Trends of aquatic alien species invasions in Ukraine

Aquatic Invasions (2007) Volume 2, Issue 3: 215-242
URL: http://www.aquaticinvasions.ru
© 2007 European Research Network on Aquatic Invasive Species

215
Research article
Trends of aquatic alien species invasions in Ukraine
Boris Alexandrov1*, Alexandr Boltachev2, Taras Kharchenko3, Artiom Lyashenko3,
Mikhail Son1, Piotr Tsarenko4 and Valeriy Zhukinsky3
1Odessa Branch, Institute of Biology of the Southern Seas, National Academy of Sciences of Ukraine
(NASU); 37, Pushkinska St, 65125 Odessa, Ukraine, e-mail: alexandrov@paco.net
2Institute of Biology of the Southern Seas NASU; 2, Nakhimova avenue, 99011 Sevastopol, Ukraine,
e-mail: boltach@ibss.iuf.net
3Institute of Hydrobiology NASU; 12, Geroyiv Stalingrada avenue, 04210 Kiyv, Ukraine, e-mail:
hydrobiol@igbibc.com.ua
4Institute of Botany NASU; 2, Tereschenkivska St, 01601 Kiyv, Ukraine, e-mail: algae@botany.kiev.ua
*Corresponding author
Received 13 November 2006; accepted in revised form 2 August 2007




Abstract
This review is a first attempt to summarize data on the records and distribution of 240 alien species in
fresh water, brackish water and marine water areas of Ukraine, from unicellular algae up to fish. A
checklist of alien species with their taxonomy, synonymy and with a complete bibliography of their first
records is presented. Analysis of the main trends of alien species introduction, present ecological status,
origin and pathways is considered.
Key words: alien species, ballast water, Black Sea, distribution, invasion, Sea of Azov

Introduction
The range of organisms of different taxonomic
groups varies with time, which can be attributed
to general processes of phylogenesis, to changes
in the contours of land and sea, forest and
deserts, to elevation and isolation of mountains,
to changes in water bodies and water flows.
These historical activities may be enhanced or
slackened under global climate changes.
Continental drift, the elevation and lowering of
large areas of the Earth, the origin and trans-
formation of oceans, inner lake-seas have led to
the formation on the Earth, in the Paleocene and
Neocene periods of isolated water bodies with
distinct flora and fauna. As a result of active
geological processes and global climate changes
(especially in the glacial age) a natural shift and
transformation of fauna occurred. However, with
the origin of man and with the progress of
civilization including extensive travel, the
introduction of plants and animals to new areas
increased over the ages.
From the beginning of the 19th century, due to
rising technical progress, the influence of man
on nature has increased in geometrical
progression, gradually becoming comparable in
dimensions to climate impact.
In the past, even when aquatic ‘stepping
stones’ existed across continents, very often
negative environmental factors created natural

