in zones of these seas influenced by freshwater. Invasive
dreissenids in Turkish rivers are local species and subspecies. More recent locations of the dreissenid
expansion within the Ponto-Caspian region (Lake Sasyk, Baraboj River, and Sukhoj Liman Basin for the
zebra mussel; Dniestr River Basin for the quagga mussel) are described. One empty valve of Dreissena
polymorpha andrusovi was found in the Taganrog Bay.

Key words: alien species, Dreissena bugensis, Dreissena polymorpha, Dreissena presbensis, molluscs,
native range, Ponto-Caspian relics

Introduction
Freshwater dreissenids have attracted attention
for a long time. Two species of freshwater
Dreissena: Dreissena polymorpha (Pallas, 1771)
and Dreissena bugensis (Andrusov, 1897) have
been indicated as invaders in different continents
(Kharchenko 1995, Bobat 2004, Orlova 2004,
Therriault et al. 2005, Gelembiuk et al. 2006,
May et al. 2006, Molloy et al. 2007). Interest in
both species is enormous: and the bibliography
on their ecology and impact according to
Schloesser et al. (1994) numbers thousands of
papers.
While the overwhelming majority of publi-
cations on Dreissena concerns invasive range,
information on distribution within their native
range is limited. Even in a special review on the
distribution of dreissenids in Ukraine (Kharchen-
ko 1995) there is almost no data present on their

M.O. Son, Nat ive range o f the zebra and quagga musse l s

175
native range. Knowledge about borders of
dreissenid native range and nearby expansion is
very important for both studies and prediction of
their widespread expansion.
For many years, all recent freshwater dreisse-
nids, except D. bugensis (quagga mussel), were
recorded as D. polymorpha polymorpha (zebra
mussel). According to this conception the native
range of D. polymorpha polymorpha includes the
Ponto-Caspian Basin and the ancient lakes of the
Balkan Penninsula and Asia Minor. A taxonomic
revision of Dreissena by Lvova and Staro-
bogatov (1982) showed that Dreissena from
Lake Ohrid (Republic of Macedonia) is a
separate species, Dreissena stankovici Lvova et
Starobogatov 1982. Later this species was
identified from nearby Lake Prespa (Gelembiuk
et al. 2006). Recent investigations showed what
D. stankovici is actually a synonym of Dreissena
presbensis (Kobelt, 1915), which occurs in
Balkan lakes as another Balkan endemic dreisse-
nid – Dreissena blanci (Westerlund, 1890)
(Albrecht et al. 2007). According this work
dreissenids from the Balkan lakes – Ohrid,
Prespa, Mikri Prespa, Skutari, Dojran, Trichonis
Vegoritis, Amvrakia, Pamvotis – are the above-
mentioned Balkan endemics.
Molecular revision of dreissenids shows that
those from some other regions form separate taxa
(Gelembiuk et al. 2006, May et al. 2006):
Dreissena polymorpha anatolica (Bourguignat,
1884) (Anatolian tectonic lakes), Dreissena
polymorpha gallandri (Locard, 1893) (lakes of
Marmara Sea Basin), and Dreissena caputlacus
(Schütt, 1993) (Golbasi tectonic basin).
These studies did not include some regions
with autochthonous populations of freshwater
dreissenids: rivers and relic lakes along the
Bulgarian Black Sea Coast (Hubenov 2005),
Lake Volvi and the Aliakmon River from the
environs of Thessaloniki in the Aegean Sea
Basin (Greece) (Zarfdjian 2000).
Our study on dreissenids from Lake Volvi
(collection of Zoological Institute of Russian
Academy of Sciences, St.-Petersburg; coll. A.
Yu. Karatayev, N 40°37´50" - 40°41´50", E
23°21´10" - 23°36´40"; 27 October 1995)
showed these to be D. presbensis (Figure 1).
According to labels, the late Starobogatov
intended further revisions in the identification of
these speci-mens.
Data on D. polymorpha from Bulgarian Black
Sea Coast was not used in taxonomic revisions,
and will be considered as D. polymorpha
polymorpha unless disproved. So, the native

