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CHOROLOGY
The transformer species of the Ukrainian Polissya
Vira V. Protopopova1, Myroslav V. Shevera1*, Oleksandr O. Orlov2
& Sergiy M. Panchenko3
1M. G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine, Tereschenkivska 2, 01601 Kiev, Ukraine
2G. M. Vysotskiy Polissky Branch of Ukrainian Scientic Research Institute of Forestry and Agro-Forest Мelioration, State Agency of Forest
Resourses of Ukraine and National Academy of Sciences of Ukraine, Neskorenykh, 2, 10004, vill. Dovzhyk, Zhytomyr District, Zhytomyr
Region, Ukraine,
3Desniansko-Starogutsky National Nature Park, Novgorod-Siversky, 62, 41000, Seredyna-Buda, Sumy Region, Ukraine
* corresponding author (e-mail: shevera.myroslav@gmail.com)
Abstract: The investigation results of the transformer species participation (Echinocystis lobata (Michx.) Torr. & A. Gray,
Heracleum sosnowskyi Manden., Impatiens glandulifera Royle, I. parviora DC., Reynoutria japonica Houtt., Robinia pseu-
doacacia L.) in different plant communities of the Ukrainian Polissya (Forest zone of Ukraine) are presented. All the above-
mentioned species are strong edicators in the region that can signicantly change important species composition parameters
of communities and character of landscape.
Key words: alien plants, transformer species, Forest zone, Ukraine
DOI 10.1515/biorc-2015-0020 Submitted 23.02.2015, Accepted 17.08.2015
1. Introduction
By the end of the 20th century, invasions of alien
organisms, including plants, were widely realized as
one of major global threats for biodiversity (Pyšek et
al. 1995; Mooney & Cleland 2001; Reichard & White
2001; Kowarik 2002; Protopopova et al. 2002, 2003;
Chornesky & Randall 2003; Davis 2003; Sax & Gaines
2003, etc.).
Ukrainian Polissya region (Forest zone of Ukraine)
for a long time has been characterized by a low level of
vegetation adventization, mostly due to its natural and
climatic conditions: large percentage of the area covered
with forests and wetlands, absence of large urban and
industrial centers, relatively poorly developed transpor-
tation network, and other factors. Following major land
reclamation conducted during the mid-20th century, the
situation changed signicantly. The urbanization and
emergence of industrial centers over the last decades of
the 20th century facilitated penetration of many species
of alien plants, which currently tend to be spreading
actively.
The alien fraction ora in the left-bank regions of
Polissya is specic; the species composition is more
diverse and alien species are generally more wide-
spread. Here, the inuence of the Forest-Steppe zone
and eastern continental Eurasiatic regions is observed.
Alien species prevail in the orogenetic spectrum of
the Eastern Mediterranean and Irano-Turanian origin.
The right-bank regions of Polissya have species of
the Central European origin, including species con-
nected with South European mountain areas, Western
Mediterranean species, species from the Balkans and
Caucasus region, as well as numerous mesophytes of
the Northern American origin. The distribution of spe-
cies on the territory of Polissya has been inuenced by
drainage reclamation with substantial changes caused
in the aboriginal species composition and structure of
natural ecosystems.
The most negative changes are caused by invasive
species, especially transformers species (according
to Richardson et al. (2000). These species changed
phytocoenotic environment of plant communities
(regime of soil and air moisture, biochemical com po-
sition of soil, light, etc.), its oristic composition and
structure.
The aim of the presented work was to study the
transformer species of plants in the Ukrainian Polissya
region. The objectives of the study included: (i) dening
the group of transformer species, (ii) reconstructing
Biodiv. Res. Conserv. 39: 7-18, 2015
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the history of their cultivation or introduction and
evaluating the current distribution and coenotic con-
nement of species.
2. Material and methods
The presented work continues the research direction
we carried out on the regional level (Protopopova et al.
2009, 2010, 2012, 2014). The study is based on original
data obtained through routine research in 2010-2014,
the analysis of literature data and examination of the
collections of the Herbaria of M. G. Kholodny Institute
of Botany, National Academy of Sciences of Ukraine
(KW), Ivan Franko Lviv National University (LW),
Taras Shevchenko Kiev National University (KWU),
Taras Shevchenko Chernigiv National Pedagogical
University (CHU), Lesya Ukrainka East-European Na-East-European Na-
tional University (LUU) and Desniansko-Starogutsky
National Nature Park (DSR). Comparative morphologi-
cal and geographical methods were used in the present
investigations. The nomenclature of species follows
the Checklist of vascular plants of Ukraine (Mosyakin
& Fedoronchuk 1999). The transformer groups were
determined according to Richardson et al. (2000) as:
“a subset of invasive plants which change the character,
condition, form or nature of ecosystems over a substan-
tial area relative to the extent of that ecosystem”. The
participation of the transformer species in the plant com-
munity was analyzed according to the Braun-Blanquet
classication.
