Поширення Trifolium rubens в Україні та сучасний стан популяцій у Закарпатській області
Trifolium rubens is a Central European forest-steppe plant species reaching its eastern distribution limit in Ukraine and listed in the Red Book of Ukraine (2009).Our study was carried out in 2016–2020 and aimed to analyze the distribution of T. rubens in Ukraine, to identify the composition of phyt...
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M.M. Gryshko National Botanical Garden of the NAS of Ukraine
2021
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Plant Introduction| _version_ | 1860145096351023104 |
|---|---|
| author | Gnatiuk, Alla Gaponenko, Mykola Loya, Vlasta Gaponenko, Andrew |
| author_facet | Gnatiuk, Alla Gaponenko, Mykola Loya, Vlasta Gaponenko, Andrew |
| author_sort | Gnatiuk, Alla |
| baseUrl_str | https://www.plantintroduction.org/index.php/pi/oai |
| collection | OJS |
| datestamp_date | 2023-08-26T20:39:20Z |
| description | Trifolium rubens is a Central European forest-steppe plant species reaching its eastern distribution limit in Ukraine and listed in the Red Book of Ukraine (2009).Our study was carried out in 2016–2020 and aimed to analyze the distribution of T. rubens in Ukraine, to identify the composition of phytocoenoses with T. rubens, and to evaluate the current state of its populations in the Zakarpattia region of Ukraine. The distribution of this species was analyzed based on data obtained from the field surveys, literature, online resources, and herbaria. In total, 78 localities of T. rubens were analyzed. Trifolium rubens was found to be distributed in Zakarpattia, Lviv, Ivano-Frankivsk, Ternopil, Rivne, Khmelnytskyi, Chernivtsi, and Vinnytsia regions of Ukraine. Two of these regions (Khmelnytskyi and Chernivtsi) were not mentioned before in the Red Book of Ukraine. The presence of T. rubens in Volyn, Zhytomyr, Cherkasy, and Kyiv regions is doubtful and not confirmed by recent data. Most localities of this species are known from Zakarpattia and Lviv regions.Three new localities were found in the Beregovo district of the Zakarpattia region in the foothills of the Volcanic Carpathians. The floristic compositions of the plant communities and ontogenetic structure of T. rubens populations in Beregovo and Mukachevo districts of the Zakarpattia region were investigated. Investigated populations are isolated with a dispersed disposition of plants on large areas. Trifolium rubens occurs here in meadow-steppe and meadow areas, on hillsides, and also participate in mesoxerophilous secondary (semi-natural) shrub communities in anthropogenically transformed areas. Often such populations are situated along pathways, on forest edges and fallow lands, and associated with ecotones of Trifolio-Geranietea sanguinei class. All studied populations were incomplete with the domination of generative shoots. |
| doi_str_mv | 10.46341/PI2020042 |
| first_indexed | 2025-07-17T12:53:48Z |
| format | Article |
| fulltext |
© The Authors. This content is provided under CC BY 4.0 license.
Plant Introduction, 89/90, 72–82 (2021)
RESEARCH ARTICLE
Distribution of Тrifolium rubens in Ukraine and the current state of its
populations in Zakarpattia
Alla Gnatiuk 1, *, Mykola Gaponenko 1, Vlasta Loya 1, Andrew Gaponenko 2
1 M.M. Gryshko National Botanical Garden, National Academy of Sciences of Ukraine, Tymiryazevska str. 1, 01014 Kyiv, Ukraine;
gnatiukalla@gmail.com
2 National University of Life and Environmental Sciences of Ukraine, Heroiv Oborony str. 15, build. 4, 03041 Kyiv, Ukraine
Received: 13.11.2020 | Accepted: 09.04.2021 | Published online: 28.04.2021
Abstract
Trifolium rubens is a Central European forest-steppe plant species reaching its eastern distribution limit in
Ukraine and listed in the Red Book of Ukraine (2009).
Our study was carried out in 2016–2020 and aimed to analyze the distribution of T. rubens in Ukraine, to
identify the composition of phytocoenoses with T. rubens, and to evaluate the current state of its populations
in the Zakarpattia region of Ukraine. The distribution of this species was analyzed based on data obtained
from the field surveys, literature, online resources, and herbaria. In total, 78 localities of T. rubens were
analyzed. Trifolium rubens was found to be distributed in Zakarpattia, Lviv, Ivano-Frankivsk, Ternopil,
Rivne, Khmelnytskyi, Chernivtsi, and Vinnytsia regions of Ukraine. Two of these regions (Khmelnytskyi and
Chernivtsi) were not mentioned before in the Red Book of Ukraine. The presence of T. rubens in Volyn,
Zhytomyr, Cherkasy, and Kyiv regions is doubtful and not confirmed by recent data. Most localities of this
species are known from Zakarpattia and Lviv regions.
Three new localities were found in the Beregovo district of the Zakarpattia region in the foothills of the
Volcanic Carpathians. The floristic compositions of the plant communities and ontogenetic structure of
T. rubens populations in Beregovo and Mukachevo districts of the Zakarpattia region were investigated.
Investigated populations are isolated with a dispersed disposition of plants on large areas. Trifolium rubens
occurs here in meadow-steppe and meadow areas, on hillsides, and also participate in mesoxerophilous
secondary (semi-natural) shrub communities in anthropogenically transformed areas. Often such
populations are situated along pathways, on forest edges and fallow lands, and associated with ecotones
of Trifolio-Geranietea sanguinei class. All studied populations were incomplete with the domination of
generative shoots.
