143
Key words: habitats, petrophytic 
communities, classification, 
chasmophytic plants, ferns, 
bryophytes, ecological parameters, 
threats. 
Ključne besede: habitati, petrofitske 
združbe, klasifikacija, hazmofiti, 
praproti, mahovi, ekološki dejavniki, 
grožnje.
Corresponding author:
Iuliia Vasheniak
E-mail: arrhenatherum@gmail.com
Received: 8. 9. 2022
Accepted: 4. 4. 2023
Rare shady chasmophytic habitat 
communities (8210) in Ukraine
Yakiv Didukh1 , Illia Chorney2 , Vasyl Budzhak3 , Iuliia Vasheniak4
,
5 , 
Olesya Bezsmertna6
,
7 , Ruslan Gleb8 & Svitlana Nyporko1 
Abstract
The characteristics of shaded chasmophytic habitats on calcareous rocks in for-
est conditions are described, where the forest canopy primarily acts as a shading 
factor that determines microclimate, humidity and light levels. These communi-
ties can be found in deciduous forests of the Carpinion betuli, Fagion sylvaticae 
and Tilio-Acerion alliances, where they are confined to steep calcareous rock faces 
and cliffs. Syntaxonomically, these communities belong to the Polypodietea class 
(Ctenidio-Polypodietalia vulgaris order, Ctenidio-Polypodion vulgaris alliance) and to 
four informal communities: Isothecium alopecuroides-Polystichum braunii, Porella 
platyphylla-Asplenium ruta-muraria, Pseudanomodon attenuatus-Chrysosplenium 
alternifolium and Pedinophyllum interruptum-Polystichum aculeatum. We evaluated 
indices of environmental parameters according to the synphytoindication method 
using Didukh’s scales. An assessment of the threat impacts and indicators of 
sociological significance shows that the habitat belongs to a rare type (II class), and 
therefore requires specific conservation measures.
Iz vleček
Opisali smo značilnosti zasenčenih hazmofitskih habitatov na karbonatnih skalah 
v gozdovih, kjer gozd predstavlja dejavnik zasenčenosti in s tem določa mikro-
klimatske, vlažnostne in svetlobne značilnosti rastišča. Take združbe najdemo v 
listopadnih gozdovih zvez Carpinion betuli, Fagion sylvaticae in Tilio-Acerion, kjer 
so omejene na strmih apneniških skalah in klifih. Sintaksonomsko te združbe 
uvrščamo v razred Polypodietea (red Ctenidio-Polypodietalia vulgaris, zveza Ctenidio-
-Polypodion vulgaris) in štiri združbe: Isothecium alopecuroides-Polystichum braunii, 
Porella platyphylla-Asplenium ruta-muraria, Pseudanomodon attenuatus-Chrysosple-
nium alternifolium in Pedinophyllum interruptum-Polystichum aculeatum. Okoljske 
spremenljivke smo ovrednostili s sinfitoindikacijsko metodo z Didukhovimi indi-
katorji. Ocena ogroženosti in indikatorji sociološkega pomena kažejo, da habitat 
uvrščamo kot redek tip (razred II) in potrebuje posebne mere varovanja.
DOI: 10.2478/hacq-2022-0022 22/2 • 2023, 143–160
1 M.G. Kholodny Institute of Botany of the National Academy of Sciences of Ukraine, Kyiv, Ukraine.
2 Yuriy Fedkovych Chernivtsi National University, Chernivtsi, Ukraine.
3 Institute for Evolutionary Ecology of the National Academy of Sciences of Ukraine, Kyiv, Ukraine.
4 Vasyl’ Stus Donetsk National University, Vinnytsia, Ukraine.
5 Masaryk University, Brno, Czech Republic.
6 Taras Shevchenko Kyiv National University, Kyiv, Ukraine.
7 Tsumanska Pushcha National Nature Park, Kivertsi, Volyn region, Ukraine.
8 Carpathian Biosphere Reserve, Rakhiv, Ukraine. 

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Introduction
Petrophytic habitats and floristically poor plant commu-
nities have generally received much less attention than 
forests and grasslands, but they have recently become the 
subject of more precise analysis. They have been consid-
ered, inter alia, to be “ecological islands”, protected from 
the competition of characteristic species and the influ-
ence of various external factors (fire, human activity, 
grazing, climate change) (Pigott & Walters, 1954; Lar-
son, 2000; Theurillat & Guisan, 2001; Lavergne et al., 
2004; Speziale & Ezcurra, 2012; Tomaselli et al., 2019). 
They have also been interpreted as “ecological traps”, 
or unique ecological niches that provide conditions for 
specific, narrowly adapted species (Larson, 2000, To-
maselli et al., 2019). Nevertheless, they contribute to an 
increase in γ-floristic and phytosociological diversity and 
are thereby protected and included within the habitat 
definitions listed in the EU Habitats Directive (Council 
Directive 92/43/EEC) as 8210 “Calcareous rocky slopes 
with chasmophytic vegetation” or 8220 “Siliceous rocky 
slopes with chasmophytic vegetation”. The EUNIS clas-
sification system allocates these habitats to a separate 
class (H) (Rodwell et al., 1998). In the latest summary 
of the European vegetation classification (Mucina et 
al., 2016), communities of this habitat are divided into 
eight classes and are given high syntaxonomic status. 
The approaches of the classification of biotopes (Palae -
arctic Habitat, CORINE, EUNIS) and plant communi-
ties (EuroVegChecklist) used do not match, so from the 
point of view of developing a new classification, these 
issues need to be resolved.
Communities of this type are not large, are often frag-
mented, and characterized by habitat mosaics because of 
typological, topological and regional differences. Many 
questions thus arise, such as the delimitation of syntaxa 
and their place in the classification system, the place of 
the bryophyte component reflected in their name etc. 
At the same time, due to the paucity of species in the 
floristic composition, the emphasis in the classification 
process is not only on floristic criteria but on the specif-
ics of the ecological conditions. The essential aspect in 
this context is the question of the rocky habitat assess-
ment forming in shady conditions. According to Didukh 
et al. (2016), a habitat is an ecosystem at a topological 
level characterized by interactions between biotic and 
abiotic components that ensure structure, organization 
and integrity over an extended period and matter cycle, 
with metabolism, energy transformation, pedogenesis, 
biota existence at the population level, reproduction, 
species evolution and adaption to particular conditions. 
Petrophytic communities in forests should therefore be 
interpreted as a habitat type, different from surrounding 
forest habitats. 
Their existence is not directly related to the forest com-
munities: the forest canopy here acts as a shading factor 
that determines the peculiarities of the microclimate, hu-
midity etc., which are different from those of open rock 
communities. The floristic composition, structure and 
organization of these habitats differ from forest habi-
tats, and they are the habitat of specific species within 
the boundaries of the forest communities (Chorney at al., 
2014). These chasmophyte species of rock communities 
develop and reproduce on shallow soils – nudilitic lepto-
sols (IUSS Working Group WRB, 2015), and are not re-
lated to the successional process of the forest ecosystems. 
Notably, this type of habitat forms in specific orographic 
conditions: with the presence of rocks and outcrops of 
calcareous bedrock and is not characteristic of the forest 
habitat type within which it occurs. According to data 
recorded for shaded rocks (Sádlo & Chytrý, 2009), the 
temperature is positive from April to the end of Novem-
ber, and the summer temperature does not exceed 15° C. 
Communities of distinctive and specific species compo-
sition (6–13 species of spermatophytes and 4–7 species 
of bryophytes) develop in shady and wet conditions, in 
contrast to those of open rock habitats. 
All the available evidence indicates that evolutionary 
processes are weak for this habitat type; successional de-
velopment here actually slows down, and when commu-
nities degrade, their recovery is probably also very slow 
(Sádlo & Chytrý, 2009). For example, all ferns are poly-
ploids in these habitats (2n=72, 143, 144, 148. 160, 168) 
due to the evolutionary process (Onete, 2012; Chorney 
et al., 2014). 
This habitat combines chasmophyte (Polypodietea 
class) and bryophyte communities: epilithic (Ctenidi-
etea mollusci class) and epiphytic-epigeic (Neckeretea 
complanatae class), which are reflected in the name of 
the Ctenidio-Polypodietalia vulgaris order. On the other 
hand, the bryophyte component of the habitat is consid-
ered independently, although in the composition of this 
habitat they co-exist as a whole system (Gapon, 2013). 
Many questions thus require complex, systematic analy-
sis, which determines the purpose of this publication. 
Considering these questions, we aimed to investigate the 
phytosociological and ecological peculiarities of shady 
chasmophytic habitats in Ukraine. To achieve this aim, 
we addressed the following questions: i) how is a shady 
chasmophytic habitat described syntaxonomically? ii) 
which ecological conditions are characteristic of shady 
chasmophytic habitats? iii) to what threats and risks are 
shady chasmophytic habitats exposed, and how can they 
be protected?

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Study area
Our research covered 5 localities from the Pre-Carpathian 
region within the Ivano-Frankivsk and Chernivtsi regions 
and Transcarpathian region (Figure 1), where the habitat 
type occurs on steep rocky slopes or vertical rock faces of 
northern or western exposure, on ridges or large calcar-
eous rocks and cliffs, shaded by a canopy of deciduous 
trees. The limiting factor that ensures constant atmos-
pheric wet and cool summer temperatures is the dense 
shading by the deciduous tree canopy (mesophytic scio-
phytic conditions) (Didukh et al., 2016).
The Pre-Carpathian region (Prut-Dniester confluence) 
is situated in the western part of Ukraine and geologi-
cally combines Precambrian bedrocks (gneisses, granites 
and argillites), Silurian and Devon bedrocks (limestones, 
dolomites and marls), alluvial clays and loess loams. The 
geomorphology of the Prut-Dniester confluence consists 
of ancient and developed valleys of the Dniester and Prut 
Rivers and their tributaries. The soil cover in the Prut-
Dniester confluence is podzolic grey soils, sod-podzolic 
soils, podzolic chernozems, wet-meadow soils and shal-
low and skeletal soils on rocks and outcrops of calcareous 
bedrocks with deciduous forests, meadows, dry meadow 
steppes and chasmophytic communities. The climate of 
the Prut-Dniester confluence is moderately continental 
with mild winters and a relatively warm, wet summer 
(the average temperature for January is -5.1 °C, the av-
erage temperature for July is 18.7 °C, and the range in 
annual precipitation is from 500 to 700 mm (Heren-
chuk, 1973, 1978). 
The T ranscarpathian region is also situated in the west-
ern part of Ukraine and is characterized by a mountain-
ous relief. This region geologically combines Proterozoic 
(gneisses and crystalline shales), Cretaceous (flysches) and 
Sarmatian bedrocks (clays, sandstones, limestones and 
conglomerates). The hydrography of the region is provid-
ed by the Tysa, Uzh, Latorytsia rivers and their tributar-
ies. The soil cover in the region is formed by burozems, 
meadow burozems, shallow and skeletal soils on rocks 
and outcrops of sandstones, limestones and conglomer-
ates with deciduous and coniferous forests, sub-alpine 
meadows and chasmophytic communities. The climate of 
the T ranscarpathian region is moderately continental that 
strongly depends on temperature fluctuations (the aver-
age temperature for January is -4.8 °C, the average tem-
perature for July is 17.1 °C). The range in annual precipi-
tation is approximately 1122 mm (Herenchuk, 1981).
Figure 1: Locations of shady chasmophytic habitat communities in Ukraine.
Slika 1: Lokacije zasenčenih hazmofitskih združb v Ukrajini.

