Distribution and population status of rare plant species in the Marmarosh Mountains (Ukrainian Carpathians)
The article presents the results of multiple surveys of habitats of rare plant species in the Marmarosh Mts, Ukrainian Carpathians, and depicts their exact location and population parameters. Along with the analysis of earlier literature and herbarium data, this enabled to reveal some spatio-tempora...
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irk-123456789-1785002021-02-20T01:26:51Z Distribution and population status of rare plant species in the Marmarosh Mountains (Ukrainian Carpathians) Kobiv, Y. Prokopiv, A. Nachychko, V. Borsukevych, L. Helesh, M. Геоботаніка, екологія, охорона рослинного світу The article presents the results of multiple surveys of habitats of rare plant species in the Marmarosh Mts, Ukrainian Carpathians, and depicts their exact location and population parameters. Along with the analysis of earlier literature and herbarium data, this enabled to reveal some spatio-temporal patterns in their distribution, namely habitat characteristics and trends in population dynamics, which allowed to determine their conservation status. Numerous facts of decline and extinction, as well as examples of positive dynamics in populations of the studied species were described. Cold-dependent species (Cerastium cerastoides, Primula minima, Veronica bellidioides) proved to be vulnerable to climate change particularly at the lower limit of their distribution. Another factor of impact is decrease of grazing, which triggers vegetation succession and consequent replacement of some low-competitive rare plants (Genista tinctoria subsp. oligosperma, Koeleria macrantha subsp. transsilvanica) with shrubs or tussock graminoids. However, another group of rare species (Heracleum sphondylium subsp. transsilvanicum, Gentiana punctata, Jovibarba globifera subsp. preissiana, Veronica fruticans) benefits from the decrease of anthropogenic impact. У статті подано результати обстежень оселищ рідкісних видів рослин у Мармароських горах (Українські Карпати) та описано їхнє точне розташування й популяційні параметри. У результаті аналізу попередніх літературних і гербарних матеріалів встановлено деякі просторовочасові закономірності їхнього поширення, зокрема особливості оселищ і динаміки популяцій, що уможливило визначення їхнього природоохоронного статусу. Виявлено численні факти регресування чи зникнення, а також приклади позитивної динаміки популяцій досліджених видів. Холодостійкі види (Cerastium cerastoides, Primula minima, Veronica bellidioides) виявилися вразливими до кліматичних змін, зокрема на нижній межі їхнього поширення. Іншим фактором впливу є послаблення випасання, що спричинило сукцесію рослинності і, як наслідок, витіснення деяких низькоконкурентних рідкісних рослин (Genista tinctoria subsp. oligosperma, Koeleria macrantha subsp. transsilvanica) чагарничками або дернинними злаками. Натомість, інша група рідкісних видів (Heracleum sphondylium subsp. transsilvanicum, Gentiana punctata, Jovibarba globifera subsp. preissiana, Veronica fruticans) позитивно реагує на зменшення антропогенного впливу. Представлены результаты обследования местопроизрастаний редких видов растений в Мармарошских горах (Украинские Карпаты) и описаны их точное расположение и популяционные параметры. В результате анализа предыдущих литературных и гербарных данных установлены некоторые пространственно-временные закономерности их распространения, в частности особенности местообитаний, тенденции динамики популяций, что дало возможность определить их природоохранный статус. Выявлены многочисленные факты их регрессирования или исчезновения, а также примеры положительной динамики популяций исследованных видов. Холодоустойчивые виды (Cerastium cerastoides, Primula minima, Veronica bellidioides) оказались уязвимыми к климатическим изменениям, в частности на нижней границе их распространения. Еще одним влияющим фактором является ослабление выпаса, вызвавшее сукцессию растительности и последующее вытеснение некоторых низкоконкурентных редких растений (Genista tinctoria subsp. oligosperma, Koeleria macrantha subsp. transsilvanica) кустарничками или дерновинными злаками. Однако, другая группа редких видов (Heracleum sphondylium subsp. transsilvanicum, Gentiana punctata, Jovibarba globifera subsp. preissiana, Veronica fruticans) положительно реагирует на уменьшение антропогенного влияния. 2017 Article Distribution and population status of rare plant species in the Marmarosh Mountains (Ukrainian Carpathians) / Y. Kobiv, A. Prokopiv, V. Nachychko, L. Borsukevych, M. Helesh // Український ботанічний журнал. — 2017. — Т. 74, № 2. — С. 163-176. — Бібліогр.: 35 назв. — англ. 0372-4123 DOI: https://doi.org/10.15407/ukrbotj74.02.163 http://dspace.nbuv.gov.ua/handle/123456789/178500 en Український ботанічний журнал Інститут ботаніки ім. М.Г. Холодного НАН України |
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Геоботаніка, екологія, охорона рослинного світу Геоботаніка, екологія, охорона рослинного світу |
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Геоботаніка, екологія, охорона рослинного світу Геоботаніка, екологія, охорона рослинного світу Kobiv, Y. Prokopiv, A. Nachychko, V. Borsukevych, L. Helesh, M. Distribution and population status of rare plant species in the Marmarosh Mountains (Ukrainian Carpathians) Український ботанічний журнал |
description |
The article presents the results of multiple surveys of habitats of rare plant species in the Marmarosh Mts, Ukrainian Carpathians, and depicts their exact location and population parameters. Along with the analysis of earlier literature and herbarium data, this enabled to reveal some spatio-temporal patterns in their distribution, namely habitat characteristics and trends in population dynamics, which allowed to determine their conservation status. Numerous facts of decline and extinction, as well as examples of positive dynamics in populations of the studied species were described. Cold-dependent species (Cerastium cerastoides, Primula minima, Veronica bellidioides) proved to be vulnerable to climate change particularly at the lower limit of their distribution. Another factor of impact is decrease of grazing, which triggers vegetation succession and consequent replacement of some low-competitive rare plants (Genista tinctoria subsp. oligosperma, Koeleria macrantha subsp. transsilvanica) with shrubs or tussock graminoids. However, another group of rare species (Heracleum sphondylium subsp. transsilvanicum, Gentiana punctata, Jovibarba globifera subsp. preissiana, Veronica fruticans) benefits from the decrease of anthropogenic impact. |
format |
Article |
author |
Kobiv, Y. Prokopiv, A. Nachychko, V. Borsukevych, L. Helesh, M. |
author_facet |
Kobiv, Y. Prokopiv, A. Nachychko, V. Borsukevych, L. Helesh, M. |
author_sort |
Kobiv, Y. |
title |
Distribution and population status of rare plant species in the Marmarosh Mountains (Ukrainian Carpathians) |
title_short |
Distribution and population status of rare plant species in the Marmarosh Mountains (Ukrainian Carpathians) |
title_full |
Distribution and population status of rare plant species in the Marmarosh Mountains (Ukrainian Carpathians) |
title_fullStr |
Distribution and population status of rare plant species in the Marmarosh Mountains (Ukrainian Carpathians) |
title_full_unstemmed |
Distribution and population status of rare plant species in the Marmarosh Mountains (Ukrainian Carpathians) |
title_sort |
distribution and population status of rare plant species in the marmarosh mountains (ukrainian carpathians) |
publisher |
Інститут ботаніки ім. М.Г. Холодного НАН України |
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2017 |
topic_facet |
Геоботаніка, екологія, охорона рослинного світу |
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http://dspace.nbuv.gov.ua/handle/123456789/178500 |
citation_txt |
Distribution and population status of rare plant species in the Marmarosh Mountains (Ukrainian Carpathians) / Y. Kobiv, A. Prokopiv, V. Nachychko, L. Borsukevych, M. Helesh // Український ботанічний журнал. — 2017. — Т. 74, № 2. — С. 163-176. — Бібліогр.: 35 назв. — англ. |
series |
Український ботанічний журнал |
work_keys_str_mv |
AT kobivy distributionandpopulationstatusofrareplantspeciesinthemarmaroshmountainsukrainiancarpathians AT prokopiva distributionandpopulationstatusofrareplantspeciesinthemarmaroshmountainsukrainiancarpathians AT nachychkov distributionandpopulationstatusofrareplantspeciesinthemarmaroshmountainsukrainiancarpathians AT borsukevychl distributionandpopulationstatusofrareplantspeciesinthemarmaroshmountainsukrainiancarpathians AT heleshm distributionandpopulationstatusofrareplantspeciesinthemarmaroshmountainsukrainiancarpathians |
first_indexed |
2025-07-15T17:01:52Z |
last_indexed |
2025-07-15T17:01:52Z |
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1837733158033817600 |
fulltext |
163Укр. бот. журн., 2017, 74(2)
Introduction
The Marmarosh (or Maramureş – in Romanian
transcription) Mountains are situated on the Ukrainian-
Romanian border and belong to the Inner Carpathians.