Figure 1. Some important records of dreissenids in their
native range: the most eastern record of Dreissena
presbensis (Kobelt, 1915) (round); Dreissena polymorpha
polymorpha (Pallas, 1771) in a zone of the Black Sea
freshened by the Danube (ellipse); D. polymorpha
polymorpha in streams entering the Dniestr Liman
(rectangular); D. polymorpha polymorpha from a steep
small stream, the Berezan River (triangle); native locality of
Dreissena bugensis (Andrusov, 1897) in the Dniepr Delta
(square).
range of D. polymorpha polymorpha is limited
by the Northern Black Sea Region and the
Caspian Basin.
With rare exceptions, only dreissenids from
the Northern Black Sea Region are known as
invasive species. Expansion of other dreissenids
is localized in nature and mainly of concern to
Turkey. Local dreissenids invaded many reser-
voirs in the Turkish Euphrates River Basin,
Aegean, Mediterranean, and Black Sea coasts
(Bobat 2004), but data on the expansion of
separate species and subspecies are absent. As is
generally known, the dreissenids which invaded
the Seyhan Dam Reservoir were D. caputlacus
and D. polymorpha anatolica (Gelembiuk et al.
2006). According to Bobat (2004) at least some
cases of dreissenid expansion in the Turkish
Black Sea Basin (Lakes Acarlar in the Sakarya
Basin and dams on the Kizilirmak Basin)
concerned D. polymorpha gallandri (native to
the surroundings of Istanbul and Lake Uluabat in
the Marmara Sea Basin).
A dreissenid invasion of unknown origin (true
zebra mussel or autochthonous Balkan dreisse-
nids) was reported from Greek reservoirs
(especially Tavropos, Kremasta, and Kastraki),
where invaders were noted as D. polymorpha
(Economou et al. 1991, Petridis and Sims 1993,
Conides et al. 1995).
Among Caspian dreissenids, we found one
empty valve of Dreissena polymorpha andrusovi
(Brusina in Andrusov, 1897) in the western part
of Taganrog Bay (Figure 2). The subspecies is

M.O. Son , Nat ive range o f the zebra and quagga musse l s
176
wide-spread in the Northern Caspian Sea. This
shell might have been being dropped as empty
valve with building sand or by a ship.

Figure 2. Valve of Dreissena polymorpha andrusovi
(Brusina in Andrusov, 1897) from the Taganrog Bay, scale
= 1 cm (Photo: M. O. Son)
Material and Methods
This paper is based on material collected by the
author as well as on literature data. Original
material is deposited in the mollusk collection of
the Odessa Branch Institute of Biology of the
Southern Seas (OB IBSS). All collected material
was fixed in 70% ethanol. Danube, Dniepr,
Dniestr, Don, and Volga river basins, small
rivers and reservoirs of the Azov and Black sea
coasts were investigated annually from 1999 to
2007. Field research was undertaken according
to standard methods (Zhadin 1965).
Results and Discussion
Borders between native and invasive ranges of
the zebra mussel
The modern native range of D. polymorpha
polymorpha includes different habitats in the
Ponto-Caspian Region.
In the rivers of the Caspian Sea Basin (Volga
and Ural deltas) D. polymorpha polymorpha
occurs in biotopes of river channel and wetlands
(Pirogov 1974, Pirogov et al. 1994).
In the Azov-Black Sea Basin (where estuarian
biotopes are characterized by higher diversity,
than in the Caspian Sea Basin) the native range
of the zebra mussel has a complicated
disjunctive structure. This kind of native range
causes high geographical variability in D.
polymorpha polymorpha (Figure 3).
The zebra mussel was noted from Bulgarian
relic lakes: Varnensko, Beloslavsko, Mand-
rensko, Bourgasko, and also in the Rivers
Kamchia and Veleca (Hubenov 2005).

