Сучасний стан інтродукційної ценопопуляції Gymnospermium odessanum на ботаніко-географічній ділянці “Степи України” Національного ботанічного саду імені М. М. Гришка НАН України
Gymnospermium odessanum is a rare relict endemic plant species. The research was carried out during G. odessanum flowering in 2019–2021 at the botanical-geographical plot “Steppes of Ukraine” of the M.M. Gryshko National Botanical Garden, National Academy of Sciences of Ukr...
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M.M. Gryshko National Botanical Garden of the NAS of Ukraine
2022
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Plant Introduction| _version_ | 1860145118816763904 |
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| author | Gritsenko, Victoria |
| author_facet | Gritsenko, Victoria |
| author_sort | Gritsenko, Victoria |
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| description | Gymnospermium odessanum is a rare relict endemic plant species. The research was carried out during G. odessanum flowering in 2019–2021 at the botanical-geographical plot “Steppes of Ukraine” of the M.M. Gryshko National Botanical Garden, National Academy of Sciences of Ukraine (NBG). Ecological and coenotic conditions of introduction at the NBG differ from natural habitats of the species and are not optimal for its vegetation. However, this species demonstrated ecological-coenotic plasticity and, over the decades, has formed a stable homeostatic introduction coenopopulation here. As of 2021, the area of introduction coenopopulation of G. odessanum at the NBG was 2,075 m2. It comprises 412 individuals of this species (including 40 seedlings, 241 juvenile, 45 immature, 40 virginal, 45 generative, and 1 sub-senile plants). Coenopopulation fragments with a high density of G. odessanum individuals are rare here, so the average density is low – only 0.2 individuals per 1 m2. In 2021, in the spectrum of age states, the total percentage of pregenerative individuals was very high and reached 88.8 %; the share of generative individuals was 10.9 %.In general, the age structure of the introduction coenopopulation of G. odessanum is characterized by long-term (2010–2021) stability. The spatial distribution of individuals in the introduction coenopopulation is of two kinds – random and in groups. This is due to combined myrmecochoric and barochoric propagation. Also due to myrmecochory, this coenopopulation tends to spread the area. Compared to natural coenopopulations, the introduction coenopopulation of G. odessanum at the NBG is characterized by a larger area, a much significant number, and, at the same time, a low average density of individuals. However, like in most of natural populations, its age spectrum is left-sided.The conducted research testifies the successful formation of the introduction coenopopulation of G. odessanum in the meadow-steppe cultural phytocoenosis of the NBG. This introduction coenopopulation is an example of a successful multi-year scientific experiment and effective ex situ protection and preservation of G. odessanum on the northern border of the Right Bank Forest-Steppe of Ukraine, far beyond the natural range of this endemic plant species. |
| doi_str_mv | 10.46341/PI2021017 |
| first_indexed | 2025-07-17T12:54:01Z |
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| fulltext |
Plant Introduction, 93/94, 3–17 (2022)
© The Authors. This content is provided under CC BY 4.0 license.
RESEARCH ARTICLE
The current state of the introduction coenopopulation of Gymnospermium
odessanum on the botanical-geographical plot “Steppes of Ukraine” at
the M. M. Gryshko National Botanical Garden of the NAS of Ukraine
Victoria Gritsenko
M.M. Gryshko National Botanical Garden, National Academy of Sciences of Ukraine, Tymiryazevska str. 1, 01014 Kyiv, Ukraine;
gritsenkoviktoria@gmail.com
Received: 06.12.2021 | Accepted: 23.12.2021 | Published online: 14.01.2022
Abstract
Gymnospermium odessanum is a rare relict endemic plant species. The research was carried out during
G. odessanum flowering in 2019–2021 at the botanical-geographical plot “Steppes of Ukraine” of the
M.M. Gryshko National Botanical Garden, National Academy of Sciences of Ukraine (NBG). Ecological and
coenotic conditions of introduction at the NBG differ from natural habitats of the species and are not
optimal for its vegetation. However, this species demonstrated ecological-coenotic plasticity and, over
the decades, has formed a stable homeostatic introduction coenopopulation here. As of 2021, the area
of introduction coenopopulation of G. odessanum at the NBG was 2,075 m2. It comprises 412 individuals
of this species (including 40 seedlings, 241 juvenile, 45 immature, 40 virginal, 45 generative, and 1 sub-
senile plants). Coenopopulation fragments with a high density of G. odessanum individuals are rare here,
so the average density is low – only 0.2 individuals per 1 m2. In 2021, in the spectrum of age states, the
total percentage of pregenerative individuals was very high and reached 88.8 %; the share of generative
individuals was 10.9 %.
In general, the age structure of the introduction coenopopulation of G. odessanum is characterized by
long-term (2010–2021) stability. The spatial distribution of individuals in the introduction coenopopulation
is of two kinds – random and in groups. This is due to combined myrmecochoric and barochoric
propagation. Also due to myrmecochory, this coenopopulation tends to spread the area. Compared to
natural coenopopulations, the introduction coenopopulation of G. odessanum at the NBG is characterized
by a larger area, a much significant number, and, at the same time, a low average density of individuals.
However, like in most of natural populations, its age spectrum is left-sided.
The conducted research testifies the successful formation of the introduction coenopopulation of
G. odessanum in the meadow-steppe cultural phytocoenosis of the NBG. This introduction coenopopulation
is an example of a successful multi-year scientific experiment and effective ex situ protection and
preservation of G. odessanum on the northern border of the Right Bank Forest-Steppe of Ukraine, far
beyond the natural range of this endemic plant species.
Keywords: Gymnospermium odessanum, rare endemic species, introduction coenopopulation, stable age structure, ex situ protection
https://doi.org/10.46341/PI2021017
UDC 58.084 : 582.675.3 (477-25)
Funding: The work has been realized within the following research program of the Department of Natural Flora Department of
the M.M. Gryshko National Botanical Garden, National Academy of Sciences of Ukraine 2020–2024 “Botanical and geographical
principles of protection of floristic diversity and the formation of the introduction populations of plants” (state registration number
0120U000174).