They have a very diverse geological structure with
prevailing crystalline bedrock and rather frequent
limestone outcrops. This significantly differs from the
vast outer part of the Ukrainian Carpathians dominated
by mostly uniform acidic sandstone flysch (Kondracki,
1989). The Marmarosh Mts are the steepest in the
Ukrainian Carpathians (Kruhlov, 2008), therefore
rock outcrops are quite common here. That mountain
range is the second highest (after the Chornohora)
massif in the Ukrainian Carpathians with two summits
of Mt. Pip Ivan exceeding 1900 m a.s.l. This territory
makes an isolated alpine "environmental island". In
the Marmarosh Mts some rare and endemic plants are
confined to the eastern edge of their ranges and do not
occur elsewhere in the Ukrainian Carpathians. All the
above reasons make these mountains unique in terms
of their biodiversity and landscape value, therefore their
considerable part was included into the Carpathian
Biosphere Reserve. The flora and vegetation of the
Marmarosh Mts have been studied since the late
19th century. The most precise old chorological data
concerning distribution of rare species were published
by H. Zapałowicz (1889), I. Klášterský (1929–1931),
V. Krist (1935), and M. Deyl (1940). Floristic studies
during the Soviet period, i.e. in the 1950–1980s, were
rather incomplete because of the limited access to
that frontier territory, which remained one of the most
understudied regions of the Ukrainian Carpathians.
Though the "Flora of the Ukrainian SSR" (Flora RSS
Ucr., 1950–1965), as well as V.I. Chopyk's (1976) and
K.A. Malynovski's (1980) monographs contain much
information on the flora of the Marmarosh Mts, most
publications from the last decades lack the exact data
on the localities of rare species and the status of their
populations in the region.
The aim of this study is to fill that gap, i.e. to specify
their distribution in the Marmarosh Mts and compare
contemporary data with those contained in literature
and herbaria in order to reveal possible dynamic patterns.
Special attention was paid to population parameters of
the rare species, their viability and conservation status.
We indicated the exact location of their habitats to set a
baseline for future monitoring.
©Yu. KOBIV, A. PROKOPIV, V. NACHYCHKO,
L. BORSUKEVYCH, M. HELESH, 2017
doi: 10.15407/ukrbotj74.02.163
Distribution and population status of rare plant species
in the Marmarosh Mountains (Ukrainian Carpathians)
Yuriy KOBIV1,2, Andriy PROKOPIV2, Viktor NACHYCHKO2, Liubov BORSUKEVYCH2, Myroslava HELESH2
1Institute of Ecology of the Carpathians, National Academy of Sciences of Ukraine
4, Kozelnytska Str., Lviv 79026, Ukraine
ykobiv@gmail.com
2Botanical Garden of Ivan Franko National University of Lviv
44, Cheremshyny Str., Lviv 79014, Ukraine
botsad@franko.lviv.ua
Kobiv Y., Prokopiv A., Nachychko V., Borsukevych L., Helesh M. Distribution and population status of rare plant species in the
Marmarosh Mountains (Ukrainian Carpathians). Ukr. Bot. J., 2017, 74(2): 163–176.
Abstract. The article presents the results of multiple surveys of habitats of rare plant species in the Marmarosh Mts, Ukrainian
Carpathians, and depicts their exact location and population parameters. Along with the analysis of earlier literature and
herbarium data, this enabled to reveal some spatio-temporal patterns in their distribution, namely habitat characteristics and
trends in population dynamics, which allowed to determine their conservation status. Numerous facts of decline and extinction,
as well as examples of positive dynamics in populations of the studied species were described. Cold-dependent species (Cerastium
cerastoides, Primula minima, Veronica bellidioides) proved to be vulnerable to climate change particularly at the lower limit
of their distribution. Another factor of impact is decrease of grazing, which triggers vegetation succession and consequent
replacement of some low-competitive rare plants (Genista tinctoria subsp. oligosperma, Koeleria macrantha subsp. transsilvanica)
with shrubs or tussock graminoids. However, another group of rare species (Heracleum sphondylium subsp. transsilvanicum,
Gentiana punctata, Jovibarba globifera subsp. preissiana, Veronica fruticans) benefits from the decrease of anthropogenic impact.
Keywords: climate change, extinction, population dynamics, locality, Carpathians
https://doi.org/10.15407/ukrbotj74.02.163
mailto:ykobiv@gmail.com
mailto:botsad@franko.lviv.ua
164 Ukr. Bot. J., 2017, 74(2)
In small populations their total census was estimated
(Elzinga et al., 2009).
Species abundance in Tables 1, 2 is presented
according to J. Braun-Blanquet (1964) scale.
Exact site locations were determined in WGS-84
system with Garmin eTrex GPS navigator with the
accuracy of measurements within 10 metres.
Content of calcium and magnesium in soil was
measured complexometrically and pH(H
2
O) value
electrometrically (Arinushkina, 1970).
Nomenclature of vascular plants follows "Flora
Europaea" (1964–1980), with modern updates for
endemics by J. Kliment et al. (2016), and M.O. Hill
et al. (2006) – for bryophytes.
Results and Discussion
The habitats of rare species are mostly confined to sites
with special ecological conditions, geological structure
or landforms, which can be considered as "hot spots"
of their occurrence (Ziman et al., 2009). For instance,
in the Ukrainian Carpathians those are usually alpine,
rocky, marshy or calcareous localities (Kobiv, 2010).
Although the presented data on the rare species are
not fully comprehensive, they are supposed to depict
properly their distribution and population status in the
target region.
Materials and Methods
The results presented below were obtained during field
surveys carried out in 2002, 2006, 2010, and 2016 in the
Marmarosh Mts, Ukrainian Carpathians. The studied
area stretches about 15 km along the Ukrainian-
Romanian border (Figure).
Thorough analysis of the previous publications and
herbarium data on rare species had been performed
before surveying their known localities in situ.
The map of the target area with locations of the
surveyed habitats of rare species is provided in the Figure
where their numeration corresponds to Tables 1–3.
The species were considered as rare following
K.A. Malynovski et al. (2002) compendium.
Density of populations was estimated on sets of
square plots placed randomly using 0.5 × 0.5 m² or 0.1 ×
0.1 m² frames with 10-fold repetition (Harper, 1977).
Studied localities of rare species in the Marmarosh Mts. 1, 2 ... 30 – locality numbers
165Укр. бот. журн., 2017, 74(2)
A small population of an East-South-Carpathian
endemic, Heracleum sphondylium L. subsp.
transsilvanicum (Schur) Brummitt (= H. palmatum
Baumg.), is situated in a steep hollow near the summit
of Mt. Nenieska. Its core zone is confined to the tall-
forb community, which covers about 30 m² (locality
1). Density of flowering undividuals of H. sphondylium
subsp. transsilvanicum is 1.8 specimens per 1 m². This
aclonal species is capable only of seed reproduction.
All the mentioned species are restricted to the
subalpine zone within 1700–1800 m a.s.l. In terms of
occurrence of the rare plants (for instance, Achillea
oxyloba subsp. schurii, Allium montanum, Aster
alpinus, Astragalus australis subsp. krajinae, Draba
siliquosa, Erigeron alpinus, Leontopodium alpinum,
Primula halleri), the rocks of Mt. Nenieska have most
resemblance to saxicolous sites from the Svydovets Mts
that apparently can be explained by similar geological
structure.
The next cluster of rare species’ localities is the
highest part of the Marmarosh Mts, i.e. Mt. Pip
Ivan with adjacent Mt. Gropa, Mt. Shcherban
(aka Zherban), and Mt. Polonynka built mostly of
crystalline bedrock – gneisses, metamorphic schists and
amphibolites. Considerable area is covered with alpine
vegetation there. Snowbed habitats that harbor highly
specialized prostrate cold-adapted species are most
characteristic of the alpine environment. One of such
chionophilous species is Salix herbacea L. We found
its two small populations (localities 10, 12) confined to
snowbeds on the highest elevations at the ridge on Mt.
Pip Ivan on poorly insolated N and NW aspects. Due to
clonal growth S. herbacea forms dense mats, while its
patches cover small areas of 15 and 4 m².