Figure 3. Conchological diversity of the zebra mussel from
different river basins, scale = 1 cm; A: Dniepr River; B:
Danubian lakes; C: Berezan River (Photo: M.O. Son)
In eastern Bulgarian lakes the zebra mussel
was situated within the Sinoie-Razim lagoonal
complex (Romania) connected with Danube
Delta.
In the Danube Basin it occupied riverine lentic
habitats and associated estuarine reservoirs –
Kagul, Kugurluj, Yalpug, Katlabug, and Kitaj
(Grossu and Paladian 1956, Milashevich 1908,
new data). Along the Danube channel it was
located as a native species up to the Middle
Danube (Mordukhay-Boltovskoy 1960). We
found it also in a zone of the Black Sea,
freshened by the Danube (shallows Kurilskije).
The zebra mussel occurred in branches of the
Dniestr Delta and in the main channel of the
Lower and Middle Dniestr, and in estuarine
reservoirs – Dniestr Liman and Kuchurgan
Liman. In the Middle Dniestr we observed the
largest specimens of the zebra mussel (length of
the shell = 49 mm) found in all native habitats
(Figure 4).

Figure 4. Giant zebra mussel from the Middle Dniestr, scale
= 1 cm (Photo: M. O. Son)
Along the coasts of the Dniestr Liman,
D. polymorpha polymorpha was found in streams
(the first records of Ponto-Caspian bivalves in
this type of habitat). It now seems to have
disappeared from the floodplain lakes of the
Dniestr, where it was recorded in last century
(Grinbart 1967). This is probably connected with
disimproved oxygen levels in these reservoirs. In
the Dniestr Liman the zebra mussel was recorded

M.O. Son, Nat ive range o f the zebra and quagga musse l s

177
in the northern (freshwater) and central
(brackishwater) parts and in springs in its basin.
In the Tiligul Liman, where D. polymorpha
polymorpha was identified in the early twentieth
century (Grinbart 1967) it disappeared after a
rise in salinity when the liman was connected by
channel with the Black Sea.
In the Berezan Liman, the zebra mussel occurs
in brackish water in central and northern parts of
the liman (Grinbart 1955). We found it also in
the Lower Berezan River.
In the Southern Bug River and Dniepr River
the zebra mussel was located in the lower and
middle stretches including a zone upstream of
rapids (up to Vinnitsa City and Kiev according-
ly) (Polishchuk 1978, Krasheninnikov 1929).
This species also occurred in their large tributa-
ries: Ingul River, Ingulets River, and Saxagan
River (tributary of the Ingulets River) (Polish-
chuk 1978). All these streams were situated on
locations of former large brackishwater resevoirs
or coastal bays that are established on paleonto-
logical data (Polishchuk 1978). The zebra mussel
is a dominant species in estuaries of these rivers,
which form an estuarine complex – the Dniepr-
Bug Liman.
On the Northern Azov Sea Coast it was noted
in the Molochnaya River (Lubyanov 1954) and
in the mouth of the Kalmius River (Mordukhay-
Boltovskoy 1960). At present, this mollusc is
absent in the last river; this absence is probably
connected with the construction of a giant
industrial complex in this place.
The zebra mussel was widespread in the
estuarine system of the Don River, including the
brackishwater Taganrog Bay of the Azov Sea,
Mius Liman, Yejsk Liman, and the Don Delta;
along the Don River its native range spread
upstream to Voronezh City (Mordukhay-
Boltovskoy 1960). It also was recorded from the
Manych River (tributary of the Don River) and
in large reservoirs of the Don and the Manych
(Ust-Manych, Proletarskoye reservoir, and
Tsimlyansk reservoirs) (Zhulidov et al. 2004).
In the nineteenth century the zebra mussel was
recorded in the Severskij Donets River (tributary
of the Don River) (Krynicki 1837). Later this
species was not recorded in this river until
construction of the Dniepr-Donbass Channel.
Probably it was occasionally acclimatization.
In the Eastern Don River, the zebra mussel
lives in the Kuban River from its mouth to
Krasnodar City (Rosen 1911) and in the estua-
rine reservoirs: Ahtanizovskij Liman (Puzanov
1929) and Dolgij Liman (Povazhnyj and Semin
2005).
These basins in the Black Sea Region are the
most eastern native locations of dreissenids’
population. According to Mordukhay-Boltovskoy
(1960) dreissenids have not been found in the
relic lakes of the Caucasian Region (Abrau and
Paleostomi) where other Ponto-Caspian species
are known. Now the zebra mussel is a major
species in Lake Abrau (M. V. Nabozhenko,
personal communication).
In the time of acclimatization of aquatic
invertebrates into Georgian reservoirs (Hram-
skoje and Tkibulskoje) the zebra mussel was
introduced there (Sergeyeva 1968) but has not
since been recorded.
During the twentieth century, range expansion
for the zebra mussel was described in a number
of channels and reservoirs, associated with the
Lower Dniepr Basin (Kahovka Meliorative Sys-
tem, Ingulets Channel, North Crimean Channel,
etc). This spread took place only within artificial
systems and has not impacted on natural
ecosystems. Penetration of this species to new
Ponto-Caspian Rivers began at the end of the
twentieth century (Figure 5).