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Materials and methods
The subject of this research was calcareous cliffs and rocks 
that occur within the deciduous forests of the south-
eastern and south-western macro-slopes of the Ukrainian 
Carpathians (Table 1). These are confined to altitude lim-
its of 250 ± 20 to 850 ± 20 m a.s.l., respectively. The rock 
faces and slopes range from a height of 1.5 m to 5–10 m, 
with slopes up to 90°, with narrow rock protrusions up 
to 30 cm. 
We recorded 46 phytosociological relevés with a uni-
form plot size of 4 m
2
, using the Braun-Blanquet ap-
proach (Braun-Blanquet, 1964) and the recommenda-
tions of Chytrý & Otýpková (2003) for the plot size of 
rocky vegetation, which we collected from Ukraine in 
2018–2019, with specific vascular plants, bryophytes 
and epigeic lichens. We identified habitats by the pres-
ence of chasmophyte ferns (Asplenium  L.,  Cystopteris 
Bernh., Polypodium L.), a dense cover of mosses that grow 
on rocks with steep slopes, and cliffs with ledges up to 
30 cm wide (such ledges can accumulate soil and have 
a different floristic composition). We recorded different 
types of slopes and focused on the typical conditions of 
the habitat. The distribution map was created with the 
help of MapInfo Professional software (MapInfo, 2007). 
The nomenclature for the vascular plant species is ac-
cording to the Euro+Med Database checklist (http://
www.emplantbase.org/), while bryophytes are according 
to Hodgetts et al. (2020), and lichens are according to 
the Index Fungorum (http://www.indexfungorum.org/).
Syntaxa nomenclature followed Mucina et al. (2016), 
Stodiek (1937), Cain & Sharp (1938), Herzog & Höfler 
(1944), Duda (1951) and Redžić et al. (2002). 
The data were analysed in JUICE software (Tichý, 
2002). We used a TWINSPAN modified algorithm 
(Roleček et al., 2009): three pseudospecies cut level -0, 
5 and 25, and Whittaker’s method for distinguishing the 
clusters. Diagnostic species for individual clusters were es-
timated based on the phi-coefficient (threshold of fidelity 
at more than 30%), tested by Fisher’s exact test (p≥0.01) 
(Chytrý et al., 2002), with the sizes of all groups stand-
ardized to an equal size (Tichý & Chytrý, 2006). We used 
a threshold of constancy of more than 50% for the de-
termination of highly constant species, and for constant 
species of more than 25%. 
Phytoindication scales of Y. Didukh (Didukh, 2011; 
Didukh et al., 2021) are comparable to other environ-
mental indicator values (Ellenberg) and reflect ampli-
tude indices of the species characterized by the following 
dimensions: ecological indicator of soil humidity (Hd 
– 23 grades), ecological indicator of variability of damp-
ing (fH – 11 grades), ecological indicator of soil acidity 
(Rc - 15 grades), ecological indicator of total salt regime 
(Sl – 19 grades), ecological indicator of carbonate content 
(Ca – 13 grades), ecological indicator of nitrogen con-
tent (Nt – 11 grades), ecological indicator of soil aeration 
(Ae – 15 grades), ecological indicator of thermoregime 
(Tm – 17 grades), ecological indicator of humidity of 
microclimate (Om – 23 grades), ecological indicator of 
continentality (Kn – 17 grades), ecological indicator of 
cryoregime(Cr – 15 grades) and ecological indicator of 
light in the community (Lc – 9 grades). 
DCA analysis (Detrended Correspondence Analysis) 
was conducted to assess the impact of environmental fac-
tors on plant communities. Phytoindication values were 
passively projected onto a DCA graph. We used the R 
program (R Core T eam, 2021) to visualize the DCA anal-
ysis. The cyclogram of the quantitative indices of leading 
environmental parameters dispersion and determination 
of ecological valence of the communities was conducted 
using the application of environmental parameters’ calcu-
lations (Didukh & Budzhak, 2020).
Habitat-level identification was performed using the 
EUNIS-ESy Expert System (Chytrý et al., 2020) to iden-
tify each relevé in the phytosociological table (Table 2). 
The impact of threats and sociological assessment of the 
habitats were conducted using a four-categories impact 
scale (Table 4) and ultimately summing them (Table 3) 
(Didukh, 2014; Didukh et al., 2018). By calculating the 
sum of points, the indices (in percentages) of the stability 
degree (Stability) S = (S*1 – 3.99) × 8.33 and value de-
gree (Value) V= (V*1 – 9.99) ×3.33 were evaluated. Based 
on these indices, the corresponding classes were selected: 
I – > 80%, II – 61–80, III – 41–60, IV – 21–40, and 
V – < 21%. 
Results
Shady chasmophytic communities occur from 250–
500 m a.s.l., confined to forests of the Carpinion betuli 
and Tilio-Acerion alliances. The dominant tree species 
are Acer platanoides, A. pseudoplatanus, Carpinus betu-
lus, Quercus robur and Tilia cordata. In Transcarpathia, 
such communities occur within an altitudinal range of 
520–850 (1020) m a.s.l., in beech forests of the Fagion 
sylvaticae alliance. The cover of vascular plants fluctu-
ates from 20 to 50%, the cover of bryophytes is 50–80% 
and the cover of the bare surface of rocks is 10–40%. At 
the foot of the rocks, and on more or less horizontal sur-
faces, there are beds of shrubs dominated by Sambucus 
nigra, rarely by Euonymus verrucosa, as well as seedlings 
of Acer platanoides or Fagus sylvatica trees, the roots of 
which penetrate very deeply and disrupt the structure of 
the calcareous rocks. The syntaxonomical peculiarity of 

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Rare shady chasmophytic habitat communities in Ukraine
these communities is the presence of chasmophyte ferns 
with winter-green fronds (Asplenium trichomanes, A. ruta-
muraria, A. viridis, Phyllitis scolopendrium and Polypodi-
um vulgare). These chasmophyte species develop rhizomes 
that fill screes and rock depressions, thus contributing to 
soil formation (Fabiszewski et al., 1997). 
Description of the communities 
(Table 1, Table 2, Figure 3 a-d)
We obtained four well-differentiated clusters using a 
modified TWINSPAN algorithm (Figure 2), which could 
be interpreted as informal communities.
this community is endangered due to climbing and tour-
ist activity (camping sites, fireplaces, litter accumulation) 
in the T ranscarpathian region (Table 3). 
Cluster 2. Porella platyphylla-Asplenium ruta-
muraria community
Floristical characteristics. The Porella platyphylla-
Asplenium ruta-muraria community is characterized by 
the following diagnostic species: Anomodon viticulosus, 
Cardaminopsis arenosa, Homalothecium philippeanum, 
Porella platyphylla. Sagina procumbens, Seseli libano-
tis. The species Exerthotheca crispa and Hedera helix are 
dominant in the communities. The total species cover is 
70–90%, based on bryophytes and overlapped by vascu-
lar plants. 
Distribution and threats. This community forms 
loose mats on thin soils that are rich in nitrogen com-
pounds, covering partially shaded calcareous rocks 
mainly on western, northern and north-western slopes 
and occupying an area of almost 40–60%, where the rest 
of the surface is exposed rock at an elevation from 232 
to 657 m a.s.l. The community is found in the Pre-Car -
pathian (Chernivtsi region, Ukraine) and the Transcar-
pathian regions (Ukraine). According to the sociologi-
cal assessment of the habitat and impact of threats, this 
plant community is an endangered habitat due to tourist 
activity and has a high constancy of synanthropic spe-
cies (Chelidonium majus, Geranium robertianum, Urtica 
dioica) (Table 3).
Cluster 3. Pseudanomodon attenuatus-
Chrysosplenium alternifolium community 
Floristical characteristics. The Pseudanomodon atten-
uatus-Chrysosplenium alternifolium community is char-
acterized by the following diagnostic species: Campan-
ula rapunculoides, Chiloscyphys polyanthos var. rivularis, 
Conocephalum salebrosum, Cystopteris fragilis, Lamium 
galeobdolon. The species Conocephalum salebrosum and 
Pseudanomodon attenuatus are dominant in the com-
munities. The total species cover is 60–80%, based on 
bryophytes and overlapped by vascular plants.
Distribution and threats. This community occurs 
in forests, mainly on shaded steep calcareous cliffs (75–
90°), primarily on northern, north-western, and south-
eastern slopes of the Transcarpathian region (Ukraine) 
at an elevation of 536–1014 m a.s.l. and is formed on 
dry nudilitic leptosols. According to the sociological as-
sessment of the habitat and impact of threats, this plant 
community is influenced by climbing and the mining 
of limestones in illegal quarries in the Transcarpathian 
region; we therefore decreased the threat impact index (3 
points) (Table 3).
 