The dot maps of distribution of Echinocystis lobata,
Heracleum sosnowskyi, Impatiens glandulifera and
Reynoutria japonica in the region were prepared based
on herbarium and eld expedition data.
The satellite images from the RapidEye and
World
View-2 used in the paper for Echinocystis lobata
were obtained during the “Avoidance of greenhouse
gas emissions by restoration and sustainable mana-
gement of peatlands in Ukraine” project, 2009-2012.
(BMU-№: 09_III_009_UKR_K_Moor Renaturierung),
executed by the Ukrainian Society for the Protection
of Birds (USPB) in a partnership with the Institute for
Commu nity Development (ICDU), the Royal Society
for the Protection of Birds (RSPB, UK) and the Michael
Succow Foundation (Germany) with nancial support
from the Federal Ministry of Environment, Nature
Conservation and Nuclear Safety (Germany) through
KfW Entwicklungsbank within the International Initia-
tive Government of Ukraine and Germany to reduce
the negative effects of climate change. Based on their
spectral characteristics, the areas that distinguish from
the surrounding vegetation were highlighted on the
satellite images and their geographical coordinates were
determined. Using GPS, such areas were subjected to
deciphering of plant communities; investigation of
oristic composition determining trends of succession
and photographing was also conducted.
3. The studied area
The Ukrainian Polissya is a physiographic province
located in the south-west part of mixed forest zone, on
the border with the forest-steppe zone. The Polissya
region is characterised by a low relief, extensive
hydrological system, wide, swampy river valleys, high
groundwater level and prevalence of turf-podzol and
swamp soils. The highest participation in plant cover of
this region have pine and mixed forests, meadows and
swamps. The area has also a high share of agricultural
lands. It is occupied by different geological structures of
Russian Platform. The west part of Ukrainian Polissya
is situated on the northern part of Galytsko-Volynska
depression, middle part – on the north-west part of the
Ukrainian crystalline shield and its slope, and eastern
part – in the Dnipro-Donetsk depression. Landscapes
of these Polissyan areas are characterized by nu merous
landforms, such as: river valleys and outwash, moraine-
outwash and moraine plains. These landforms prevail
The transformer species of the Ukrainian PolissyaVira V. Protopopova
Fig. 1. Map of Ukrainian Polissya
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Biodiv. Res. Conserv. 39: 7-18, 2015
in the modern relief of Polissya. The region has a
temperate-continental climate with warm and humid
summer and mild, cloudy winter, and it is divided into
5 botanical and geographical regions (Fig. 1).
Volynian Polissya is a at undulating plain, located
to the north of the Pripyat River valley. This region is
characterized by numerous swamps, deposits of marl
and chalk, turf-podzol soils and lakes of karst origin.
Vegetation is dominated by broad-leaved pine forests.
Zhytomyr Polissya is located on the Ukrainian crys-Ukrainian crys-
talline shield. It is distinguished by the signicant devel-
opment of denudation forms of relief, well-developed
topographic features and small waterlogged areas. Its
southern and central parts are located at the altitude of
200–250 m a.s.l., northern part – 150–200 m a.s.l., and
Slovenchansko-Ovruchsky ridge – 315 m a.s.l. The
region has numerous outputs of crystalline rocks and
sand-dominated clay-sandy and sod-podzolic soils. In
vegetation prevail oak-pine, birch-pine, pine, hornbeam-
oak and oak forests with high species diversity.
Kiev Polissya occupies an area of the Middle Dni-
pro region, located on the Ukrainian crystalline shield
in the Dnipro-Donetsk depression. Its topographic
features are poorly developed and a high groundwater
level promotes waterlogging. Among soils prevail turf-
podzol soils, while vegetation is dominated by oak-pine
and pine forests, as well as pine plantations. Meadows,
swamps and shrubby vegetation are distributed along
the region’s river valley.
Chenigiv Polissya occupies an area located in the
Dnipro-Donetsk depression. It is a low relief area – a
moraine-outwash, slightly undulating plain with small
loess islands, dissected by the Dnipro, Desna and Snow
River valleys. The region is dominated by poor and
medium turf-podzolic soils, occupied by mixed forests.