Keywords: Trifolium rubens, chorology, populations, Ukraine
https://doi.org/10.46341/PI2020042
UDC 633.325:(477):(477.87)
Authors’ contributions: Gnatiuk A. and Gaponenko M. conceived the investigation and its main conceptual ideas, designed the study,
and analyzed the data. Loya V. and Gnatiuk A. collected the data from the herbaria, and made a map. Gnatiuk A., Gaponenko M.,
Loya V., and Gaponenko A. realized the field research, collected data, took photos, collected herbarium, identified plant samples,
composed species lists, and realized a final approval of the manuscript. Gnatiuk A., Gaponenko M., and Loya V. wrote the manuscript
аnd worked on its critical revisions.
Funding: The work has been conducted within the research program of the M.M. Gryshko National Botanical Garden № 387-РН
“Development of scientific bases for conservation, enrichment and efficient use of resources of valuable introducers and rare plants
under the influence of climate change in Ukraine” (state registration number 0117U000816) and at the authors’ own expense.
Competing Interests: The authors declared no conflict of interest.
https://creativecommons.org/licenses/by/4.0/
https://orcid.org/0000-0001-5001-971X
https://orcid.org/0000-0003-1513-1065
https://orcid.org/0000-0001-5450-9433
https://orcid.org/0000-0003-4215-2216
Plant Introduction • 89/90 73
Distribution of Тrifolium rubens in Ukraine and state of its populations in Zakarpattia
Introduction
Trifolium rubens L. is a Central European
forest-steppe plant species. It is also present
in Southern Europe, including southern Spain,
Italy, and Eastern Europe. Some species’
findings were reported from Belarus (Meusel
& Jäger, 1992). Also, a few localities are known
from the Baltic region (Rašomavičius, 2007)
and the south of Thrace (Meusel & Jäger, 1992).
In Ukraine, T. rubens is a rare species
(Didukh, 2009), reaching its eastern
distribution limits. It is mostly distributed
in the right-bank part of Ukraine, where it
occurs on dry meadows, meadow-steppes,
forest edges, and growths among shrubs.
Didukh (2009) indicated that T. rubens occurs
in Zakarpattia and Volyno-Podilia of Ukraine
only. However, the preliminary chorological
research showed that it probably has a broader
distribution range.
In Ukraine, Т. rubens mainly participates in
the plant communities Festuco-Brometea Br.-
Bl. et Tx. ex Soó 1947, Molinio-Arrhenatheretea
Tx. 1937, and Trifolio-Geranietea sanguinei
T. Müller 1961. It is also a diagnostic species
for association Primulo veris-Agrostietum
capillaris Uhliarová et Janišová 2014. Beside
this, T. rubens participates in the thermophilic
communities of ecotones, forest edges,
and shrubby association Antherico ramosi-
Geranietalia sanguinei Julve 1993 (Didukh,
2009; Dubyna et al., 2019).
In the adjacent countries, T. rubens is
associated with some more communities. For
example, in Poland it is listed for the order
Quercetalia pubescentis Br.-Bl. (1931) 1932
(Matuszkiewicz, 2001). Trifolium rubens is also
mentioned as an indicator of thermophilic
Trifolio-Geranietea communities in Slovakia
(Ružičková et al., 1996).
Usually, the populations of T. rubens
in Ukraine are small, with sporadically
distributed or joined in small groups (from a
few individuals to several dozens) plants. These
populations are mainly situated on the slopes
of the southern and south-eastern exposures
(Ralo, 2002; Onyshchenko & Andrienko
2012; Dmytrash, 2015; Dmytrash-Vatseba &
Shumska, 2016; Konishchuk et al., 2017).
This study was aimed to clarify the
distribution of T. rubens in Ukraine and
its participation in plant communities. It
also aimed to evaluate the current state of
T. rubens populations in the Zakarpattia region
of Ukraine.
Material and methods
The study was carried out in 2016–2020.
Trifolium rubens distribution analysis was
based on the data from the field surveys,
literature, Internet resources, and examination
of the herbarium vouchers from UU, KW,
LWS, КWHA, CHER, MW, MNR, MSUD, UPU,
SOF, UM, KWHU, KWU (see Thiers (2016) for
acronyms), and KW-museum (herbarium of
the National Museum of Natural History of the
NAS of Ukraine, Kyiv).
Populations were studied in the Zakarpattia
region in June (flowering period) and August
(fruiting period). In particular, the populations
in the low mountain range of the Beregovo
volcanic hills and on the mountain Lovachka
were investigated in the following localities:
1) Beregovo; 2) Zatyshne 3) Muzhievo, and
4) Mukachevo (Figs. 1 & 2).
Trifolium rubens is a herbaceous perennial
polycarpic. Shoot system оf T. rubens is
represented by elongated monocyclic
shoots. The degree of development and
number of shoots reflect individuals’ state
and prospects of seed reproduction in the
population. Establishing the real age structure
of T. rubens populations in natural habitats, as
in some other perennial species, is difficult.
It is impossible to do this without damaging
plants and soil surface, especially in dense
communities. Such damage is unacceptable for
studying rare plant species. In this case, a ratio
of the number of generative and vegetative
shoots may be a key for population monitoring
(Kagalo et al., 2012). Hence, in each locus, the
shoots of four age stages: virginal, generative,
subsenile, and senile (Gnatiuk et al., 2020),
were registered and investigated.
In each population, the vegetation
composition was evaluated following Braun-
Blaquet (1964). The scientific names of the
plants are provided according to POWO (2021),
the names of the vegetation communities
– according to Dubyna et al. (2019) аnd
Solomakha (2008).
Revised specimens
Vinnytsia region. Nemyriv district: Antsipolivka, 08.06.2009,
Vashenyak (KW), (Didukh et al., 2010).