Another chionophilous species is Cerastium
cerastoides (L.) Britton, which inhabits bottoms of large
snowbeds in the glacial cirques of Mt. Pip Ivan. It occurs
mostly on gravel sites with poor floristic composition
and low abundance of other herbaceous plants (locality
8). Cerastium cerastoides can form mats up to 100–
150 cm² with 15–20 flowering shoots. However,
flowering is not common and most individuals are
vegetative and stunted. The species is rare now and we
found only two its small populations restricted to 1745–
1820 m a.s.l. altitudinal range. It is remarkable that
according to M. Deyl (1940) it was much more abundant
in the late 1930s being recorded in 19 localities within
1600–1910 m a.s.l. Moreover, earlier H. Zapałowicz
(1889) had reported on its still lower locality at 1580
m a.s.l. Apparently, decline of C. cerastoides is caused
Therefore, our results are presented with regard to such
hotspots.
Mt. Nenieska. This site is remarkable for its ca. 500 m
long steep rocky north-eastern slope with calcite-
rich sandstone bedrock. It is a habitat of many rare
saxicolous species, most of which are calciphilous
(Table 1). One of them, Leontopodium alpinum (L.)
Cass., we discovered in two localities (#3, 4), where
only about a dozen of its flowering individuals occur.
Local rangers confirm that the population number has
decreased significantly during the last decades, because
the plants have been illegally dug up by souvenir
hunters. That has undermined the population viability
and only some solitary individuals have survived in the
most inaccessible rocky loci. Therefore, the species is
critically endangered.
The rocks of Mt. Nenieska harbor an East-
Carpathian endemic Astragalus australis (L.) Lam.
subsp. krajinae (Domin) Domin, which is restricted
to steep slopes (localities 2, 3) where the density of its
flowering individuals is about 0.3 specimens per m².
The population is maintained by seed recruitment.
Another rare saxicolous species, Aster alpinus L.,
is abundant in the same localities. Its clones with
numerous flowering shoots may cover up to 200 cm².
The rocks are inhabited by a highly viable population
of Primula halleri J.F. Gmel. The average density of its
flowering individuals is 0.05 specimens per m². The only
way for recruitment of this short-lived species is seed
reproduction.
Other noteworthy alpine rare taxa are: Cerastium
alpinum L. subsp. lanatum (Lam.) Asch. & Graebn.,
Erigeron alpinus L., Minuartia verna (L.) Hiern. subsp.
gerardii (Willd.) Graebn., and Ranunculus thora L.,
which occur here on a rather large area of about 5
hectares though they do not exhibit high density
(localities 2–5). Scarce patches of sward vegetation
harbor Achillea lingulata Waldst. & Kit.
Steep shady and moist rocks provide habitats for a
rare East-South-Carpathian endemic Achillea oxyloba
(DC.) Sch. Bip. subsp. schurii (Sch. Bip.) Heimerl. Due
to clonal growth it forms dense patches up to 200–300
cm² with numerous flowering shoots (localities 2, 5).
Much more abundant are Hieracium villosum
Jacq. which forms dense clones on rocky shelves and
Botrychium lunaria (L.) Sw. mostly confined to loci of
almost barren gravel. Rhodiola rosea L. that occupies
both types of these habitats is also rather frequent but
subjected to severe damage due to illegal extirpation of
its rhizomes for medicinal purposes.
166 Ukr. Bot. J., 2017, 74(2)
Table 1. Floristic composition in localities of rare plant species in the hotspots of Mt. Nenieska and Mt. Pip Ivan with adjacent
area (names of rare species in boldface)
Number of locality 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17
Location Mt. Nenieska
Mt.
Gropa
Mt. Pip Ivan
Mt.
Polonynka
Date of relevé (day, month, year)
21
.0
7.
06
21
.0
7.
06
21
.0
7.
06
22
.0
7.
06
22
.0
7.
06
22
.0
7.
06
22
.0
7.
06
26
.0
7.
16
25
.0
7.
92
28
.0
7.
16
25
.0
7.
02
26
.0
7.
16
26
.0
7.
16
27
.0
6.
16
27
.0
6.
16
27
.0
7.
02
28
.0
7.
16
Area of relevé (m²) 25 100 25 25 25 100 100 25 100 4 100 15 25 10 15 25 2
Elevation (m a.s.l.)
17
95
17
30
17
85
17
75
17
50
15
30
18
65
17
50
18
35
18
55
18
10
19
25
17
90
19
10
19
30
15
15
16
05
Exposition NE NE NE NE NE E E NE NE NW NE N NE E NE NW NE
Inclination (°) 40 45 50 50 50 2 20 15 30 40 20 30 15 50 15 3 70
Vegetation cover (%) 100 70 40 70 40 90 90 50 50 60 80 70 59 50 70 100 25
Achillea lingulata · · + · · · + · · · · · · · · · ·
Achillea oxyloba subsp. schurii · + · · + · · · · · · · · · · · ·
Achillea strictag · · + · · · + · · · + · · + · · ·
Allium montanum · + · · · · · · · · · · · · · · ·
Alnus viridis · + · · 1 · · · · · · · · · · · ·
Anemone narcissifolia · · · · · · + · · · · · · · + · ·
Anthemis cretica subsp. pyrethriformis · · · · · · + · · · · · · + + · ·
Anthoxanthum alpinum · · + · · + + · + · · 2 · · + · ·
Arnica montana · · · · · · + · · · · · · · + · ·
Aster alpinus · 1 + · · · · · · · · · · · · · ·
Astragalus australis subsp. krajinae · + + · · · · · · · · · · · · · ·
Botrychium lunaria · · · + · · · · · · · · · · · · ·
Calamagrostis villosa 1 2 · · · · 2 · · · · · · · · 2 ·
Campanula alpina · · · · · · · · + 1 · · · · + · ·
Campanula carpatica · · · · · · · · · · · · · · · · 2
Campanula kladniana · · + · · · · · + · + · + 1 + · ·
Cardaminopsis halleri · · · · + · · · · · · · · · · · ·
Carduus kerneri · + · · · · · · · · · · · + · · ·
Carex limosa · · · · · 2 · · · · · · · · · · ·
Carex pauciflora · · · · · 2 · · · · · · · · · · ·
Carex sempervirens · 1 · · 1 · 4 · 2 1 2 · · 3 3 · ·
Cerastium cerastoides · · · · · · · + · · · · · · · · ·
Cerastium lanatum · + + + · · · · · · · · · · · · ·
Cirsium erisithales · + + + · · · · · · · · · · · · ·
Deschampsia caespitosa · · · · · + · 3 1 · 1 + · · · 1 ·
Draba siliquosa · · + · · · · · · · · · · · · · ·
Erigeron alpinus · · + + · · · · · · · · · · · · ·
Festuca airoides · · 1 4 · · · · · 2 · · · · · · 3
Festuca carpatica · · + · · · · · · · · · · · · · ·
Festuca picta · + 1 1 · · 2 + 1 · · 1 · · 2 · ·
Festuca rubra · · 2 · · · · · · · · · · · · 4 ·
Galium anisophyllon · · + + · · · · · · · · · + · · ·
Galium erectum subsp. suberectum · + + · · · · · · · · · · · · · ·
Gentiana acaulis · · · + · · · · + · · · · · + · ·
Gentiana asclepiadea · + · · · · · · · · + · · · · · ·
Gentiana lutea · · · · · · · · · · 1 · · · · · ·
Gentiana punctata · · · · · · · · · · + + · · · · ·
Heracleum palmatum 2 · · · · · · · · · · · · · · · ·
Hieracium alpinum · · · · · · · · · + · + · · · · ·
Hieracium umbellatum · · · · · · · · · · · · · + · · 1
Hieracium villosum · + · + · · · · · · · · · · · · ·
Homogyne alpina · · · · · + + · · + · · · · · 1 ·
Huperzia selago · · · · · · · · · + · + · · · · ·
Hypericum richeri subsp. grisebachii · + · · · · · · · · + + · 1 + + ·
Hypochoeris uniflora · · · · · · · · · · · · · + + · +
