275
A Mediterranean element of the vegetation: 
Junco maritimi-Cladietum marisci – a new 
association for Ukraine
Abstract
Cladium mariscus (L.) Pohl (Cyperaceae) is a rare species in Europe considered 
by several authors to be a relict of the early Holocene period. It is listed in the 
Red Data Book of Ukraine, Annexes of the Habitat Directive and the Bern 
Convention. Communities with domination of this species are included in the 
Green Data Book of Ukraine. Substantial differences in major ecological factors 
for Cladium mariscus communities in the western (carbonate bogs) and the 
southern (marshes and floating swamps of the northern Black Sea) regions of 
Ukraine were shown. The author carried out comparisons of relevés characterizing 
different communities with Cladium mariscus within Europe. Based on the results 
of TWINSPAN analysis, four associations were identified, confirmed by floristic 
indices and ecological data: Cladietum marisci Allorge 1921, Soncho maritimi-
Cladietum marisci (Br.-Bl. & O. de Bolòs 1957) Cirujano 1980, Dorycnio recti-
Cladietum marisci Gradstein & Smittenberg 1977 and Junco maritimi-Cladietum 
marisci (Br.-Bl. & O. de Bolòs 1957) Géhu & Biondi 1988. Thus, in addition 
to the association Cladietum marisci, a new one was indicated for Ukraine, Junco 
maritimi-Cladietum marisci.
Iz vleček
Cladium mariscus (L.) Pohl (Cyperaceae) je v Evropi redka vrsta in številni avtorji 
jo uvrščajo med relikte iz zgodnjega holocena. Uvrščena je na rdeči seznam 
Ukrajine, Habitatno direktivo in Bernsko konvencijo. Združbe, v katerih je vrsta 
dominantna, so uvrščene na zeleni seznam Ukrajine. Med združbami z vrsto 
Cladium mariscus v Ukrajini obstajajo znatne razlike v rastiščnih dejavnikih 
med zahodnimi (karbonatna barja) in južnimi (mokrišča in plavajoča močvirja 
ob severnih obalah Črnega morja) regijami. Avtorica je naredila primerjavo 
vegetacijskih popisov različnih združb s to vrsto v Evropi. Na osnovi rezultatov 
TWINSPAN analize je ugotovila štiri asociacije, ki jih je potrdila s florističnimi 
indeksi in ekološkimi podatki: Cladietum marisci Allorge 1921, Soncho maritimi-
Cladietum marisci (Br.-Bl. & O. de Bolòs 1957) Cirujano 1980, Dorycnio recti-
Cladietum marisci Gradstein & Smittenberg 1977 in Junco maritimi-Cladietum 
marisci (Br.-Bl. & O. de Bolòs 1957) Géhu & Biondi 1988. Ugotovila je tudi, 
da v Ukrajini poleg asociacije Cladietum marisci obstaja tudi asociacija Junco 
maritimi-Cladietum marisci.
Key words: Cladium mariscus 
communities, associations, 
marshland, swamp, Europe, Junco 
maritimi-Cladietum marisci.
Ključne besede: združbe z vrsto 
Cladium mariscus, asociacije, 
mokrišče, močvirje, Evropa, Junco 
maritimi-Cladietum marisci.
Received: 26. 3. 2019
Revision received: 21. 1. 2020
Accepted: 18. 5. 2020
1 M. G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine, 2 Tereschenkivska St., 01004, Kyiv, Ukraine. E-mail: anasta3Kz@gmail.com
Anastasia Davydova1 
DOI: 10.2478/hacq-2020-0010 19/2 • 2020, 275–291

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a new association for Ukraine
Introduction
The genus Cladium P . Browne comprises 3 species which 
are typical for temperate, subtropical and partly tropi-
cal regions (Plants of world…, 2020). One of them is 
C. mariscus (L.) Pohl, distributed in Eurasia and accepted 
in the wide sense to consist of an uncertain number of 
infraspecific taxa. In Ukraine, two species from this genus 
(C. mariscus (L.) Pohl and C. martii (Roem. et Schult.) K. 
Richt.) are indicated by I. M. Danylyk (Danylyk 2011, 
2012). The key for their determination proposed by Da-
nylyk was compiled from T. V. Egorova`s paper (1976), 
who accepted these taxa at subspecies level as C. mariscus 
subsp. mariscus and C. mariscus (L.) Pohl subsp. martii 
(Roem & Schult.) T.V. Egorova.