Alexandrov et al., Trends of aquatic alien species invasions in Ukraine


216
obstacles hindering the spread of different
species of aquatic bacteria, fungi, plants and
animals. For example water salinity serves as a
barrier impeding the entry of hydrobionts from
the Mediterranean to the Black Sea and Sea of
Azov. However, aquatic habitats with salinities
of 5-26‰ similar to the Black Sea and therefore
suitable for brackish organisms are widely
distributed on the Earth (Vinogradov 1986).
Thus as these areas are geographically isolated,
spread by man can lead to successful
colonization of new comers. So the Black Sea
has high risk in naturalization of exotic species
in comparison with other areas of world ocean.
Once the natural obstacle can be overcome,
species can spread over a new territory and
expand its range. Often this occurs with the help
of intentional or accidental human introductions.
As an example of accidental introduction
(simultaneously with intentional introduction of
valuable species) data can be obtained that of the
36 species brought to the Black Sea from the
USA, only 4-6 have been applied in fisheries and
in angling and 10 for aquarium fishes. The rest,
which penetrated into natural water bodies, are
to some degree dangerous for aboriginal fauna
(Zaitsev and Ozturk, 2001). The most common
way of accidental introduction of new species is
via ocean-going ships that transfer organisms
through natural barriers. According to the
International Marine Organisation (IMO) 80% of
world cargoes are transported by shipping.
Annually, about 85 000 ships carry 3-10 billion
tons of ballast water in which more than 3,000
species of algae, invertebrates and fish have been
recorded (IMO Bulletin 1998).
The dispersal of species is accompanied by a
simultaneous decrease in native species diversity
in the entire biosphere and in separate locations
unifying the genetic fund of the planet. The
successful invasive species inevitably leads to
heavy competition squeezing out weaker often
native species, resulting in a decline of
biological diversity (Convention on Biological
Diversity: June 5, 1992, Rio de Janeiro, Brasil).
The Black Sea – as a brackish water basin,
semi-isolated from the Mediterranean Sea has
been closely studied for biological invasions.
Not only has a high biodiversity of alien species
been established – more than 140 – but also their
substantial impact on the marine ecosystem
(Gomoiu et al. 2002; Alexandrov 2004) is
evident. The invasion of the comb jelly
Mnemiopsis leidyi Agassiz, 1865 was characte-
rized as an ecological disaster (GESAMP 1997).
The risk of entry of alien organisms into the
Black Sea has been promoted by the dredging of
shipping channels. These include the Danube-
Main-Rhine Canal, connecting the Black and
North Seas in 1836, and the Volga-Don Canal
connecting it with the Caspian Sea in 1952.
An important factor simplifying the natura-
lization of alien species in the Black Sea is the
instability of its ecosystem. Wide scale
eutrophication, observed first in the northwestern
sea in the early 1970s, spread almost all over the
coastal shelf especially close to river estuaries
and large cities. These changes in the trophic
status of the sea led to a decline in biological
diversity (losses of immunity due to a reduction
in the number of predators) the freeing of
ecological niches and their occupation by highly
productive alien species (Alexandrov and Zaitsev
1998).
Of all the lacustrine states in the Black Sea
basin, Ukraine, due to its physical-geographic
position and economic development, has all the
conditions necessary for becoming the main
testing area for control the introduction of
aquatic alien species.
The aim of this paper is to illustrate, with the
help of data from the literature, the chronology
of biological invasions of freshwater, brackish
and marine species in Ukraine ranging from
unicellular algae to fish, with synonyms
included.
Methodology
All species of aliens can be divided into two
groups:
1) Distant aliens – species which never
previously inhabited the Black Sea–Azov
basin, In other words these have a different
origin, and were introduced into the water
bodies of Ukraine unintentionally or
intentionally with human help.
2) Neighbouring aliens – inhabitants of the
Black Sea – Azov basin, or adjacent areas
expanding by chance within Ukranian waters
or with spread accelerated by human
activities (this is mostly freshwater species).
Alien species have subsequently been grouped
into four categories namely casual, invasive,
established and cryptogenic (questionable).
• Casual: In this paper alien is used in the
sense of CIESM: Alien species are identified