Juncus trifidus · · · · · · · · 2 2 · · 3 1 · · ·
Juniperus communis subsp. alpina · · · · · · · · 2 · · · · · + · 2
Knautia dipsacifolia · + + · · · · · · · · · · · · · ·
Laserpitum alpinum · · + · · · · · + · + · · + · + 1
167Укр. бот. журн., 2017, 74(2)
Number of locality 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17
Leontopodium alpinum · · + + · · · · · · · · · · · · ·
Leucanthemum vulgare subsp. alpicola · · · · · · · · + · · · · · + · ·
Linum extraaxillare · · + + · · · · · · · · · · · · ·
Lotus corniculatus · · 1 + · · · · · · · · · + + · ·
Luzula luzuloides · 2 2 1 1 · + · 2 · 3 · · · · · ·
Ligusticum mutellina · · · · · + + · · · + 2 2 · · · ·
Minuaria verna subsp. gerardii · · + + · · · · · · · · · · · · ·
Narcissus angustifolius · · · · · · · · · · · · · · · + ·
Nardus stricta · · · · · + · + · · · · · · · · ·
Pedicularis hacquetii · + · · + · · · · · · · · · · · ·
Phyteuma orbiculare · + + · · · · · · · · · · · · · ·
Phyteuma vagneri · · · · · · + · + · + · · + + + ·
Poa granitica subsp. disparilis · · + · · · · 2 · · · 1 3 · + · ·
Polygala amara subsp. brachyptera · · · + · · · · · · · · · · · · ·
Polygonum bistorta · · · · · + · · · · + · · · · + ·
Potentilla aurea · · + · · · + · · · · · + · + · ·
Potentilla crantzii · · · · · · · · · · · · · 1 · · ·
Primula elatior · + · + + · · · · · · · · · · · ·
Primula halleri · + + + · · · · · · · · · · · · ·
Primula minima · · · · · · · · · + · · · · · · ·
Pulsatilla alba · · · · · · 1 · + 1 · · · + + · ·
Ranunculus thora · + + · + · · · · · · · · · · · ·
Rhodiola rosea · + · + · · · · · · · · · · · · ·
Rhododendron myrtifolium · · · · · · 1 · 1 4 · · · · + · ·
Rumex carpaticus + · · · · · · · · · + · · · · · ·
Salix herbacea · · · · · · · · · 2 · 4 · · · · ·
Saxifraga paniculata · · · + 1 · · · · · · · · · · · ·
Scabiosa lucida subsp. barbata · + + · · · · · · · · · · 1 + · ·
Scorzonera rosea · · · · · · + · · · + · · · · + ·
Sedum alpestre · + + · · · · · · · · · 4 + + · +
Sempervivum carpathicum subsp. carpathicum · · · · · · · · + · · · · · + · ·
Silene dubia · + + · · · · · · · · · · · · · ·
Silene vulgaris · + + + · · · · · · · · · · + · ·
Soldanella hungarica · · · · · + · · · · · 2 · · · · ·
Solidago virgaurea subsp. minuta · · · · · · · · + · + + · · + · ·
Thymus alpestris · + + + · · · · · · + · · + + · 1
Vaccinium myrtillus · · + · · + · · 1 · · + 2 1 + · ·
Vaccinium vitis-idaea · · · · · · · · · 1 · · · · + · ·
Valeriana tripteris + + · · + · · · · · · · · · · · ·
Veronica baumgartenii · · · · + · · · · + · · · · · · ·
Veronica bellidioides · · · · · · · · · · · · · · + · ·
BRYOPHYTES
Polytrichum sexangulare · · · · · · · 3 · 1 · · · · · · ·
Racomitrium heterostichum · · · · · · · 4 · 3 · · · · · · ·
Sphagnum russowii · · · · · · · · · 2 · 3 · · · · ·
Tortella tortuosa · 2 · 2 · · · · · · · · · · · · 3
Other species occurring in a single locality (number in brackets) – Adenostyles alliariae (1): 2; Alchemilla sp. (5): +; Asplenium
septentrionale (17): +; Caltha laeta (6): +; Campanula serrata (17): +; Cardaminopsis halleri (5): +; Carex atrata (2): +;
C. canescens (6): 1; C. echinata (6): 2; C. nigra (6): 1; C. ornithopoda (17): +; C. rostrata (6): 3; Carlina acaulis (4): +; Centaurea
mollis (2): +; Cicerbita alpina (1): 1; Cirsium waldsteinii (1): 2; Clematis alpina (2): +; Dianthus compactus (7): +; Eriophorum
polystachyon (6): 3; E. vaginatum (6): +; Geranium sylvaticum subsp. alpestre (1): +; Gymnadenia conopsea (15): +; Leontodon
croceus (15): +; Lilium martagon (2): +; Luzula alpino–pilosa (12): +; L. sudetica (6): +; Molinia caerulea (6): 2; Parnassia
palustris (3): +; Picea abies (16): 2; Polygonum viviparum (5): +; Pulmonaria filarszkyana (2): +; Ranunculus carpaticus (1):
+; Sedum telephium subsp. fabaria (14): +; Senecio nemorensis (1): +; Symphytum cordatum (2): +; Thesium alpinum (14): +;
Vaccinium gaultherioides (15): +; Veratrum lobelianum (12): +; Viola biflora (5): +; V. declinata (7): +; Cetraria islandica (10):
1; Dicranum scoparium (10): 3; Grimmia anodon (17): 1; Homalothecium philippeanum (17): 2; Hylocomium splendens (10): 3;
Lophozia excisa (13): 4; Pohlia nutans (13): 3; Polytrichastrum alpinum (12): 3; Pseudoleskeella nervosa (17): 1; Sphagnum sp. (6): 4.
Location of the sites: 1 – 47°57′44,9′′N 24°27′53,0′′E; 2 – 47°57′54,2′′N 24°27′21,6′′E; 3 – 47°57′56,9′′N 24°27′18,6′′E; 4 –
47°57′59,1′′N 24°27′17,0′′E; 5 – 47°58′07,0′′N 24°27′09,7′′E; 6 – 47°54′58,2′′N 24°20′58,0′′E; 7 – 47°55′05,4′′N 24°20′20,6′′E;
8 – 47°55′19,3′′N 24°19′48,7′′E; 9 – 47°55′39,5′′N 24°16′41,0′′E; 10 – 47°55′15,8′′N 24°19′48,6′′E; 11 – 47°55′83,0′′N
24°19′38,9′′E; 12 – 47°55′27,6′′N 24°19′37,4′′E; 13 – 47°55′35,4′′N 24°19′38,0′′E; 14 – 47°55′33,0′′N 24°19′31,4′′E; 15 –
47°55′35,2′′N 24°19′30,5′′E; 16 – 47°54′27,2′′N 24°17′01,1′′E; 17 – 47°54′20,6′′N 24°16′39,5′′E.
168 Ukr. Bot. J., 2017, 74(2)
past data (Klášterský, 1931; Deyl, 1940) shows that the
species was much more abundant there in the 1930s,
and the lowermost limit of its distribution has shifted
about 100 m upwards in the altitudinal terms since then.
This proves that currently V. bellidioides is at the brink
of extinction in Ukraine. Such dynamics is in line with
the decline of the species in the Sudetes (Szczęśniak
et al., 2011) at comparatively low elevations, as well as
upward shifts of its altitudinal limits in the Alps (Pauli
et al., 2007; Frei et al., 2010), which apparently have
resulted from climate change.
Another cold-adapted species, Luzula spicata L., has
also demonstrated vulnerability to climate change and
consequent decline in the Alps (Pauli et al., 2007; Frei
et al., 2010). It was reported from the very top of Mt.
Pip Ivan at the end of the 19th century by H. Zapałowicz
(1889), but we failed to find it in the Marmarosh Mts.
Most probably, L. spicata, which was documented in
several highest massifs of the Ukrainian Carpathians in
the late 19th – early 20th century, has become completely
extinct in the region by now, because it has not been
confirmed anywhere in Ukraine since World War II.
We also did not manage to reconfirm the occurrence
of another chionophylous species, Saxifraga carpatica
Sternb., which is documented by an old herbarium
specimen from Mt. Pip Ivan (LWS 46140).
According to the European compendia on
ecological indicator values (Ellenberg et al., 1992;
Zarzycki et al., 2002), the above mentioned species,
which show declining trends in their distribution in
the Marmarosh Mts (Cerastium cerastoides, Luzula
spicata, Primula minima, Saxifraga carpatica, Veronica
bellidioides), belong to the most cold-adapted plants
in the Carpathian flora. Their Marmarosh habitats are
restricted to the lowermost limits of their ecological
ranges and the ongoing climate change has an adverse
impact on them.