The latter subspecies has actually been described from 
Spain as Isolepis martii Roem. et Schult. (Egorova 1976). 
Tzvelev (1966) advanced the idea of a divided area of the 
genus Cladium in view of the evolutionarily younger C. 
mariscus represented in Europe. The areas of distribution 
of two older close species, C. grossheimii Pobed. and C. 
medwedewii (Meinsh.) Grossh., are situated along the 
Black Sea (Ukraine, Abkhazia) and near the Caspian Sea 
(Azerbaijan, Iran), respectively. Both these species are 
confined to coastal areas (Tzvelev 1966), but now they 
are mainly synonymized with C. mariscus subsp. martii 
which some other authors have considered to be a syn-
onym of C. mariscus subsp. mariscus (Castroviejo et al. 
2007, Verloove 2012). 
According to the Red Data Book of Ukraine, C. mari-
scus is protected on the territory of Shatsk National Na-
ture Park in the Volyn region and C. mariscus ssp. martii 
– in “Dzharylhatskyi” National Nature Park in the Kher-
son region (Andrienko et. al. 2009). In the Green Data 
Book of Ukraine, the communities of “Cladieta marisci” 
(association Cladietum (marisci) phragmitosum (australis), 
Cladietum marisci purum, Cladietum (marisci) caricosum 
(acutiformis), Cladietum (marisci) schoenosum (ferruginei) 
(Andrienko-Malyuk & Dubyna 2009) are indicated. Cla-
dium mariscus has had the status of a relict species con-
firmed by various palaeoecological studies (Pokorný et al. 
2010, Gałka & Tobolski 2012, Hájková et al. 2013). 
In addition to the inconsistency of taxonomical status 
of Cladium mariscus, there are several problems with the 
syntaxonomy of its communities in Ukraine. This species 
is diagnostic for the association Cladietum marisci Allorge 
1921 (Allorge 1921, Dengler et al. 2004). Currently, two 
isolated fragments of this species distribution are known 
from the territory of Ukraine – in western regions on cal-
careous fens and in southern regions on salt depressions 
between dune slacks. I therefore decided to compare the 
existing relevés from different locations in Ukraine with 
a protologue to discover significant differences based on 
ecological factors. There have previously been suggestions 
for classifying variants based on various ecological factors 
(Buczek 2005, Borsukevych 2008). However, these data 
concern only western Ukrainian communities on calcare-
ous fens and peatlands. 
The main objective was to conduct syntaxonomic 
analysis of Ukrainian plant communities dominated by 
Cladium mariscus and compare them with similar ones 
from other European regions. In addition, the author at-
tempted to study the features of ecological differentiation 
for these communities and to find differences in their flo-
ristic composition.
Material and methods
The studied area
The research territory was “Dzharylhatskyi” National Na-
ture Park located in the southern part of Ukraine. This 
area includes the island of Dzharylhach in the Black Sea 
and continental areas, which are mainly represented by 
forest cultures and ruderal plant communities. The island 
of Dzharylhach is 42 km long, 4.6 km at its widest (Arda-
matskaya et al. 2000). This island is an elevated sand-shell 
spit formed at the beginning of the Quaternary period 
by the accumulation of bottom sediments (Ardamatskaya 
et al. 2000). Between 700 and 1700 years ago, it was the 
eastern part of a single spit Dzharylhach-Tendra (Pravo-
torov 1967). The most represented soil types are sands 
and salt soils. The main climatic features are a predomi-
nance of eastern and north-eastern winds, relatively low 
humidity of the air, low cloudiness, slight precipitation 
and relatively large daily and annual amplitudes of air 
temperature fluctuations. The territory is located in the 
zone of hot-summer humid continental climate (Peel et. 
al. 2007). The average annual air temperature is + 10 °C. 
The lowest temperatures are observed in January (aver-
age monthly temperature -2.6 °C) and the highest values 
are recorded in July (+22.9 °C). Usually 270–345 mm 
of precipitation falls annually (Ardamatskaya et al. 2000, 
Shaposhnikova 2017).
There is only one type of vegetation in the studied 
area according to the Map of the Natural Vegetation of 
Europe – western and central Pontic desert steppes (Bohn 
et al. 2004).