Novgorod-Siversky Polissya is an extreme eastern
part of the Ukrainian Polissya. It occupies the northern
part of south-western slope of the Voronezh crystalline
massif. The climate is more continental than in other
parts of the Ukrainian Polissya and the region has the
lowest percentage of waterlogged areas. The specic
features of this region are numerous chalk outputs and
development of karst phenomena. The main types of
forests are pine-mixed and oak-hornbeam forests. They
are characterised by high species diversity.
The Polissya region of Ukraine occupies the southern
part of Polissian Lowland of the East European Plain.
The land area is approximately 113 thousand km² (19%
of the territory of Ukraine). It is divided by the Dnipro
River into the Right-bank Polissya (western part) and
the Left-bank Polissya (eastern part). The Right-bank
Polissya (Wolynian Polissya, Zhytomyr Polissya and
Kiev Polissya) differs from the Left-bank Polissya
(Chernigiv and Novgorod-Sivers’k Polissya) (Popov
et al. 1968).
4. Results
Depending on the determined invasive potential of
alien species, six transformers were dened and char-
acterized in detail, which are common for both regions
of Ukrainian Polissya: Echinocystis lobata (Michx.)
Torr. & A. Gray, Heracleum sosnowskyi Manden., Im-
patiens glandulifera Royle, I. parviora DC., Reynou-
tria japonica Houtt., Robinia pseudoacacia L. Among
the determined transformers, the following types were
identied: therophytes (3 species), phanerophyte (1),
hemicryptophyte (1), and geophyte (1). All species
are neophytes judging by their time of immigration,
and ergasiophytes – by the way of introduction; four
species were introduced into the region during the
second half of the 20th cent., and two – by the end of
19th cent., but their active distribution and pervasion
into natural ecosystems had begun at the end of the 20th
cent. and was characterized by a relatively short period
of naturalization. The participation of the transformer
species in the anthropogenic, semi-natural and natural
ecosystems and plant communities of Polissya and
their inuences on the aboriginal ora were analyzed.
Echinocystis lobata (Michx.) Torr. & A. Gray – spe- spe-
cies of the North American origin, which is common
in the North-east of the United States (reaching Texas
and Florida) and also the neighboring areas of Canada
(49°-55°N) (Gleason & Cronquist 1991; Vinogradova
et al. 2010), mostly along river banks, moist areas and
oodplain shrubs.
Its transformative impact is caused by entwining
bushes (highly ramied stem can reach the length of 4-6
m), therefore shading the area is signicant as well as
increasing its air humidity level, making other natural
inhabitants of the area, especially photophilic ones, drop
out of vegetation which causes signicant depletion of
phytodiversity of oodplain ecosystems.
Its high levels of seed production (one plant is able
to produce 40-160 seeds), high germination rate – up
to ~ 70 % (Vinogradova et al. 2010) promotes further
spreading of the species and its rapid growth which,
together with the simplicity of the conditions required
for growth, leads to displacement of other species of
the grass community.
The species was cultivated in Europe since the
middle of the 19th century (Romania) and it was found
growing wild in 1904. Then after 1923, it was also
found in Ukraine (Dnister source) and in 1938 – in
Transcarpathian region (Protopopova & Shevera 2014).
In Polissya it is mainly found to be part of sparse tree
and shrub communities growing on the relatively rich
and moist soils (Fig. 2). It prefers natural and semi-
natural riverside and ooded areas, and can be often
found along river banks, ditches, shrub-willow thickets,
along drainage channels, on drained eutrophic swamps
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Fig. 2. Distribution of Echinocystis lobata in Ukrainian Polissya
Fig. 3. Fragment of RapidEye satellite image of extensive Romen
swamp, between the villages of Ponory, Korіnetske and Lypove in
the Talalayivsky District of Chernigiv Region (09.08. 2010), with
participation of Echinocystis lobata. The pixel size – 5 × 5 m
and areas formed after peat extraction. The species is a
component of Alnetea glutinosae and Galio-Urticetea
plant communities (Lukash 2008).
As it was shown during the execution of the
«Avoidan ce of greenhouse gas emissions by restoration
and sustainable management of wetlands in Ukraine»
Project, E. lobata formed dense thickets (projected
coverage was 80-100%) on the studied wetland areas
of Chernihiv Polissya (Romen and Udai wetlands, on
the border with Forest-Steppe zone) which can be easily
distinguished among other vegetation even with remote
sensing on space images.
The following fragments of satellite images of the
wetlands are matched with the ground-level photos
of the thickets. Figure 3 clearly shows that E. lobata
occupies the narrow strip along the drainage channel
(1 km length and 10-30 m width), covering the riparian
vegetation and the shrub willows (Fig. 4).