74 Plant Introduction • 89/90
A. Gnatiuk, M. Gaponenko, V. Loya, A. Gaponenko
Volyn region. Volodymyr-Volynskyi (Visjulina, 1954).
Zakarpattia region. Beregovo district: Beregovo, 15.06.
1947, Bilyk (UU), (Visjulina, 1954); 16.06.1953, Tovt, (UU);
Beregovo, Zolota mountain, 16.06.1968, Chopyk (KW);
Muzhiyivsk, 11.06.1957, Uliganets (UU); Bene, Chillagash
Tetev mountain, 15.07.1960, Tovt (UU); Kuklabanya,
09.06.1948, Malynovskyi (LWS); Velykobereznyanskyi district:
Maly Berezny, 07.07.1956, Khymynets (UU); Uzhok, Kinchyk
mountain, 26.08.1955, Chopyk (KW); Maly Berezny,
Dilok mountain, 25.06.1954, Rudenko, (UU); Vynohradiv
district: Kholmovets, 12.06.1956, Chubirko (UU); Chopyk,
16.07.1963 (KWHA); Mukachevo district: Mukachevo, near
the monastery 11.06.1947, Grin (KW), (Visjulina, 1954);
Mukachevo vicinity, Lovachka mountain, (Margittai,
1911); 11.07.2018, Loya (KWHA); Mukachevo, Galish
mountain, 21.05.2020, Loya (KWHA); Perechyn district:
Plishka mountain, 04.08.1955 (Chopyk-KW); 09.07.1954,
Tovt (UU); 05.06.1956, Tovt (UU); 03.08.1958, Tovt (UU);
25.07.1958, Simchera (UU); Sinatoria, 03.07.1956, Chopyk
(KW); Rakhiv district: Dilove, Polonskaya, 03.07.1956, Tovt
(UU); Rakhiv vicinity, Lysa Gora mountain, 26.06.1995,
Kucheryava (KWU); Uzhhorod district: Kholmountainsi,
11.06.1947, Bilyk (KW); 06.03.1947, ? (UU), (Visjulina,
1954); 23.07.1958, Kovalenko (UU); 29.06.1956, Tovt
(UU); 28.07.1987, Tasenkevych (LWS); Antalovetska
Polyana mountain, 24.07.1958, Ilnytska (UU); Antalovtsi,
25.06.1955, Tovt (UU); Nevytsky castle, 24.07.1958,
Zotyk (UU); Onokivtsi, 01.07.1968, Bartok (UU); Hlyboke,
08.07.1960, Karaeva (MW).
Ivano-Frankivsk region. Halytskyi district: Bovshev, Kasova
Hora mountain, 13.7.1973, Kukovytsya; 49.226342,
24.696633, 23.06.2015, N. Sytschak (UkrBIN, 2021);
10.06.1969, Shelyag-Sosonko, Kukovytsya (KW), (Shumska
& Dmytrash, 2012); 08.06.2000, Kuzyarin (LWS); Tyaziv,
04.1960, Doroshenko (CHER); Rohatyn district: Velyki Goldy
(Luchyntsi) (Shumska & Dmytrash, 2012); Tysmenytsia
district: Uzin, urochysche Zhdymyr (Shumska & Dmytrash,
2012); Kolodiyivka, urochysche Pidgoroddya (Shumska
& Dmytrash, 2012); urochysche Simlin (Shumska &
Dmytrash, 2012); urochysche Hareva (Shumska &
Dmytrash, 2012); urochysche Gorozhanka (Shumska &
Dmytrash, 2012).
Lviv region. Lviv city, place Kryvchytsi (former village
of Kryvchytsi), 07.1856, Łobarzewski (LWS); Lviv city,
park Znesinnya, 49.847987, 24.076442, 20.07.2020,
Voityk (UkrBIN, 2021); north-eastern vicinity of Lviv
city, urochysche Khomets, 20.06.1986, Zelenchuk (LW);
Brody district: Buchyna, Makitra mountain, 13.07.1997,
Kuzyarin (LWS); 49.967495, 25.189118 (Batochenko,
2019); 50.035774 25.264379, 30.08.2019, Yurechko
(Yurechko, 2020; UkrBIN, 2021); Ponykovytsya, Makitra
mountain, 15.08.1954, Shelest (KWHA), 05.07.1987
(KW); Boratyn, Krugla mountain, 11.07.1987, Zelenchuk
(LW, МW); 50.002236, 25.147742, 30.07.2019, Yurechko
(Yurechko, 2020; UkrBIN, 2021); Vydra urochysche,
49.970372 25.180636, 04.08.2017, Yurechko (UkrBIN,
2021); Tetylkivtsi, 49.959404, 25.361032 (Batochenko,
2019); Verbivchyk, 49.893272, 25.283389 (Batochenko,
2019); Zolochiv district: Chervone, Lysa Gora mountain,
BA
DC
Figure 1. Тrifolium rubens in Beregovo (A), Zatyshne (B), Muzhievo (C), and Mukachevo (D) populations.
Plant Introduction • 89/90 75
Distribution of Тrifolium rubens in Ukraine and state of its populations in Zakarpattia
BA
DC
E
Figure 2. General distribution of Тrifolium rubens
in Ukraine (A) and its investigated populations
in Beregovo (B), Zatyshne (C), Muzhievo (D),
and Mukachevo (E). Numbers indicate the
investigated fragments (loci).