Competing Interests: The author declares no conflict of interest.
https://creativecommons.org/licenses/by/4.0/
https://orcid.org/0000-0002-1783-6977
4 Plant Introduction • 93/94
V. Gritsenko
Introduction
Gymnospermium Spach (Berberidaceae
Juss.) is a small genus (7–12 species) of early
flowering tuberous herbs, distributed from
east China to the Balkans (Rosati et al., 2018).
Gymnospermium odessanum (DC.) Takht.
is a relict endemic species with a narrow
disjunctive distribution range (Scherbakova
& Novosad, 2018). It is listed as a vulnerable
species in the Red Book of Ukraine (Kritska &
Novosad, 2009) and also protected by the Red
Book of the Republic of Moldova (Kolomiychuk
& Popova, 2018). This species is distributed
in Dobrogea region of Moldova and North-
Western part of the Black Sea region of Ukraine
(Kritska & Novosad, 2009; Scherbakova &
Novosad, 2018). It is also reported as a phyto-
rarity for the Steppe zone of Ukraine (Sobko et
al., 2013).
As a result of phylogenetic analyzes, it
was found that G. odessanum is sister to
G. peloponnesiacum (Phitos) Strid from South
Greece. Both species are, in turn, sister to
G. altaicum (Pallas) Spach (Barina et al., 2017;
Rosati et al., 2018). However, despite small
number of representatives, the taxonomy
of the genus is not fully clear and well-
established yet. In particularly, recently Rosati
et al. (2018) discribed a new subspecies of the
genus (G. scipetarum Paparisto & Qosja ex
E. Mayer & Pulevic subsp. eddae Rosati, Farris,
Fascetti & Selvi) for the flora of Italy.
In Ukraine, G. odessanum has been
extensively investigated in the nature
(Visjulina, 1953; Litvinenko, 1986, 2010;
Melnyk, 1994, 2000; Scherbakova, 2005,
2008, 2010, 2011; Kritska & Novosad, 2009;
Litvinenko & Shcherbakova, 2009; Kritska,
2012; Scherbakova & Barmak, 2013; Smetana
et al., 2014; Trotner, 2016, 2017). This species
has also been studied in Dobrogea (Doroftei
& Mierla, 2007) and Moldova, in general
(Izverskaya et al., 2013, 2017). In particular,
is interesting that Izverskaya et al. (2017)
predicted displacement of G. odessanum in
the Dniester river basin in Moldova to the
north due to climate change.
Scherbakova & Novosad (2018)
comprehensively investigated the distribution,
ecological and coenotic peculiarities, and
some aspects of G. odessanum protection.
After that, additional information about the
distribution and ecology of G. odessanum in
Ukraine was provided by Krasova et al. (2019),
Ponomareva et al. (2019), and Shiryaeva et al.
(2019, 2020).
Krasova et al. (2016) introduced
G. odessanum on the dump of the mining
quarry to restore phytodiversity in the
anthropogenic landscapes of Kryvbas, which
were decommissioned. Pavlenko et al. (2020),
during a survey of post-mining quarry and
dump complexes in the area of Kryvyi Rih,
found that G. odessanum is naturalized here as
a result of reclamation.
Gymnospermium odessanum is an
ornamental spring ephemeroid growing and
ex situ protected in several botanical gardens
of Ukraine (Mashkovska, 2015; Scherbakova
& Novosad, 2018). Namely, it is cultivated
at the M.M. Gryshko National Botanical
Garden, National Academy of Sciences of
Ukraine (NBG); Donetsk Botanical Garden,
National Academy of Sciences of Ukraine
(Krokhmal & Netsvetov, 2015); Botanical
Garden of Kherson State University;
Botanical Garden of V.N. Karazin National
University of Kharkiv; Botanical Garden of
I.I. Mechnikov National University of Odesa
(Yermolaieva, 2010); Botanical Garden of Ivan
Franko National University of Lviv; Kryvyi
Rih Botanical Garden, National Academy of
Sciences of Ukraine (Trotner, 2016; Krasova
et al., 2019); O.V. Fomin Botanical Garden of
Taras Shevchenko National University of Kyiv
(Peregrym et al., 2014).
In the Department of Natural Flora of
the NBG, G. odessanum was introduced
and partially studied on the botanical-
geographical plots “Steppes of Ukraine”
(Borodina, 1972, 1976) and “Forests of the
plain part of Ukraine” (Melnyk, 1994),
and the collection plot “Rare plants of
the Ukrainian flora” (Sobko & Borodina,
1987; Sobko & Gaponenko, 1996). It was
found that G. odessanum formed a stable
homeostatic introduction coenopopulation
on the botanical-geographical plot “Steppes
of Ukraine” of the NBG (Gritsenko, 2009).
Previously, only brief information on this
introduction coenopopulation was known
(Gritsenko, 2010, 2012, 2014, 2017a, 2019a, b;
Gritsenko et al., 2017). Thus, there was a need
for a more detailed and in-depth study of this
introduction coenopopulation.
In 2021, the phenology of G. odessanum was
studied on the botanical-geographical plot
Plant Introduction • 93/94 5
The current state of the introduction coenopopulation of Gymnospermium odessanum
“Steppes of Ukraine” of the NBG (Gritsenko,
2021). Gymnospermium odessanum grows here
outside its natural range (far to the north). The
introduction conditions at the NBG, which is
located on the northern border of the Right
Bank Forest-Steppe of Ukraine, significantly
differ from those in the natural habitats of
the species in the North-Western Black Sea
region.