We managed to find the locality of an arctic-alpine
saxicolous species Potentilla crantzii (Crantz) Beck
ex Fritsch (locality 14) on a steep rocky slope at the
NW summit of Mt. Pip Ivan, which was reported by
M. Deyl (1940) but has not been reconfirmed ever since.
The area of the site is only about 10 m². The species
produces compact clones with numerous subprostrate
flowering shoots.
An East-South-Carpathian endemic Anthemis
cretica L. subsp. pyrethriformis (Schur) Govaerts (=
A. carpatica Waldst. & Kit. ex Willd. subsp. pyrethriformis
(Schur) Prodan) is restricted to the high-mountain
zone of Mts Gropa, Pip Ivan and Shcherban. This is
mainly by shrinkage of its snowbed habitats due to
decreasing thickness and duration of snowpack that
has been described in the Carpathians recently (Micu,
2009). Interestingly, M. Deyl (1940) admitted that
some largest snowbeds on Mt. Pip Ivan even persisted
throughout some growth seasons in the 1930s, though
that has never happened in the last decades.
The aforementioned chionophylous species are
adapted to long-lasting snow cover, short growth period
and are confined to snowbed communities that belong
to the class Salicetea herbaceae Br.-Bl. 1948. They are
low-competitive and most viable on moss-dominated or
barren patches with initial soil. As a result of warming-
induced succession they are being gradually replaced
by more competitive sward-forming cold-tolerant
graminoids, for instance Poa granitica Br.-Bl. subsp.
disparilis (Nyár.) Nyár. (= P. deylii Chrtek & V. Jirásek).
The latter taxon, which is endemic to the Eastern and
Southern Carpathians, is rather common in the alpine
zone of Mt. Pip Ivan.
Primula minima L. is also a cold-adapted low-
statured species, which forms patches of 25–40 rosettes
per 100 cm². It occurs at saxicolous alpine habitats up to
the highest elevations, while the lowest locality that we
found on Mt. Shcherban is on 1750 m a.s.l. Interestingly,
M. Deyl (1940) reported the species locality situated
100 m lower, while H. Zapałowicz (1889) – as low as
on 1580 m a.s.l.
A frequent component of many alpine communities
is Sedum alpestre Vill. It is noteworthy that in one of the
glacial cirques of Mt. Pip Ivan (locality 13) this species
dominates in the snowbed community and forms dense
mats that cover up to 0.25 m², which is uncommon for
other regions of the Carpathians.
It is remarkable that in population replenishment
of all the aforementioned alpine cold-tolerant species
vegetative reproduction markedly prevails over seed
recruitment. That provides for their better persistence
in harsh climatic conditions.
One of the rarest and most endangered species,
Veronica bellidioides L., occurs on the ridge at the NW
summit of Mt. Pip Ivan, which is its only locality still
remaining in the Ukrainian Carpathians (Kobiv, 2009).
The area of its saxicolous habitat is merely about 15 m²
(locality 15). Monitoring of the population showed that
the number of flowering individuals has decreased in
14 years from 25 to only 3 specimens by 2016, which is
critical, because seed recruitment is crucial for further
survival of the species. Moreover, the density of its
seedlings has decreased dramatically. Analysis of the
169Укр. бот. журн., 2017, 74(2)
mostly on the Romanian side. The species forms lax
clones with about 1.5 flowering stems per 100 cm².
Gentiana punctata L., a Central European alpine
species, is very common in the Marmarosh Mts. Due to
clonal growth it may form clumps up to 3 m² with 8–10
flowering shoots per 1 m².
As follows from the above data, the massif of Mt.
Pip Ivan with the adjacent territory is a remarkable
hotspot, where localities of many rare alpine species are
concentrated on a comparatively small area (Figure).
Another unique area comprises Mt. Petros and
Mt. Berlebashka (aka Latundur). A series of alpine
calcareous species occur here (Table 2), because
limestone bedrock lies at a considerable altitude, which
is unusual for the Ukrainian Carpathians. One of them
is Genista tinctoria L. subsp. oligosperma (Andrae) Jáv.,
an East-South-Carpathian endemic, which does not
occur anywhere else in the Ukrainian Carpathians.
It is a mat-forming dwarf shrub, 10–15 cm high. The
mats cover up to 300 cm² and contain 20–40 flowering
shoots. It is restricted to only two small localities (#22,
28) in the subalpine zone. The area of each of them is
ca. 150–200 m². However, as follows from M. Deyl's
(1940) monograph, the taxon inhabited a larger range
of localities in the 1930s, which leads to an assumption
that it has declined significantly by now.
The same habitats harbor Jovibarba globifera (L.)
J. Parnell subsp. preissiana (Domin) Holub, a West-
East-Carpathian subendemic. Besides one locality on
Mt. Berlebashka, its largest metapopulation inhabits
the rocks on Mt. Petros and consists of several scattered
subpopulations. They are replenished mainly by stolon
growth. Numbers of rosettes of J. globifera subsp.
preissiana ranges within 5–35 specimens in each
subpopulation, while the percentage of flowering ramets
may reach up to 22%.
Similar metapopulation pattern and habitat
requirements refer to other clonal high-mountain
saxicolous calcareous species, Veronica fruticans Jacq.
and Potentilla crantzii, which often occur together.
Altitudinal range of Jovibarba globifera subsp. preissiana,
V. fruticans, and P. crantzii on the rocks of the S and E
aspects of Mt. Petros is 1530–1755 m a.s.l. (localities
18, 19, 21, 22). Monitoring carried out in selected
localities showed considerable increase in the numbers
of flowering individuals of J. globifera subsp. preissiana
and V. fruticans (2.6–4 and 2.8 times, respectively) in
14 years. This can be caused by the obvious decrease of
grazing during that period.
the only area in Ukraine where it occurs. Apparently,
A. cretica subsp. pyrethriformis forms a metapopulation
comprised of a number of local populations scatthered
within a rather narrow stripe along the Ukrainian-
Romanian border. This suffruticose plant occurs on
rocky and grassland habitats mostly in the alpine zone.
Its highest density (up to 20 flowering individuals per
m²) was noted alongside of the tourist path (locality 7)
in the gaps among the heath or herbaceous vegetation,
which result from moderate trampling. Such gaps
provide suitable microhabitats for seed recruitment
of A. cretica subsp. pyrethriformis (Kobiv, 2012). Its
lowermost locality is situated at 1520 m a.s.l. at the SW
foot of Mt. Shcherban, where the density of individuals
has decreased by 1.8 times over the 14-year monitoring
period.
Rhododendron myrtifolium Schott & Kotschy, a
narrow-range high-mountain species, has a similar
distribution pattern. It is very abundant in the massif
of Mt. Pip Ivan and occurs up to the highest elevations
(localities 10, 15).
A large metapopulation of a pan-Carpathian endemic
Sempervivum carpathicum Wettst. ex Prodan subsp.
carpathicum (= S. montanum L. subsp. carpathicum
(Wettst. ex Prodan) Wettst. ex A. Berg.) inhabits rocky
outcrops in the high-mountain zone of Mt. Pip Ivan
within about 5-hectare area. The species produces mats
of numerous rosettes that may cover up to 300 cm².
The population is sustained primarily by vegetative
reproduction provided by stolon growth.
Gentiana lutea L., a high-mountain Central European
species, is widely spread in glacial cirques on NE slopes
of Mt. Pip Ivan and Mt. Gropa within 1600–1900 m
a.s.l. The species occurs either in the open grassland
habitats or at the edge of scrub vegetation. The largest
population on Mt. Pip Ivan (locality 11) covers ca.
4 hectares and includes 3.5–5.5 thousand flowering
individuals. Density of flowering shoots in clusters may
reach 4 specimens per m².
On the saddle between Mt. Shcherban and Mt.
Polonynka (locality 16) we found Narcissus angustifolius
Curtis, a species protected by the Bern Convention.
Its population, which numbers ca. 100 flowering
individuals occupies about 30 m².
Not far from that site, on the rocks at the W summit of
Mt. Polonynka we found a small locality of Campanula
carpatica Jacq., a pan-Carpathian endemic (locality
17). It occurs on an area of only about 2 m², apparently
being a fragment of the large metapopulation situated
170 Ukr. Bot. J., 2017, 74(2)
Table 2. Floristic composition in localities of rare plant species in the hotspots of Mts Petros-Berlebashka and the Bilyi Potik Valley
(names of rare species in boldface)
Number of locality 18 19 20 21 22 23 24 25 26 27 28 29 30
Location Mt. Petros Mt. Berlebashka
Bilyi Potik
Valley
Date of relevé (day, month, year)
28
.0
7.