Alexandrov et al., Trends of aquatic alien species invasions in Ukraine


217
as having been recorded only once (no more
than twice for fishes) in the scientific
literature: they are presumed to be not
established in a river basin.
http://www.ciesm.org/online/atlas/index.htm
• Invasive: These are the introduced species
that, having overcome biotic and abiotic
barriers, can propagate away from their area
of initial introduction through the production
of fertile offspring without any reference to
impact (Richardson et al. 2000). In many
definitions the term invasive is also
considered as established species which are
agents of change and threaten native
biological diversity (IUCN 2002) or as
species that threaten the diversity or
abundance of native species, the ecological
stability of infested ecosystems, economic
activities dependent on these ecosystems
and/or human health (EPA 2001). It is in this
last meaning that the term IAS (Invasive
Alien Species is used here).
• Established (synonym: naturalized): These
include an introduced or feral population of
a species. Species established in the wild
with free-living, self-maintaining and self-
perpetuating populations unsupported by and
independent of humans (European
Commission 2004).
• Cryptogenic species are species with no
definite evidence of their native or
introduced status according to Carlton
(1996). Species whose probable introduction
has occurred prior to 1800, i.e. has not been
witnessed, have also been included in our
compiled list. Often these species are
excluded from lists of aliens or included
among the established ones. In this review
we considered it best to separate them.
Physical and Geographical Features
Among the Black Sea countries Ukraine has the
longest shoreline – 4,431 km making up 36.7%
of the total (Shujskiy 1989).
Ukraine has the greatest shelf area as opposed
to its territorial waters (to 200m isobaths), equal
to 55,750 km2 or 57% of the total Black Sea
shelf (Zaitsev 1992).
The dense indentations of the shore line and
the high number of water body types (lagoons,
bays, embayments, limans, lakes) provide
various habitats for aquatic organisms. These
aquatic habitats are of varying salinity (0.2-300
‰), depth (0.8-36.0 m), substrate and vegetation
(Zaitsev and Alexandrov 1998). Ukraine has 17
large limans with a total area of 1952 km2, 8
bays –1770 km2, 19 coastal wetlands – 6350 km2
(Conservation… 2003). A liman is the former
mouth of a river now covered with sea water.
This definition is very close to a estuary, but as a
tide-less sea, the Black Sea does not have
estuaries.
The largest rivers of the Black Sea (Danube,
Dnieper, Dniester) make up more than 50% of
the total river runoff into the Black Sea
(Nikolaenko and Reshetnikov 1991), and their
catchments create a zone of maximum biological
productivity in most of the shelf of Ukraine. In
total, Ukraine has about 73,000 rivers. The
highest number are present in the western
mountainous area including the Dniester and
Danube basins. The density of the river network
there is 1–1.5 km/km2. In the Crimea, rivers are
relatively rare, with many areas lacking natural
water flow. The density of the Crimean river
networks is 0.22 km/km2.
The continental waters of the Ukraine pertain
to two of the largest zoogeographical areas, i.e.
Palearctic and Ponto-Caspian brackish areas
(Starobogatov 1970). The Palearctic includes
North Eurasia and Northwest Africa. Within the
boundaries of Ukraine, there are two zoogeo-
graphic provinces: the Baltic and Danube-Don.
The former includes the rivers of the Visla basin
and Shatskie lakes. The Ponto-Caspian area
includes the Caspian Sea, limans and the lower
reaches of the rivers of the Black and Azov Seas.
According to some data a number of relict lakes
in Macedonia, Greece, Italy, Iran, Iraq, Central
and Western Turkey, i.e. Eastern Mediterranean
coast and Asia Minor (Wilke et al. 2007) should
be included.
The Crimean peninsula could be included in
the Ponto-Caspian area. Up until the 20th
century it lacked the relict Ponto-Caspian fauna
as zoogeographically it is not a peninsula, but an
island and its isolation from the lower basins of
the Dnieper and Don occurred before the
introduction of Ponto-Caspian fauna. The Crimea
can actually be considered to be a faunal isolate
of the European-Siberian subarea of Palearctic.
However, the Crimean hydrofauna contains a
number of Asian elements. The Crimea itself was
never considered in large zoogeographical
classifications (Figure 1).

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218
Results and Discussion
On the base of collected information 169 species
of distant aliens and 71 neighbouring aliens have
been registered in aquatic areas of the Ukraine
(Annex). Check-list of aquatic alien species of
Ukraine have been compiled according to
available data published before May 2007. For
this reason some recent data are not included in
the results of our investigations. For example,
this includes information about distant (mystery
blenny Parablennius incognitus Bath, 1968,
golden goby Gobius xanthocephalus Heymer and
Zander, 1992; Syngnathus acus Linnaeus, 1758;
chameleon goby Tridentiger trigonocephalus
Gill, 1859) and neighbouring (bleak Alburnus
alburnus Linnaeus, 1758; carp C. carpio; pike
Esox lucius Linnaeus, 1758; goby Gobius
ophiocephalus var. Ninni 1938; black goby G.
niger jozo Linnaeus, 1758; ruffe Gymnocephalus
cernuus Linnaeus, 1758; chub Leuciscus
cephalus Smitt, 1895; sand goby Neogobius
fluviatilis fluviatilis Pallas, 1814; racer goby N.
gymnotrachelus Kessler, 1857; round goby N.
melanostomus Pallas 1811; European perch
Perca fluviatilis Linnaeus, 1758; tubenose goby
Proterorhinus marmoratus Pallas, 1814; rodeus
Rhodeus sericeus amarus Berg, 1949; roach
Rutilus rutilus Linnaeus, 1758) alien species in
coastal and inland water of Crimea that have
been firstly registered during 2002-2006
(Boltachev еt al. 2007).
Analysis of distant species registration
In relation to salinity many of the encountered
species are brackish organisms encountered both
in the sea and in freshwater bodies. However,
59% of the registered species were attributed to
the coastal ecosystem of the Black and Azov
seas, 29% to brackish and 16% to freshwater
areas. Most of the distant alien species (23
species of fish) were registered in the period
from the beginning of 1950 to 1975 due to wide
scale introduction of valuable varieties of fish in
the water bodies of Ukraine and neighboring
countries (Russian, Moldova, Belarus). Of the
number of aliens observed in that period, 44%
relate to introductions. However, of the above
mentioned 23 fish species brought to Ukraine,
only 4 species became naturalized. These were
able to reproduce in new conditions without the
help of man. These species include spotted silver
carp A. nobilis, silver carp H. molitrix and grass
carp C. idella among the freshwater hydrobionts


Figure 1. Ukrainian part of the Black Sea
and haarder L. haematocheila in the marine
species (see Annex).
The process of successful introduction is
linked with a number of biological, physical and
chemical factors which at present cannot be
accurately forecasted. Of the total number of
registered species of distant aliens, only 30% (or
50 species) were naturalized, and today are part
of the food web of Ukranian aquatic ecosystems
able to support their populations. Almost half of
the species (41% or 69 species) were discovered
no more than twice and in small amounts, and
that is why they are placed in the casual category
(Figure 2).