Dataset
We used 172 relevés made by different authors according 
to the Braun-Blanquet approach (Braun-Blanquet 1964, 
Westoff & van der Maarel 1973) for critical analysis and 

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a new association for Ukraine
construction of the classification scheme. Six of them were 
made by the author on the research territory and five of 
them are unpublished relevés from Dzharylhach island by 
D. V. Dubyna. Other relevés were published in various 
papers and were carried out in other regions: 12 – in the 
western region of Ukraine (Datsyuk & Andrienko 2013, 
Chorna 2013) and 151 – in other countries of Europe: 
Great Britain (Wheeler 1980: 1 relevé), Greece (Theocha-
ropoulos et al. 2006: 18), Croatia (Stančić et al. 2010: 15), 
Crete (Gradstein & Smittenberg, 1977: 14), France (Al-
lorge 1921: 1), Hungary (Lájer 2006: 6), Ireland (Mooney 
& O’Connell 1990: 14), Italy (Biondi et al. 2006: 11, 
Taffetani 2011: 2, Lastrucci et al. 2017: 8, eVeg (Géhu 
& Biondi 1988): 5, eVeg (Géhu & Biondi 1994): 1), 
Netherlands (Verhoeven 1992: 19), Poland (Buczek 2005: 
18), Spain (Braun-Blanquet & de Bolos 1957: 5, Cirujano 
1980: 5, Rivas-Martínez et al. 1980: 5, Conesa 1991: 2), 
Switzerland (Dengler et al. 2004: 1) (Table 3, 4). 
Data analysis
The classification of the vegetation was conducted by 
Modified TWINSPAN (Roleček et al. 2009), implement-
ed in the software package JUICE 7.0.102 (Tichý 2002). 
T o clarify the limiting ecological factors, statistical analysis 
in the STATISTICA 7.0 software package and JUICE was 
used. In order to assess the impact of ecofactors and rela-
tionship of communities to the main ecological factors, a 
method of synphytoindication was used with the calcula-
tion for each species of vascular plants in all communities 
(Didukh 2011). Ellenberg indicator values were used for 
this purpose (Ellenberg et al. 1991). For each species in 
the relevés, quantitative (point) indicators are calculated 
for the six most important environmental factors.
The world map of Köppen-Geiger climate classification 
(Peel et. al. 2007) was used for the confirmation of the 
association distribution depending on temperature.
The combined synoptic table shows the combination 
of percentage frequency and modified fidelity index phi 
coefficient (Tichý 2002). The table shows species with phi 
values of frequency over 10. The phi fidelity index was 
used for diagnostic species identification. Non-essential 
values of fidelity (less than 0.001) were removed on the 
basis of Fischer’s exact test. The fidelity threshold for the 
allocation of diagnostic species is at least 25%, for highly 
diagnostic species – 50%. 
The Czekanowski-Sørensen coefficient was used for the 
calculation of floristic similarity: 
b a
c
Ksc
+
=
2
,
where a – number of species in first community; b – num-
ber of species in second community; c – number of spe-
cies common to the two communities. The limit values of 
Figure 1: Map of the research area: A – general location in eastern Europe, B – location within the southern part of Ukraine (green points – locali-
ties of Cladium mariscus).
Slika 1: Zemljevid raziskovanega območja: A – splošna lokacija v vzhodni Evropi, B – lokacija v južnem delu Ukrajine (zelene točke – lokacije 
vrste Cladium mariscus).

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a new association for Ukraine
this factor are from 0 to 1, Ksc = 1 absolute coincidence 
of floristic lists, Ksc = 0 they have no common species.
The Stugren-Radulescu coefficient of species composi -
tion similarity is calculated by the formula
c b a
c b a
Ksr
+ +
− +
=
3
.
This coefficient ranges from -1 to +1, the range from 1 
to 0 indicates a similarity, the range from 0 to +1 – shows 
a difference of floristic lists (Kostina 2013).
For comparison, I analyzed floristic lists (only the spe-
cies composition) from the relevés and identified clusters 
based on floristic composition.
The names of syntaxa were used in accordance with the 
International Code of Phytosociological Nomenclature 
(Weber et al. 2000). The higher syntaxonomic units are 
given in the latest prodromes of Europe (Mucina et al. 
2016) and Ukraine (Dubyna & Dziuba 2019). Names of 
vascular plant species are used in accordance with the re-
cent Ukrainian nomenclatural checklist (Mosyakin & Fe-
doronchuk 1999). Only some plant names with doubtful 
taxonomical status are cited with reference to the relevant 
sources (e.g. Mavrodiev et al. 2015). The names of the 
species in the synoptic table are given in accordance with 
the authors of the original works.