Figure 2: Dendrogram based on the modified TWINSPAN algorithm 
classification. Cluster 1 – Isothecium alopecuroides-Polystichum braunii 
community (8 relevés); Cluster 2 – Porella platyphylla-Asplenium 
ruta-muraria community (16 relevés); Cluster 3 – Pseudanomodon 
attenuatus-Chrysosplenium alternifolium community (17 relevés); 
Cluster 4 – Pedinophyllum interruptum-Polystichum aculeatum 
community (5 relevés). 
Slika 2: Dendrogram na osnovi algoritma klasifikacije TWINSPAN. 
Klaster 1 – združba Isothecium alopecuroides-Polystichum braunii 
(8 popisov); Klaster 2 – združba Porella platyphylla-Asplenium ruta-
muraria (16 popisov); Klaster 3 – združba Pseudanomodon attenuatus-
Chrysosplenium alternifolium (17 popisov); Klaster 4 – združba 
Pedinophyllum interruptum-Polystichum aculeatum (5 popisov). 
Cluster 1. Isothecium alopecuroides-Polystichum 
braunii community 
Floristical characteristics. The Isothecium alopecu-
roides-Polystichum braunii community is characterized by 
the following diagnostic species: Ctenidium molluscum, 
Dentaria glandulosa, Homalia trichomanoides, Isothesium 
alopecuroides, Plagiomnium cuspidatum, Polystichum brau-
nii. The species Ctenidium molluscum, Homalia trichoma-
noides, Pseudanomodon attenuatus are dominant in the 
communities. The total cover of the species is 60–90% 
and the cover of cryptogam species is 60%.
Distribution and threats. This community occurs on 
Marmarosh calcareous cliffs and conglomerates (Tran-
scarpathian region) of north-eastern and north-western 
slopes on which dry nudilitic leptosols are formed at an 
elevation of 233–307 m a s.l. According to the sociologi-
cal assessment of the habitat and the impact of threats, 

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Cluster 4. Pedinophyllum interruptum-Polystichum 
aculeatum сommunity
Floristical characteristics. The Pedinophyllum inter-
ruptum-Polystichum aculeatum community is character-
ized by the following diagnostic species: Asarum europae-
um, Campanula carpatica, Fissidens dubius, Pedinophyllum 
interruptum, Scopolia carniolica, T amobryum alopecurum,. 
The species Conocephalum salebrosum and Hedera helix are 
dominant in the communities. The total species cover is 
60–95%, based on bryophytes and overlapped by vascu-
lar plants.
Distribution and threats. This community occurs on 
steep Marmarosh calcareous cliffs (45–90°), mainly on 
southern and south-western slopes of the T ranscarpathian 
region (Ukraine) at an elevation of 519–574 m a.s.l. and 
formed on dry nudilitic leptosols. According to the socio-
logical assessment of the habitat and impact of threats, 
this plant community is influenced by the colonization 
of forest species (Aegopodium podagraria, Asarum euro-
paeum, Scopolia carniolica) in the Carpathian Biosphere 
Reserve (Transcarpathian region); we therefore decreased 
the threat impact index (3 points) (Table 3). 
Number of relevés 8 16 17 5
Cluster 1 2 3 4
Isothecium alopecuroides-Polystichum braunii community
Isothecium alopecuroides 63 6 . .
Plagiomnium cuspidatum 63 . . .
Ctenidium molluscum 63 19 . 20
Homalia trichomanoides 50 . . 20
Acer platanoides (juv.) 38 6 . 20
Polystichum braunii 38 . . 20
Rubus idaeus 38 . . .
Eurhynchium striatum 38 19 . .
Porella platyphylla-Asplenium ruta-muraria community
Campanula trachelium . 50 6 .
Porella platyphylla . 44 . .
Fagus sylvatica (juv.) 25 38 . .
Asplenium ruta-muraria . 38 18 .
Sagina procumbens 13 38 . .
Galium intermedium 25 31 24 .
Exsertotheca crispa 25 31 . 20
Pseudanomodon attenuatus-Chrysosplenium alternifolium
community
Pseudanomodon attenuatus 88 50 94 40
Conocephalum salebrosum . . 71 40
Chrysosplenium alternifolium . . 53 40
Campanula rapunculoides . . 35 20
Pedinophyllum interruptum-Polystichum aculeatum community 
Polystichum aculeatum 13 19 . 80
Asarum europaeum 25 . 12 80
Campanula carpatica . 6 . 60
Poa nemoralis 13 6 29 60
Pedinophyllum interruptum . 6 . 60
Lunaria rediviva . 19 6 60
Fissidens dubius . . . 40
Veronica urticifolia . 6 . 40
Carex montana . 13 . 40
Pulmonaria obscura . . . 40
Scopolia carniolica . . . 40
Senecio nemorensis . . . 40
Number of relevés 8 16 17 5
Cluster 1 2 3 4
Radula complanata 25 13 . 40
Salvia glutinosa . 25 18 40
Aegopodium podagraria . . 6 40
Moehringia trinervia . 13 6 40
Galium transcarpaticum . 6 . 40
Alleniella complanata . 19 . 20
Ctenidio-Polypodion vulgaris alliance, 
Ctenidio-Polypodietalia vulgaris order, Polypodietea class
Phyllitis scolopendrium 88 88 100 100
Polypodium vulgare 88 50 59 60
Asplenium trichomanes 75 94 65 60
Hedera helix 75 19 29 80
Cystopteris fragilis 13 63 76 40
Dryopteris filix-mas 75 31 6 60
Lamium galeobdolon 25 25 82 80
Other vascular plants in alphabetical order
Cardaminopsis arenosa . 50 . 60
Chelidonium majus . 38 29 20
Geranium robertianum 25 50 47 80
Hepatica nobilis . 31 24 40
Hylotelephium maximum . 31 18 40
Mercurialis perennis . 25 6 20
Mycelis muralis 13 75 . 40
Sambucus nigra (juv.) . 31 41 40
Urtica dioica 13 25 12 20
Valeriana sp. . 19 . 20
Other cryptogam species in alphabetical order
Anomodon viticulosus 25 63 12 60
Chiloscyphus polyanthos var. rivularis . . 18 .
Homalothecium philippeanum . 25 . .
Lepraria membranacea 13 31 . 40
Metzgeria conjugata 63 . . 40
Mnium stellare . . 24 20
Plagiochila porelloides 50 6 6 40
Thamnobryum alopecurum 50 13 . 60
Table 1: Synoptic table of shadow chasmophytic communities, indicating the constancy of species expressed by their percentage 
frequency in the respective four clusters. Species are ranked by decreasing constancy and phi coefficient values are not shown, 
although species with a phi- value greater than 0.25 are accepted as differential for the syntaxa. Species of each syntaxon with a 
constancy of 15% or less, as well as other taxa with a constancy of 15% or less, are not shown in the table.
Tabela 1: Sinoptična tabela zasenčenih hazmofitskih združb. Prikazana je stalnost vrst s frekvenco v odstotkih v štirih klastrih. 
Vrste so razvrščene glede na padajočo stalnost, vrednosti fi koeficienta niso prikazane. Kot razlikovalne vrste smo upoštevali tiste, 
ki imajo fi koeficient večji od 0,25. Vrste s stalnostjo, manjšo od 15% v posameznem sintaksonu in vse vrste z stalnostjo manjšo od 
15%, niso prikazane v tabeli.

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Rare shady chasmophytic habitat communities in Ukraine
Proposed syntaxonomical scheme
We did not find relevant associations within the Ctenidio-
Polypodion vulgaris alliance, Ctenidio-Polypodietalia vul-
garis order, Polypodietea class. However, we did not find 
all diagnostic bryophyte species to describe them as bry-
ophyte associations. We therefore we propose a syntaxo-
nomical scheme of the distinguished clusters interpreted 
as informal communities. 
Figure 3: Pictures of shady chasmophytic habitat communities: 
a) Velyka Ugolka, T ranscarpathia region, Ukraine, photo by Bezsmertna 
Olesya; b) Prut-Dniester confluence, Chernivtsi region, photo by 
Budzhak Vasyl; c) Pre-Carpathia, Ivano-Frankinsk region, photo by 
Didukh Yakiv; d) right bank of Dniester River, Chernivtsi region, 
Ukraine, photo by Didukh Yakiv
Slika 3: Slike zasenčenih hazmofitskih habitatnih tipov:  
a) Velyka Ugolka, regija T ranscarpathia, Ukrajina, fotografija Olesya 
Bezsmertna; b) sotočje rek Prut-Dniester, regija Chernivtsi, fotografija 
Vasyl Budzhak; c) Pre-Carpathia, regija Ivano-Frankinsk, fotografija 
Yakiv Didukh; d) desni breg reke Dnester, Chernivtsi region, Ukraine, 
fotografija Yakiv Didukh.
Cl. Polypodietea Jurko et Peciar ex Boscaiu, Gergely et 
Codoreanu in Ratiu et al. 1966
Ord. Ctenidio-Polypodietalia vulgaris Jurko et Peciar ex 
Boscaiu, Gergely et Codoreanu in Ratiu et al. 1966
All. Ctenidio-Polypodion vulgaris S. Brullo et al. 
2001
1)  Isothecium alopecuroides-Polystichum brau-
nii community 
2)  Porella platyphylla-Asplenium ruta-muraria 
community 
3)  Pseudanomodon attenuatus-Chrysosplenium 
alternifolium community 
4)  Pedinophyllum interruptum-Polystichum acu-
leatum community
a
b
c
d

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Didukh et al.
Rare shady chasmophytic habitat communities in Ukraine
Ecological characteristics of the 
shadow chasmophytic habitat 
communities
We assessed the ecological factors’ indices according to 
the synphytoindication methodology (Didukh, 2011). 
According to humidity indices, these communities can be 
interpreted as meso-hygromesophyte and well-wetted; ac-
cording to the variability of dampness indices, these com-
munities can be interpreted as hydro-hemicontrasthopho-
bic with uniform steady or slightly uniform hydration; 
according to the aeration indices, hemiaerophobic with 
uniform aerated substrata; according to the acidity indi-
ces – neutrophilic with pH from 6.5 to 7.1; according 
to the total salt regime in soil indices, semi eutrophic; 
according to the carbonate content indices these commu-
nities are acarbonatophilic, despite the carbonate richness 
of the bedrocks; according to the nitrogen content indi-
ces, heminitrophilic – nitrophilic, with high concentra-
tions of nitrate ions. According to the thermoregime in-
dices these communities are submesothermic; according 
to the cryoregime indices, hemicryophytic; according to 
the continentality indices, hemioceanic; according to the 
humidity of the climate indices, subombrophytic. Such 
climatic characteristics of this habitat correlate with the 
indicators of the Carpinion betuli and Fagion sylvaticae 
alliances (Didukh et al., 2016), They are similar to for-
est habitats, but deviate from zonal ones and reflect their 
preferences to shading by a wooded canopy.
Based on the calculation of ecological indicators’ values, 
cyclograms illustrate the environmental niches of shady 
chasmophytic habitat communities, which are character-
ized by a narrow amplitude of environmental factors and 
are typical for stenotopic species. The most limited range 
of tolerance is prominent (Figure 4 a-d): the soil humid-
ity (Hd) (mainly Isothecium alopecuroides-Polystichum 
braunii community and Pedinophyllum interruptum-Pol-
ystichum aculeatum community), the total salt regime (Sl) 
(all distinguished communities), carbonate content (Ca) 
(mainly Pedinophyllum interruptum-Polystichum aculea-
tum community) and acidity (Rc) (mainly Pedinophyllum 
interruptum-Polystichum aculeatum community).