Less frequently, it creates large thickets, 250-300 m in
diameter, covering the old peat extraction areas after some
reclamation. On the Udai wetland, E. lobata is spread
mostly between the regular drainage channels on the old
peat extraction spots forming nearly monospecies thickets,
20-150 m in diameter, with the projective coverage of
90-100 % (Figs. 5-6), suppressing all other species.
The distribution assessment of the species across
the wetland showed that it hindered renaturalization of
the wetland vegetation, created sustained plant com-
munities with the succession different from the planned
rehabilitation directions, quickly expanded onto new
areas and - on the whole – created signicant problems
for rewetting.
On the left bank Polissya, the E. lobata was common
in Fraxino-Alnetum (the frequency of occurrence was
10-15%), Chelidonio-Robinietum (1-5%), Salicetum
pentandro-cinereae Passarge 1961 (30-40%).
Heracleum sosnowskyi Manden – the species of
Caucasian origin; in its natural habitat, it grows in the
middle and upper forest zones of mountains, mainly on
the forest glades and edges (Mandenova 1951). In its
secondary habitat during the initial stages of expansion,
the species is spreading through open, mainly anthropo-
genic or semi-natural ecotopes: eld edges, abandoned
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Biodiv. Res. Conserv. 39: 7-18, 2015
Fig. 5. Fragment of WorldView-2 (DigitalGlobe) of the satellite
image of extensive Uday swamp, between the villages of Mona-
sterysche and Korshaky in the Ichnya District of Che rnigiv Region
(02.08. 2010), with participation of Echinocystis lobata. The pixel
size – 2 × 2 m
places of culture, road sides, along the drainage chan-
nels, in the meadows, especially disturbed ones, around
farms in ruderal ecotopes, in gullies and ravines, etc.
Among the natural ecotopes during this period, it grows
in oodplains. During the second period of expansion,
this species appears under the forest canopy, creating
dense thickness even in oak forests and spruce forests.
Such situation now is observed in some localities in
Belorussia (Laman et al. 2009), although it prefers better
lighted areas. At the same time, the essential feature of
this species is its high ability to continue enlarging the
areas created during the previous period.
Its high germination ability (above 90%), early
germination, high viability and rapid growth, high
population density, early and stretched blooming
periods, self-pollination, high content of biologically
active substances (furanocoumarins), effective ways
of seed dispersal etc. cause great negative impact on
phytodiversity (Dalke & Chadin 2008).
The species had been cultivated in Russia since
1947 and was well known in Ukraine starting from
1960s-1970s in western and northern forested regions
(Protopopova & Shevera 2014). The plant was grown
in Polissya since 1960s as a silage crop at the Zhytomyr
regional bee farm (Tokar 1975). Intensive distribution
in the western regions of Ukraine inclu ding Polissya
was observed in late 1980s – early 1990s (Fig. 7).
In Eastern Polissya, H. sosnowskyi was found along
roadsides, highways, train tracks, abandoned cultural
places, fallows, forested oodplains with mosaic vegeta-
tion, willow dominated areas, forests with tree canopy
of low density, forest edges as a component of the
Artemisietea vulgaris Lohm., Prsg. et R. Tx. in R. Tx.
1950, Agropyretea repentis Oberd., Th. Müll. et Görs
in Oberd. et al. 1967 plant communities (Lukash 2008).
The situation is similar in the central part of Polissya.
As for the Western Polissya, oodplain localities are
prevailing (rivers Styr, Horyn’, Ustia, Ikva), where
thickets of considerable length (250-500 m) and width
are formed.
Fig. 4. Thickets of Echinocystis lobata along the watercourse, as shown by the arrow in Fig. 3
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Fig. 6. Continuous thickets of Echinocystis lobata on the old peat extraction site, as shown by the arrow in Fig. 5
Fig. 7. Distribution of Heracleum sosnowskyi in Ukrainian Polissya
Sometimes H. sosnowskyi is already a component of
the herb-dwarf-shrub layer of the dense hornbeam-oak
forests, where its projective coverage can reach up to 80%.
In the future, the number of such localities of this species
in dense forests in the region is expected to be increasing.
H. mantegazzianum Sommier & Levier has a similar
inuence on the plant communities – another species of
the Caucasian origin. Both species of Heracleum were
often cultivated together in Polissya region, their wild
populations tend to consist of plants of both species as
well and, possibly, these species can give hybrids, since
the plants now often show some signs not typical for
their parental forms.