There are two specific directions of plant
introduction – within and outside the natural
range, both of which are strongly argued
(Gaponenko & Gnatiuk, 2016) and can be
successfully realized. In recent years, in
both of dirrections, the experimental studies
of introduction populations of different
rare plant species were conducted at the
Department of Natural Flora of the NBG
(Melnyk et al., 2018; Gritsenko, 2020). In
particular, it was found that the introduction
coenopopulation of the rare early-spring
ephemeroid Crocus reticulatus Steven ex
Adam at the NBG (which is, by the way,
located within the species’ natural range) was
in the phase of logistic growth in 2020. This
confirmed the success of the introductory
experiment (Gritsenko, 2020).
Hence, it was worthing to find out whether
the introduction of G. odessanum has been
also fruitful. We suggested this an example of
successful multi-decade scientific experiment,
and effective ex situ protection and
preservation of G. odessanum on the northern
border of the Right-Bank Forest-Steppe of
Ukraine.
Material and methods
The studies were conducted during the time
of G. odessanum flowering in 2019–2021 at
the botanical-geographical plot “Steppes
of Ukraine” of the NBG. Gymnospermium
odessanum was introduced here from the
natural habitats of Odessa surroundings in
1953 and, repeatedly, in 1964 (Borodina, 1972).
These initial individuals served as a base to
form the introduction coenopopulation of
G. odessanum. Additional solitary individuals
were collected in 2002 and 2010 (Gritsenko
et al., 2017) and planted in the nursery – they
did not participate in the formation of the
main introduction coenopopulation of the
species.
In this article, for coenopopulation
description, author applies the term
‘introduction’ (‘інтродукційна’ in Ukrainian)
instead of ‘introduced’ (‘інтродукована’ in
Ukrainian), because it reflects the semantic
meaning much more correctly.
The area of the introduction
coenopopulation of G. odessanum was
calculated with a precision of 1 m2. The area’s
configuration was plotted on the satellite
images of Google Maps (2021). According to
standard methodology, phenomenological
observations of G. odessanum were performed
daily during its mass flowering (Beideman,
1974; Zaitsev, 1978). The plants were not
excavated or damaged during the research;
only aboveground parts were considered. The
age states of G. odessanum were determined
according to Litvinenko (2010). In general,
the number of generative individuals (total
amount of young, middle-aged, and old
generative plants) was counted. The study of
the coenopopulation structure was conducted
according to Uranov (1976). The average
density of individuals per 1 m2 was determined
as the individuals’ number divided by the total
area. When mapping the spatial location of
generative individuals, the total area of the
coenopopulation was subdivided into smaller
5 × 5 m plots, using cords and measuring
tape. Deviations between two numerical
data were calculated by the formulae:
Dev. = 100 – y × 100 / x, where ‘x’ is the first
datum, and ‘y’ is the second one.
Plant names were verified and provided
according to Euro+Med PlantBase (2021)
nomenclature. Species were determined
following Prokudin (1987). All photos were
captured by the author using a Canon Power
Shot SD 4000 IS digital ELPH camera.
Results and discussion
Area of the introduction coenopopulation
From a landscape perspective, the botanical-
geographical plot “Steppes of Ukraine” is an
extensive glade covering 2.5 hectares and
surrounded by forest vegetation (Fig. 1 A). The
introduction coenopopulation of G. odessanum
is located in the western part of this plot on
the open area with a flat relief (Fig. 1 A–C).
The coordinates inside of G. odessanum
coenopopulation are following – N 50.412000 °,
6 Plant Introduction • 93/94
V. Gritsenko
E 30.565062 °. The coenopopulation has
polygonal (decagonal) outlines and area of
2075 m2 (over 0.2 ha), bounded by earth roads
(Fig. 1 A).
Scherbakova & Novosad (2018) generally
characterized contemporary natural
populations of G. odessanum as local and having
small areas. Smetana et al. (2014) indicated
that 27 natural populations of G. odessanum
in the Ingulets river basin occupy areas from
0.5 m2 to 500 m2 and are confined to the
slopes of the predominantly eastern exposure
with 5–30 ° inclination. Thus, the introduction
coenopopulation of G. odessanum at the NBG,
compared to natural ones, is characterized by
significantly larger area and is scattered on the
flat relief.
Coenotic features
Gymnospermium odessanum grows on the
botanical-geographical plot “Steppes of
Ukraine” in the composition of artificially
created meadow-steppe phytocoenosis
(Gritsenko, 2017b). During the flowering time
of G. odessanum (March – beginning of April)
the soil surface is covered with a loose litter
from the remnants of various plants of the
previous year (Fig. 1 B). The projective cover of
the grass stand reaches 60 % (Fig. 1 C). In some
places, the projective cover of G. odessanum
can be up to 10 %. The height of G. odessanum
flowering shoots above the soil surface is
5–12 cm (Fig. 1 D).
In the natural coenopopulations of
G. odessanum, the projective grass cover varies
B
D
A
C
Figure 1. The introduction coenopopulation of Gymnospermium odessanum on the botanical-geographical
plot “Steppes of Ukraine” at the M. M. Gryshko National Botanical Garden of the NAS of Ukraine: A – location
and configuration (source: Google Maps; the yellow line indicates the boundaries of the coenopopulation;
the red point indicates the coordinates inside of the coenopopulation); B – the fragment (the red point
corresponds to A); C – the projective cover of the grass during the flowering time of G. odessanum and
the beginning of vegetation of cereals; D – the fragment of spring synusia with the participation of
G. odessanum (1), Colchicum bulbocodium subsp. versicolor (2) and Corydalis solida (3). Scales: A–B – 50 m;
C – 0.5 m; D – 0.1 m.