 

 


 







  
Figure 4: Cyclogram of ecological 
indicator values of shady chasmo-
phytic habitats. Acronyms of the 
ecological indicators are the same 
as in the Materials and methods 
section. The dashed line shows the 
minimum and maximum of the 
ecological indicator indices. The 
solid line shows the ecological op-
timum of the shady chasmophytic 
habitat communities.  
a) – Isothecium alopecuroides-
Polystichum braunii community, 
b) – Porella platyphylla-Asplenium 
ruta-muraria community, 
c) – Pseudanomodon attenuatus-
Chrysosplenium alternifolium 
community, d) – Pedinophyllum 
interruptum-Polystichum aculeatum 
community.
Slika 4: Ciklični graf ekoloških 
indikatorskih vrednosti zasenčenih 
hazmofitskih združb. Okrajšave 
ekoloških indikatorjev so enake 
kot v poglavju Material in 
metode. Prekinjena črta prikazuje 
najmanjšo in največjo vrednost 
indikatorja. Polna črta prikazuje 
optimum zasenčenih hazmofitskih 
združb.


 

 


 







  
b


 

 


 







  
d


 

 


 







  
a


 

 


 







  
c

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Rare shady chasmophytic habitat communities in Ukraine
Four shady chasmophytic communities were clearly 
distinguished on the DCA visualization (Figure 5). The 
main axis (DCA 1) compared to the vectors of ecological 
indicators (carbonate content, thermoregime, cryoregime 
and humidity), whereby a significant differentiation be-
tween the Isothecium alopecuroides-Polystichum braunii 
community and the Pseudanomodon attenuatus-Chrysosp-
lenium alternifolium community is observable. Although 
the second axis (DCA 2) does not show substantial differ-
ences between the communities, it is closer to the vectors 
of ecological indicators (total salt regime in the soil and 
soil humidity).
In the Prodrome of the Vegetation of Europe, the clas-
sification of these communities has undergone signifi-
cant changes (Mucina et al., 2016). The chasmophytic 
vegetation of rocks and screes belongs to 8 classes, 36 or-
ders and 146 alliances. However, at the same time, many 
questions arise regarding the delimitation of syntaxa and 
their place in the classification system, especially when 
the bryophyte component, which plays a significant role 
in certain classes and has high diagnostic value, is not 
taken into account. We must emphasize the division 
of two classes Asplenietea trichomanis and Polypodietea. 
This division is based on the chemical characteristics of 
substrata, humidity and shading. The Polypodietea class, 
which consists of 3 orders and 13 alliances and represents 
communities of shady rocks and crevices, with character-
istic species: chasmophyte, sciophyte plants and a high 
cover of bryophytes, which occupy rock surfaces and 
crevices where a small amount of detritus accumulates 
(Tomaselli et al., 2019). 
Finding the syntaxonomical affiliation of the bryophyte 
component generally causes discussion (Cain & Sharp, 
1938). On the one hand, if the bryophyte communities 
are interpreted as a part of the whole, synusia or mero-
coenos, within the shady chasmophytic habitat, then the 
opinion of Berg et al. (2019) is fully justified. Following 
this claim of Berg et al. (2019), bryophyte communi-
ties should not be considered separate classes at the level 
of holocoenoses, which can be reflected in their name 
by introducing the appropriate suffixes. On the other 
hand, Mucina et al. (2016) claimed that synusia (micro-
communities) are elements within plant associations or 
separate syntaxonomic units; we therefore preliminarily 
consider the distinguished clusters to be informal com-
munities within the Ctenidio-Polypodion vulgaris alliance, 
Ctenidio-Polypodietalia vulgaris order, Polypodietea class 
(Table 1). 
Moreover, we did not aim to investigate the bryophyte 
communities as separate syntaxonomical units and used a 
traditional approach to describe the vascular plants’ com-
munities. Based on this, we omitted some species that 
could be diagnostic in some bryophyte associations.
The first cluster was interpreted as the Isothecium 
alopecuroides-Polystichum braunii community, which 
had related connections with the association Tortello-
Ctenidietum mollusci (Gams 1927) Stodiek 1937, which 
was described by Stodiek (1937) with the presence of 
diagnostic bryophyte species (Ctenidium molluscum, 
Isothecium alopecuroides, Plagiomnium cuspidatum) that 
belong to the alliance Ctenidion mollusci Ştefureac 1941, 
order Ctenidietalia mollusci Hadač et Šmarda in Klika et 
Hadač 1944, class of Ctenidietea mollusci von Hübsch-
mann ex Grgić 1980. On the other hand, there is a 














	
 

Figure 5: Detrended Correspondence Analysis (DCA) of shady 
chasmophytic habitat communities. 1 – Isothecium alopecuroides-
Polystichum braunii, 2 – Porella platyphylla-Asplenium ruta-muraria, 
3 – Pseudanomodon attenuatus-Chrysosplenium alternifolium, 
4 – Pedinophyllum interruptum-Polystichum aculeatum. Acronyms of 
the ecological indicators are the same as in the Materials and methods 
section.
Slika 5: Korespondenčna analiza z odstranjenim trendom (DCA)  
zasenčenih hazmofitskih združb. 1 – Isothecium alopecuroides-
Polystichum braunii, 2 – Porella platyphylla-Asplenium ruta-muraria, 
3 – Pseudanomodon attenuatus-Chrysosplenium alternifolium, 
4 – Pedinophyllum interruptum-Polystichum aculeatum. Okrajšave 
ekoloških indikatorjev so enake kot v poglavju Material in metode. 
Discussion
Syntaxonomical revision
Chasmophytic communities are often a mosaic in struc-
ture, due to variations in the microrelief. They typically 
have a lower level of floristic richness, with a combination 
of different taxa (vascular plants, bryophytes, lichens) and 
their syntaxonomy is described differently (Valachovič, 
1995; Sádlo & Chytrý, 2009). 

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Rare shady chasmophytic habitat communities in Ukraine
significant presence of vascular plants (Acer planatoides 
(juv.), Denatria glandulosa, Polystichum braunii) and the 
absence of numerous diagnostic species (Brachythecium 
glareosum, Ditrichum flexicaule, Fissidens dubius) of the 
Ctenidion mollusci alliance (Ştefureac, 1941; Klika, 1948; 
Ragulina & Orlov, 2021).
We classified the second cluster as the Porella platyphyl-
la-Asplenium ruta-muraria community, with the presence 
of ferns (Asplenium ruta-muraria, Phyllitis scolopendrium, 
Polypodium vulgare) and spermatophytes (Campanula 
trachelium, Fagus sylvatica, Sagina procumbens) as the di-
agnostic species. It should be added that the distinguished 
cluster has two diagnostic species (Porella platyphylla and 
Exsertotheca crispa) of the bryophyte association of Homa-
lothecio sericei – Porelletum platyphyllae Stormer 1938 ex 
Duda 1951, which was described by Duda (1951) within 
the alliance Neckerion complanatae Šmarda et Hadač ex 
Klika 1948, order Neckeretalia complanatae Jezek et Von-
dracek 1962 and class Neckeretea complanatae Marstaller 
1986. However, since we have diagnostic species of the 
association (Ragulina & Orlov, 2021), we did not observe 
numerous diagnostic species of the Neckerion complana-
tae alliance (Metgeria conjugata, Sciuro-hypnum populeum, 
Thamnobryum alopecurum). 
In the third cluster, there were some diagnostic bryo-
phyte species (Pseudanomodon attenuates, Conocepha-
lum salebrosum), which could lead us to the association 
Anomodontetum attenuati Cain & Sharp 1938, which 
has also been considered within the alliance Neckerion 
complanatae, order Neckeretalia complanatae and class 
Neckeretea complanatae (Peciar 1965). The fourth cluster, 
designated the Pedinophyllum interruptum-Polystichum 
aculeatum community, has the features of Pedinophylle-
tum interrupti Herzog & Höfler 1944, which was consid-
ered by Vadam (1983). However, it cannot currently be 
treated as the Pedinophylletum interrupti association since 
we do not have a clear group of diagnostic bryophyte spe-
cies (Cololejeunea calcarea, Fissidens pusillus, Pedinophyl-
lum interruptum). Moreover, we did not find evidence of 
the presence of the Pedinophylletum interrupti association 
in Ukraine (Ragulina & Orlov, 2021). 
Additionally, it must be noted that the high constancy 
of Phyllitis scolopendrium within all distinguished com-
munities may indicate Asplenio-Phyllidetum scolopendrii 
Redžić et al. 2002, first described by Redžić et al. (2002) 
and confirmed within the Carpathian Mountains (Besky-
dy Mountains) by Świerkosz (2004). However, it can 
only be confirmed after collecting appropriate data and 
providing a comprehensive analysis within the Carpatho-
Pannonian region in Central Europe.
Ecological assessment of shady 
chasmophytic communities
Calcareous rocks in the Pre-Carpathian and Transcar-
pathian regions obviously have various chemical com-
positions of rocks (thin-bedded limestones, sandstones, 
flysches and conglomerates): a different content of car-
bonate anions, calcium/magnesium cations and soil acid-
ity indices that have apparently (Hnylko et al., 2015) gen-
erated a differentiation of the shady chasmophytic habitat 
communities (Didukh et al., 2016).
The distribution of species within the shady chasmo-
phytic habitat communities is irregular – they grow in 
local spots, mats, and have vascular plants that occur in 
small beds and patches, determined by the range of vari-
ation of the microrelief (Sádlo & Chytrý, 2009). Moreo-
ver, they are heated unequally, which also causes unequal 
transpiration and humidity loss on the different faces of 
rocks (Larson et al., 2000; Păşcuţ, 2018). The cryo- and 
thermoregimes, as the microclimate indices, also caused 
a differentiation of the plant communities in the micro-
refugia formed on the rock cliffs (Garcia et al., 2020).
Conservation status of the shady 
chasmophytic habitat communities
Shady chasmophytic habitats are protected in accord-
ance with Resolution 4 of the Bern Convention (Coun-
cil of Europe, 2018) as H3.252 Communities of shady, 
cool, often moist rockfaces of the Alps and neighbour-
ing regions, of the Carpathians, of the Jura, the Hercyn-
ian ranges, the British Isles, with many ferns, including 
[Cystopteris fragilis], [Cystopteris regia], [Asplenium viride], 
[Asplenium scolopendrium], [Asplenium trichomanes] and 
with [Carex brachystachys] and 8210 Calcareous rocky 
slopes with chasmophytic vegetation in accordance with 
NATURA 2000 (Table 3). 
According to the classification of Ukrainian habitats 
(UkrBiotop), the equivalent of this habitat has been de-
scribed for the Crimea Mountains: H:2.114 Shaded chas-
mophyte communities with a complex of lichens and bry-
ophytes (Didukh et al., 2011). Thereafter Didukh et al. 
(2018) considered a single habitat K5.2 “Chasmophyte 
and bryophyte communities of shaded calcareous cliffs”, 
which combined communities of the Polypodietea class, 
Ctenidio–Polypodion vulgare alliance, which is distributed 
in the Carpathian Mountains and adjacent lowlands of 
Ukraine (Table 3).
It must be added that the results of the ESy-EUNIS 
expert system evaluation (Table 2) showed undefined 
expectations: many relevés were identified with a ques-
tion mark “?”. On the other hand, the last cluster has 