Figure 5. Directions of nearby expansion of the zebra
mussel within North Black Sea Region: 1) Lake Sasyk; 2)
Baraboj River; 3) Sukhoy Liman Basin; 4) Ingulets Channel;
5) Kahovka and North Crimean Channel
Expansion in the Danube-Dniestr Meliorative
system
In the 1960s the creation of a common
meliorative system was planned between the
Danube and Dniepr. The building of the Danube-
Dniestr Meliorative system was planned as the
first stage of its creation. The marine gulf –
Sasyk Lake (in this project – a reservoir in the
Danube-Dniestr Meliorative system) was desali-
nated by separating it from the Black Sea and by
building the Danube-Sasyk Channel, which

M.O. Son , Nat ive range o f the zebra and quagga musse l s
178
connected Lake Sasyk and the Danube River.
This remedial plan included a large lagoonal
complex Tuzly Limans between the Danube and
Dniestr basins.
In 1981 the project began, but irrigation by
mineralized water made a catastrophic impact on
local soils and the project was disrupted.
As result of the desalination of Lake Sasyk
and its connection with the Danube many
freshwater species invaded this reservoir.
D. polymorpha polymorpha is widespread in this
reservoir and found along all its length (more
than 30 km).
Both the zebra mussel and other Ponto-
Caspian bivalves have a very specific impact on
the ecosystem of Lake Sasyk: their empty shells
make enormous deposites which have caused
shallowing and transformation of a southwestern
part of this reservoir into a swamp.
The next penetration of Danubian species was
through Lake Sasyk into the higher part of the
Danube-Dniestr Meliorative system (channels
and transformed Rivers Kogilnik and Sarata) is
this is complex, because introduced species in
Lake Sasyk include mainly species with high
sensivity to hypoxia. This interferes with their
penetration to the above-mentioned streams
which have a variation in oxygen concentration.
Expansion in the Lower Dniestr Meliorative
System
The Baraboj River was connected by channels
with the Dniestr River to the Lower Dniestr
Meliorative System in the second half of the
twentieth century. During this time the upper
part of the Baraboj River was turned into the
Baraboj Reservoir. This reservoir has two
applications: as part of a meliorative system and
as a cooler of a planned power plant. Before this
building, the Baraboj River was temporarily dry
and many species, occurring in large rivers,
would not live in this river.
The population of zebra mussels there was
formed as a result of water transferring from the
Odessa Channel (water-mine with pumping
station on the Dniestr River near Belyajevka
City) to the Baraboj Reservoir and Baraboj
River. Now this species is found in the Baraboj
Reservoir and in a little pond in the river mouth.
Invasion to the Sukhoy Liman Basin
A population of D. polymorpha polymorpha was
recorded in the Sukhoy Liman Basin in July
2006 in a small rapid stream. This stream is the
remains of the Gross-Libental River (the middle
section) which flowed into the Sukhoy Liman.
The connection between the Gross-Libental
River and Sukhoy Liman was disrupted by the
building of a road. Now the lower part of the
river is a small floodplain. The total sampled
population consisted of juvenile specimens only
(some at pediveliger stage). Adult shells were
not found. Molluscs occurred on the stones and
on other hard substrates.
In the Black Sea Region, this record is a
unique case of dreissenid invasion to a small
stream. Upstream of the sampling site, a limnic
reservoir feeds water to the stream via pipework
and hence the stream flow is artificially sped up.
Range expansion in the human-made rheophilic
biotopes of small stream was also noted for other
species.
In this biotope we also found the exotic
mudsnail Potamopyrgus antipodarum (Gray,
1843) and an aquarium snail Physella
heterostropha (Say, 1817). Both these species
were also found in all small rivers of the Sukhoy
Liman Basin (Son, 2007a). In similar biotopes in
the Baraboj River we found the rheophilic
Viviparus vivparus (Linnaeus, 1758) and the
Ponto-Caspian species Euxinipirgula lincta
(Milashevitch, 1908), both of which are non-
indigenous for this river (Son, 2007a).
To summarise these facts we conclude that
this type of biotope is characterized by high
invasibility. Similar biotopes probably play an
important role in the expansion of rheophilic
species into steep droughted rivers.
The native range of the quagga mussel
The question on chronology of the quagga
mussel invasion history is extremely confusing.
The data published till now on its distribution
testifies, that its native range is in a lower stretch
of the Southern Bug, whence it has been
described, and its expansion in the Dniepr and
moving on the cascade of reservoirs date from
the 1930-1940s (Tseeb et al. 1966, Zhuravel
1967).
The designation of its native range as “a lower
stretches of the Southern Bug and Dniepr rivers”
in some English-written literature was based on a
vagueness of the term “Dniepr-Bug Liman”. This
name is modular for estuaries of the Dniepr and
Southern Bug rivers. D. bugensis has been
described from the Lower Southern Bug, and the
introduction of the concept “Dnepr-Bug Liman”