Plant Introduction • 93/94 7
The current state of the introduction coenopopulation of Gymnospermium odessanum
Nr Plant species Group Phenophase
1 Achillea millefolium L. N I
2 Adonis vernalis L. In III
3 Adonis volgensis DC. In III
4 Arrhenatherum elatius (L.) J. Presl & C. Presl A I
5 Bromopsis inermis (Leyss.) Holub N I
6 Calamagrostis epigejos (L.) Roth N I
7 Colchicum bulbocodium Ker Gawl. subsp. versicolor (Ker Gawl.) K. Perss. In VI
8 Corydalis solida (L.) Clairv. N IV
9 Dactylis glomerata L. N I
10 Delphinium sergii Wissjul. In I
11 Elytrigia intermedia (Host) Nevski In I
12 Elytrigia repens (L.) Nevski N I
13 Ficaria calthifolia Rchb. In IV
14 Ficaria verna Huds. N IV
15 Gagea liotardii (Sternb.) Schult. & Schult. f. N IV
16 Gagea minima (L.) Ker Gawl. N IV
17 Gagea pusilla (F. W. Schmidt) Sweet N IV
18 Gymnospermium odessanum (DC.) Takht. In V
19 Muscari neglectum Guss. ex Ten. In III
20 Ornithogalum boucheanum (Kunth) Asch. In I
21 Ornithogalum fimbriatum Willd. In III
22 Paeonia tenuifolia L. In I
23 Papaver orientale L. A I
24 Pulsatilla pratensis (L.) Mill. In III
25 Ranunculus illyricus L. In I
26 Scilla bifolia L. N IV
27 Scilla siberica Haw. In III
28 Tulipa biebersteiniana Schult. & Schult. f. In I
29 Veronica hederifolia L. N V
30 Viola odorata L. N II
Table 1. Spring synusiae during the flowering of Gymnospermium odessanum on the botanical-geographical
plot “Steppes of Ukraine” at the NBG in 2019–2021.
Note. Groups: N – native plant species; In – introduced plant species; A – alien plant species. Phenophases:
I – beginning of vegetation; II – beginning of vegetation and budding; III – budding; IV – budding and
beginning of flowering; V – mass flowering; VI – completion of flowering.
in a wide range from 20 % to 95 % (Smetana
et al., 2014). Gymnospermium odessanum often
dominates in the spring synusia (Scherbakova
& Novosad, 2018). Krasova et al. (2016)
observed a projective cover of G. odessanum
reaching 50–60 %. The height of G. odessanum
in nature is 5–20 cm (Kritska & Novosad, 2009)
or 10–15 cm (Trotner, 2017).
In the species composition of the artificially
created phytocoenosis (Table 1), introduced
plant species represent 50.0 % (15 species),
native – 43.3 % (13 species), alien – 6.7 %
8 Plant Introduction • 93/94
V. Gritsenko
(two species). At the same time, many plant
species (13 species, 43.3 %) have not begun
vegetation and therefore are not visible in
the spring synusiae yet. In particular, cereals
have just started their growth (Table 1; Fig. 1 C).
From other hand, at this time, Colchicum
bulbocodium Ker Gawl. subsp. versicolor (Ker
Gawl.) K. Perss. already finishes its flowering
(Table 1; Fig. 1 D). In this spring synusia, all
plants are herbaceous (Table 1). Of course,
such species composition of the spring synusia
is possible only in introduction conditions and
does not occur in natural phytocoenoses.
In Ukraine, the natural habitats of
G. odessanum are located on the limestones,
loess-limestones and granite slopes, ravines
and gullies, in the phytocoenoses of shrub
steppes, on the open areas of steppe slopes
adjacent to shrubby thickets, and forest glades
(Scherbakova, 2005; Kritska & Novosad, 2009;
Kritska, 2012). Smetana et al. (2014) indicated
a wide range of coenotic conditions for the
G. odessanum existence, particularly shrubby,
steppe, and rocky vegetation. Solomakha
(2015) noted that in the Ukrainian Northern
Black Sea region, G. odessanum occurs in the
forest and shrub communities. Scherbakova
& Novosad (2018) indicate that sylvatic
communities (i.e., natural ravine forests,
thickets of shrubs, and secondary forest
phytocoenoses) are optimal for the existence
of G. odessanum in Ukraine. In Moldova, this
species occurs exclusively in light oak forests.
It does not spread to open areas in Moldova,
and therefore it is considered there a typical
forest representative (Izverskaya et al., 2013).
Thus, the coenotic conditions of
G. odessanum habitats under introduction
on the botanical-geographical plot “Steppes
of Ukraine” differ from those in nature.
This confirms the opinion of Scherbakova &
Novosad (2018), who declared broad ecological
and coenotic plasticity of G. odessanum in
Ukraine.
Phenology features during Gymnospermium
odessanum flowering
During the flowering of G. odessanum, the
following phenophases were distinguished: the
beginning of flowering, the mass flowering,
and the completion of flowering. The mass
flowering of G. odessanum in 2019 lasted eight
days, in 2020 – seven days, in 2021 – nine days
(Fig. 2). The earliest mass flowering occurred
in March 2020, the latest – in early April 2021
(Fig. 2). The beginning of mass flowering
in 2020 and 2021 differed significantly and
resulted in 24 days of delay. This may be due
to the significant differences in spring weather
during these years. In 2021, the entire growing
season of G. odessanum lasted from March 24
to May 22 and amounted to 60 days (Gritsenko,
2021). At the same time, mass flowering took
up to 15 % of the entire vegetation period.
The flowering of G. odessanum occured
in March-April both in the conditions of
introduction at the NBG, and in nature
(Visjulina, 1953; Kritska & Novosad, 2009;
Trotner, 2017). During G. odessanum flowering,
the composition of plant species in the spring
synusia and phenophases of these plants were
the same in 2019–2021 (Table 1). At different
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Dates
Days of mass flowering in
2020 2019 2021
Figure 2. Dynamics of Gymnospermium odessanum mass flowering in the introduction coenopopulation at
the NBG in 2019–2021.
Plant Introduction • 93/94 9
The current state of the introduction coenopopulation of Gymnospermium odessanum
dates of G. odessanum flowering, an associated
shift of phenophases of other plant species in
the spring synusia was observed. Therefore,
the phenophases of these plants in different
years were the same, too (Table 1).