18.06.1956, Bukhalo (KW); 49.800963, 24.714529
(Batochenko, 2019); Tserkovna mountain, 07.07.1974,
Shelyag-Sosonko, Kukovytsya, Didukh (KW); Stinka,
49.771803, 24.738519 (Batochenko, 2019); Khmelevo
(Ralo, 2002); 19.07.1988, Iljinska (KW); Zolochiv, “nad
Kraglem” 12.07.1938, Mądalski (LWS); Derevyanky
urochysche, 49.824592, 25.062689, 10.07.2020, Yurechko
(UkrBIN, 2021); Luka urochysche, 49.794502, 25.01998,
27.09.2017, Yurechko (Ralo & Yurechko, 2019; UkrBIN,
2021); Zhovkva district: Mervychi, Kamin mountain,
30.06.1999, Kuzyarin (LWS); Zhydachiv district: Zhuravno,
1857, Łobarzewski (LWS); 29.07.1936, Mądalski (LWS);
Peremyshlyany district: Borschiv, 16.07.1933, ?, (LWS);
Pustomyty district: Derevach 06.1855; 1857, Łobarzewski
(LWS); Sokal district: ?, 1911, Szafer (LWS); Skomorokhy,
1869, Rehmann (LWS); Staryi Sambir district: Dobromyl,
Meznetska mountain, 27.05.1967, Boyko (LWS).
Rivne region. Radyvyl district: Druzhba, 1859; 1858,
Gomolinski (LWS); Berezne district: “Nadsluchanska
Shveytsariya”, the right bank of the river Sluch,
Andriyenko (KW). Mlyniv district: Vladyslavivka, urochysche
Hraboveshchyna (Didukh et al., 2010; Lohvynenko,
2012); Kurylykha, 50.562694, 25.7545 (Shtogryn et al.,
2020). Shumsk district: 50.142583, 26.159583; 50.126444,
26.066889 (Shtogryn et al., 2020).
Ternopil region. Berezhany district: Kuryany, Holytsya
mountain, 27.06.1976, Zaverukha (KW), (Yavorivskyi &
76 Plant Introduction • 89/90
A. Gnatiuk, M. Gaponenko, V. Loya, A. Gaponenko
Stakhurska, 2017); Kremenets district: Pidlisne, urochysche
Vakatsy, 20.07.2007, 18.06.2009, 24.07.2010, Hlinska
(KWHA), 30.06.2014, Krytska, Novosad, Shcherbakova
(KW-museum); Novosilky, 29.06.1958, Klokov (KW);
Raslavka, 49.958393, 25.465275 (Batochenko, 2019);
Maslyatyn mountain, 2006, Pronyuk, Oliyar (MNR);
50.137694, 26.092528 (Shtogryn et al., 2020); Pochayiv
[?] (Visjulina, 1954); Zboriv district: Ratyshchi, Ratyska
mountain, 1875, Schauer (LWS).
Khmelnytskyi region. Kamyanets-Podilskyi district:
urochysche Sovyi Yar, 06.07.1949, Kuznetsova (KW),
(Visjulina, 1954); Nizhyn, 13.07.1964, Krutskevych (KWHA);
Sataniv (Visjulina, 1954).
Chernivtsi region. Khotyn district: Polyana, 26.06.1958,
Kozak (CHER); Vyzhnytsa district: urochysche Velyki Luzhky,
15.07.1977, Shterma (CHER).
Results and discussion
Distribution of Trifolium rubens in Ukraine
Basing on the analysis of herbarium material
and published data, we have confirmed that
T. rubens is distributed only in the right-bank
part of Ukraine (Fig. 2 A). It is registered from
Vinnytsia, Zakarpattia, Ivano-Frankivsk, Lviv,
Rivne, Ternopil, Khmelnytsky, and Chernivtsi
regions. Although according to the Red
Book of Ukraine (Didukh, 2009), T. rubens
is not mentioned for the Khmelnytsky and
Chernivtsi regions. Most locations are known
from the Zakarpattia and Lviv regions. The
current presence of T. rubens in the Volyn,
Zhytomyr, Cherkasy, and Kyiv regions is
doubtful and/or has no recent confirmation.
For example, there is no recent confirmation
of the presence of T. rubens in the Kyiv
region. The herbarium voucher “Zhytomyr
region. Novograd-Volynskyi district, village
Mala Tsvilya, June 13, 2000, Orlov” deposited
at the KW was re-identified by Y.P. Didukh
as T. medium L., and we agree with that new
identification. The report of T. rubens from
Uman (Visyulina, 1954) was not confirmed too
– there are no herbarium vouchers from this
region yet.
Populations of Trifolium rubens in Zakarpattia
region
In the investigated areas, the fragments of oak
forests and shrubs occurred. The slopes were
partially terraced and planted with vineyards,
gardens or used as arable fields or pastures.
Population 1. The population of T. rubens in
Beregovo (Figs. 1 & 2) was confined to a plant
community with rich floristic composition
and mosaic structure. In this population,
five fragments (loci) were studied (Table 1).
The population was numerous, with about
300 plants. Monodominant fragments of
the community were formed mainly by
Саlamagrostis, Trifolium, Coronilla, and
Rubus species. Plants of T. rubens occurred on
the slopes of the western and north-western
exposures, and the relatively flat upper part
of the hills. Some individuals grew on the
northern slope among trees and shrubs, and
some – along the trail. The herb layer was 30–
60 cm high and has 80–90 % of the projective
cover. The shrub layer reaches up to 2 m high.
Sometimes Quercus petraea Liebl., Populus
tremula L., Castanea sativa Mill. overgrew
between shrubs of Cornus sanguinea L.,
Prunus spinosa L., Chamaecytisus supinus (L.)
Link, C. austriacus L., Rhamnus cathartica L.,
Rosa canina L. agg., and Rubus species
(Table 2).