M.O. Son, Nat ive range o f the zebra and quagga musse l s

179
in the twentieth century has resulted in the
beginning of a designation of this species locus
typicus as Dniepr-Bug Liman and the subsequent
transferral to the English-written literature as
“lower stretches of the Dniepr River”. This is
absolutely inadmissible, because the locus
typicus of D. bugensis is the estuarine Southern
Bug – Bug Liman (expanded river mouth),
whereas the estuarine system of the Dniepr River
includes two parts – Dniepr Delta – the expanded
river mouth and Dniepr Liman (part of Dniepr-
Bug Liman) – the freshened sea gulf between the
Bug Liman and Dniepr Delta.
For this reason, the occurrence of D. bugensis
in the Dniepr Delta may be considered as the
beginning of the quagga mussel’s expansion into
the Dniepr Basin.
However malacological investigations of the
Dniepr and Southern Bug basins at the time of
the quagga mussel’s description were fragmen-
tary and the two similar species concerned could
not simply be differentiated.
In Lindgolm's article (1908), the first complex
malacological investigation in the given region,
D. bugensis was not sampled at all in the south
of the Russian Empire.
While this author was processing collection of
Zoological Institute of Russian Academy of
Sciences, D. bugensis was found among them;
this was collected by Brauner in the Dniepr Delta
(Lake Beloje, surrounding Kherson City). The
sample year was absent from the label but
Lindgolm (1908) indicated it as 1899 in a publi-
cation. Thus, D. bugensis was already present in
the Lower Dniepr practically at the same time,
when it had been described from the Southern
Bug. This raises doubts as to whether this
species is alien to the Dniepr Basin.
References on recent native localities of the
quagga mussel in the Ingulets River are
consistent with ambiguity in reporting by
Polishchuk for the Soviet malacological meeting.
This was reprinted by Malacological Review
(Polishchuk 1978). In the original text it states
clearly that D. bugensis was recorded in
geological sediments of the Ingulets River.
The quagga mussel invasion within the Ponto-
Caspian region
The quagga mussel has grasped, within Ponto-
Caspian region, some large river basins and has
expanded its distribution to the Dnepr Basin.
In the Dniepr Basin, the quagga mussel is
widespread in the Dniepr Cascade of Reservoirs
which occupy almost all the Ukrainian part of
the Dniepr River and the associated artificial
systems of hydrotechnical buildings (Kahovka
Meliorative System, Ingulets Channel, North
Crymean Channel, etc).
In the Don River Basin (first record in the
1980) it occurs in the Lower and Middle Don
River, Manych River and large reservoirs of the
Don and Manych (Ust-Manych, Proletarskoye
reservoir, and Tsimlyansk reservoirs) (Zhulidov
et al. 2004, 2005).
In the Volga River Basin (first record – 1981)
it occurs in the Volga River (including Volga
Delta) and Volga Cascade of Reservoirs (Orlova
et al. 2004, Zhulidov et al. 2005).
The quagga mussel also penetrates to
freshened zones of the Azov and Caspian seas. In
the Northern Caspian Sea it was recorded as