Figure 2. Ratio between main categories of distant alien
species: 1 - Established (naturalized), 2 – Invasive, 3 –
Casual, 4 – Cryptogenic. Total number of distant alien
species is 169.

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219
In total, most of the registered number
recorded in Ukranian water bodies are distant
aliens (Figure 3), with the majority of the species
being either planktonic or fouling organisms
(Figure 4).

Figure 3. Main sources of alien species introduction into
Ukrainian waters: 1 – Ponto-Caspian, 2 – Atlantic, 3 – SE
Asia, 4 – N America, 5 – Other, 6 – Indo-Pacific, 7 –
Europe, 8 – Atlantic, 9 – Indo-pacific, 10 – Atlantic Pacific,
11 – Pacific, 12 – Holarctic.

Figure 4. Ratio of life forms of distant alien species
(biological composition of distant alien species of Ukrainian
aquatic ecosystems): 1 – Plankton, 2 – Fouling, 3 – Benthos,
4 – Nekton, 5 – Parasites
This confirms the key role of shipping in
transferring aliens. Usually, benthic animals
entered Ukranian water bodies as result of ships’
hull fouling. The maximum number of recorded
aliens occurred before the early 1970s (Figure
5). It is possible to explain the subsequent
decline due to the widespread introduction of
hull antifouling coatings. For example, in the
1970s, the overwhelming majority of ships’ hulls
were coated with tributyltin (TBT), which
protected the hulls from fouling for 18 to 24
months. Since then the number of cases of
recorded planktonic organisms invasions has
risen, due to ballast water exchange in tanker
fleets (see Figure 5). The total tonnage of tankers
in the 1980s made up 42% of the world total and
determined their priority in cargo turnover
(Marine encyclopedic reference… 1986).

Figure 5. Biological structure of distant alien species
introduced into Ukrainian aquatic ecosystems: 1 – fish, 2 -
phytoplankton, 3 – zooplankton, 4 – zoobenthos
Since 1973, subsequent to the adoption of
MARPOL -73/78, a measure to prevent pollution
from ships, conditions for survival of hydro-
bionts in ballast tanks were much better due to
isolation from oil products. This was another
reason why the probability of transferring
planktonic hydrobionts to new ecosystems has
increased.
When characterizing the long-term dynamics
of alien introduction records into Ukranian water
bodies, two maxima have been noted (Figure 6).

Figure 6. Long-term changes in distant alien species
introduction

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220
One of them, in the 1960-1970s, is connected
with the introduction of fish in freshwater bodies
and also with the establishment of new marine
ports, increasing the risk of the introduction of
aliens into the Black Sea. The second maximum
has been recorded since the beginning of the new
millenium and can be explained by two factors.
On one hand, it can be explained by a large
increase in ballast water transportation especially
by the oil tanker fleet. On the other hand, there
also has been a rise in scientific interest into the
problems of biological pollution in the world’s
oceans resulting in a more comprehensive study
of its biological structure. This can be illustrated
by the increasing number of publications in the
electronic abstract version of ASFA. In the past
decade the number of publications on alien
species and invasion has increased by 6-7 times
(Figure 7).