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Rare shady chasmophytic habitat communities in Ukraine
been assigned as R55 (Lowland moist or wet tall herb and 
fern fringe dominants) and R57 (Herbaceous forest clear-
ing vegetation) habitats, although the elements of these 
habitats are present in shady chasmophytic communities. 
It should be mentioned that these shady chasmophytic 
habitat communities are described as H32c (Temperate 
lowland to montane base rich inland cliff) according to 
the modern EUNIS classification (Chytrý et al., 2020); 
however, the group of characteristic species of H32c is 
missing in our results. 
The main threat to this habitat’s existence is deforesta-
tion, which increases light levels, warming and a moisture 
deficiency of the substrata, and eutrophication, as a result 
of which nitrophilic species invade these communities 
(Didukh et al., 2016). Moreover, in our opinion, some 
other kinds of threats could include climbing, tourist ac-
tivity and mining of dolomites and limestones (Świerkosz 
et al., 2011; Świerkosz & Reczynska, 2012), when, due to 
mechanical disturbance, the restoration of the communi-
ties is very slow (Didukh et al., 2012).
The shady chasmophytic habitat is thus rare (Table 3) 
and has limited distribution, with a low potential for res-
toration. These communities are sensitive to the influence 
of anthropogenic factors and require targeted protection 
measures, which must be taken into account when man-
agement plans for habitat protection are created.
Acknowledgments
We warmly appreciate Neil Lockhart and Martin Cree-
gen for proofreading the text and improving the English 
spelling. We are also grateful to Dr. Marina Ragulina 
and Dr. Vitalii Virchenko for the determination of the 
bryophyte difficult groups. We are grateful to the Editor-
in-Chief Urban Šilc, Milan Chytrý and two anonymous 
referees for their valuable comments and suggestions. 
Notably, we want to express our gratitude to the Armed 
Forces of Ukraine for providing the opportunity for us to 
undertake our research activity in safety.
Yakiv Didukh  https://orcid.org/0000-0002-5661-3944
Illia Chorney  https://orcid.org/0000-0002-1382-9112
Vasyl Budzhak  https://orcid.org/0000-0002-7754-6437
Iuliia Vasheniak  https://orcid.org/0000-0003-1020-3007
Olesya Bezsmertna  https://orcid.org/0000-0001-8764-8878
Svitlana Nyporko  https://orcid.org/0000-0003-3451-0319
References
Braun-Blanquet, J. (1964). Pflanzensoziologie. Grundzüge der 
Vegetationskunde. Springer.
Cain, S. A., & Sharp, A. J. (1938). Bryophytic unions of certain forest 
types of the Great Smoky Mountains. American Midland Naturalist, 
20, 249–301.
Chorney, I., Budzhak, V., Tokaryuk, A., & Volutsa, O. (2014). 
Khorolohiya rarytetnoho bioriznomanittya Bukovyny. Chastyna I. 
Arkhehoniaty. DrucArt.
Chytrý, M., Tichý, L., Holt, J., & Botta-Dukát, Z. (2002). 
Determination of diagnostic species with statistical fidelity 
measures. Journal of Vegetation Science, 13, 79–90. https://doi.
org/10.1111/j.1654-1103.2002.tb02025.x 
Chytrý, M., Tichý, L., Hennekens, S.M., Knollová, I., Janssen, J.A.M., 
Rodwell, J.S., Peterka, T., Marcenò, C., Landucci, F ., Danihelka, J., 
Hájek, M., Dengler, J., Novák, P ., Zukal, D., Jiménez-Alfaro, B., 
Mucina, L., Abdulhak, S., Aćić, S., Agrillo, E., Attorre, F ., Bergmeier, 
E., Biurrun, I., Boch, S., Bölöni, J., Bonari, G., Braslavskaya, T., 
Bruelheide, H., Campos, J.A., Čarni, A., Casella, L., Ćuk, M., 
Ćušterevska, R., De Bie, E., Delbosc, P ., Demina, O., Didukh, Y., 
Dítě, D., Dziuba, T., Ewald, J., Gavilán, R.G., Gégout, J.-C., Giusso 
del Galdo, G.P ., Golub, V., Goncharova, N., Goral, F ., Graf, U., 
Indreica, A., Isermann, M., Jandt, U., Jansen, F ., Jansen, J., Jašková, 
A., Jiroušek, M., Kącki, Z., Kalníková, V., Kavgacı, A., Khanina, 
L., Korolyuk, A.Yu., Kozhevnikova, M., Kuzemko, A., Küzmič, 
F ., Kuznetsov, O.L., Laiviņš, M., Lavrinenko, I., Lavrinenko, O., 
Lebedeva, M., Lososová, Z., Lysenko, T., Maciejewski, L., Mardari, 
C., Marinšek, A., Napreenko, M.G., Onyshchenko, V., Pérez-Haase, 
A., Pielech, R., Prokhorov, V., Rašomavičius, V., Rodríguez Rojo, 
M.P ., Rūsiņa, S., Schrautzer, J., Šibík, J., Šilc, U., Škvorc, Ž., Smagin, 
V.A., Stančić, Z., Stanisci, A., Tikhonova, E., Tonteri, T., Uogintas, 
D., Valachovič, M., Vassilev, K., Vynokurov, D., Willner, W ., Yamalov, 
S., Evans, D., Palitzsch, Lund M., Spyropoulou, R., T ryfon, E., & 
Schaminée, J.H.J. (2020). EUNIS Habitat Classification: expert 
system, characteristic species combinations and distribution maps of 
European habitats. Applied Vegetation Science, 23, 648–675. https://
doi.org/10.1111/avsc.12519
Council of Europe. (2018). Revised Annex I to Resolution 4 (1996) of 
the Bern Convention on Endangered Natural Habitat Types using the 
EUNIS Habitat Classification (Adopted by the Standing Committee on 30 
November 2018). https://rm.coe.int/16807469e7 
Didukh, Y. (2011). The ecological scales for the species of Ukrainian flora 
and their use in synphytoindication. Phytosociocentre. 
Didukh, Y., Phytsailo T., Korotchenko, I., Iakushenko, D., 
Pashkevych, N., & Alioshkina, U. (2011). Biotopy lisovoyi ta 
lisostepovoyi zon Ukrayiny. PP Makros. 
Didukh, Y. (2014). Assessment of stability and risk of loss of 
ecosystems. Scientific notes of National University of Kyiv-Mohyla 
Academy, 158. Biology and ecology, 54–60.
Didukh, Y., Chorney, I., & Khodosovtsev, O. (2018). K5.2 
Chasmophyte and moss communities of shaded carbonate rocks. In: 
Kuzemko, A., Didukh, Y., Onyshchenko, V. et al. (Eds.). National 
habitat catalogue of Ukraine. PP Klymenko.
Didukh, Y., Chorney. I., Budzak, V. et al. (2016). Climatogenic 
Changes of Plant Life of the Ukrainian Carpathians. DrucArt.