Fruiting of G. odessanum in nature
occurs from the middle of April to early May
(Trotner, 2017). Ripening and dispersion of
seeds of this species in the introduction
conditions occur in May (Gritsenko, 2021).
The seeds germinate in the spring of next
year. Vegetative reproduction is not usual for
G. odessanum (Litvinenko & Shcherbakova,
2009; Scherbakova & Barmak, 2013).
Ontogenetic aspect
Up to date, several generations of introduced
G. odessanum were formed at the botanical-
geographical plot “Steppes of Ukraine”. During
G. odessanum flowering, seedlings, as well as
juvenile, immature, virginal, generative, and
sub-senile individuals were observed (Fig. 3).
In the introduction conditions, the principal
parameters of the aboveground plant parts
of G. odessanum corresponded to those
registered in nature (Litvinenko, 2010).
Among the generative individuals, the most
frequent were those with one flowering shoot
(Fig. 3 E; Table 2). However, plants with two or
three flowering shoots occasionally occurred
too (Fig. 1 D; Fig. 3 F; Table 2). Plants with
four, five, and six flowering shoots were quite
rare (Table 2). In nature, usually occur plants
with one-three flowering shoots, sometimes
– with five or six flowering shoots (Trotner,
2017). Although, Litvinenko (2010) reported
that generative individuals of G. odessanum
in natural conditions could form up to 12
flowering shoots.
According to our long-term observations
(2001–2021), the individuals of G. odessanum
in the introduction conditions can persist
in the generative state for over twenty
years. Litvinenko (2010) indicated that the
ontogenesis of G. odessanum typically lasts
about 30 years, and the plants spend most
of their life in a mature generative state.
Thus, all individuals in the introduction
coenopopulation of this species at the NBG
today are direct descendants of initially
introduced plants.
BA C D
E F G
Figure 3. Gymnospermium odessanum individuals: A – seedling; B – juvenile plant; C – immature plant;
D – virginal plant; E–F – a generative plant with one (E) and three (F) flowering shoots; G – sub-senile plant.
Scales: A–D – 2 cm; E–G – 10 cm.
10 Plant Introduction • 93/94
V. Gritsenko
Age structure
In the spectra of age states, the percentage
of pregenerative individuals (i.e., seedling,
juvenile, immature, and virginal) in 2019–
2021 was very high (89.5 % in 2019, 89.2 % in
2020, and 88.8 % in 2021). For this period, the
percentage of generative individuals was low
(10.0 % in 2019, 10.3 % in 2020, and 10.9 % in
2021). The percentage of sub-senile individuals
for this period was even much lower (0.5 % in
2019, 0.5 % in 2020, and 0.3 % in 2021; Fig. 4).
In previous years (2010–2017), the overall
percentage of pregenerative individuals of
ontogenesis was also very high (about 90 %),
and the rate of generative individuals was low
(about 10 %) (Gritsenko, 2010, 2014, 2017a).
Thus, the age structure of the introduction
coenopopulation of G. odessanum is
characterized by long-term stability.
In the natural populations of G. odessanum,
there are present individuals of all age states
(Scherbakova, 2005; Kritska & Novosad, 2009;
Scherbakova & Novosad, 2018). The age spectra
in the natural populations of this species
are mostly left-sided, and only occasionally
they are right-sided (Scherbakova, 2010;
Scherbakova & Novosad, 2018). In particular, in
the natural coenopopulations of G. odessanum
in the Ingulets river basin, the share of
generative individuals was high (about 50 %),
and age spectra were right-sided (Smetana
et al. 2014). The age spectra of the introduction
coenopopulation of G. odessanum at the NBG
in 2010–2021 were left-sided, like in the most
of natural populations.
Spatial distribution and dissemination modes
The spatial distribution of particular
individuals in the introduction
coenopopulation of G. odessanum at
the NBG is of two kinds – random and
grouped. As a result of seed propagation
and different dissemination modes, single
individuals and groups of individuals of
different age states occur within the same
coenopopulation. Gymnospermium odessanum
is a myrmecochorous plant (Litvinenko &
Shcherbakova, 2009; Scherbakova & Barmak,
2013). Random distribution of its solitary
individuals of different age states in the
Number of
flowering shoots
per plant
Generative individuals, number / %
2019 2020 2021
1 32 / 78.0 32 / 76.2 34 / 75.6
2 4 / 9.8 5 / 11.9 5 / 11.1
3 4 / 9.8 3 / 7.1 4 / 8.9
4 0 / 0 1 / 2.4 1 / 2.2
5 1 / 2.4 1 / 2.4 0 / 0
6 0 / 0 0 / 0 1 / 2.2
Total 41 / 100 42 / 100 45 / 100
Table 2. The number of generative individuals of
Gymnospermium odessanum with a different number
of flowering shoots in 2019–2021.
61 52 40
228 228 241
40 43 45
38 41 40
41 42 45
2 2 1
0%
10%
20%
30%
40%
50%
60%
70%
80%
90%
100%
2019 2020 2021
Co
nt
rib
ut
io
n
Years
Number of
individuals
Subsenile
Generative
Virginal
Immature
Juvenile
Seedling
Figure 4. Age structure of Gymnospermium odessanum introduction coenopopulation at the NBG in
2019–2021.
Plant Introduction • 93/94 11
The current state of the introduction coenopopulation of Gymnospermium odessanum
introduction coenopopulation may be a result
of myrmecochory. The spatial distribution of
generative individuals illustrates their bivariate
(random and grouped) nature (Fig. 5 А). Many
pregenerative individuals were located near
the maternal generative individuals (Fig. 5 B),
evidencing the barochory. Individuals of
different age states differed in height. As a
result, the leaf blades of adjacent individuals
were located one above other, and formed a
B
A
Figure 5. Spatial distribution of Gymnospermium odessanum individuals in the introduction
coenopopulation at the NBG. А – boundaries of the coenopopulation (1), generative individuals in
2021 (2), and the fragment with the maximum density of generative individuals (3); scale – 50×50 m.