Figure 7. Dynamics of scientific publications on invasive
species. Source: Cambridge Scientific Abstract; key words 1
– “invasions”, 2 – “alien species”.
Due to more intense biological studies in
Ukraine the list of alien species was extended
adding marine fungi – 7 species, infusorians – 6,
parasites – 3 and unarmored Dynophyta – 11
species (see Annex). Due to the study of new
previously un-studied systematic groups, 3% (or
5 species) of the aliens discovered were placed
in the category “Cryptogenic” (see Figure 2)
The greatest numbers of distant species of
freshwater hydrobionts were introduced and
dispersed in the Ponto-Caspian basins. A small
part moved up the Dnieper to the mid Dnieper –
Danube province. Another fraction was introdu-
ced into northwestern Ukraine (Baltic province)
and the Crimea. The Annex illustrates some
species introduced into the upper Dnieper
cascade from North Europe. These species are
characteristic of the Baltic province. If the
Ukrainian water bodies of the Baltic basin were
closely studied, they would have most likely
have been found there. Probably, these species
entered the Dnieper basin through a system of
channels connecting it with the Baltic. In this
case, if borders are not taken into consideration,
then the introduction is similar to that in Annex
2 (nearby aliens).
Separate attention is necessary to focus on
aquarium species. Subsequent introductions to
waterbodies may occur unintentionally when
emptying aquariums. One of the famous
examples is killer algae Caulerpa taxifolia (M.
Vahl) C. Agardh, 1817 that penetrated into
Mediterranean coastal ecosystems from the
aquarium of Monaco Oceanographic Institute.
Most of the cultivated species in aquaria are
of tropical or subtropical origin. As a rule, they
survive in natural conditions of the area under
study, being sensitive to lower temperatures.
However, in the warm season they are
encountered close to populated areas. Examples
are aquarium fish like guppies and platies
encountered episodically in the Dnieper Basin
(Novitskij 2005), and piranha discovered in the
summer, 2006, in Lake Kasyanka (Dnieper
Basin) near Dnepropetrovsk. At present this
invasion is being studied by specialists from
Dnepropetrovsk National University (R.
Novitskij, personal communication).
However, some of the aquarium species are
able to form long-term populations in artificial
water bodies (in cooling ponds) – Ampullaria
sp., Biomphalaria glabrata Say, 1818,
Melanoides tuberculata Müller, 1774; or even in
the natural biotopes of most southern regions of
Ukraine – Craspedacusta sowerbyi Lankester
1880, Ferrissia fragilis Tryon, 1863), Physella
heterostropha (Say, 1817) (Protasov et al. 1981,
Son 2007a).
It is evident that most of the aquarium species
of the populations formed in open water bodies
are functional species (bloodfluke planorb
B. glabrata, pewter physa Ph. heterostropha)
and not cultivated decorative species which clean
the aquarium of “aquarium weeds“ entering from
one aquarium to another with sediments or plants
(red-rimmed melania M. tuberculata, freshwater
jellyfish C. sowerbyi Lankester 1880, freshwater
limpet F. fragilis. All of these species have very
a high fertility rate. The aquarium species of
gastropods have asexual reproduction (Son
2007a). These specific ecological properties
facilitate successful introductions.

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221
Analysis of neighbouring alien records
The main feature of neighbouring species, which
differentiate them from distant species is the
high percentage of naturalized species present –
90% (Figure 8), and also the fact that most of the
species records (68%) pertain to benthic
organisms (Figure 9). The chronology of
introducing species shows that their penetration
into the water bodies of the Ukraine was linked
with the wide scale melioration in 1950s, due to
the withdrawal of runoff from the largest rivers
of the Black Sea basins for agriculture purposes.

Figure 8. Ratio between main categories of neighbouring
alien species: 1 - Established (naturalized), 2 – Invasive, 3 –
Casual. Total number of nearby alien species is 71.

Figure 9. Ratio of life forms of neighbouring alien species
in Ukrainian aquatic ecosystems: 1 – Plankton, 2 – Fouling,
3 – Benthos, 4 – Nekton
The subsequent introduction of hydrobionts in
the late 1950s was not comparable in scale with
the previous one (Figure 10). The dominant
neighbouring species originated as Ponto-
Caspian relicts, which due to the artificial
integration of river systems have widespread in
the Black – Azov sea basin (Figure 11).
When considering neighbouring aliens, cases
of species introduction from adjacent
biogeographically areas, such as the Ponto-
Caspian into the European - Siberian area
through the Dnieper reservoir cascade and inner
waters of Crimea, should be of first priority. It is
quite clear that first of all many freshwater,
brackish Ponto-Caspian relicts were introduced,
which were lacking previously from the
European – Siberian sub-area. Hydrobionts were
able to overcome the zoogeographical border
along the Dneiper river bed after the removal of
the barrier between Dnipropetrovsk and
Zaporozhie following construction of a dam in
Kichkas and after activating river shipping,
dredging channels, setting up a reservoirs and
after intentional introductions. This led to further
distribution in the European-Siberian sub area:
mid Dnieper (from it to the Don, then into the
Volga) and upper Dnieper (from it the Baltic
basin, then to West Europe and Britain).

Figure 10. Long-term changes of neighbouring alien species
introduction into Ukrainian aquatic ecosystems

Figure 11. Main donor areas for neighbouring alien species
introduction into the Ukrainian part of the Black Sea: 1-
Ponto-Caspian relics, 2 – Black Sea, 3 - Mediterranean Sea,
4 - European water bodies, 5 – Holarctic, 6 - Cosmopolitan.