22/2 • 2023, 143–160
154
Didukh et al.
Rare shady chasmophytic habitat communities in Ukraine
Didukh, Y., Kagalo, O., & Prots, B. (Eds.). (2012). Biotopy (oselyshcha) 
Ukrainy: naukovi zasady yikh doslidzhennya ta praktychni rezultaty 
inventaryzatsii. Materialy robochoho seminaru. Kyiv, 21–22 bereznya 
2012 roku. M.G. Kholodny Institute of Botany. National Academy of 
Sciences of Ukraine.
Didukh, Y., Kuzemko, A., & Vakarenko, L. (2018). Otsinka 
znachymosti ta kadastr ridkisnykh biotopiv Ukrainy dlia zberezhennia 
bioriznomanittia ta monitorynhu dovkillia. Klasyfikatsiia roslynnosti 
ta biotopiv Ukrainy. M.G. Kholodny Institute of Botany. National 
Academy of Sciences of Ukraine.
Didukh, Y., & Budzhak, V. (2020). Prohrama dlya avtomatyzatsii 
protsesu rozrakhunku balnykh pokaznykiv ekolohichnykh faktoriv: 
metodychni rekomendatsii. Chernivtsi National University.
Didukh, Y. (2021). Climate Change Assessment Based on 
Synphytoindication Method. In M. Lackner et al. (Eds.). 
Handbook of Climate Change Mitigation and Adaptation. Springer 
Science+Business Media, LLC, part of Springer Nature 2021. https://
doi.org/10.1007/978-1-4614-6431-0_137-1
Duda J. (1951). Spolecenstva Bryophyt na piskovcovych skalach v 
Beskydach. Prirodoredecky Sbornik ostravskeho Kraje, 12, 323–334.
Euro+Med. (2006-2020). Euro+Med PlantBase – the information 
resource for Euro-Mediterranean plant diversity. http://ww2.bgbm.org/
EuroPlusMed/query.asp .
Fabiszewski, J., Koła, W., & Kwiatkowski, P . (1997). Phyllitis 
scolopendrium (L.) Newm. in the Sudeten Mountains. Annales Silesiae, 
27, 49–62. 
Gapon, S.V. (2013). Methodological aspects of the moss vegetation 
studies. Ukrainian Botanical Journal, 70(3), 392–397. https://doi.
org/10.15407/ukrbotj70.03.392
Herzog, T., & Höfler, K. (1944). Kalkmoosgesellschaften um Golling. 
Hedwigia, 82 (1/2), 2–90. 
Herenchuk, K.I. (1973). Pryroda Ivano-Frankivskoi oblasti. Vyshcha 
shkola.
Herenchuk, K.I. (1978). Pryroda Chernivetskoi oblasti. Vyshcha shkola.
Herenchuk, K.I. (1981). Pryroda Zakarpatskoi oblasti. Vyshcha shkola.
Hill, M. (1979). TWINSPAN – A FORTRAN program for arranging 
multivariate data in an ordered two-way table by classification of the 
individuals and attributes. Cornel University, Ithaca.
Hnylko, O., Krobicki, M., Feldman-Olszewska, A., & Iwanczuk, J. 
(2015). Geology of the volcano-sedimentary complex of the Kamyanyi 
Potik Unit on Chyvchyn Mount (Ukrainian Carpathians): preliminary 
results. Geological Quarterly, 59 (1), 145–156. http://dx.doi.
org/10.7306/gq.1220 
Hodgetts, N. G., Söderström, L., Blockeel, T. L., Caspari, S., Ignatov, 
M. S., Konstantinova, N. A., Lockhart, N., Papp, B., Schröck, 
C., Sim-Sim, M., Bell, D., Bell, N. E., Blom, H. H., Bruggeman-
Nannenga, M. A., Brugués, M., Enroth, J., Flatberg, K. I., Garilleti, 
R., Hedenäs, L., Holyoak, D. T., Hugonnot, V., Kariyawasam, I., 
Köckinger, H., Kučera, J., Lara, F ., & Porley, R. D. (2020). An 
annotated checklist of bryophytes of Europe, Macaronesia and Cyprus. 
Journal of Bryology, 42 (1), 1–116. https://doi.org/10.1080/03736687.
2019.1694329 
IUSS Working Group WRB. (2015). World Reference Base for Soil 
Resources 2014, update 2015 International soil classification system for 
naming soils and creating legends for soil maps. World Soil Resources 
Reports № 106. FAO. 
Klika, J. (1948). Rostlinna sociologie (Fytocoenologie). Melantrich.
Larson, D.W., Matthes, U., & Kelly, P .E. (2000). Cliff ecology. Pattern 
and process in cliff ecosystems. Cambridge University Press.
Lavergne, S., Thompson, J.D., Garnier, E., & Debussche, M. (2004). 
The biology and ecology of narrow endemic and wide-spread plants: a 
comparative study of trait variation in 20 congeneric pairs. Oikos, 107, 
505–518. https://doi.org/10.1111/j.0030-1299.2004.13423.x 
MapInfo. (2007). MapInfo Professional v. 9.5. Pitney Bowes MapInfo. 
https://mapinfo-professional.software.informer.com/9.5/ 
Mucina, L., Bültmann, H., Dierßen, K., Theurillat, J.-P ., Dengler, J., 
Čarni, A., Šumberová, K., Raus, T., di Pietro, R., Gavílan García, R., 
Chytrý, M., Yakushenko, D., Schaminée, J.H.J., Bergmeier, E., Santos 
Guerra, A., Daniëls, F .J.A., Ermakov, N., Valachovic, M., Pignatti, S., 
Rodwell, J. S., Pallas, J., Capelo, J., Weber, H.E., Lysenko, T., Didukh, 
Y.P ., Capelo, J., Weber, H., Solomesh, A., Dimopolous, P ., Aguiar, 
C., Freitag, H., Hennekens, S.M., & Tichý, L. (2016). Vegetation of 
Europe: hierarchical floristic classification system of vascular plant, 
bryophyte, lichen and algal communities. Applied Vegetation Science, 
19(1), 3–264. https://doi.org/10.1111/avsc.12257 
Onete, M. (2012). Assessment and conservation status of ferns from 
Buila-Vânturaria National Park. Oltenia Studii şi comunicări Ştiinţele 
Naturii, 28(2), 144–148. 
Păşcuţ, C.G. (2018). Phytocoenological study of rocky vegetation from 
Codru-Moma Mountains based on ecological indices. Annals of the 
University of Oradea, Fascicle: Environmental Protection, 30, 145–156. 
https://doi.org/10.5281/zenodo.4362296
Peciar, V. (1965). Epiphytische Moosgesellschaften der Slowakei. Acta 
Botanica Facultatis Rerum Naturalium Universitatis Comenianae, 12, 
371–471.
Pigott, C.D., & Walters, S.M. (1954). On the interpretation of the 
discontinuous distributions shown by certain British species of open 
habitats. Journal of Ecology, 42, 95–116.
R Core Team. (2021). R: A language and environment for statistical 
computing. R Foundation for Statistical Computing. Vienna, Austria. 
https://www.R-project.org/
Ragulina, M.E., & Orlov, O.L. (2021). Mokhova roslynnist skelnykh 
vykhodiv lisovogo zakaznyka “Chortova skelya” ta yiyi antropogenna 
dunamika. Naukovi Zapysky Derzhavnogo Pryrodnychogo Muzeiu, 37, 
137–142.
Redžić, S. (Ed.). (2002). Evaluation of Natural Values of Environment of 
Brcko District. Center of ecology and natural resources of the Faculty of 
Science University of Sarajevo & Counterpart International.
Rodwell, J.S., Schaminée, J.H.J., Mucina, L., Pignatti, S., Dring, J., 
& Moss, D. (1998). The Scientific Basis of the EUNIS Habitat 
Classification. Report to the European Topic Centre on Nature 
Conservation. Unit of Vegetation Science, Lancaster
Roleček, J., Tichý, L., Zelený, D., & Chytrý, M. (2009) Modified 
TWINSPAN Classification in Which the Hierarchy Respects Cluster 

22/2 • 2023, 143–160
155
Didukh et al.
Rare shady chasmophytic habitat communities in Ukraine
Heterogeneity. Journal of Vegetation Science, 20, 596–602. https://doi.
org/10.1111/j.1654-1103.2009.01062.x 
Sádlo, J., & Chytrý, M. (2009). Vegetace skal, zdí a stabilizovaných 
sutí (Asplenietea trichomanis). Vegetation of rocks, walls and stable 
screes. In: Chytrý M. (Ed.). Vegetace České republiky. 2. Ruderální, 
plevelová, skalní a suťová vegetace. Academia. 
Speziale, K.L., & Ezcurra, C. (2012). The role of outcrops in the 
diversity of Patagonian vegetation: Relicts of glacial palaeofloras. Flora, 
207, 141–149. https://doi.org/10.1016/j.flora.2011.09.002 
Stodiek, E. (1937). Soziologische und ökologische Untersuchungen 
an den xerotopen Moosen und Flechten des Muschelkalkes in der 
Umgebung Jenas. Repertorium specierum novarum regni vegetabilis. 
Beiheft, 99, 1–46. 
Ştefureac, T. I. (1941). Cercetări sinecologice şi sociologice asupra 
Bryophytelor din codrul secular Slătioara (Bucovina). Analele 
Academiei Române, Memoriile Secțiuni & Ibreve; Ştiințifice, 3(16), 
1–197.
Świerkosz, K. (2004). Notes on the syntaxonomy of the Asplenietea 
trichomanis class in Poland. Polish Botanical Journal, 49(2), 203–213.
Świerkosz, K., Reczyńska, K. Kulpiński, K., & Tyc, A. (2011). Udział 
gatunków obcych w zbiorowiskach z klasy Asplenietea trichomanis (Br.-
Bl. in Meier & Br.-Bl. 1934) Oberd. 1977 w południowej Polsce. Acta 
Botanica Silesiaca, 6, 97–114.
Świerkosz K., & Reczynska K. (2012). Wapienne ściany skalne 
ze zbiorowiskami Potentilletalia caulescentis. In: W. Mróz (Ed.). 
Monitoring siedlisk przyrodniczych. Przewodnik metodyczny. GIOŚ.
Theurillat, J.-P ., & Guisan, A. (2001). Potential impact of climate 
change on vegetation in the European Alps. Climatic Change, 50, 
77–109. https://doi.org/10.1023/A:1010632015572 
Tichý, L. (2002). JUICE, software for vegetation classification. 
Journal of Vegetation Science, 13, 451–453. https://doi.
org/10.1111/j.1654-1103.2002.tb02069.x 
Tichý, L., & Chytrý, M. (2006). Statistical determination of diagnostic 
species for site groups of unequal size. Journal of Vegetation Science, 17, 
809–818. https://doi.org/10.1111/j.1654-1103.2006.tb02504.x 
Tomaselli, M., Foggi, B., Carbognani, M., Gennai, M., & Petraglia, 
A. (2019). The rock-face vegetation in the northern Apennines and 
neighbouring mountain areas, from the coast line to the highest 
summits. Phytocoenologia, 49(1), 7–70. https://doi.org/10.1127/
phyto/2018/0117 
Vadam, J.-C. (1983). Les groupements muscinaux des escarpements et 
rochers calcaires des environs de Montbéliard (Doubs). Ann. Sc. Univ. 
Franche-Comté, Besançon, Biol. Vég.,4(4), 55—96.
Valachovič, M. 1995: Asplenietea trichomanis (Br.-Bl. In Meier et Br.-
Bl. 1934) Oberd. 1977. In: M.Valachovič (Ed.). Rastlinné spoločenstvá 
Slovenska. 1. Pionierskavegetácia, Veda.