B – individuals of different age states, including seedling (p), and juvenile (j), immature (im), virginal
(v), and generative (g) plants; the circles indicate the phytogenic fields; scale – 10 cm.
12 Plant Introduction • 93/94
V. Gritsenko
vertical spatial pattern. Hence, phytogenic
fields of individuals of different age states
overlaped (Fig. 5 B).
Number and density of individuals
In 2019–2021, the total number and average
density of G. odessanum individuals did not
change significantly. The total number of
individuals of different age states was 410 in
2019, 408 – in 2020, and 412 – in 2021 (Fig. 4).
The most abundant were juvenile individuals,
while sub-senile ones were very rare (Fig. 4).
The number of individuals of other age
states (seedlings, immature, virginal, and
generative individuals) was not large and was
commensurate with each other (Fig. 4). The
largest number of seedlings was observed in
2019, the smallest – in 2021 (Fig. 4).
The number of seedlings and sub-
senile individuals in the introduction
coenopopulation at the NBG in 2019–2021
was characterized by the most extensive
variation range. Simultaneously, juvenile,
immature, virginal, and generative individuals
demonstrated much lower variation ranges
(Table 3).
Natural populations of G. odessanum
predominantly consist of a pretty high
number of individuals (Scherbakova, 2005).
In particular, natural coenopopulations of
this species in the basin of Ingulets river
have from several tens to several hundreds of
individuals (Smetana et al., 2014). Therefore,
the number of individuals in the introduction
coenopopulation of G. odessanum is
significant compared with many natural
coenopopulations.
Considering the density of individuals in the
introduction coenopopulation of G. odessanum
at the NBG, the following aspects were
noted. In several different fragments of the
coenopopulation, the density of individuals
was high and reached up to 25 individuals per
1 m2 (Fig. 5 B). At the same time, there were
many fragments within the coenopopulation
with low density (from one to several
individuals) or even zero presence. Hence,
the average density of G. odessanum here
was only 0.1975 individuals per 1 m2 in 2019,
0.1966 individuals per 1 m2 in 2020, and 0.1985
individuals per 1 m2 in 2021. Considering that
fragments with a high density of G. odessanum
individuals are rather rare, its average density
within coenopopulation is relatively low.
Trotner (2017) also observed uneven
distribution of G. odessanum individuals in
natural conditions. However, the density
was high for most part of the population.
Natural populations of G. odessanum are
generally characterized by high density
Age states Number of
individuals, %
Deviation,
%
Number of
individuals, %
Deviation,
%
Number of
individuals, %
Deviation,
%
2019 2020 2019 2021 2020 2019
Seedling 14.9 12.8 +14.09 14.9 9.7 +34.90 12.8 14.9 -16.41
Juvenile 55.7 55.9 -0.36 55.7 58.5 -5.03 55.9 55.7 +0.36
Immature 9.6 10.5 -9.38 9.6 10.9 -13.54 10.5 9.6 +8.57
Virginal 9.3 10.0 -7.53 9.3 9.7 -4.30 10.0 9.3 +7.00
Generative 10.0 10.3 -3.00 10.0 10.9 -9.00 10.3 10.0 +2.91
Subsenile 0.5 0.5 0.00 0.5 0.3 +40.00 0.5 0.5 0.00
2020 2021 2021 2019 2021 2020
Seedling 12.8 9.7 +24.22 9.7 14.9 -53.61 9.7 12.8 -31.96
Juvenile 55.9 58.5 -4.65 58.5 55.7 +4.79 58.5 55.9 +4.44
Immature 10.5 10.9 -3.81 10.9 9.6 +11.93 10.9 10.5 +3.67
Virginal 10.0 9.7 +3.00 9.7 9.3 +4.12 9.7 10.0 -3.09
Generative 10.3 10.9 -5.83 10.9 10.0 +8.26 10.9 10.3 +5.50
Subsenile 0.5 0.3 +40.00 0.3 0.5 -66.67 0.3 0.5 -66.67
Table 3. The number of individuals of different age states in the introduction coenopopulation of
Gymnospermium odessanum at the NBG in 2019–2021 (pairwise comparison).
Plant Introduction • 93/94 13
The current state of the introduction coenopopulation of Gymnospermium odessanum
(Scherbakova, 2005) and comprise several
dozen individuals per 1 m2 (Kritska & Novosad,
2009). Thus, Shiryaeva et al. (2020) reported
15–25 individuals per 1 m2. Krasova et al. (2016)
counted up to 170 individuals of the species per
1 m2 and registered its high projective cover.
Similarly, Smetana et al. (2014) indicated from
1 to 44 G. odessanum plants per 0.25 m2 for 27
natural habitats. Scherbakova & Novosad (2018)
also noted the high number and density of
G. odessanum individuals under conditions of
ecological-coenotic optimum. Of course, the
NBG conditions are far from such optimum.
Development trends of introduction
coenopopulation
Gymnospermium odessanum on the botanical-
geographical plot “Steppes of Ukraine”
successfully reproduces and settles for
decades. Introduction coenopopulation of
G. odessanum here is steady in time (long-
term existence) and space (occupy the stable
area). In this coenopopulation, the species
demonstrates a constant tendency to preserve
its stable position in the meadow-steppe
cultural phytocoenosis and, probably, to
further area extension due to myrmecochory.
Conclusions
Ecological and coenotic conditions at the
NBG differ from those in the natural habitats
of G. odessanum and are not optimal for this
species. It grows here on the botanical-
geographical plot “Steppes of Ukraine”
in the composition of artificially created
meadow-steppe phytocoenosis. However,
Gymnospermium odessanum demonstrated
high eco-coenotic plasticity and, over
the decades, has formed a homeostatic
introduction coenopopulation here. Today all
contemporary G. odessanum individuals in the
introduction coenopopulation at the NBG are
descendants of the initially introduced plants.