In forested and forest edge areas of Eastern Polissya,
H. mantegazzianum displaces the following species
from the grass level: Calamagrostis epigeios (L.)
The transformer species of the Ukrainian PolissyaVira V. Protopopova
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Biodiv. Res. Conserv. 39: 7-18, 2015
Fig. 8. Distribution of Impatiens glandulifera in Ukrainian Polissya
Roth, Agrostis vinealis Schreb., Convallaria majalis
L., Chamaerion angustifolium (L.) Holub, Gnaphalium
sylvaticum L., Euphorbia cyparissias L., Galium verum
L., Veronica spicata L. etc.; and in the ruderalized
psammophytic and meadow Artemisietea vulgaris and
Agropyretea repentis communities, it displaces both
the natural meadow species (e.g. Festuca pratensis
Huds., Dactylis glomerata L.) and the synanthropic
species (like Elytrigia repens (L.) Nevsks, Melilotus
albus Medik., Pastinaca sylvestris Mill., Daucus carota
L., etc.), and also alien species (Lupinus polyphyllus
Lindl., Cichorium intybus L., Centaurea diffusa Lam.,
etc.), with the progective coverage reaching up to 95%
in some places.
Less commonly, it inhibits grass communities
formed within forest clearings, classied as Epilo-
bietea angustifolii Tx. et Prsg. ex von Rochow 1951
and the transformed forest cоenoses – Calamagrostio
arundinaceae-Pinetum Shevchyk et V. Sl. 1996 (Lukash
2008).
Impatiens glandulifera Royle – the species of West-
Himalayan origin (Beerling & Perrins 1993), is often
located along river banks, eld edges and undisturbed
moist forests (Valentine 1971) at the altitude of 1800-
4000 meters, and in Pakistan – up to 4300 m (Nasir 1980).
At its secondary area, one plant is able to produce
from 500 to 2500 seeds, has wide range of habitats,
yet prefers moist areas with rich soils and sparse grass
coverage (Markov et al. 1997). Factors limiting the
expansion of this species include moisture scarcity and
low temperatures (Beerling & Perrins 1993). In Europe,
the species tends to form thickets with 40, sometimes
up to 70 plants/m2, in Central Russia – 200-700
plants/m2
(Wadsworth et al. 2000; Vinogradova et. al.
2010; Vinogradova & Kuklina 2012) that impede local
ora development, suppressing not only grass species,
but tree-species as well. As a result, instead of mixed
grass associations, plant communities dominated by I.
glandulifera are formed, for example the Impatienti-
Calystegium com. association found in the Czech
Republic (Vinogradova et. al. 2010).
In Europe (England), this species has been cultivated
since 1838 and the rst references of its cultivation in
Ukraine go back to the 1930s, rst wild plants were
mentioned in 1938 in Osii and Han’kovytsia villages
(Transcarpathian region) and in 1939 – in Mykhailivka
vil. (Khmelnitsky region) (Protopopova & Shevera
2014). It was also cultivated in Polissya, including the
Left-bank Polissya; wild forms found growing in 1980s.
The species can sporadically spread in the region at
present (Fig. 8).
In Polissya, the species can be found on river banks,
tall grass eutrophic wetlands and swampy thickets,
alder forests, forested oodplains of streams and rivers,
moist forests with high and low density tree strand, on
forest edges, along roadsides and near houses in ruderal
ecotopes as a part of Phragmito-Magnocaricetea Klika
in Klika et Novak 1941, Mollinio-Arrhenatheretea R.
Tx. 1937, Alnetea glutinosae Br.-Bl. et R. Tx. 1943 ex
Westhoff et al. 1946, Bidentetea tripartiti Tx., Lohm.
et Prsg. in Tx. 1950 communities.
In some places, it occupies areas 2 km × 10-30 m
(Lukash 2008). In black alder forests of Zhytomyr
Polissya, the population density of I. glandulifera ranges
from 10 to 40 plants/m2 with the projective coverage
between 50 and 100%. In open ruderal habitats, the
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Fig. 9. Distribution of Reynoutria japonica in Ukrainian Polissya
population density is somewhat lower – 10-30
plants/m2;
however, the average plant is much taller –
up to 2.3 m.
On the left bank Polissya, I. glandulifera was very
rarely noted near settlements in Carici elongatae-
Alnetum W. Koch. 1926 ex Tx. 1931 ass. cenoses.