02
28
.0
7.
02
28
.0
7.
02
03
.0
7.
10
03
.0
7.
10
28
.0
7.
02
26
.0
7.
16
28
.0
7.
02
04
.0
7.
10
04
.0
7.
10
26
.0
7.
16
29
.0
7.
16
24
.0
7.
02
Area of relevé (m²) 25 5 100 50 100 25 10 100 100 25 100 25 25
Elevation (m a.s.l.)
15
55
15
30
17
65
17
55
17
10
16
35
16
15
15
45
15
30
15
70
16
45
60
0
58
5
Exposition NE E N S S SW SW SE E E S E SE
Inclination (°) 20 40 30 30 40 40 5 3 10 30 40 15 40
Vegetation cover (%) 30 30 100 50 60 40 95 95 95 70 60 100 30
Achillea stricta · · · + 1 · · · · · + · ·
Acinos alpinus · · · · · · · · + + · · ·
Aconitum anthora · · · · · + · · · · · · ·
Anthoxanthum alpinum · · · · 1 · · 1 · · 1 · ·
Aruncus sylvestris · · · · · · · · · · · + 2
Asplenium trichomanes · · · · · · · · · · · + +
Asplenium viride · · · · · + · · · + · · ·
Athyrium distentifolium · · · · · · · · · · · 2 2
Avenula pubescens · · · · · · · · + + · · ·
Bellardiochloa violacea · · · · 2 · · · · · · · ·
Calamagrostis arundinacea 2 1 · · 1 · · · · · · · ·
Calamagrostis villosa · · 4 · · · · · · · 2 · ·
Campanula carpatica · 2 · · · + · · · · · · 2
Campanula kladniana · · · + · · · · · · 1 · ·
Campanula serrata · · · · + · · · · · + · ·
Carduus kerneri · · · + + + · · · · + · ·
Carex pauciflora · · · · · · · 3 · · · · ·
Carex sempervirens · · · 1 1 · · · 2 2 · · ·
Carlina acaulis · · · · · · · · + · + · ·
Chrysosplenium alternifolium · · · · · · · · · · · + ·
Cortusa matthioli · · · · · · · · · · · · +
Cotoneaster integerrimus · 1 · 1 · · · · · · · · ·
Cruciata glabra · · · · + 1 · · + + · · ·
Dactylorhiza cordigera · · · · · · · + · · · · ·
Dactylorhiza fuchsii · · · · + · · · · · · · ·
Daphne mesereum · · · · · + · · + + · · ·
Deschampsia caespitosa · · + · · · 2 2 + · · · ·
Dianthus carthusianorum subsp. tenuifolius + · · + · · · · · · + · ·
Euphorbia carpatica · · · · · · · · + + · · ·
Fagus sylvatica · · · · · · · · · · · 3 2
Festuca airoides 3 2 · 1 2 · · · · 2 3 · ·
Festuca rubra 2 · · · · · · · 2 3 · · ·
Festuca saxatilis · · · · 1 · · · · · · · ·
Fragaria vesca · · · · · · · · + + · · ·
Genista tinctoria subsp. oligosperma · · · · + · · · · · 1 · ·
Gentiana acaulis · · · · + · · · · · · · ·
Gentiana lutea · · 1 · · · · · · · · · ·
Helianthemum nummularium subsp. grandiflorum · · · + · · · · · · · · ·
Heracleum carpaticum · · · · · · 1 · · · · · ·
Hieracium umbellatum · · · · · · · · · + + · ·
Hypochoeris uniflora · · · · + · · · · · + · ·
Jovibarba globifera subsp. hirta + + · + · · · · · · + · ·
Juniperus communis subsp. alpina 2 2 1 2 1 2 2 · + + 1 · ·
Laserpitum alpinum + · + + + · · · · + + · ·
171Укр. бот. журн., 2017, 74(2)
Number of locality 18 19 20 21 22 23 24 25 26 27 28 29 30
Linum extraaxillare · · · · + · · · · 1 · · ·
Lotus corniculatus · · · + · · · · + · · ·
Luzula luzuloides · · 1 + 1 · 3 · · · 2 · ·
Nardus stricta · · · · · · · 1 · · 1 · ·
Parnassia palustris · · · · · · · + · + · · ·
Phyllitis scolopendrium · · · · · · · · · · · 2 ·
Phyteuma orbiculare · · · · · · · · + + · · ·
Phyteuma vagneri · · · · + · · · · · + · ·
Picea abies 2 2 + · · 2 1 · · · · · ·
Poa nemoralis 1 1 · · 2 2 · · · 3 · · ·
Polygala amara subsp. brachyptera · · · · · · · · + + · · ·
Potentilla aurea · · · + + · · · · · · · ·
Potentilla crantzii 1 + · 1 + + · · · + · · ·
Prunella vulgaris · · · · · · + · + + · · ·
Rhododendron myrtifolium · · · · · · · · · · + · ·
Saxifraga adscendens · · · · · · · · + · · · ·
Saxifraga paniculata · 1 · 1 · 1 · · · · · · ·
Scabiosa lucida subsp. barbata · · · + + + · · 1 + · · ·
Scopolia carniolica · · · · · · · · · · · 1 +
Scorzonera rosea · · · · + · · · · · + · ·
Sedum telephium subsp. fabaria 1 · · · · 1 · · · · · · ·
Silene dubia · + · + + · · · · + + · ·
Stellaria holostea · · · · + · · · · · + · ·
Stellaria nemorum · · · · · · + · · · · 1 ·
Thymus alpestris 2 2 · 1 2 1 + · + 2 1 · ·
Vaccinium myrtillus 1 1 1 1 2 2 · · · · 1 · ·
Vaccinium vitis-idaea · · · · + · · · · · + · ·
Veronica fruticans + · · 1 + · · · · · · · ·
Veronica officinalis · · · · · · + · · + · · ·
Veronica urticifolia · · · · · + + · · · · · 2
Viola declinata · · · · · · · · · + + · ·
BRYOPHYTES
Ctenidium molluscum · · · · · 2 · · · · 2 · 2
Ditrichum flexicaule 1 1 · 2 2 · · · · · 2 · ·
Rhytidiadelphus triquetrus · 1 · 3 2 · · · · · · · 2
Tortella tortuosa 1 1 · · · 2 · · · · 3 · 2
Other species occurring in a single locality (number in brackets) – Alchemilla monticola (26): +; Alnus viridis (22): +; Aposeris
foetida (22): +; Asplenium septentrionale (19): +; Astrantia major (26): 1; Bellis perennis (26): +; Caltha laeta (25): +; Campanula
abietina (24): +; C. glomerata (21): +; Cardaminopsis arenosa (23): +; Carex canescens (25): 2; C. echinata (25): 2; C. nigra
(25): 1; Chrysosplenium alternifolium (29): +; Clematis alpina (30): 1; Corylus avellana (29): 3; Crepis conyzifolia (21): +;
Cystopteris fragilis (19): +; Digitalis grandiflora (30): 1; Dryopteris carthusiana (23): 2; Epilobium collinum (23): +; E. roseum
(29): +; Eriophorum polystachyon (25): 3; Galeobdolon luteum (29): 1; Geranium robertianum (29): +; Glechoma hederacea (29):
1; Hieracium auranthiacum (24): +; H. pilosella (27): +; H. richeri subsp. grisebachii (20): +; Hypericum maculatum (27): +;
Impatiens noli–tangere (29): 2; Juncus conglomeratus (25): 1; J. trifidus (28): 2; Knautia dipsacifolia (26): 1; Lamium maculatum
(29): +; Leucanthemum vulgare subsp. alpicola (21): +; Lilium martagon (21): +; Ligusticum mutellina (25): +; Lunaria rediviva
(29): 1; Luzula sudetica (25): 1; L. sylvatica (24): +; Lysimachia nummularia (29): +; Melampyrum herbichii (21): +; Molinia
caerulea (25): 2; Oxalis acetosella (24): +; Petasites albus (29): 2; Phleum alpinum (24): +; Polypodium vulgare (30): 2; Potentilla
erecta (25): +; Ranunculus nemorosus (22): +; Rubus caesius (29): 2; R. idaeus (23): +; Salvia glutinosa (30): 2; Sambucus nigra
(29): 2; Sedum atratum (18): +; Senecio papposus (22): +; Solidago virgaurea subsp. minuta (20): +; Spiraea chamaedryfolia
(30): 1; Stachys sylvatica (29): +; Symphytum tuberosum (23): +; Tanacetum subcorymbosum subsp. clusii (21): +; Thesium
alpinum (18): +; Trifolium repens (26): +; Urtica dioica (29): 2; Vaccinium gaultherioides (22): +; Valeriana tripteris (18): +;
Veronica serpyllifolia (27): +; Plagiochila asplenioides (21): 1; Neckera crispa (30): 1; Sphagnum sp. (25): 4. Location of the
sites: 18 – 47°57′30,5′′N 24°20′23,5′′E; 19 – 47°57′33,5′′N 24°20′30,5′′E; 20 – 47°57′46,5′′N 24°20′17,9′′E; 21 – 47°57′42,4′′N
24°20′11,8′′E; 22 – 47°57′40,0′′N 24°20′10,9′′E; 23 – 47°57′35,9′′N 24°20′05,6′′E; 24 – 47°57′34,3′′N 24°20′04,5′′E;
25 – 47°57′16,9′′N 24°19′38,8′′E; 26 – 47°56′51,9′′N 24°19′04,9′′E; 27 – 47°57′54,3′′N 24°19′01,7′′E; 28 – 47°57′06,3′′N
24°18′58,3′′E; 29 – 47°56′08,9′′N 24°15′26,1′′E; 30 – 47°55′49,4′′N 24°15′22,0′′E.