Shoots were concentrated in the groups
(clumps) by 5–50. Each clump occupied an
area of about 0.5–2.0 m2. The average density
of shoots was 28.4 ± 1.3 per 1 m2. In particular,
5.2 ± 1.0 per 1 m2 – for virginal shoots, 21.2 ± 1.4
per 1 m2 – for generative, 0.4 ± 0.4 per 1 m2 –
for subsenile, and 3.6 ± 1.0 per 1 m2 – for senile.
Like all others, this population was incomplete.
Shoots in the generative state predominated
(Table 1). There were also some separate
generative plants at a distance 20–25 m from
each other. Neither seedlings nor juvenile
plants were found.
Plants were scattered irregularly on the
territory. Occasionally, T. rubens plants
co-dominated there together with Rubus
fruticosus L. agg., Coronilla varia L., Lotus
corniculatus L., and Clinopodium vulgare L.
According to the diagnostic species
(Agrimonia eupatoria L., Campanula
persicifolia L., Clinopodium vulgare, Geranium
sanguineum L., Melampyrum nemorosum L.,
Origanum vulgare L., Peucedanum cervaria
(L.) Lap., Trifolium medium, Vincetoxicum
hirundinaria Medik., and others – see Table 2),
communities with the participation of T. rubens
should be attributed to the order Origanetalia
Th. Müller 1962 class Trifolio-Geranietea
sanguinei Th. Müller 1961 (Solomakha, 2008).
Plant Introduction • 89/90 77
Distribution of Тrifolium rubens in Ukraine and state of its populations in Zakarpattia
Population
Number of shoots per 1 m2
Virginal Generative Subsenile Senile Total
1. Beregovo
locus 1 4 ± 0.5 17 ± 1.3 5 ± 1.2 0.0 26
locus 2 6 ± 0.9 10 ± 1.3 4 ± 1.1 2 ± 0.1 22
locus 3 10 ± 1.3 45 ± 1.5 3 ± 1.0 0.0 58
locus 4 4 ± 0.5 16 ± 1.3 0.0 0.0 20
locus 5 2 ± 0.1 8 ± 1.5 6 ± 1.0 0.0 16
Mean ± SD 5.2 ± 1.0 19.2 ± 1.4 3.6 ± 1.0 0.4 ± 0.4 28.4 ± 1.3
Total (%) 26 (18.3) 96 (67.6) 18 (12.7) 2 (1.4) 142 (100.0)
2. Zatyshne
locus 1 3 ± 0.5 9 ± 1.5 2 ± 0.1 0.0 14
locus 2 1 ± 0.1 26 ± 1.4 4 ± 0.6 0.0 31
locus 3 0.0 16 ± 0.5 1 ± 0.1 0.0 17
Mean ± SD 1.3 ± 0.3 17.0 ± 1.7 2.3 ± 0.4 0.0 20.6 ± 0.6
Total (%) 4 (6.3) 51 (82.5) 7 (11.2) 0 (0.0) 41 (100.0)
3. Muzhieve
locus 1 13 ± 0.0 75 ± 0.0 3 ± 0.0 0.0 91
locus 2 8 ± 0.0 62 ± 0.0 4 ± 0.0 0.0 74
Mean ± SD 10.5 ± 0.0 68.5 ± 0.0 3.5 ± 0.0 0.0 82.5 ± 0.0
Total (%) 21 (12.7) 137 (83.0) 7 (4.3) 0 (0.0) 165 (100.0)
4. Lovachka mountain
locus 1 13.1 ± 0.4 20 ± 1.1 0.0 0.0 33
locus 2 16.3 ± 1.3 11.1 ± 0.9 0.0 0.0 27
Mean ± SD 14.4 ± 0.9 15.6 ± 1.0 0.0 0.0 30.2 ± 0.0
Total (%) 29 (48.7) 31 (51.5) 0 (0.0) 0 (0.0) 60 (100.0)
Table 1. Characteristics of investigated Trifolium rubens populations.
Population 2. Near the village Zatyshne,
T. rubens occurred on gentle slopes of
southeastern and northern exposures
with an inclination of about 20–30 ° among
shrubs, along paths, and, sometimes, in
the 10–15-meter ecotone of the oak forest.
A rich floral composition characterized
this community due to the penetration of
species from adjacent plant communities
with the formation of mosaic complexes and
due to the lack of shading. The community
consisted of deciduous shrubs with significant
participation of xerothermic species. The
shrub layer was up to 2 m high. The herb layer’s
height was 30–60 cm, and its projective cover
reached 80–100 %. Calamagrostis epigejos
(L.) Roth, C. arundinacea (L.) Roth, Trifolium
rubens, T. medium, Melampyrum nemorosum,
Clinopodium vulgare, Inula salicina L., and
Vicia cracca L. predominated here (Table 2).
Plants of T. rubens were represented by
small dense clumps (1–3 m2). In each clump
were 2–8 generative individuals. An average
density of shoots was 31 per 1 m2. In this
population, only three loci were found with 41
shoots (Table 1).
Population 3. In T. rubens population near
the village Muzhievo, two clumps formed by
generative shoots were found. The population
situated in the upper part of the gentle slope of
the eastern exposure near thickets of shrubs
with Robinia pseudoacacia L., Rosa canina
agg., Rubus fruticosus agg., and Vitis vinifera L.
78 Plant Introduction • 89/90
A. Gnatiuk, M. Gaponenko, V. Loya, A. Gaponenko
Population 1 2 3 4
Tree layer
Castanea sativa Mill. + . . +
Populus tremula L. 1 + . .