Alexandrov et al., Trends of aquatic alien species invasions in Ukraine


222
It is known that separate species of hydrobionts
with local areas in Europe as the cladocerans
Cercopagis pengoi (Ostroumov, 1891), Limno-
cletodes behning (Borutzky, 1926), Para-
leptastacus spinicaudata (T. Scott et A. Scott,
1895), and the amphipods Rivulogammarus
kischineffensis (Schell.), Synurella ambulans
(Müller, 1846) (see Annex) appeared in the
Dnieper cascade also at the time of introduction
of Ponto-Caspian fauna. These species, which
have very wide natural habitat, are made up of
several interrupted localities. This is characte-
ristic to Ukraine as in the lower Dniester and
Danube where there are many separate relict
localities of species, the main ranges of which
are in South or West Europe. Possibly, these
species were introduced into the Dnieper water
reservoir simultaneous with the mass introduc-
tion of Ponto-Caspian species as food resource
for fish.
Besides being spread along the Dnieper, they
penetrated to the Crimea. After the opening of
the North-Crimean canal and as a result of
introduction of food invertebrates in some water
reservoirs in the peninsula, some species with a
wide range of dispersal as crustaceans Daphnia
cucullata Sars, 1862, Leptodora kindtii Focke,
1844, Mesocyclops leucarti Claus, 1857 and
molluscs: river snail Viviparus viviparus Lin-
naeus, 1758; European fingernailclam Sphaerium
corneum Linnaeus, 1758, S. rivicola Lamarck,
1818 were able to enter with representatives of
the Ponto-Caspian fauna. Despite this there are
some species, with a main range located in the
European-Siberian sub area, which also entered
the Crimea as Ponto-Caspian fauna, at the
southern border of their distribution. The
situation differs for three molluscs species
Fagotia danubialis Bourguignat, 1884; gravel
snail Lithoglyphus naticoides Pfeiffer, 1828 and
freshwater nerite Th. fluviatilis. Their dispersal
was first noted only in the Crimea but they,
together with other species similar to the
representatives of Ponto-Caspian fauna, have
actively spread in Europe (see Annex). Often
they are confused with Ponto-Caspian as most of
the areas (lower Dnieper, South Bug, Dniester
and Danube) coincide. However, in contrast to
representatives of the Ponto-Caspian fauna, they
have not been formed in the Sea of the Sarmat,
but in rivers and are absolutely freshwater in
nature. L. natcoides inhabits an isolated part of
the area in the Baltic, while F. danubialis is
found in the eastern tributaries of Pripyat. As for
Th. fluviatilis, it pertains to a special
zoogeographical group of the most ancient
representatives of Ponto-Caspian fauna. It was
formed like the zebra mussel D. polymorpha in
the Eastern Mediterranean basins, but then
migrated to the northwestern Black Sea from
which many new species emerged - classical
Ponto-Caspian. In origin it is like the zebra
mussel as it is traditionally classified as a Ponto-
Caspian species. However, in Annex this species
is noted as European as in the summary table, its
affinity as a species is not related to the
zoogeographical group, but to its natural range.
The classification of the molluscs in the
European species category is attributed to the
time of paleoinvasion Th. fluviatilis has kept to
its localities in the Baltic, as well as in Spain. At
present, it is actively spreading in Europe. In the
Ukraine it has been inhabiting limans and rivers
(in contrast to classical representatives of the
Ponto-Caspian fauna it is native in upper
stretches of the large rivers) and also in several
marine bays (Odessa, Egorlitsky, Tendrovsky)
earlier forming the avandelta of the Dnieper (Son
2005).. However, the marine part of its range
does not border with the Chernaya River estuary.
Evidently, it has been introduced to the Crimea
like other species during faunal introductions
from the lower Dnieper to mountainous water
reservoirs.
Economic prerequisites in biological invasions
The presence of an indented coastline, deep
water bays and limans promoted the establish-
ment of a maximum amount of sea ports
following intensive agricultural and industrial
development in the Ukraine (see Figure 1), the
largest of which are Odessa, Yuzhny, Ilyichovsk.
In 2004 their annual turnover was more than a
half of the sum total of 19 marine ports of
Ukraine and amount to >60 mln. t (M t). By the
way it is necessary to stress on the fact that main
volum of shipping transportation in the Black
Sea comes through Ukraine. For example at the
same period (2004) cargo traffic of two main
ports of Russian Federation (Novorossisk and
Tuapse) was 89.8 M t, largest Black Sea ports in
Constanza (Romania) – 50.43 M t, in Georgian
ports (Poti, Supsa and Batumi) – 20.55 M t, in
Bulgarian ports (Varna and Burgas) – 13.35 M t.
Black Sea ports of Turkey (Istanbul, Darince,
Zonguldag and Samsun) develop very quickly
(Tokman 2005).