Impatiens parviorа DC. – is a species of Central
Asian origin (Nasir 1980; Beerling & Perrins 1993),
natural inhabitant of the hazelnut-maple forests of
Tien Shan and Pamir-Alai, where it growth in the
grass layer with Brachypodium sylvatica (Huds) P.
Beauv., Poa nemoralis L., Geum urbanum L. It prefers
rich well-aerated soils but can also grow on relatively
poor ones (Pobedimova 1949; Ovchinnikov 1971;
Vinogradova et al. 2010; Protopopova et al. 2010).
The species re produces with seeds; is chasmogamous,
or less frequently – cleistogamous; thus providing
polli nation independent of the presence of insect
pollina tors (Vinogradova et al. 2010).
As a secondary forest habitat, due to its high
competitive ability and seed production (up to 10000
seeds per plant), early and long germination and
fruiting periods (aprox. 3 months dissem ination period),
effective ways of seed dispersal (autho-mechanochorus),
formation of great phytomass etc., it suppresses species
of natural ora through altering light regimes, moisture
and temperature, replaces the aboriginal Impatiens noli-
tangere L., suppresses other species of grass layer of
the deciduous and coniferous-deciduous forests. Thus,
expansion of I. parviora leads to transformation of
vegetation.
In Europe (Switzerland), the plant had been culti-
vated since 1831, while in Ukraine – the cultivation
was recorded in 1895 in Dubliany (Lviv region) and the
species grew wild – in 1908 in Lviv and the Carpathians
(Protopopova 1973; Protopopova & Shevera 2014).
In Polissya region, the species is registered in both
low and high stand density forests, on forest edges,
in younger forest plantations, forest nurseries, forest
parks, disturbed natural forests and forest edges, as
well as along roadsides, train tracks, ruderal areas, as
part of the Querco-Fagetea Br.-Bl. et Vlieger in Vlieger
1937, Galio-Urticetea Passarge ex Kopecky 1969,
Robinietea Jurko ex Hadač et Sofron 1980 plant com- plant com-
munities (Orlov & Yakushenko 2005; Lukash 2008).
The species, in the Kiev urban forest, is characte rized
by the following parameters, e.g.: constancy (100%),
population density (29.2 plants/m2) exceeding those
of other species by 10 or more, and according to R.
Burda (2012) the I. parviflora is a transformer of
forest herbaceous cover. In forests of Kiev and vicin-
ity, the species is characterized by high ecological and
phenotypical variability, including population level.
The relative intensity of lighting and soil moisture are
main limited factors of the functioning of the species
populations (Golivets 2014). In pine, pine-oak and
hornbeam-oak forests of Zhytomyr Polissya, its popu-
lation density varies from 20 to 70 plants/m² with the
projective cover between 50 and 100%. On the left-bank
Polissya, I. parviora participates in the following plant
communities: Chelidonio-Robinietum Hadač et Sofron
1963 (the frequency of occurrence is 25-30%), Fraxino-
Alnetum W. Matuszkiewicz 1952 (10%), Mercurialo
perrenis-Quercetum roboris Bulokhov et Solomeshch
2003 (2%), Galeobdolono lutei-Carpinetum Shevchik
et al. 1996 em. Onyschenko et Sidenko 2002 and in the
The transformer species of the Ukrainian PolissyaVira V. Protopopova
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Biodiv. Res. Conserv. 39: 7-18, 2015
secondary associations of Lathyro nigri-Quercetum
roboris Bulokhov et Solomeshch 2003 (Potentillo
albae-Quercetum roboris Bulokhov 1991), which are
r
e presented by oak- and pine-hazel forests, and in the
ruderal communities and erosion-controling forest
cultures of Chelidonio-Robinietum (30-40%) asso-
ciation.
Reynoutria japonica Houtt. is a species of the
East Asian origin [Komarov & Grigor’ev 1936; Vino-
gradova et al. 2010] (southern Primorye of Far East
and Sakhalin, Southern Kuriles, Japan, Korea, most of
China, Taiwan). It can be found in China on mountain
slopes, valleys and eld edges; in Japan – it grows at
altitudes of up to 1500 m above the sea level, is a pioneer
species to cover volcanic deserts and open areas with
bare ground. It creates huge phytomass which leads
to changes of illumination regime, shading and later
displacing phytophillous aboriginal plant species.
This species was introduced in Europe in 1825:
originally brought from China by the London agricul-
tural society, later seedings were mailed from Siebold
in Leiden (Netherlands).
The data about the beginning of cultivation of the
species in Ukraine is absent; in the wild state – the
plants were recorded in 1929 in Rakhiv (Transcar-
pathian region) (Protopopova & Shevera, 2014) where
they probably arrived from Romania.