Prunus avium (L.) L. . + . .
Prunus cerasus L. . + . .
Quercus petraea (Matt.) Liebl. + + . 1
Robinia pseudoacacia L. . . + +
Shrub layer
Cornus sanguinea L. 1 1 . .
Prunus spinosa L. + + . .
Rhamnus cathartica L. + . . .
Rosa canina s.l. 2 1 + .
Rubus fruticosus s.l. . 2 + .
Sambucus ebulus L. . 1 . .
Vítis vinifera L. . . + .
Herb layer
Achillea millefolium L. 1 1 . .
Agrimonia eupatoria L. + + . .
Ajuga reptans L. . . . +
Allium oleraceum L. 1 . . .
Allium scorodoprasum L. + + . .
Anthericum ramosum L. . . . +
Artemisia absinthium L. + . . +
Astragalus glycyphyllos L. 1 + . .
Berteroa incana (L.) DC. . + . .
Betonica officinalis L. . + . .
Calamagrostis arundinacea (L.) Roth 1 2 . .
Calamagrostis epigejos (L.) Roth 1 1 2 2
Campanula bononiensis L. + . . .
Campanula cervicaria L. . + . .
Campanula glomerata L. . + . .
Campanula persicifolia L. + . . .
Chamaecytisus austriacus (L.) Link + . . .
Chamaecytisus supinus (L.) Link + . . .
Clinopodium vulgare L. 2 1 . 1
Cota tinctoria (L.) J. Gay . . . +
Dactylis glomerata L. 1 1 . 1
Dianthus carthusianorum L. 1 + . .
Digitalis grandiflora Mill. + + . .
Population 1 2 3 4
Echium vulgare L. + . . .
Elymus repens (L.) Gould 1 1 1 .
Erigeron annuus (L.) Pers. 1 1 1 +
Euphorbia cyparissias L. . . . +
Festuca ovina L. + . . .
Festuca pratensis Huds . . + .
Filago arvensis L. + . . .
Fragaria vesca L. + . . .
Galatella linosyris (L.) Rchb.f. . + . .
Galium mollugo L. + + . .
Galium verum L. + . . +
Genista tinctoria L. + + . +
Geranium sanguineum L. + . . .
Geum urbanum L. + + . .
Hypericum perforatum L + + . .
Inula acuminate DC. + + . .
Lamium galeobdolon (L.) L. + . . .
Lathyrus niger (L.) Bernh. . . . 1
Lathyrus pratensis L. 1 + . .
Lathyrus sylvestris L. + . . .
Lathyrus tuberosus L. + . . .
Leucanthemum vulgare Lam. . 1 . .
Lotus corniculatus L. 2 2 1 .
Medicago falcata L. . . . .
Medicago lupulina L. . 1 . .
Melampyrum nemorosum L. + 1 . .
Melica transsilvanica Schur + . . .
Melilotus albus Medik. + . . .
Melilotus officinalis (L.) Lam. . + . .
Origanum vulgare L. + 1 . +
Pentanema germanicum (L.) D. Gut.
Larr. et al.
+ . . +
Pentanema hirtum (L.) D. Gut. Larr. et al. + . . .
Peucedanum cervaria (L.) Lap. 1 . . .
Plantago media L. . 1 . .
Potentilla inclinata Vill. + . . .
Potentilla recta L. . 1 . .
Ranunculus acris L + . . .
Rumex confertus Willd. . 1 1 .
Table 2. Floristic composition of the plant communities in studied populations of Trifolium rubens.
Plant Introduction • 89/90 79
Distribution of Тrifolium rubens in Ukraine and state of its populations in Zakarpattia
Population 1 2 3 4
Securigera varia (L.) Lassen 2 2 . .
Sedum maximum (L.) Suter + . . .
Silene latifolia Poir. + . . .
Stachys annua (L.) L. + 1 . .
Symphytum officinale L. . + . .
Tanacetum corymbosum (L.) Sch. Bip. + 1 . .
Tanacetum vulgare L. + 1 . .
Teucrium chamaedrys L. . + . .
Torilis japonica (Нoutt.) DC. + . . .
Trifolium arvense L. + 1 . .
Trifolium dubium Sibth. 1 . . .
Trifolium medium L. 1 2 . +
Trifolium montanum L. + 1 . +
Trifolium pannonicum Jacq. . + . .
Trifolium pratense L. + . . .
Trifolium repens L. . 1 . .
Trifolium rubens L. 1 1 1 1
Verbascum lychnitis L. 1 . . .
Veronica spicata L. + 1 . .
Vicia cracca L. 2 2 1 .
Vicia lathyroides L. + . . .
Vicia villosa Roth . + 1 .
Vincetoxicum hirundinaria Medik. . . . +
Viola arvensis Murr. + . . .
Viscaria vulgaris Röhl. 1 . . .
Table 2. Continued.
participation (Table 2). Lotus corniculatus,
Calamagrostis epigeos, and Erigeron annuus
(L.) Pers were co-dominants of the herb layer
together with T. rubens. In this population, the
smallest number of clumps and the highest
density of shoots were registered. The clumps
area did not exceed 1 m2.
Population 4. The population from the top of
Lovachka mountain is located along the trail at
the edge of the oak forest at the 295 m a.s.l. on
a slope with an inclination of about 30 °. The
tree layer formed by Quercus petraea (Matt.)