Alexandrov et al., Trends of aquatic alien species invasions in Ukraine


223
In the past 5 years (2001-2006) the total
turnover of Ukrainian ports has increased 1.6
fold. Simultaneously, the volume of ballast water
exchange increased; this is determined by the
numbers of export-import operations. For
example, in the port of Odessa in 2001, it
increased 17% in comparison to the previous
period, while in the port of Ilyichovsk it dropped
by 54%. It has been estimated that in 2001 alone,
more than 11 M t of ballast waters were
discharged in ten of the largest ports of Ukraine
(Savusin 2002).
Another pathway for the introduction of alien
species are the estuarine ports Ust-Dunaisk (near
the Danube), Belgorod-Dnestrovsk (near the
Dniester), the ports of Nikolaev and Kherson
(near the Dneipro-Bug Liman). Although their
cargo turnover is less than those mentioned
previously, there is a greater risk of transferring
alien species from a more dense marine environ-
ment to a less dense freshwater when ballast
waters are discharged. This is evident in the high
number of acclimatised species discovered in the
Danube delta (Alexandrov 2004). It should be
noted that most of the above mentioned ports,
e.g. Ust-Dunaisk, Belgorod-Dnestrovskiy, Ilyich-
ovsk and Yuzhny have been established in the
past 50 years during the period 1958-1980. This
should be taken into conside-ration when
analyzing the chronology of invasions.
The Northern Crimean canal has become a
powerful man-made source of biological
invasions for the freshwater water hydrobionts of
the Crimea.
The first Dneiper water came to the Crimean
Peninsula in 1963. Having filled up 8 water
reservoirs it is supplied to the cities of Kerch,
Feodosiya, Simferopol and Sevastopol with
maximum consumption of 300 m3/sec.
The source of many biological invasions is the
intentional introduction by man of valuable fish
species for consumption. In addition to spreading
naturalized fish e.g. haarder Liza haematocheila
Temmnick and Schlegel, 1845 (=Mugil soiuy
Basil and Europeski, 1855) and silver carp
Hypophthalmichthys molitrix Valenciennes, 1844
which took place in the Black Sea basin, it
allowed the simultaneous entry of other
undesirable organisms. For example, some
parasites, free-living algae, invertebrates and

even fish as stone moroko Pseudorasbora parva
Temminck and Schlegel, 1846 penetrated into
the Black Sea. Fish introductions were carried
out mostly in freshwater bodies. The existing
network of rivers and streams in Ukraine is
closely tied with those in the neighboring
Moldova Republic and the Russian Federation,
which should be taken into considerations in
terms of fish breeding. During the Russian
Empire, work for the introduction of species was
conducted 150 years ago under the guidance of
the Russian Society for Acclimatization founded
in 1857. Before the disintegration of the USSR
almost 250 introductions of 35 fish and 13
invertebrate species were performed annually.
Before 2000 almost 100 introductions of 16 fish
and 2 invertebrate species have been carried out
in Russia (Stroganov and Zadoenko 2000). As an
example, the river nerite Theodoxus fluviatilis
Linnaeus, 1758, belongs to a special zoogeo-
graphical group of the most ancient Ponto-
Caspian fauna representatives. Similar to zebra
mussel Dreissena polymorpha Pallas, 1771 they
evolved in the basins of the Eastern Mediter-
ranean, but migrated to the Ponto-Caspian area
producing many new classical Ponto-Caspian
species (Gelembiuk et al. 2006). Introductions of
fish from the Russian Federation were also
carried out. In Moldova, the introduction of fish
was carried out in three stages. In the first stage
in 1950-1961 there were organized introductions
of Lake Chud whitefish Coregonus lavaretus
maraenoides Poljakow 1874, European carp
Cyprinus carpio Linnaeus 1758, zander
(European pike-perch) Stizostedion lucioperca
Linnaeus 1758, eastern bream Abramis brama
Linnaeus 1758, roach Rutilus rutilus heckeli
Nordmann 1840 and various sturgeons (Acipen-
seridae). In the second stage, 1961-1974, fishes
were introduced from Chinese water bodies:
silver carp H. molitrix, spotted silver carp
Aristichthys nobilis Richardson 1846, grass carp
Ctenopharyngodon idella Valenciennes 1844. In
the third stage, 1974-1990, valuable varieties of
fish from the freshwater bodies of the USA:
smallmouth buffalo Ictiobus bubalus Rafinesque
1818, bigmouth (common) buffalo Ictiobus
cyprinellus Valenciennes 1844, black buffalo
Ictiobus niger Rafinesque 1818, channel catfish
Ictalurus punctatus Rafinesque 1818, paddlefish
(spadefish) Polyodon spathula Walbaum 1792
were introduced (Lobchenko 1999).
Acknowledgements
We are grateful to Dr. Frances Lucy (Institute of
Technology Sligo, Ireland) for her comments and
English editing. This study has been supported