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Relevé 
ID
Date (year/
month/day)
Latitude Longitude Relevé area 
(m2)
Altitude  
(m)
Aspect 
(degrees)
Slope 
 (degr ees)
Cover total 
(%)
Cover herb 
layer (%)
Cover moss 
layer (%)
Author
1 29.07.2018 49.002238 24.877798 8 263 45 40 90 40 90 BV
2 29.07.2018 49.000647 24.835605 10 283 45 30 80 70 80 ChI
3 28.05.2008 48.496594 27.437630 15 233 180 80 80 30 ChI
4 28.07.2018 49.021633 24.784517 – 287 350 80 60 30 50 DY
5 28.07.2018 49.021633 24.784517 – 287 360 80 80 20 80 DY
6 28.07.2018 49.021633 24.784517 – 287 270 90 70 20 70 DY
7 28.07.2018 49.021633 24.784517 – 287 360 90 70 20 70 DY
8 29.07.2018 48.990813 24.857666 – 307 360 80 80 50 40 DY
9 29.07.2018 48.990813 24.857666 – 307 270 70 50 40 40 DY
10 29.07.2018 48.990813 24.857666 – 247 360 80 90 45 80 DY
11 29.07.2018 49.134067 24.700400 – 232 320 80 60 40 40 DY
12 28.07.2018 49.021633 24.784517 – 284 360 90 70 35 70 DY
13 29.07.2018 48.990813 24.857666 – 247 360 80 85 20 80 DY
14 28.07.2018 49.021308 24.768802 10 273 – – 80 40 80 ChI
15 12.07.2013 48.529253 24.446425 14 233 45 50 60 60 50 ChI
16 28.07.2018 49.021633 24.784517 – 284 360 80 70 30 70 DY
17 29.07.2018 48.990813 24.857666 12 307 180 60 50 50 10 DY
18 01.08.2018 47.942574 24.115118 10 486 120 90 90 20 50 VI/BO/RG
19 01.08.2018 47.940410 24.111568 10 521 320 90 95 20 55 VI/BO/RG
20 01.08.2018 47.936740 24.104383 10 426 300 90 95 40 55 VI/BO/RG
21 31.07.2018 47.950290 24.091400 10 657 290 87 80 20 60 VI/BO/RG
22 31.07.2018 47.950330 24.090490 10 645 275 90 70 15 55 VI/BO/RG
23 01.08.2018 47.938880 24.109010 10 439 20 90 95 40 55 VI/BO/RG
24 01.08.2018 48.458617 26.103283 – 334 360 80 70 40 40 VI/BO/RG
25 31.07.2018 47.950170 24.089620 10 638 280 20 80 20 60 VI/BO/RG
26 31.07.2018 47.950450 24.083850 10 635 350 90 80 20 60 VI/BO/RG
27 31.07.2018 47.950290 24.090760 10 651 310 85 90 20 70 VI/BO/RG
28 03.08.2019 48.266350 23.634320 10 564 80 5 60 60 0 VI/BO/RG
29 03.08.2019 48.263900 23.629190 10 1014 300 90 60 20 40 VI/BO/RG
30 03.08.2019 48.266880 23.629290 10 598 120 90 60 20 40 VI/BO/RG
31 05.08.2019 48.256430 23.677520 10 827 200 90 80 20 60 VI/BO/RG
32 05.08.2019 48.256550 23.677540 10 807 210 85 80 30 50 VI/BO/RG
33 05.08.2019 48.256430 23.677500 10 846 210 90 80 5 75 VI/BO/RG
34 05.08.2019 48.256430 23.677870 10 842 223 75 70 10 60 VI/BO/RG
35 05.08.2019 48.256970 23.678410 10 835 270 90 60 5 55 VI/BO/RG
36 05.08.2019 48.256300 23.677300 10 840 140 90 70 20 50 VI/BO/RG
37 03.08.2019 48.266350 23.634320 10 557 100 90 80 20 60 VI/BO/RG
38 04.08.2019 48.268990 23.633700 10 565 160 30 80 30 50 VI/BO/RG
39 03.08.2019 48.264850 23.642230 10 754 290 90 80 20 60 VI/BO/RG
40 03.08.2019 48.270900 23.629260 10 602 300 90 70 20 50 VI/BO/RG
41 03.08.2019 48.257540 23.531370 10 536 10 90 60 20 40 VI/BO/RG
42 04.08.2019 48.268970 23.630950 10 547 270 90 90 20 70 VI/BO/RG
43 04.08.2019 48.269050 23.633440 10 547 250 87 95 10 85 VI/BO/RG
44 04.08.2019 48.276870 23.640530 10 557 140 90 90 10 80 VI/BO/RG
45 05.08.2019 48.268910 23.633920 10 519 240 90 80 2 78 VI/BO/RG
46 03.08.2019 48.267120 23.629600 10 574 40 45 60 20 40 VI/BO/RG
Table 1: Geographical references of shady chasmophytic habitat communities in Ukraine.
Tabela 1: Geografski podatki za zasenčene hazmofitske združbe v Ukrajini.
Acronyms: 
BV – Budzhak Vasyl, ChI – Chorney Illia, DY – Didukh Yakiv, VI – Vasheniak Iuliia, BO – Bezsmertna Olesya, RG – Gleb Ruslan
Appendix

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Number of relevés 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46
Esy-EUNIS expert system ? ? ? ? ? ?
C22a
? ? ?
R55
R55
V ? ? ?
R55
? ? ?
R55
? ? ?
S35
? ? ?
C21a
? ? ? ? ? ? ? ? ?
V39
? ?
R57
R55
R55
R57
R55
Syntaxa ID 1 2 3 4
Isothecium alopecuroides-Polystichum braunii community 
Isothecium alopecuroides 3 3 + . 2 . 2 . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Plagiomnium cuspidatum . 1 . + + . + 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ctenidium molluscum + . . r 5 5 . 2 5 5 + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Homalia trichomanoides . 3 . + . . 3 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 .
Acer platanoides . . + . + . . 1 . . . . + . . . . . . . . . . . . . . . + . . . . . . . . . . . . . r . . .
Polystichum braunii . . . . 1 1 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . .
Rubus idaeus 1 . . . . + 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Eurhynchium striatum + 2 . . 2 . . . . . . . + . . . . . . . + 2 . . . . . . . . . . . . . . . . . . . . . . . .
Porella platyphylla-Asplenium ruta-muraria community
Campanula trachelium . . . . . . . . . . . + 1 . + . + . + . 1 . 1 1 . . . . . . . . . 1 . . . . . . . . . . . .
Porella platyphylla . . . . . . . . . . . r . + . . . . + + r . r 2 . . . . . . . . . . . . . . . . . . . . . .
Fagus sylvatica (juv.) . . . . + . . 1 . . + + . . . . r . . . + + r . . . . . . . . . . . . . . . . . . . . . . .
Asplenium ruta-muraria . . . . . . . . 1 1 . + . . . + 1 . . . . . + . . . . . . . 3 . 1 . . . . . . . 1 . . . . .
Sagina procumbens 1 . . . . . . . . 2 . 1 2 . . . 1 + . . . 1 . . . . . . . . . . . . . . . . . . . . . . . .
Galium intermedium + . . . 1 . . . 1 . 1 1 . . . . 2 . . . . . r . . . . . . . . . 1 . 1 . . . 1 1 . . . . . .
Exsertotheca crispa 2 . + . . . . . . . + . . 4 . 4 . 5 . . . 5 . . . . . . . . . . . . . . . . . . . 2 . . . .
Pseudanomodon attenuatus-Chrysosplenium alternifolium community
Pseudanomodon attenuatus 3 1 3 + + . + 3 . . + + + . 3 . . . 2 3 + . + . 5 5 5 5 5 5 5 5 3 1 5 4 5 . 5 5 5 . . + 4 .
Conocephalum salebrosum . . . . . . . . . . . . . . . . . . . . . . . . 5 2 2 . 2 2 1 5 . . . 3 3 . 3 1 2 4 . . . 4
Chrysosplenium alternifolium . . . . . . . . . . . . . . . . . . . . . . . . 2 . 5 1 1 1 . 3 . . . . . 5 3 . 2 . . . 1 +
Campanula rapunculoides . . . . . . . . . . . . . . . . . . . . . . . . r . . . . . 1 1 1 . 2 . . . . 1 . 1 . . . .
Pedinophyllum interruptum-Polystichum aculeatum сommunity
Polystichum aculeatum . . . . 1 . . . . . + . + . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . 1 . 1 1 +
Asarum europaeum . . . 1 . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . 3 . . 1 . . . . . 1 2 . 1 1
Campanula carpatica . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . + 2
Poa nemoralis . . . . . . . + . . . . . . . . . + . . . . . . . 1 . . . . 5 2 . . . . . . 3 . 1 1 . . + 1
Pedinophyllum interruptum . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . + . . + +
Lunaria rediviva . . . . . . . . . . . . 1 . . . + . . . . . + . . . . . . . . 1 . . . . . . . . . 1 1 2 . .
Fissidens dubius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . 1 .
Veronica urticifolia . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . + . . 1 .
Table 2: Phytosociological table of shady chasmophytic habitat communities. Species are arranged according to decreasing phi-values and according to decreasing overall 
constancy for non-diagnostic species
Tabela 2: Fitocenološka tabela zasenčenih hazmofitskih združb. Vrste so urejene po padajočih vrednostih fi koeficienta, ne diagnostične vrste pa po padajoči stalnosti.

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Carex montana . . . . . . . . . 1 . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . + +
Pulmonaria obscura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . + .
Scopolia carniolica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . + .
Senecio nemorensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . + .
Radula complanata 1 . 2 . . . . . . . . . . . . . . . . . + . + . . . . . . . . . . . . . . . . . . . . + r .
Salvia glutinosa . . . . . . . . . . . + 1 . . + . . . . + . . . 1 . . 1 . . r . . . . . . . . . . r . . . +
Aegopodium podagraria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 . . . . . . . . . . . 1 r . . .
Moehringia trinervia . . . . . . . . . . . . . . + . . . . . . . . 1 . . . . . . . . . . . . . . . r . 1 . . . +
Galium transcarpaticum . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . +
Alleniella complanata . . . . . . . . . . . . . 2 . 2 . . . . . . . 1 . . . . . . . . . . . . . . . . . 2 . . . .
All. Ctenidio-Polypodion vulgaris, Ord. Ctenidio-Polypodietalia vulgaris, Cl. Polypodietea
Phyllitis scolopendrium 1 2 2 3 . 1 1 2 1 . + 1 2 3 1 1 2 1 + 2 2 . 1 1 2 5 4 2 5 3 2 2 3 3 3 4 4 3 2 3 3 1 4 2 1 2
Polypodium vulgare + 2 + 2 2 1 . + + 2 1 . . . . 1 . + . 1 . 1 . 1 5 . . 2 . . r 5 1 . 2 . 1 3 3 5 . 1 . . 1 1
Asplenium trichomanes + 1 2 . 1 1 . 1 2 2 + + . 1 2 1 1 1 1 + 1 1 r 1 . 1 1 1 . . 3 2 . 3 5 1 1 . 1 . 5 1 . . 2 2
Hedera helix 1 1 1 1 1 . . 1 3 . . . . 5 . . . . . . 2 . . . 3 . . . . 2 . . 5 . . 1 . . . 1 . . 5 1 4 1
Cystopteris fragilis . . . . . . 1 . . . 1 1 + + . + 1 + . . 1 + + . 2 1 3 1 1 . 1 1 . 2 . 1 1 . 1 r 1 r . . . 1
Dryopteris filix-mas 2 1 1 . 2 1 . + . . + . . + . . . . 1 1 . . . 1 . . . . . . . . . . 1 . . . . . . . 1 + . +
Lamium galeobdolon . . 1 + . . . . . . . . . . . + 1 . 1 1 . . . . 3 1 5 5 1 1 3 5 2 1 3 . 1 . . 1 5 . 5 1 2 1
Other vascular plants in alphabetical order
Cardaminopsis arenosa . . . . . . . . . . . . r + + 1 r . + . r . r . . . . . . . . . . . . . . . . . . 1 . . r 1
Chelidonium majus . . . . . . . . . . . r + + . + . . . . + . + . . . 5 . . . 2 . . . . . . . 2 3 3 1 . . . .
Geranium robertianum r . . 1 . . . . . . + 1 r . + . 1 r . . r . 1 . 2 2 1 . . . 2 . . . . 1 + 5 5 . . 1 r . + 1
Hepatica nobilis . . . . . . . . 1 . . 1 1 . . . 1 . . . . . + . . . . . . . . . 1 1 1 . 1 . . . . . 2 . . +
Hylotelephium maximum . . . . . . . . + 1 . . 1 . . . 1 . . . . . 1 . . + . . . . . . . . . . . . 3 1 . + . . + .
Mercurialis perennis . . . . . . . . . . + . . . . . . . . 1 1 . + . . . . . . . . . 3 . . . . . . . . . . . . +
Mycelis muralis + . . . . . . . . . + + + . + + 1 + 1 + 1 . + + . . . . . . . . . . . . . . . . . r . . . +
Sambucus nigra (juv.) . . . . . . . . . . . + 2 . . . + . . . + . r . 5 . 3 . + 1 . 2 . . 1 . . . . 2 . + . + . .
Urtica dioica . . . + . . . . . . . + + . . . + . . . + . . . . 1 . . . . . . . . . . . . . . r . . 1 . .
Valeriana sp. . . . . . . . . . . . + + . . . + . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . .
Other cryptogam species in alphabetical order
Anomodon viticulosus . . 5 + . . . . . . + + + 2 3 . . . 2 2 + . + 3 . . 1 . . . . . . . . . . . 1 . . 2 . + . 3
Chiloscyphus polyanthos var. rivularis . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . 1 . 1 . . . . . . . . . . . .
Homalothecium philippeanum . . . . . . . . . . . . . . 2 . . . 1 1 . . . 1 . . . . . . . . . . . . . . . . . . . . . .
Lepraria membranacea . . . . . + . . . . . . . + . + . + . . . + r . . . . . . . . . . . . . . . . . . + . . . +
Metzgeria conjugata . . + + . 1 3 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 r
Mnium stellare . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . 1 . 1 . . . . 2 . . . . . + .
Plagiochila porelloides r . r . . 1 . r . . . . . + . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . 1 +
Thamnobryum alopecurum . . 3 + . . + 1 . . . . . 1 . . . 5 . . . . . . . . . . . . . . . . . . . . . . . 1 . + . 2