In Polissya region, R. japonica grows on former
places of cultivation, along roadsides, train tracks,
forest edges, fallows, parks, cemeteries, forest edges.
Active distribution of the species in Polissya regions
was observed since 1990s (Fig. 9).
The species is a diagnostic species of the Reynoutrie-
tum japonicae Görs et Müller in Görs 1975 and a
component of several plant communities: Artemisietea
vulgaris Lohm. et Prsg. & R. Tx. in R. Tx. 1950 em.
Lohm. & al. 1962, Galio-Urticetea Passarge ex Ko-
pecký, and also Plantaginetea majoris R. Tx. et Prsg. in
R. Tx 1950, Stellarietea media R. Tx., Lohm. et Prsg.,
1950 classes (Lukash 2008).
Robinia pseudoacacia L. – species of the North
American origin, in its natural habitat (from Pennsyl-
vania south to Georgia and west towards Iowa and
Oklahoma), it grows in mixed forests of various
broadleaf species (Gleason & Cronquist 1991).
Unpretentious to soils, it, however, prefers light,
well-aerated soils, loamy or with admixture of lime.
The plant is photophilous, drought- and frost-resistant
(withstands up to -40° С). Its annual growth at young
age
is 1-1.2 m/year, starting from the age of 10 years
– 25-50 cm/year
and starting from the age 40 years
– approx. 20 cm/year. The seed reproduction is less
effective (seed viability is 90-100 %, with 10-20 %
germination) than vegetative, seeds germinate quickly
and the seedlings rapidly develop, which ensures their
competitive ability. The species enriches soil with ni-
trogen, thus, suppressing growth of natural inhabitants
which could not withstand high nitrogen levels and
causing vast rooting of weeds, nitrophilous species
instead, such as Urtica dioica L., Cannabis sativa L.
etc.
The root system covers an area of 15 m2, forms
numerous shoots from roots which quickly develops
creating tight undergrowth. When it prevails in the
forest stands, it can cause a certain impact on the surface
of the soil acting like a strong edicator. High levels of
nitrogen released from fallen leaves in upper soil levels
during certain vegetation periods and high illumination
level provide a dominant position for the nitrophilous
herbaceous plants (Protopopova et. al. 2014). There is
also literature data about allelopathic impact of this
species on aboriginal vegetation through the inuence
of some phenolic compounds and their derivatives in
the soil (Vinogradova & Kuklina 2012).
In Europe (England), the species was cultivated since
the 17th and in Ukrai ne – since the 18th century (Palimp-
sestov 1855), as for Polissya region – the exact date is
unknown. According to Schmalhausen (1895), the plant
was cultivated but without any specic guidelines and
according to Flora of Ukrainian SSR (Visiulina 1954)
– it was cultivated as common species throughout the
whole country.
In Polissya region, the plant acts as a transformer
species, mainly in pine forests. Seedlings of R. pseudo-
acacia tend to appear on recreational and gra zing terri-
tories or in places of total tree felling, esp. where the
moss-coverage is disturbed; they develop quickly and
replace the undergrowth of Pinus sylvestris and sup-
press other forest species replacing them with weeds,
Chelidonium majus. According to Lukash (2008), R.
pseudoacacia forms such ecological-coenotic series
in pine forests of Eastern Polissya: disturbed coenosis
of Peucedano-Pinetum W. Matuszkiewicz 1962 with
numerical juvenile instances of R. pseudoacacia →
depleted coenosis of Peucedano-Pinetum with dense
spinneys of R. pseudoacacia → transition cenoses be-
tween Peucedano-Pinetum and Chelidonio-Robinietum
→ typical Chelidonio-Robinietum.
On the left-bank Polissya, R. pseudoacacia is widely
used for creation of forest cultures of shelterbelts
on agricultural land sites and on cuttings of oak-pine
and deciduous forests. It is a characteristic spe- It is a characteristic spe-
cies of Chelidonio-Robinietum (the frequency of
occurrence is 100%) (Panchenko 2013) and noted in
Mercurialo perrenis-Quercetum roboris (the frequency
of occurrence is 1%), Querco roboris-Pinetum sylve-
stris (W. Matuszkiewicz 1981) J. Matuszkiewicz 1988
(the frequency of occurrence is 1%), and Agrostio
vinealis-Salicion acutifoliae Bulokhov 2005 (single
indivi duals).
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16
There are four more species changing the structure
of plant communities of the region, but as they more
or less occupy limited areas, we only consider them to
be potential transformers: Amelanchier spicata (Lam.)