Results and Discussion 
Cladium in Europe: two species or 
one? 
Cladium mariscus and C. martii are morphologically very 
similar species. Their main features are: C. martii – broad 
panicle with diffused strongly branched lateral inflores-
cences, up to 20 cm in length with numerous elongated 
twigs, diffused clusters with 3–7(10) spikelets; C. mari-
scus – narrow and compact inflorescence with 5–20(30) 
spikelets aggregated into dense clusters (Egorova 1976, 
Danylyk 2011, 2012) (Figure 2).
Both C. mariscus and C. martii are geographically iso-
lated so they may be treated as subspecies of C. mari-
scus. However, our field studies did not confirm a clear 
geographical correlation with morphological features in 
Cladium mariscus populations. For example, photographs 
1 and 2 were taken on Dzharylhach Island (Ukraine) 
in different locations about 1200 m apart. Photo 3 was 
taken in the Lviv region at the same time. According to 
the current determination keys (Egorova 1976, Danylyk 
2011, 2012), plants from photographs 2 and 3 should be 
treated as C. mariscus subsp. mariscus and the plant from 
photograph 1 – as C. martii or C. mariscus subsp. martii.
In addition, herbarium specimens in KW, KWHA 
and MW from Germany, Italy, Poland, Czech Republic, 
Latvia, Estonia, Lithuania, Ukraine, Russian Federation, 
Belarus, Kazakhstan, Kyrgyzstan, Turkey, Azerbaijan and 
Georgia were analyzed and a similar situation was found: 
in many different locations both morphotypes (“south-
ern” with diffused inflorescences – Cladium martii-mor-
photype – and “western” with compact inflorescences – 
Cladium mariscus-morphotype) were identified. Based on 
these data, I treat Cladium mariscus as one polymorphic 
species and accept it in the wide sense to include C. martii 
as a heterotypic synonym of C. mariscus. This solution has 
also been confirmed by several other authors (Castroviejo 
et al. 2007, Verloove 2012).
Figure 2: Morphological differences between Cladium mariscus and C. 
martii (from Egorova 1976, modified by the author): 1 – C. mariscus: 
1a – spikelet, 1b – fruit, 2 – C. martii.
Slika 2: Morfološke razlike med vrstama Cladium mariscus in C. martii 
(iz Egorova 1976, spremenjeno): 1 – C. mariscus: 1a – klasek, 1b – 
plod, 2 – C. martii.

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a new association for Ukraine
Figure 3: Polymorphism of Cladium mariscus s.l.: 1– photo by D. A. 
Davydov (4 July 2017), 2 – photo by A. O. Davydova (6 July 2017), 
both – on Dzharylhach Island, 3 – photo by A. A. Kuzemko (23 June 
2017) near Zolochivka River in the Lviv region: 1a, 2a, 3a  – enlarge-
ments of lateral inflorescences and spikelets, 1, 2, 3 –general views of 
inflorescences of different individuals.
Slika 3: Polimofizem vrste Cladium mariscus s.l.: 1– foto D. A. Davy-
dov (4 julij 2017), 2 – foto A. O. Davydova (6 julij 2017), oboje – na 
otoku Dzharylhach, 3 – foto A. A. Kuzemko (23 junij 2017) ob reki 
Zolochivka v regiji Lviv: 1a, 2a, 3a – povečava stranskih cvetov in 
klaskov, 1, 2, 3 – splošni pogled na cvetove različnih primerkov.
Communities with Cladium 
mariscus in Ukraine
Cladium mariscus is distributed on calcareous wetlands 
in western Ukraine and on the banks of freshwater and 
salty reservoirs of the northern Black Sea Coast (An-
drienko-Malyuk & Dubyna 2009). Based on the dif-
ferences in ecological conditions of these communities, 
it would be logical to ensure that it is correct to merge 
relevés into the same association.
Figure 4: A general view of Cladium mariscus communities on 
Dzharylhach Island (photo by A. O. D    avydova).
Slika 4: Videz združbe z vrsto Cladium mariscus na otoku Dzharyl-
hach (foto A. O. Davydova).

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A Mediterranean element of the vegetation: Junco maritimi-Cladietum marisci – 
a new association for Ukraine
Cladium mariscus was found by the author in 6 loca-
tions (Figure 1) in the central part of Dzharylhach Island. 