Liebl., Robinia pseudoacacia, and Castanea
sativa. Calamagrostis epigejos, Dactylis L.,
Trifolium rubens, Clinopodium vulgare, and
Lathyrus niger (L.) Bernh dominated in the
herb layer, the projective cover of which
reached about 50%. Two loci of this population
were studied. Only generative and virginal
shoots were found in clumps. Totally, 145
shoots were recorded in this population. The
average density was 28.4 ± 5.0 shoots per 1 m2;
in particular, 27 ± 3.6 vegetative and 26.8 ± 2.9
generative shoots per 1 m2. Neither senile nor
subsenile shoots were detected.
Hence, the density of virginal shoots in the
studied populations varied from 1 to 8 per 1 m².
The most abundant were generative shoots
(17–68 per 1 m2 on average). The number of
subsenile shoots was 0–3 per 1 m2, and senile
– 0–2 per 1 m2 (Fig. 3). The total number of
shoots in the studied loci ranged from 41 to 165
per 1 m2.
Conclusions
Trifolium rubens is distributed only in the
right-bank part of Ukraine. There are 78
confirmed localities of this species in Ukraine.
The presence of T. rubens in Volyn, Zhytomyr,
Cherkasy and Kyiv regions is doubtful and has
no recent confirmations.
Studied populations in Zakarpattia region
were associated with meadow-steppe areas
with the participation of shrubs. They were
often located on slopes and associated with
xerophile semi-natural shrub communities
in areas with anthropogenic impact. They
were often situated along pathways, on forest
Figure 3. The ontogenetic spectrum of Trifolium
rubens populations. 1–4 –populations’ numbers;
v – virginal, g – generative, ss – subsenile, and
s – senile shoots.
80 Plant Introduction • 89/90
A. Gnatiuk, M. Gaponenko, V. Loya, A. Gaponenko
edges, and associated with ecotones between
forest or shrubs and herb communities from
the class Trifolio-Geranietea sanguinei. In
studied localities, T. rubens participated in
communities with Rubus fruticosus agg.,
Сoronilla varia, Lotus corniculatus,
Clinopodium vulgare, Trifolium medium,
Melampуrum nemorosum, Inula salicіna, Vicia
cracca, and Сalamagrostis species.
Apparently, metapopulational structure
with the dispersed disposition of plants
on the large areas is characteristic for the
T. rubens populations. The distances between
loci in populations varied from 10 to 100 m.
All studied populations were incomplete
with the advantage of generative shoots
(17–68 shoots per 1 m2 or 51.7–82.5 % from
the total number of shoots). The absence of
senile shoots evidences normal populations’
conditions.
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Plant Introduction • 89/90 81
Distribution of Тrifolium rubens in Ukraine and state of its populations in Zakarpattia
Поширення Trifolium rubens в Україні та сучасний стан популяцій у Закарпатській
області
Алла Гнатюк 1, *, Микола Гапоненко 1, Власта Лоя 1, Андрій Гапоненко 2
1 Національний ботанічний сад імені М.М. Гришка НАН України, вул. Тимірязєвська, 1, Київ, 01014,
Україна; gnatiukalla@gmail.com
2 Національний університет біоресурсів і природокористування України, вул. Героїв оборони, 15,
корпус 4, Київ, 03041, Україна
Тrifolium rubens – це центральноєвропейський лісостеповий вид рослин, який в Україні знаходиться
на східній межі поширення і включений до Червоної книги України (2009).
Дослідження проводили протягом 2016–2020 років. Метою дослідження було проаналізувати
поширення цього виду в Україні, описати умови місцезростань та оцінити сучасний стан його
популяцій у Закарпатській області. Для вивчення поширення виду було використано результати
польових досліджень, дані з літературних та онлайн джерел, а також відомості з гербаріїв. Загалом
було проаналізовано 78 місцезнаходжень T. rubens в Україні. В результаті чого було встановлено, що
T. rubens трапляється в Закарпатській, Львівській, Івано-Франківській, Тернопільській, Хмельницькій,
Рівненській, Чернівецькій та Вінницькій областях. Дві з цих областей (Хмельницька і Чернівецька) не
наводилися раніше у Червоній книзі України. Присутність природних популяцій T. rubens у Волинській,
Житомирській, Черкаській та Київській областях є сумнівним і не має сучасного підтвердження.
Найбільше місцезнаходжень цього виду відомо з Закарпатської та Львівської областей.
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82 Plant Introduction • 89/90
A. Gnatiuk, M. Gaponenko, V. Loya, A. Gaponenko
Виявлено три нові місцезнаходження Т. rubens у Вулканічних Карпатах та їхньому передгір’ї.
Зокрема, описано флористичний склад фітоценозів та оцінено онтогенетичний стан популяцій
у Берегівському та Мукачівському районах Закарпатської області. Ці популяції ізольовані з
дисперсним розташуванням особин на великих площах. Trifolium rubens трапляється тут на лучно-
степових і лучних ділянках, на схилах пагорбів, а також у складі мезоксерофільних вторинних
(напівприродних) чагарникових угрупованнь на антропогенно трансформованих ділянках. Часто
популяції цього виду розташовані вздовж стежок, на узліссях, перелогах у формаціях класу Trifolio-
Geranietea sanguinei. Усі досліджені популяції неповностанові з переважанням генеративних пагонів.