172 Ukr. Bot. J., 2017, 74(2)
in 2016, whereas C. carpatica is more common on the
rocks in the forest zone.
On the same limestone bedrock nearby the above-
mentioned habitat we found a small population
of Heracleum carpaticum Porcius, an East-South-
Carpathian endemic (locality 24). It occupied only ca.
10 m² in 2016 and contained 6 flowering and 19 vegetative
individuals. This aclonal species is replenished only due
to seed reproduction. The habitat conditions are rather
unusual for that alpine species, which is confined to a
forest glade there at a comparatively low elevation.
Another East-South-Carpathian endemic, Silene
nutans L. subsp. dubia (Herbich) Zapał., is rather com-
mon on Mt. Petros and Mt. Berlebashka and forms a
large metapopulation. Helianthemum nummularium (L.)
Mill. subsp. grandiflorum (Scop.) Schinz & Thell.,
a high-mountain Central European subspecies, was
found only in one locality (#21) on Mt. Petros.
A large population of Gentiana lutea stretches along
steep hollows on the N and NE slopes of Mt. Petros near
its summit (locality 20). The population is confined
to 1660–1770 m a.s.l., occupies about 5000 m² and
numbers 1.5–2.0 thousand flowering individuals.
Polygala amara L. subsp. brachyptera (Chodat) Hayek,
a narrow-range montane taxon, occurs in some calcium-
rich saxicolous or grassland sites (localities 4, 26, 27).
The last noteworthy "hotspot" of the rare species is
the Bilyi Potik Valley, which is remarkable for numerous
cliffs of marbelized limestones situated in the beech
forest zone at much lower elevations in comparison with
the listed above localities. Campanula carpatica is very
abundant on such cliffs and smaller outcrops and forms
lax mats, with up to 3 flowering shoots per 100 cm².
Cortusa matthioli L., a saxicolous chasmophite,
inhabits shady rock fissures. Its largest population
(locality 30) contains over 100 flowering individuals. The
species number is limited by the availability of suitable
microhabitats and low seed productivity (Kobiv, 1999).
Some publications (Shushman, 2008) and herbarium
data indicate that a very rare saxicolous species,
Sempervivum marmoreum L., also occurs on the cliffs
near the Bilyi Potik stream, but we failed to find it.
Tall-forb communities beneath the rocks harbor
Phyllitis scolopendrium (L.) Newman and Scopolia
carniolica Jacq. Their average density is 1.2 and 0.4
individuals per 1 m², respectively (locality 29).
Calcicolous flora of these habitats is typical of the
belt of Pieniny and Marmarosh limestone cliffs, which
stretches along the border between the Inner and Outer
Carpathians.
We also discovered other positive examples of some
rare species dynamics, namely establishment of their
new populations on limestone bedrock on the S slope
of Mt. Berlebashka. Thus, in 2010 we found a few
individuals of Potentilla crantzii on anthropogenic
outcrops of limestone that had appeared on the side-
slope of a recently constructed road (locality 27). The
species was missing there in 2002, whereas its quite
viable population has developed at the site by 2016.
Most probably, P. crantzii has dispersed from its large
«mainland» population on Mt. Petros situated ca. 2 km
eastwards.
A newly established population of Acinos alpinus (L.)
Moench (= A. baumgartenii (Simk.) Klokov) was first
found near that site on a pasture (locality 26) in 2010.
It occupied about 500 m² area then, but has expanded
to ca. 2000 m² by 2016. However, the average density
of flowering individuals of A. alpinus has not changed
significantly remaining within 1.1–1.3 specimens
per m².
Quite opposite dynamics was revealed for Saxifraga
adscendens L., an arctic-alpine species that occurred in
the same locality in 2001, but was not found there in the
later surveys.
The same refers to Koeleria macrantha (Ledeb.)
Schult. subsp. transsilvanica (Schur) A. Nyár., an East-
South-Carpathian endemic, which was reported from
the S slope of Mt. Petros by M. Deyl (1940), but was never
confirmed later. That author also indicated the altitude
of the locality and published its phytosociological
relevé, with a very peculiar floristic composition, which
enabled to locate the site correctly (locality 22) and to
make sure that K. macrantha subsp. transsilvanica has
become extinct in its only known Ukrainian habitat. Its
extinction could be caused by considerable decrease of
grazing and consequent overgrowth of its habitat with
more competitive graminoids or shrubs.
Nevertheless, Festuca saxatilis Schur, another East-
South-Carpathian endemic, has persisted in the same
locality, which is the only one we managed to find in
the region. This saxicolous species forms dense tufts
and occurs within about 25 m². That locality is also
inhabited by a large population of Bellardiochloa
violacea (Bellardi) Chiov.
A limestone rock in the spruce forest at the SW foothill
of Mt. Petros provides a habitat for some rare calcareous
species, e.g. Aconitum anthora L. (= A. jacquinii Rchb.),
Campanula carpatica, Symphytum tuberosum L. (locality
23). The first of them is very rare in the region and its
population numbered only 14 flowering individuals
173Укр. бот. журн., 2017, 74(2)
frontier regions of the Ukrainian Carpathians are:
Festuca saxatilis, Jovibarba globifera subsp. preissiana,
Sempervivum marmoreum, Veronica fruticans. Some of
the listed species, for instance, Veronica bellidioides,
are critically endangered, while Koeleria macrantha
subsp. transsilvanica, has aready become extinct
in the Marmarosh Mts. The localities of these taxa
in the Marmarosh Mts are confined to the north-
eastern limit of their distribution in Central Europe
and their extinction causes contraction of the species’
geographical ranges and possible irreversible loss of
biodiversity, because some peripheral populations
have a unique gene pool (Lesica, Allendorf, 1995). As
follows from the above results, many rare species of the
Marmarosh Mts are narrow-range and the mentioned
facts of decline or extinction are most worrying for the
endemics (Genista tinctoria subsp. oligosperma, Koeleria
macrantha subsp. transsilvanica). Decline of cold-
adapted alpine species (Cerastium cerastoides, Luzula
spicata, Primula minima, Saxifraga carpatica, Veronica
bellidioides) has taken place mainly at the lower limit of
their ecological range and can be explained by climate
change.
Another factor that affects rare species is
considerable decrease of grazing in the region, which
has been observed lately. Along with climate change it
triggers natural succession of vegetation in the habitats
of some rare species and their replacement by highly
competitive tussock graminoids or shrubs. For that
reason, mild grazing regime that still remains in some
open habitats of saxicolous or chionophylous species,
e.g. on Mts Nenieska, Pip Ivan, Petros, Berlebashka
is favorable for their persistence, because it mitigates
adverse changes in these localities. This refers mostly
to low-statured poorly competitive species (e.g. Acinos
alpinus, Cerastium cerastoides, Saxifraga adscendens,
S. carpatica, Veronica bellidioides), which need gaps in
vegetation for their recruitment (Virtanen et al., 1997).