The relevés were made mainly in dune slack habitats, with 
only a few locations being situated near lakes.
At the first stage, the author analyzed published data 
devoted to the syntaxonomy of Cladium mariscus com-
munities in Ukraine (Dubyna et al. 2004, Yaschenko & 
Turich 2007, Borsukevych 2008, Kuzyarin & Zhizhin 
2008, Datsyuk & Andrienko 2013, Chorna 2013) and 
compared the floristic composition with the data of the 
original relevé of the association Cladietum marisci (Al-
lorge 1921). The floristic composition was identified for 
all relevés (from western to southern regions) (species 
occurring in >50% of relevés): Cladium mariscus, Phrag-
mites australis, Lycopus europaeus, Lythrum salicaria, 
Mentha aquatica. So-called “vikarians” were established 
for the western and southern locations: Schoenus fer-
rugineus and S. nigricans, a group of freshwater sedges 
(Carex acutiformis, C. nigra, C. spicata) and a group 
of sedges on salt areas (C. extensa and C. distans). The 
common species from the original relevés (Allorge 1921) 
and the data from Ukraine are Eupatorium cannabinum, 
Molinia caerulea, Ranunculus lingua, Schoenoplectus 
tabemaemontani.
Comparison of Ukrainian and 
European communities
The authentic Cladietum marisci relevés were carried out 
in the north-western part of France, on the limestone pla-
teau on the right bank of the Seine (Allorge 1921). Ac-
cordingly, the protologue of this syntaxon refers to conti-
nental freshwater habitats, not coastal areas, which differ 
in both ecological conditions and floristic composition. 
The group of coastal Cladietum marisci relevés is charac-
terized by poor species composition, lacking a moss layer 
and the presence of halophytic species.
The next step was therefore to create a database of 
vegetation relevés with Cladium mariscus. It included se-
lected relevés from various publications and represented 
by various associations, subassociations and variants with 
domination of Cladium mariscus. Twenty-two floris-
tic lists from Europaean communities with C. mariscus 
were analysed in STATISTICA and three large clusters 
were identified combining similar floristic lists: cluster A 
(freshwater and mostly flooded communities with a large 
number of aquatic species and mosses, Cladietum marisci 
sensu stricto); cluster B (communities with low saliniza-
tion, Soncho maritimi-Cladietum marisci) and cluster C 
(more salinized communities, Junco maritimi-Cladietum 
marisci) (Figure 5).
At the lower levels, the clusters were distributed with 
an aberration: each of three large clusters has uncharac-
teristic inclusions of untypical communities, since the 
distribution of the floristic lists was calculated by Euclid-
ean distances, Ward’s method, which provides an accurate 
distribution at the highest levels. However, since the main 
task was to re-distribute the array of floristic data, the au-
thor focused on the allocation of three main clusters.
The next step was to apply the modified TWINSPAN 
algorithm for the allocation of individual clusters of rel-
evés (Roleček et al. 2009). At this stage, 162 relevés were 
used for analysis. I selected the most ecologically appro-
priate distribution of communities to six clusters at the 
association level (Figure 6). Clusters 1 and 2 represent 
the association Dorycnio recti-Cladietum marisci Grad-
stein & Smittenberg 1977 (Gradstein & Smittenberg 
1977). The distribution into two clusters is due to the 
presence of two clearly separated variants of the asso-
ciation – for Crete (typical) and for continental Europe. 
Based on the last critical revisions, the typical Dorycnio 
recti-Cladietum marisci association belongs to Central 
and Eastern Mediterranean tall-herb vegetation of Do-
rycnio recti-Rumicion conglomerati, Convolvuletalia sepi-
um and Epilobietea angustifolii (Gradstein & Smittenberg 
1977, Mucina et al. 2016).
The third cluster is Junco maritimi-Cladietum marisci 
(Br.-Bl. & O. de Bolòs 1957) Géhu & Biondi 1988 de-
scribed from Italy (eVeg database); the fourth one – Son-
cho maritimi-Cladietum marisci (Br.-Bl. & O. de Bolòs 
1957) Cirujano 1980 described from Spain. There are 
now two opinions about the classification of associations: 
various authors have synonymized them using the priority 
name Soncho maritimi-Cladietum marisci (eVeg database, 
Figure 5: Dendrogram based on floristic lists of communities with 
Cladium mariscus as dominant species: A – communities with a large 
number of aquatic species and mosses; B – communities with low 
salinization, C – more salinized communities. 