recently as 1994-1997 (Orlova et al. 2004). In
the Azov Sea it lives in the Taganrog Bay – a
brackishwater part of sea which is an estuary of
the Don River (see Annex).
More recently, the quagga mussel invasion has
expanded in the western Ponto-Caspian rivers:
Dniestr and Danube.
Invasion in the Danube River was recorded
during recent years and is known from some
local findings in the Lower (except delta) and
Middle Danube (Micu and Telembici 2004, Popa
and Popa 2006).
A very intensive expansion of the quagga
mussel has taken place in the Dniestr Basin. The
first record of D. bugensis in the Dniestr Basin
was the invasion of the Dniestr Reservoir
(Middle Dniestr) in 1992-1993 (Kharchenko,
1995). Now it occupies a number of habitats in
the Dniestr River and Dniestr estuarine system.
The quagga mussel has been recorded in the
main channel of the Middle and Lower Dniestr
River, branches of the Dniestr Delta, the
northern part of the Dniestr Liman and streams
entering to the Dniestr Liman (see Annex).
Unique specimens of quagga mussel were
recorded in the estuarine reservoir – Kuchurgan
Liman (Filipenko and Lejderman 2006).
Biotopical distribution of dreissenids in the
Azov-Black Sea Region, ecological barriers and
vectors of invasions
Both species of dreissenids occur in the Black
Sea region in various habitats, where hypoxia is
absent (see Table 1). We also noted an interes-
ting way of supporting dreissenids’ populations
in the shallows along coasts of the Dniestr

M.O. Son , Nat ive range o f the zebra and quagga musse l s
180
Liman, where hypoxia, periodically takes place.
Number of dreissenids accumulate in the
streams, which enter the estuary, and in the
shallows near the streams’ mouths (Figure 6).
Until 2007 we found only zebra mussel in these
habitats, but in 2007 massive populations of the
quagga mussel were located in the four streams
along the southern coast of the Dniestr Liman.
Table 1 Biotopical distribution of the zebra mussel and
quagga mussel in their native and invasive range within
Ponto-Caspian Region
D. polymorpha
polymorpha D. bugensis
Habitats
Native
range
Range
of
invasion
Native
range
Range
of
invasion
River channel
and branches of
deltas
+ + + +
Rapids + - - -
Tributaries + - - -
Streams + + - +
Freshened
zones of seas + - - +
Freshwater
parts of
estuaries
+ + - +
Brackishwater
parts of
estuaries
+ + + +
Small rivers + + - -
Artificial
reservoirs + + - +
Lakes + + + +
Table 1 shows the significant extension of the
quagga mussel’s biotopical distribution in the
range of invasion. Characteristically, the
biotopical distribution of both dreissenids in the
range of invasion is very similar, whereas in the
native range D. polymorpha polymorpha
occurred in more kinds of habitats than
D. bugensis.
The widespread invasion of the dreissenids is
caused by a diversity of successful vectors.
Dreissenids can use different modes of spread
in different stages of their life cycle as they have
a planktonic larval stage (trochophora, veliger),
settling benthic stages (pediveger, juvenile
mollusc), and stationary attached stage (adult
mollusc attached by byssus).