As of 2021, the area of introduction
coenopopulation at the NBG was 2,075 m2.
It consisted of 412 individuals (40 seedlings,
241 juvenile, 45 immature, 40 virginal, 45
generative, and one sub-senile plants).
Fragments with a high density of individuals
were rare, so the average density was low,
with only about 0.2 individuals per 1 m2.
In 2021, in the spectrum of age states, the
total rate of pregenerative individuals of
ontogenesis was high and amounted to 88.8 %,
while the share of generative individuals was
10.9 %. In general, the age structure of this
introduction coenopopulation is characterized
by long-term (2010–2021) stability. Due to
myrmecochoric and barochoric propagation,
the spatial distribution of individuals in this
introduction coenopopulation is uneven and
of two kinds – random and in groups.
Compared to natural populations, the
investigated introduction coenopopulation of
G. odessanum is characterized by a large area,
significant number, and low average density of
individuals. Like in most natural populations,
the age spectrum in this introduction
coenopopulation is left-sided.
The conducted research testifies the
successful formation of the introduction
coenopopulation of G. odessanum in the
meadow-steppe cultural phytocoenosis of the
NBG. This introduction coenopopulation is an
example of successful multi-decade scientific
experiment and effective ex situ protection and
preservation of G. odessanum on the northern
border of the Right Bank Forest-Steppe of
Ukraine, far beyond its natural range.
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16 Plant Introduction • 93/94
V. Gritsenko
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імені М. М. Гришка НАН України
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Plant Introduction • 93/94 17
The current state of the introduction coenopopulation of Gymnospermium odessanum
Gymnospermium odessanum – це раритетний реліктовий ендемічний вид. Дослідження проведені
під час цвітіння G. odessanum у 2019–2021 рр. на ботаніко-географічній ділянці “Степи України”
Національного ботанічного саду імені М. М. Гришка НАН України (НБС). Еколого-ценотичні умови
інтродукції у НБС відрізнялись від природних місцезростань і не були оптимальними. Зокрема,
G. odessanum росте на ботаніко-географічній ділянці “Степи України” у складі штучно створеного
лучно-степового фітоценозу. Втім, G. odessanum проявив еколого-ценотичну пластичність і,
упродовж десятиліть, сформував тут стійку гомеостатичну інтродукційну ценопопуляцію. Станом на
2021 р. площа інтродукційної ценопопуляції цього виду становила 2075 м2. У її складі налічувалось
412 особин G. odessanum (40 проростків, 241 ювенільних, 45 імматурних, 40 віргінільних, 45
генеративних та 1 субсенільна). Фрагменти з високою щільністю особин G. odessanum траплялися
рідко, тому середня щільність була низькою і становила 0,2 особини на 1 м2. У 2021 р. у спектрі
вікових станів загальний відсоток особин прегенеративного періоду онтогенезу був дуже високим і
становив 88,8 %, а частка генеративних особин становила 10,9 %.
В цілому, вікова структура інтродукційної ценопопуляції G. odessanum характеризується тривалою
(2010–2021 рр.) стабільністю. Просторовий розподіл особин в інтродукційній ценопопуляції
випадковий та груповий. Це пов’язано з мірмекохорним і барохорним поширенням. Ценопопуляція
має тенденцію до подальшого збільшення площі, що обумовлено мірмекохорією. У порівнянні
з природними ценопопуляціями, інтродукційна ценопопуляція G. odessanum характеризується
великою площею, значною кількістю та низькою середньою щільністю особин. Вікові спектри, як і
у більшості природних популяцій, є лівосторонніми.
Проведені дослідження свідчать про успішне формування інтродукційної ценопопуляції G. odessanum
в лучно-степовому культур-фітоценозі НБС. Ця інтродукційна ценопопуляція є вдалим багаторічним
науковим експериментом та представляє собою приклад ефективної ex situ охорони та збереження
G. odessanum на північній межі Правобережного Лісостепу України, далеко за межами природного
ареалу цього ендемічного виду.
Ключові слова: Gymnospermium odessanum, раритетний ендемічний вид, інтродукційна ценопопуляція, стабільна вікова
структура, охорона ex situ
|
| id | oai:ojs2.plantintroduction.org:article-1598 |
| institution | Plant Introduction |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2025-07-17T12:54:01Z |
| publishDate | 2022 |
| publisher | M.M. Gryshko National Botanical Garden of the NAS of Ukraine |
| record_format | ojs |
| resource_txt_mv | wwwplantintroductionorg/d9/2053b7e1028f0d4322a10937ec08e1d9.pdf |
| spelling | oai:ojs2.plantintroduction.org:article-15982023-08-26T20:38:56Z The current state of the introduction coenopopulation of Gymnospermium odessanum on the botanical-geographical plot “Steppes of Ukraine” at the M. M. Gryshko National Botanical Garden of the NAS of Ukraine Сучасний стан інтродукційної ценопопуляції Gymnospermium odessanum на ботаніко-географічній ділянці “Степи України” Національного ботанічного саду імені М. М. Гришка НАН України Gritsenko, Victoria Gymnospermium odessanum is a rare relict endemic plant species. The research was carried out during G. odessanum flowering in 2019–2021 at the botanical-geographical plot “Steppes of Ukraine” of the M.M. Gryshko National Botanical Garden, National Academy of Sciences of Ukraine (NBG). Ecological and coenotic conditions of introduction at the NBG differ from natural habitats of the species and are not optimal for its vegetation. However, this species demonstrated ecological-coenotic plasticity and, over the decades, has formed a stable homeostatic introduction coenopopulation here. As of 2021, the area of introduction coenopopulation of G. odessanum at the NBG was 2,075 m2. It comprises 412 individuals of this species (including 40 seedlings, 241 juvenile, 45 immature, 40 virginal, 45 generative, and 1 