Slika 5: Dendrogram na osnovi florističnih popisov združb z 
dominantno vrsto Cladium mariscus: A – združbe z velikim številom 
vodnih vrst; B – združbe z manjšo slanostjo, C – združbe na bolj 
slanih rastiščih. 






 

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Landucci et al. 2020) or recognize both associations in 
parallel (Taffetani 2011, Pirone 2014, Pirone et al. 2014, 
Lastrucci et al. 2017, Pedrotti 2018, Habitats Naturels 
supports de la biodiversité 2018, Habitat Italia 2018). 
Both of these associations are present on saline soils but 
Junco maritimi-Cladietum marisci is characterized by the 
presence of numerous subhalophytic diagnostic species: 
Bolboschoenus maritimus, Juncus maritimus and Carex 
extensa. The relevés from Dzharylhach Island are also in 
cluster 3. Two associations from this cluster also belong 
to different syntaxa of the higher ranks. The association 
Junco maritimi-Cladietum marisci belongs to the alliance 
Scirpion maritimi from the order Bolboschoenetalia mar-
itimi (Biondi et al. 2014, Mucina et al. 2016, Habitat Ita-
lia 2018) and the association Soncho maritimi-Cladietum 
marisci belongs to the alliance Magnocaricion elatae from 
the order Magnocaricetalia (Rivas-Martínez et al. 2001, 
Mucina et al. 2016).
Two clusters (5 and 6) are distinguished as separate 
groups: 5 – Cladietum marisci (terrestrial variant) and 6 
– Cladietum marisci (water variant). Cluster 5 includes 
relevés from western Ukraine. The problem of the 
separation of these relevés into two variants is based on 
the flooding degree, but also depends on the technique 
of plot selection. For example, relevés of the typical 
Cladietum marisci by H. A. Chorna (Chorna 2013; 
Borysova & Chorna 2011) were originally indicated as 
the association Charetum tenuispinae Dąmbska 1966 ex 
Tomaszewicz 1979 from the class Charetea intermediae 
F. Fukarek 1961 and included Schoenus ferrugineus 
and Cladium mariscus as common species. At the same 
time, European relevés combine water and coastal-
water species in the association Utriculario-Cladietum 
(Jeschke 1963) Succow in Knapp et al. 1985 (Verhoeven 
1992, Zscheile & Schubert 2010). There is also a view 
of the classification of C. mariscus communities into 
successional stages based on paleoecological research in 
Poland: lake phase, lower peatland phase, upper peatland 
phase (Gałka & Tobolski 2012). It is possible that all 
flooded communities of C. mariscus with aquatic plants 
are the starting point of successional development of a 
“typical community”. It is obvious, though, that these 
plant communities do not fit the original diagnosis in 
terms of floristic composition and ecological conditions 
(Allorge 1921; Dengler et al. 2004). 
It was interesting to calculate the floristic similarity be-
tween selected clusters, so floristic lists for these clusters 
were prepared. Cluster analysis was conducted by STA-
TISTICA combining the Dorycnio recti-Cladietum mari-
sci group into a joint cluster (Figure 7).
Figure 6: Dendrogram of “similarity-difference” between plant 
communities with Cladium mariscus as dominant species in Europe: 
1, 2 – Dorycnio recti-Cladietum marisci; 3 – Junco maritimi-Cladietum 
marisci; 4 – Cladio marisci-Schoenetum nigricantis, Soncho maritimi-
Cladietum marisci; 5 – Cladietum marisci (terrestrial variant); 
6 – Cladietum marisci (water variant). 
Slika 6: Dendrogram podobnosti med rastlinskimi združbami na 
osnovi “razlike v podobnosti” z dominantno vrsto Cladium mariscus 
v Evropi: 1, 2 – Dorycnio recti-Cladietum marisci; 3 – Junco maritimi-
Cladietum marisci; 4 – Cladio marisci-Schoenetum nigricantis, Soncho 
maritimi-Cladietum marisci; 5 – Cladietum marisci (kopenska varianta); 
6 – Cladietum marisci (vodna varianta).