Figure 6. Dreissenid, field in the stream mouth
(Photo: M. O. Son)
During these different stages dreissenids use
different modes of invasion. Downstream drift is
possible for planktonic larvae by current and for
other life stages on aquatic plants drifting
downstream (Sullivan et al. 2003, Son 2007a).
Drift is a very rapid way of spread and it helps
dreissenids penetrate through stretches which
have unfavorable conditions. This vector of
spread is not available to another highly invasive
exotic bivalve genus, Corbicula. Corbicula
fluminea (Müller, 1774) and Corbicula
fluminalis (Müller, 1774), which at the end of
the twentieth century had invaded the Lower
Danube and were widespread through most of
the Danube Delta branches, but similar to
dreissenids (both genera have marine origin) had
avoided colonizing stretches with variations in
oxygen regime. Both species of Corbicula
recorded in Europe lack a free–living larval stage
and have not penetrated into Lake Sasyk, which
is connected with various branches of the
Danube Delta (Voloshkevitch and Son 2002, Son
2007b). Contrary to this, it appears that the zebra
mussel has successfully overcome this barrier by
larval drift.
Spread by shipping is typical for planktonic
larvae via ballast water and also by biofouling by
the adult stage on the hull. Water movement
(flow and wave action) and animal vectors are
also important for spread of pediveger and
juvenile molluscs. Use of animal vectors is also
notable in adult dreissenids; including birds,
aquatic insects, amphipods, crayfish, and,
probably some other animals (Mordukhay-
Boltovskoy 1960, Dedyu 1963, McCauley and
Wehrly 2007).

M.O. Son, Nat ive range o f the zebra and quagga musse l s

181
What kinds of ecological barriers have limited
dreissenids’ spread within native range?
It is obvious, that the native range of the zebra
mussel and quagga is less than the basic
ecological limiting factors would allow.
The most important factors for dreissenids are
salinity, mineralization, and speed of current
isolation between river basins, oxygen concen-
tration, and drying (Mordukhay-Boltovskoy
1960, Spidle et al. 1995, Orlova et al. 2005).
The last two factors prevent occurrence in
some habitats, but do not limit distribution of
dreissenids along river networks of the Ponto-
Caspian Region (excluding drying rivers and salt
lakes). However in the Ponto-Caspian Region
salinity and high velocity in streams do not
usually limit dreissenids’ spread in situ.
The distribution of dreissenids within their
native range is limited by the presence of all
habitat requirements, necessary to form stable
populations.
Such as many marine species with planktonic
larvae, dreissenids can form a compound
structure of metapopulations. The concept of
metapopulation (where integral genetic system
without free interbreeding but is not mixed on
isolated populations) are discussed in detail in
the issues of the “Helgoland marine research”
(volume 56, number 4), which defined the
metapopulation concept. In the Ponto-Caspian
Region we observe some variants of spatial
structure of the dreissenids’ occurrence.
Three main genetic variants are showed in
Figure 7. First variant (A) is known in the Ponto-
Caspian Region from the Caspian Sea, Taganrog
Bay, and from a lot of estuaries in the native and
invasive ranges of the zebra mussel and quagga
mussel. In these cases we observe the “classic
population”.
Even in the desalinated parts of the sea (such
as the shallows of Kurilskije in the Black Sea)
dreissenids don’t form a metapopulation,
because transfer by sea currents along coast, not
only brings larvae to other freshened localites in
the sea, but also passes through marine habitats
which are characterized by high salinity.
A good example of the variant B occurs in the
Volga and Dniepr cascades of reservoirs by both
dreissenids (invasive ranges) or in the Lower
Danube by D. polymorpha polymorpha (native