The mitigating effect of grazing against warming-
induced decline of alpine species has been proved
experimentally (Klein et al., 2004).
On the other hand, another group of species has
benefited from the decrease of grazing. They demonstrate
their highest viability in undisturbed conditions. This
Because of steep topography, wetlands are infrequent
in the Marmarosh Mts. The largest peat bogs that cover
100–750 m² (localities 6, 25) are inhabited by Carex
limosa L. and C. pauciflora Lightf.
In many of his relevés M. Deyl (1940) mentioned
Avenula pubescens (Huds.) Dumort. subsp. laevigata
(Schur) Holub, which could imply that this narrow-
range montane plant is rather common in the Marmarosh
Mts. Apparently, that results from misidentification,
because thorough examination of the specimens, which
we collected in the region (including the localities
stated by that author) and their comparison with the
type material (Nachychko et al., 2017) proved that they
belong to A. pubescens subsp. pubescens, and the former
taxon hardly occurs in Ukraine.
Soil analysis in the selected localities of rare species
(Table 3) showed the highest pH values and content of
calcium on the limestone cliffs in the Bilyi Potik Valley.
Rocky habitats of Mt. Nenieska are also rich in calcium.
By contrast, most acidic and calcium-poor soil is in
the snowbed locality in the glacial cirque of Mt. Pip
Ivan. Almost equal content of calcium and magnesium
was noted in some localities on crystalline bedrock
on Mt. Pip Ivan and Mt. Petros, whereas the former
element prevails significantly in most soils all over
the Ukrainian Carpathians. Apparently, presence of
markedly basiphilous species in some of these localities
is considerably contributed by magnesium.
Conclusion
In biogeographical terms, the Marmarosh Mountains
are the area of remarkable biodiversity, where many
rare plants occur. Some of them, namely Anthemis
cretica subsp. pyrethriformis, Genista tinctoria subsp.
oligosperma, Potentilla crantzii and Veronica bellidioides,
do not occur elsewhere in Ukraine, because edges of
their geographical ranges slightly protrude into the
Ukrainian territory only in that area, while they are
much more abundant in the Romanian Carpathians.
That floristic specificity is caused mostly by the
geological structure of the region, which differs from
the rest of the Ukrainian Carpathians. Other rare
range-edge species that in addition to the Marmarosh
Mts have a very limited distribution in some other
Table 3. Soil parameters in selected localities of rare plant species in the Marmarosh Mts
Number of locality 2 3 5 8 9 14 15 17 21 22 28 30
рН(Н
2
О) 5.86 6.78 6.56 3.75 4.15 3.91 4.76 5.21 3.86 4.40 6.19 7.05
Content of Ca (mg-eq./100 g) 34.83 12.82 24.40 1.52 1.95 9.80 12.42 13.60 9.60 10.81 18.40 36.24
Content of Mg (mg-eq./100 g) 6.02 3.22 4.80 1.50 1.62 8.15 4.01 11.21 8.00 4.82 6.40 17.31
174 Ukr. Bot. J., 2017, 74(2)
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concerns primarily tall forbs (Heracleum sphondylium
subsp. transsilvanicum, Gentiana punctata), but also
refers to some rare saxicolous components of grassland
or dwarf-shrub communities (Jovibarba globifera subsp.
preissiana, Veronica fruticans).
Thus, various groups of rare species have different
needs for their effective conservation, which should
be carefully considered in implementation of land-use
measures in their habitats. Supposedly, their persistence
in open saxicolous and chionophilous localities could
be best provided by maintenance of mild grazing regime
to prevent overgrowth of their habitats with highly
competitive species.
By contrast with the above-mentioned extinction
events, we have revealed a few facts of colonization
of new habitats by some rare species (Acinos alpinus,
Potentilla crantzii), which happened recently in
anthropogenically disturbed localities.
Thus, populations of many rare species in the
Marmarosh Mts undergo continuing transformations
responding to changes in climatic factors and varying
anthropogenic pressure or resulting from succession of
vegetation in their habitats.
Almost all species mentioned in this article are listed
in the Red Data Book of Ukraine (Chervona knyha…,
2009). Our survey proved that some studied taxa should
be included into its next edition, namely, Koeleria
macrantha subsp. transsilvanica and Luzula spicata,
both as extinct (EX); however, a few historical localities
of the latter species in the Chornohora still need to be
thoroughly checked for its presence.
Though Potentilla crantzii is rare and confined to a
very small area in Ukraine, it is not prone to decline and
therefore does not require any conservation status.
Acknowledgements
We are very grateful to I. Rabyk, Institute of Ecology of the
Carpathians, NASU for determining the bryophytes.
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176 Ukr. Bot. J., 2017, 74(2)
Кобив Ю.1,2, Прокопив А.2, Начычко В.2, Борсукевич Л.2,
Гелеш М.2 Распространение и состояние популяций
редких видов растений в Мармарошских горах
(Украинские Карпаты). Укр. бот. журн., 2017, 74(2):
163–176.
1Институт экологии Карпат НАН Украины
ул. Козельницкая, 4, Львов 79026, Украина
2Ботанический сад Львовского национального
университета им. Ивана Франко
ул. Черемшины, 44, Львов 79014, Украина
Представлены результаты обследования местопроиз-
растаний редких видов растений в Мармарошских горах
(Украинские Карпаты) и описаны их точное расположе-
ние и популяционные параметры. В результате анализа
предыдущих литературных и гербарных данных установ-
лены некоторые пространственно-временные законо-
мерности их распространения, в частности особенно-
сти местообитаний, тенденции динамики популяций,
что дало возможность определить их природоохранный
статус. Выявлены многочисленные факты их регресси-
рования или исчезновения, а также примеры положи-
тельной динамики популяций исследованных видов.
Холодоустойчивые виды (Cerastium cerastoides, Primula
minima, Veronica bellidioides) оказались уязвимыми к кли-
матическим изменениям, в частности на нижней грани-
це их распространения. Еще одним влияющим факто-
ром является ослабление выпаса, вызвавшее сукцессию
растительности и последующее вытеснение некоторых
низкоконкурентных редких растений (Genista tinctoria
subsp. oligosperma, Koeleria macrantha subsp. transsilvanica)
кустарничками или дерновинными злаками. Однако,
другая группа редких видов (Heracleum sphondylium subsp.
transsilvanicum, Gentiana punctata, Jovibarba globifera subsp.
preissiana, Veronica fruticans) положительно реагирует на
уменьшение антропогенного влияния.
Ключевые слова: изменения климата, вымирание,
динамика популяций, локалитет, Карпаты
Кобів Ю.1,2, Прокопів А.2, Начичко В.2, Борсукевич Л.2,
Гелеш М.2 Поширення і стан популяцій рідкісних видів
рослин у Мармароських горах (Українські Карпати).
Укр. бот. журн., 2017, 74(2): 163–176.
1Інститут екології Карпат НАН України
вул. Козельницька, 4, Львів 790026, Україна
2Ботанічний сад Львівського національного
університету ім. Івана Франка
вул. Черемшини, 44, Львів 790014, Україна
У статті подано результати обстежень оселищ рідкісних
видів рослин у Мармароських горах (Українські Карпа-
ти) та описано їхнє точне розташування й популяційні
параметри. У результаті аналізу попередніх літературних
і гербарних матеріалів встановлено деякі просторово-
часові закономірності їхнього поширення, зокрема осо-
бливості оселищ і динаміки популяцій, що уможливило
визначення їхнього природоохоронного статусу. Виявле-
но численні факти регресування чи зникнення, а також
приклади позитивної динаміки популяцій досліджених
видів. Холодостійкі види (Cerastium cerastoides, Primula
minima, Veronica bellidioides) виявилися вразливими до
кліматичних змін, зокрема на нижній межі їхнього по-
ширення. Іншим фактором впливу є послаблення ви-
пасання, що спричинило сукцесію рослинності і, як
наслідок, витіснення деяких низькоконкурентних рід-
кісних рослин (Genista tinctoria subsp. oligosperma, Koeleria
macrantha subsp. transsilvanica) чагарничками або дер-
нинними злаками. Натомість, інша група рідкісних ви-
дів (Heracleum sphondylium subsp. transsilvanicum, Gentiana
punctata, Jovibarba globifera subsp. preissiana, Veronica
fruticans) позитивно реагує на зменшення антропогенно-
го впливу.
Ключові слова: зміни клімату, вимирання, динаміка
популяцій, локалітет, Карпати
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