Alexandrov et al., Trends of aquatic alien species invasions in Ukraine


224
by the European Commission Sixth Framework
Programme Integrated Project ALARM
(Assessing LArge scale environmental Risks for
biodiversity with tested Methods, contract
GOCE-CT-2003-506675), and part of the EC
FP6 Strategic Targeted Research Project DAISIE
(Delivering Alien Invasive Species Inventories
for Europe, contract SSPI-CT-2003-511202).
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229


Annex
Check-list of aquatic alien species of Ukraine)

Year of first record and
recipient region
Species Sea and
inland
saline
waters
Fresh
waters
Inland
brackish
waters or
waters
with
salinity
variation
Origin /
donor area
Vector Pathway
Impact or
potential
impact /
invasive-
ness
References
Fungi, ANAMORPHIC
Cirrenalia basiminuta
(Кaghu-Kumar and
Zainal, 1988)
2001 - -
Indo-
Pacific
Unintentional Ships Unknown Kopytina 2002
Cumulospora marina
(Schmidt, 1985) =
Vesicularia marina
(Schmidt, 1974);
Basramyces marinus
(Abdullah, Abdulkadder
and Goos, 1989)
2001 - -
Indo-
Pacific
Unintentional Ships Unknown Kopytina 2006
Cumulospora varia
(Chatmata and
Somrithipol, 2004)
2004 - - SE Asia Unintentional Ships Unknown Kopytina 2006
Fungi, ASCOMYCOTA
Lulworthia uniseptata
(Nakagiri, 1984) 2001 - - SE Asia Unintentional Ships Unknown Kopytina 2002
Savoryella lignicola
(Jones and Eaton, 1969) 2001 - -
Cosmo-
politan
Unintentional Ships Unknown Kopytina 2004
Zopfiella latipes
(Malloch and Cain,
1971)
2001 - -
Indo-
Pacific
Unintentional Ships Unknown
Zaitsev et al.
2004,
Kopytina 2006
Gloniella clavatispora
(Steinke and Hyde,
1997)
2002 - -
Indo-
Pacific
Unintentional Ships Unknown Kopytina 2004
Microalgae, BACILLARIOPHYTA
Toxonidea insignis
(Donkin, 1858) 1902 - - N Atlantic Unintentional Ships Unknown
Merezhkowsky
1902-1903
Cocconeis britannica
(Naegeli, 1849) 1902 - - N Atlantic Unintentional Ships Unknown
Merezhkowsky
1902-1903
Pinnularia trevelyana
((Donkin) Rabenh.,
1861)
1902 - - N Atlantic Unintentional Ships Unknown
Merezhkowsky
1902-1903
Bacteriastrum hyalinum
(Lauder, 1864) 1907 - - Atlantic Unintentional Ships Unknown Reingardt 1909
Pseudosolenia calcar-
avis (Sundström, 1986) 1924 - -
Atlantic,
Indo-
Pacific
Unintentional Ships Unknown Usachev 1928
Asterionellopsis
glacialis (Round,1990) 1967 - - Atlantic Unintentional Ships Unknown Senicheva 1971
Navicula finmarchica
(Cleve and Grunow,
1880)
1970 - -
N Atlantic,
Pacific
Unintentional Ships Unknown Bodeanu 1970
Achnanthes
pseudogroenlandica
(Hendey, 1964)
1984 - - Atlantic Unintentional Ships Unknown
Guslyakov and
Gerasemiuk
1984, Nevrova
2003
Undatella quadrata
((Brebisson) Paddock
and Sims, 1980)
1985 - - N Atlantic Unintentional Ships Unknown
Roschin et al.
1992
Nitzschia sigmoidea
(Nitzsch, 1817), (Smith,
1853)
1986 - - N Atlantic Unintentional Ships Unknown
Roschin et al.
1992