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1 – Isothecium alopecuroides-Polystichum braunii commu-
nity, 2 – Porella platyphylla-Asplenium ruta-muraria com-
munity, 3 – Pseudanomodon attenuatus-Chrysosplenium 
alternifolium community, 4 – Pedinophyllum interruptum-
Polystichum aculeatum community
Taxa that rarely occurred in the table
Vascular plants:
Acer pseudoplatanus (11:+; 34:+), Aruncus dioicus (28:4), 
Asplenium viride (9:1; 11:+), Athyrium filix-femina (4:+; 11:+; 
27:+), Atropa bella-donna (14:+; 24:1; 25:+), Bromopsis sp. 
(13:+), Campanula persicifolia (12:1; 28:+), Carex digitata 
(36:r; 38:r), Carex sp. (12:+; 13:+), Circaea lutetiana (27:2), 
Corylus avellana (25:1), Daphne mezereum (9:1), Dentaria 
glandulosa (2:+; 4:+), Dryopteris carthusiana (4:+), D. expansa 
(6:1), Epipactis helleborine (17:+), Equisetum palustre (43:r), 
Euonymus europaea (27:3), E. verrucosa (38:3; 45:3), Fraxinus 
excelsior (17:+), Galium aparine (31:+), G. odoratum (6:+; 
21:+; 26:r), Gentiana asclepiadea (25:+), Geranium lucidum 
(44:5), Glechoma hederacea (28:+), Hieracium murorum subsp. 
sylvularum (31:+), Impatiens noli-tangere (13:1; 17:1), Luzula 
luzuloides (6:+), Maianthemum bifolium (9:+), Melandrium 
dioicum (21:+), Melica picta (17:+), M. uniflora (15:+), Melittis 
melissophyllum subsp.carpatica (9:+), Orthilia secunda (9:+), 
Oxalis acetosella (6:1; 18:1; 36:3), Paris quadrifolia (18:+; 26:r), 
Phegopteris connectilis (6:+), Pilosella echioides (16:1; 29:+), 
Polygonatum multiflorum (17:+), Prenanthes purpurea (9:+), 
Quercus robur (10:r), Rosa pendulina (9:+), Seseli libanotis 
(12:+; 17:+; 23:+), Silene sp. (17:+; 23:+), Solidago virgaurea 
(36:1; 38:1), Sorbus aucuparia (10:+), Stachys sylvatica (11:+; 
12:+), Symphytum cordatum (27:1), Tilia cordata (5:+), Ulmus 
glabra (41:+), Valerianella dentata (16:1; 29:1), Vincetoxicum 
hirundinaria (17:+; 23:+), Viola riviniana (15:+).
Cryptogam species:
Atrichum undulatum (6:1), Brachythecium rivulare (6:1; 7:3) 
B. rutabulum (30:1), B. salebrosum (7:3), Campyliadelphus 
chrysophyllus (29:1), Campylophyllopsis calcarea (37:1), 
Cololejeunea rossetina (29:+), Conocephalum conicum (18:+), 
Dicranum scoparium (9:3), Ditrichum flexicaule (10:1), 
Eurhynchium angustirete (7:+), Homalothecium lutescens 
(11:+; 12:+; 37:3), Hypnum cupressiforme (6:1; 10:2), Lejeunea 
cavifolia (14:+; 28:+), Lescuraea incurvata (13:+; 17:+), Leucodon 
sciuroides (10:+; 12:+), Lophocolea bidentata (29:+), Mannia 
fragrans (39:1), Metzgeria furcata (2:r), M. pubescens (14:+), 
Plagiomnium rostratum (14:+), Polytrichum sp. (27:1), Porella 
arboris-vitae (28:2), Ptychostomum capillare (17:+), Rhizomnium 
punctatum (1:5; 3:+; 30:5), Sciuro-hypnum populeum (39:1), 
Thuidium recognitum (2:+; 7:1), Tortella tortuosa (11:+; 17:+).
Table 3: Impact of threats and sociological assessment of the shady chasmophytic communities.
Tabela 3: Vpliv groženj in sociološka ocena za zasenčene hazmofitske združbe.
Cluster ID
UkrBiotop (2011)
National Habitat 
Catalogue (2018)
Natura 2000
EUNIS
1
2
3
4
Sum of the threats
Stability, %
Class
5
6
7
8
9
10
11
12
13
14
Sum of the sociological 
assessment
Value, %
Class
1 H2.114 K5.2 8210 H3.252 2 2 2 4 10 66.77 II 4 4 4 4 4 4 4 3 1 2 34 79.95 II
2 H2.114 K5.2 8210 H3.252 2 2 2 4 10 66.77 II 4 4 4 4 4 4 4 3 1 2 34 79.95 II
3 H2.114 K5.2 8210 H3.252 3 2 2 4 11 77.88 II 4 4 4 4 4 4 4 3 1 2 34 79.95 II
4 H2.114 K5.2 8210 H3.252 3 2 2 4 11 77.88 II 4 4 4 4 4 4 4 3 1 2 34 79.95 II
1 – Isothecium alopecuroides-Polystichum braunii community, 2 – Porella platyphylla-Asplenium ruta-muraria community, 
3 – Pseudanomodon attenuatus-Chrysosplenium alternifolium community, 4 – Pedinophyllum interruptum-Polystichum aculeatum 
community.
1 – results of threats’ impact; 2 – scale of the negative impact; 3 – degree of the negative influence of external factors; 4 – degree and 
speed of recovery (plastic, dynamic stability); 5 – position in the successional series is the final steady climax and subclimax stages; 
6 – regional representativity; 7 – nature of the distribution; 8 – ecological amplitude (scales); 9 – environmental  parameters of dis-
tribution; 10 – presence of invasive species; 11 – degree of hemerobity, 12 – ratio of species between strategy types stress-tolerant/
expellerant species; 13 – sociological significance; 14 – synsociological state.

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Didukh et al.
Rare shady chasmophytic habitat communities in Ukraine
Name of the factor
Impact of threats
Category
4 3 2 1
Results of the threats’ 
impact
(EX) damaged habitats;  
(CR) habitats under critical 
threat of extinction
(EN) endangered habitats (VU) vulnerable habitats (TC) habitats that are not 
threatened with extinction
Scale of the negative 
impact
General (global) Regional Local Absent negative impact
Degree of the negative 
influence of external 
factors
Very powerful (destructive), 
destroyed completely, and 
recovers from pioneer stages
Significant, the structure  
of dominants is changing
Moderate, changing  
species composition
Changes are invisible, or 
biotopes are formed under 
the direct influence of an 
anthropogenic factor
Degree and speed 
of recovery (plastic, 
dynamic stability)
Very weak  
(more than 100 years)
Weak  
(more than 10 years)
Tolerable  
(more than 15 years)
Good  
(several years)
Sociological assessment
The position in the 
successional series is the 
final steady climax and 
subclimax stages
Permanent climax and  
sub-climax stages 
Long-term stages of 
endoecogenesis affecting 
change in the  
microclimate and soil
Medium-term serial 
syngenetic stages that do 
not affectchange in soil and 
microclimate characteristics
Pioneer and  
short-term stages
Regional representativity Spread within one or two 
geobotanical regions or 
districts
Spread within one 
geobotanical province
Spread within one 
geobotanical area or 
geographical zone
Spread within several 
geobotanical area or 
geographical zones
Nature of the 
distribution
Some small localities are 
known
Habitat has disjunctive 
distribution
Habitat is sporadically 
spread on the border  
of its area
Habitat is usually spread 
within its area
Ecological amplitude 
(scales)
Narrow amplitude  
(less than 5%) of all 
environmental factors
Narrow amplitude 
(less than 5%) for one 
environmental factor; 
and the amplitude is 
less than 10% for other 
environmental factors
Wider amplitude  
(less than 10%) of more 
than one environmental 
factors
Wide amplitude  
(more than 10%) of all 
environmental factors
Environmental 
parameters of 
distribution
Specific, extreme 
environmental conditions
Narrow distribution of rare 
habitat
Sporadical distribution 
within optimal conditions
Common distribution 
within optimal conditions
Presence of  
invasive species
Absence of  
invasive species
Presence of  
invasive species
Presence of  
invasive species as 
diagnostic species with 
high constancy
Invasive species are 
dominant in the habitat
Degree of hemerobity Ahemerobic and 
oligohemerobic habitats  
(ha less than 25)
Mesohemerobia habitats  
(ha=25–50)
Euhemerobia habitats  
(ha=50–75)
Polyhemerobic and 
metahemerobic habitats 
(more than 75)
Ratio of species between 
strategy types stress-toler-
ant / expellerant species
More than  
1.7
1.2–1.7 0.7–1.2 Less than  
0.7
Sociological significance Presence or dominance of 
CR, EN species or a large 
number of red-listed species
Dominant species are VU, 
LC, DD categories or red-
listed species
Presence of VU, LC, DD 
species, or a couple of red-
listed species
Absence of rare species
SYnsociological estate Habitat listed in Resolution 
4 of the Bern Convention, 
Habitat Directive and in the 
Green Book of Ukraine
Habitat is listed in the 
Green Book of Ukraine
Habitat is listed in 
Resolution 4 of the Bern 
Convention, Habitat 
Directive
Habitat is not listed and 
not protected 
Table 4: Categorization of habitat characteristics for assessing the impact of threats and sociological assessment (Didukh et al., 2018)
Tabela 4: Kategorije habitatnih značilnosti za oceno vpliva groženj in sociološko oceno.