Figure 7. Different variants of spatial structure of the
dreissenids’ occurring: A – isolated lentic reservoirs,
estuaries, freshened parts of seas; B – reservoirs and
channel downstream; C – reservoirs and small streams
upstream
A good example of the variant B occurs in the
Volga and Dniepr cascades of reservoirs by both
dreissenids (invasive ranges) or in the Lower
Danube by D. polymorpha polymorpha (native
range). In the cascades of reservoirs the
dreissenids form an integral genetic system
(metapopulation) with asymmetric genetic drift
caused by drift of larval drift downstream. A
similar situation was observed in the Ukrainian
stretch of the Danube, where the river is
connected by small channels with a series of
separate estuarine lakes, which are a “donor
fund” of the dreissenids for the main river
channel.
Variant C was found in the first investigation
of the Dniestr Liman. In some streams entering
into this estuary the zebra mussel was record
above sea level. In this case we suggest transfer
by amphipods. A similar stream habitat for the
zebra mussel was observed on the Uglich
Reservoir (Volga River). Meanwhile we have not
recorded this type of habitat for the quagga
mussel.
Acknowledgements
We are grateful to Alexey V. Korniushin for his
help in questions of dreissenid taxonomy, and to
Frances Lucy (Institute of Technology Sligo,
Ireland) for valuable comments and English
editing. This work was partly supported by the
European Commission 6th Framework Program
Integrated Project ALARM (contract GOCE-CT-
2003-506675).

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182
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184

Annex
Records of dreissenids in their range of invasion

Record coordinates
Species Location
Latitude,°N Longitude,°E
Date of record Collector
A mouth of the anonymous river
entering to the Dniestr Liman 46°20'02" 30°06'06" 03.05.2007 M. O. Son
Alexandrovskij Erik 46°40'50" 30°25'18" 04.05.2006 M. O. Son
Dniestr River, Republica
Moldova, Site 1 47°05'43" 29°04'40" 13.10.2006 M. O. Son
Dniestr River, Republica
Moldova, Site 2 47°41'43" 28°58'35" 02.08.2005 M. O. Son
Dniestr River, Ukraine, Site 1 46°27'48" 30°15'39" July 2006 M. O. Son
Dniestr River, Ukraine, Site 1 46º27'40" 30º14'58" May 2007 M. O. Son
Dniestr River, Ukraine, Site 2 46°44'34" 30°19'17" 22.08.2006 M. O. Son
Dniestr River, Ukraine, Site 3 46°25'48" 30°10'16" 08.07.2001 M. O. Son
Dniestr River, Ukraine, Site 4 46°22'28" 30°16'19" 28.04.2007 M. O. Son
Dniestr River, Ukraine, Site 5 46°42'16" 30°17'49" 05.05.2007 M. O. Son
Dniestr River, Ukraine, Site 6 46°41'15" 30°26'21" 22.08.2006 M. O. Son
Lake Mertvyj Turunchuk 46°25'00" 30°14'20" 04.05.2007 M. O. Son
Spring entering to the Dniestr
Liman 46°17'49" 30°07'58" 02.05.2007 M. O. Son
Taganrog Bay of the Azov Sea,
Taganrog City, Russia 47°12'49" 38°56'31" 08.06.2007 M. O. Son
Dreissena
bugensis
Turunchuk River (branch of
Dniestr Delta), Site 1 46°27'21" 30°11'36" 29.05.2002 M. O. Son
Dreissena
polymorpha
andrusovi
Taganrog Bay of the Azov Sea,
Ukraine 47°06'40" 38°03'52" 09.08.2004 M. O. Son
Pond in the mouth of the
Baraboj River 46°12'24" 30°33'42" 05.09.2003 M. O. Son
Baraboj Reservoir, Site 1 46°28'51" 30°20'29" 02.07.2006 M. O. Son
Baraboj Reservoir, Site 2 46°28'54" 30°20'53" 02.07.2006 M. O. Son
Small river entering to the
Sukhoj Liman 46°20'52" 30°35'43" 18.07.2006 M. O. Son
Lake Sasyk, Site 1 45°32'23" 29°38'51" 19.07.2002 M. O. Son
Lake Sasyk, Site 2 45°34'28" 29°41'44" 05.07.2002 M. O. Son
Dreissena
polymorpha
polymorpha
Danube-Sasyk Channel 45°33'13" 29°36'08" 29.04.2003 M. O. Son