Ключові слова: Тrifolium rubens, хорологія, популяції, Україна
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| id | oai:ojs2.plantintroduction.org:article-1574 |
| institution | Plant Introduction |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2025-07-17T12:53:48Z |
| publishDate | 2021 |
| publisher | M.M. Gryshko National Botanical Garden of the NAS of Ukraine |
| record_format | ojs |
| resource_txt_mv | wwwplantintroductionorg/c8/92a752b28f50bd21e1f011f22276e6c8.pdf |
| spelling | oai:ojs2.plantintroduction.org:article-15742023-08-26T20:39:20Z Distribution of Тrifolium rubens in Ukraine and the current state of its populations in Zakarpattia Поширення Trifolium rubens в Україні та сучасний стан популяцій у Закарпатській області Gnatiuk, Alla Gaponenko, Mykola Loya, Vlasta Gaponenko, Andrew Trifolium rubens is a Central European forest-steppe plant species reaching its eastern distribution limit in Ukraine and listed in the Red Book of Ukraine (2009).Our study was carried out in 2016–2020 and aimed to analyze the distribution of T. rubens in Ukraine, to identify the composition of phytocoenoses with T. rubens, and to evaluate the current state of its populations in the Zakarpattia region of Ukraine. The distribution of this species was analyzed based on data obtained from the field surveys, literature, online resources, and herbaria. In total, 78 localities of T. rubens were analyzed. Trifolium rubens was found to be distributed in Zakarpattia, Lviv, Ivano-Frankivsk, Ternopil, Rivne, Khmelnytskyi, Chernivtsi, and Vinnytsia regions of Ukraine. Two of these regions (Khmelnytskyi and Chernivtsi) were not mentioned before in the Red Book of Ukraine. The presence of T. rubens in Volyn, Zhytomyr, Cherkasy, and Kyiv regions is doubtful and not confirmed by recent data. Most localities of this species are known from Zakarpattia and Lviv regions.Three new localities were found in the Beregovo district of the Zakarpattia region in the foothills of the Volcanic Carpathians. The floristic compositions of the plant communities and ontogenetic structure of T. rubens populations in Beregovo and Mukachevo districts of the Zakarpattia region were investigated. Investigated populations are isolated with a dispersed disposition of plants on large areas. Trifolium rubens occurs here in meadow-steppe and meadow areas, on hillsides, and also participate in mesoxerophilous secondary (semi-natural) shrub communities in anthropogenically transformed areas. Often such populations are situated along pathways, on forest edges and fallow lands, and associated with ecotones of Trifolio-Geranietea sanguinei class. All studied populations were incomplete with the domination of generative shoots. Тrifolium rubens – це центральноєвропейський лісостеповий вид рослин, який в Україні знаходиться на східній межі поширення і включений до Червоної книги України (2009).Дослідження проводили протягом 2016–2020 років. Метою дослідження було проаналізувати поширення цього виду в Україні, описати умови місцезростань та оцінити сучасний стан його популяцій у Закарпатській області. Для вивчення поширення виду було використано результати польових досліджень, дані з літературних та онлайн джерел, а також відомості з гербаріїв. Загалом було проаналізовано 78 місцезнаходжень T. rubens в Україні. В результаті чого було встановлено, що T. rubens трапляється в Закарпатській, Львівській, Івано-Франківській, Тернопільській, Хмельницькій, Рівненській, Чернівецькій та Вінницькій областях. Дві з цих областей (Хмельницька і Чернівецька) не наводилися раніше у Червоній книзі України. Присутність природних популяцій T. rubens у Волинській, Житомирській, Черкаській та Київській областях є сумнівним і не має сучасного підтвердження. Найбільше місцезнаходжень цього виду відомо з Закарпатської та Львівської областей.Виявлено три нові місцезнаходження Т. rubens у Вулканічних Карпатах та їхньому передгір’ї. Зокрема, описано флористичний склад фітоценозів та оцінено онтогенетичний стан популяцій у Берегівському та Мукачівському районах Закарпатської області. Ці популяції ізольовані з дисперсним розташуванням особин на великих площах. Trifolium rubens трапляється тут на лучно-степових і лучних ділянках, на схилах пагорбів, а також у складі мезоксерофільних вторинних (напівприродних) чагарникових угрупованнь на антропогенно трансформованих ділянках. Часто популяції цього виду розташовані вздовж стежок, на узліссях, перелогах у формаціях класу Trifolio-Geranietea sanguinei. Усі досліджені популяції неповностанові з переважанням генеративних пагонів. M.M. Gryshko National Botanical Garden of the NAS of Ukraine 2021-04-28 Article Article application/pdf https://www.plantintroduction.org/index.php/pi/article/view/1574 10.46341/PI2020042 Plant Introduction; No 89/90 (2021); 72-82 Інтродукція Рослин; № 89/90 (2021); 72-82 2663-290X 1605-6574 10.46341/PI89-90 en https://www.plantintroduction.org/index.php/pi/article/view/1574/1511 Copyright (c) 2021 Alla Gnatiuk, Mykola Gaponenko, Vlasta Loya, Andrew Gaponenko http://creativecommons.org/licenses/by/4.0 |
| spellingShingle | Gnatiuk, Alla Gaponenko, Mykola Loya, Vlasta Gaponenko, Andrew Поширення Trifolium rubens в Україні та сучасний стан популяцій у Закарпатській області |
| title | Поширення Trifolium rubens в Україні та сучасний стан популяцій у Закарпатській області |
| title_alt | Distribution of Тrifolium rubens in Ukraine and the current state of its populations in Zakarpattia |
| title_full | Поширення Trifolium rubens в Україні та сучасний стан популяцій у Закарпатській області |
| title_fullStr | Поширення Trifolium rubens в Україні та сучасний стан популяцій у Закарпатській області |
| title_full_unstemmed | Поширення Trifolium rubens в Україні та сучасний стан популяцій у Закарпатській області |
| title_short | Поширення Trifolium rubens в Україні та сучасний стан популяцій у Закарпатській області |
| title_sort | поширення trifolium rubens в україні та сучасний стан популяцій у закарпатській області |
| url | https://www.plantintroduction.org/index.php/pi/article/view/1574 |
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