sub-senile plants). Coenopopulation fragments with a high density of G. odessanum individuals are rare here, so the average density is low – only 0.2 individuals per 1 m2. In 2021, in the spectrum of age states, the total percentage of pregenerative individuals was very high and reached 88.8 %; the share of generative individuals was 10.9 %.In general, the age structure of the introduction coenopopulation of G. odessanum is characterized by long-term (2010–2021) stability. The spatial distribution of individuals in the introduction coenopopulation is of two kinds – random and in groups. This is due to combined myrmecochoric and barochoric propagation. Also due to myrmecochory, this coenopopulation tends to spread the area. Compared to natural coenopopulations, the introduction coenopopulation of G. odessanum at the NBG is characterized by a larger area, a much significant number, and, at the same time, a low average density of individuals. However, like in most of natural populations, its age spectrum is left-sided.The conducted research testifies the successful formation of the introduction coenopopulation of G. odessanum in the meadow-steppe cultural phytocoenosis of the NBG. This introduction coenopopulation is an example of a successful multi-year scientific experiment and effective ex situ protection and preservation of G. odessanum on the northern border of the Right Bank Forest-Steppe of Ukraine, far beyond the natural range of this endemic plant species. Gymnospermium odessanum – це раритетний реліктовий ендемічний вид. Дослідження проведені під час цвітіння G. odessanum у 2019–2021 рр. на ботаніко-географічній ділянці “Степи України” Національного ботанічного саду імені М. М. Гришка НАН України (НБС). Еколого-ценотичні умови інтродукції у НБС відрізнялись від природних місцезростань і не були оптимальними. Зокрема, G. odessanum росте на ботаніко-географічній ділянці “Степи України” у складі штучно створеного лучно-степового фітоценозу. Втім, G. odessanum проявив еколого-ценотичну пластичність і, упродовж десятиліть, сформував тут стійку гомеостатичну інтродукційну ценопопуляцію. Станом на 2021 р. площа інтродукційної ценопопуляції цього виду становила 2075 м2. У її складі налічувалось 412 особин G. odessanum (40 проростків, 241 ювенільних, 45 імматурних, 40 віргінільних, 45 генеративних та 1 субсенільна). Фрагменти з високою щільністю особин G. odessanum траплялися рідко, тому середня щільність була низькою і становила 0,2 особини на 1 м2. У 2021 р. у спектрі вікових станів загальний відсоток особин прегенеративного періоду онтогенезу був дуже високим і становив 88,8 %, а частка генеративних особин становила 10,9 %.В цілому, вікова структура інтродукційної ценопопуляції G. odessanum характеризується тривалою (2010–2021 рр.) стабільністю. Просторовий розподіл особин в інтродукційній ценопопуляції випадковий та груповий. Це пов’язано з мірмекохорним і барохорним поширенням. Ценопопуляція має тенденцію до подальшого збільшення площі, що обумовлено мірмекохорією. У порівнянні з природними ценопопуляціями, інтродукційна ценопопуляція G. odessanum характеризується великою площею, значною кількістю та низькою середньою щільністю особин. Вікові спектри, як і у більшості природних популяцій, є лівосторонніми.Проведені дослідження свідчать про успішне формування інтродукційної ценопопуляції G. odessanum в лучно-степовому культур-фітоценозі НБС. Ця інтродукційна ценопопуляція є вдалим багаторічним науковим експериментом та представляє собою приклад ефективної ex situ охорони та збереження G. odessanum на північній межі Правобережного Лісостепу України, далеко за межами природного ареалу цього ендемічного виду. M.M. Gryshko National Botanical Garden of the NAS of Ukraine 2022-01-14 Article Article application/pdf https://www.plantintroduction.org/index.php/pi/article/view/1598 10.46341/PI2021017 Plant Introduction; No 93/94 (2022); 3-17 Інтродукція Рослин; № 93/94 (2022); 3-17 2663-290X 1605-6574 10.46341/PI93-94 en https://www.plantintroduction.org/index.php/pi/article/view/1598/1524 Copyright (c) 2022 Victoria Gritsenko http://creativecommons.org/licenses/by/4.0 |
| spellingShingle | Gritsenko, Victoria Сучасний стан інтродукційної ценопопуляції Gymnospermium odessanum на ботаніко-географічній ділянці “Степи України” Національного ботанічного саду імені М. М. Гришка НАН України |
| title | Сучасний стан інтродукційної ценопопуляції Gymnospermium odessanum на ботаніко-географічній ділянці “Степи України” Національного ботанічного саду імені М. М. Гришка НАН України |
| title_alt | The current state of the introduction coenopopulation of Gymnospermium odessanum on the botanical-geographical plot “Steppes of Ukraine” at the M. M. Gryshko National Botanical Garden of the NAS of Ukraine |
| title_full | Сучасний стан інтродукційної ценопопуляції Gymnospermium odessanum на ботаніко-географічній ділянці “Степи України” Національного ботанічного саду імені М. М. Гришка НАН України |
| title_fullStr | Сучасний стан інтродукційної ценопопуляції Gymnospermium odessanum на ботаніко-географічній ділянці “Степи України” Національного ботанічного саду імені М. М. Гришка НАН України |
| title_full_unstemmed | Сучасний стан інтродукційної ценопопуляції Gymnospermium odessanum на ботаніко-географічній ділянці “Степи України” Національного ботанічного саду імені М. М. Гришка НАН України |
| title_short | Сучасний стан інтродукційної ценопопуляції Gymnospermium odessanum на ботаніко-географічній ділянці “Степи України” Національного ботанічного саду імені М. М. Гришка НАН України |
| title_sort | сучасний стан інтродукційної ценопопуляції gymnospermium odessanum на ботаніко-географічній ділянці “степи україни” національного ботанічного саду імені м. м. гришка нан україни |
| url | https://www.plantintroduction.org/index.php/pi/article/view/1598 |
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