K. Koch, Quercus rubra L., Symphyotrichium novae-
angliae (L.) Nesom, S. lanceolatum (Willd.) Nesom.
In Polissya region, the Amelanchier spicata was
known since 1980s, growing not far from areas of
its cultivation, for example, in Korostyshiv, near the
former manor and in Zhytomyr park. It creates under
the tree canopy almost one-species bush thickets which
are formed through vast growth, suppressing and
gradually displacing aboriginal plants of pine forests
(Peucedano-Pinetum) in Korostyshiv and Zhytomyr
(Zhyto myr Region), near Sarny town, Sarny State Fore-
stry, Kostyantynivske forestry unit (Rivne Region),
where the area of thicket of the species is almost 3.0 ha,
the height of undergrowth is 0.8-1.0, closeness – 0.6-1.0.
Also several separate thickets of this species were found
in Kiev Polissya – in Ivankiv distr. of Kiev Region. On
the left-bank Polissya, e.g. Sumy Region, the species is
characterized by sporadic distribution in some forest
localities where it was previously cultivated. Birds and
mammals are the main agents of distribution of plants.
The species grows in Lathyro nigri-Quercetum roboris,
Cladonio-Pinetum Juraszek 1927 and Peucedano-
Pinetum, and also in articial plantings on agricultural
sites.
The species Quercus rubra was cultivated in
Ukraine since 1970s, tested as tree species for planta-
tion growing in Zhytomyr region. The experimental
areas which were left behind became the center of the
species’ expansion on the surrounding forest territories.
Such situation is observed in Zhytomyr State Forest r y,
Boguns’ke forestry unit. It roots on lit areas and light
forests, forest clearings and also tight oak and pine-
oak forests, creating high-density thickets and, then,
spreading on the surrounding territory.
Unlike Quercus robur L., the abundant seed years
of which have been noted in Ukrainian Polissya once
every 8-10 years, Q. rubra is fruiting annually, forming
thick undergrowth with nearly 100% coverage keeping
herbaceous as well as tree species from growth. The
species is a component of Potentillo albae-Quercetum
Libbert 1933.
Our pilot research showed that Symphyotrichium
Nees genus has a great transformer potentials in the
region. Just like other species of this genus – Sym-
phyotrichium novаe-angliae and S. lanceolatum – was
extensively cultivated throughout the region, thus often
growing wild as well. The plant, with its long rhizomes,
actively changes vegetation, creating one-species thick
curtains, occupying large areas and threatening phyto-
diversity. Besides, it has a great seed productivity; seeds
are anemochorous and can propagate on signicant
distances. The species is common along river banks,
eutrophic bogs and different types of anthropogenic
ecotopes, participating in the following plant communi-
ties: Phragmito-Magnocaricetea, Artemisietea vulgaris
and Agropyretea repentis.
5. Conclusion
All species mentioned above are strong edicators
in the Ukrainian Polissya region that can signicantly
change important parameters of environment, species
composition of communities and character of landscape.
Majority of these species is also invasive.
The transfor mer species are rarely presented in plant
communities of Vaccinio-Piceetea Br.-Bl. 1939 class on
pure sandy soil in the Region. They are most common
in rich ecological conditions of deciduous and oak-pine
forests places.
The species Robinia pseudoacacia and Amel-Amel-
anchier spicata grow in zonal pine forests, Quercus
rubra – mainly in conifer-broadleaf forests, causing
changes in plant successions. Impatiens parviora
misplaces I. noli-tangere – the aboriginal species of
wet broadleaf forests. Most of the transformer spe-
cies – Echinocystis lobata, Impatiens glandulifera,
Heracleum sosnowskyi, H. mantegazzianum, Sym-
phyotrichium species grow in natural plant communi-
ties, including riverbank willow thickets, alder forests
and oodplain meadows, suppressing the aboriginal
species and preventing initial plant communities from
renovation. Reynoutria japonica forms monodominant
communities in parks and disturbed forests near places
of former cultivation.
Acknowledgements. We express our deep gratitude to Dr.
Yuryi O. Karpenko (Taras Shevchenko Chernigiv National
Pedagogical University) and Dr. Vitalji O. Volodymerets
(National University of Water Management and Natural
Resources Use, Rivne) for information about the distri-
bution of some transformer species in Chernigiv and Volyn
Polissya and Volhynia Upland, and Ms. Oksana O. Kucher
(M. G. Kholodny Institute of Botany, NAS of Ukraine) for
technical assistance.
The transformer species of the Ukrainian PolissyaVira V. Protopopova
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17
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