According to the Czekanowski-Sørensen coefficient 
(Ksc), the most significant differences were found between 
the clusters Junco maritimi-Cladietum marisci and terres-
trial and water variants of Cladietum marisci and this sup-
ports the author’s opinion about the difference between 
communities from western and southern Ukraine. Based 
on the Stugren-Radulescu coefficient of species compo-
sition similarity (Ksr,) clusters Junco maritimi-Cladietum 
marisci, Dorycnio recti-Cladietum marisci and typical Cla-
dietum marisci have the lowest similarity, too. The two 
groups that are closest according to floristic composition 
are: the Mediterranean Junco maritimi-Cladietum marisci 
and Soncho maritimi-Cladietum marisci, and the Central 
Figure 7: Dendrogram of “similarity-difference” between the plant 
communities based on floristic composition: 1 – Dorycnio recti-
Cladietum marisci; 2 – Junco maritimi-Cladietum marisci; 3 – Soncho 
maritimi-Cladietum marisci; 4 – Cladietum marisci (terrestrial variant); 
5 – Cladietum marisci (water variant).
Slika 7: Dendrogram podobnosti med rastlinskimi združbami na 
osnovi “razlike v podobnosti” z dominantno vrsto Cladium mariscus: 
1  – Dorycnio recti-Cladietum marisci; 2 –  Junco maritimi-Cladietum 
marisci; 3 – Soncho maritimi-Cladietum marisci; 4 – Cladietum marisci 
(kopenska varianta); 5 – Cladietum marisci (vodna varianta).
  

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A Mediterranean element of the vegetation: Junco maritimi-Cladietum marisci – 
a new association for Ukraine
and Eastern European water and terrestrial variants of 
Cladietum marisci (Table 1).
Table 1: Indices of coefficients of species composition similar-
ity (1 – Dorycnio recti-Cladietum marisci; 2 – Junco maritimi-
Cladietum marisci; 3 – Soncho maritimi-Cladietum marisci; 4 
– Cladietum marisci (terrestrial variant); 5 – Cladietum marisci 
(water variant).
Tabela 1: Indeksi podobnosti vrstne sestave (1 – Dorycnio 
recti-Cladietum marisci; 2 – Junco maritimi-Cladietum marisci; 
3 – Soncho maritimi-Cladietum marisci; 4 – Cladietum marisci 
(kopenska varianta); 5 – Cladietum marisci (vodna varianta).
Ksc
Ksr 1 2 3 4 5
1 - 0.10 0.14 0.12 0.07
2 0.78 - 0.22 0.05 0.09
3 0.64 0.49 - 0,14 0.10
4 0.73 0.88 0.69 - 0.22
5 0.84 0.82 0.77 0.20 -
The floristic composition similarity between the as-
sociations Junco maritimi-Cladietum marisci and Soncho 
maritimi-Cladietum marisci was 0.49 for the Stugren-
Radulescu and 0.22 for the Czekanowski-Sørensen coeffi-
cients. The overall floristic similarity of these two clusters 
is therefore about 25%. In the author`s opinion, these 
coefficients for two associations from the same climatic 
region cannot indicate their identity.
In order to find the differentiation of the most impor-
tant ecological factors, DCA-analysis was applied using 
ecological indicator values (Ellenberg et al. 1991). These 
results show that the most ecologically isolated is the Do-
rycnio recti-Cladietum marisci association. The most im-
portant factors for Junco maritimi-Cladietum marisci and 
Soncho maritimi-Cladietum marisci are soil reaction, light 
and temperature values. The ecological amplitude of the 
terrestrial variant of Cladietum marisci almost completely 
overlaps with the amplitude of the water variant. This 
ecological differentiation is determined by the level of soil 
moisture and the continentality of the climate.
The differentiation of the associations in relation to 
the level of light showed that the highest values for this 
indicator value are observed in Dorycnio recti-Cladietum 
marisci and Junco maritimi-Cladietum marisci. Water and 
terrestrial variants of Cladietum marisci are represented 
by lower values of light, which can be explained by the 
division of association habitats into coastal territories and 
continental ones (Figure 9).
Figure 8: Results of DCA ordination of the studied communities: 1, 
2 – Dorycnio recti-Cladietum marisci; 3 – Junco maritimi-Cladietum 
marisci; 4 – Soncho maritimi-Cladietum marisci; 5 – Cladietum marisci 
(terrestrial variant); 6 – Cladietum marisci (water variant). Values: 
MOIST – moisture; NUTR – nutrients; REACT – soil reaction; 
LIGHT – light; TEMP – temperature; CONT – continentality.
Slika 8: Rezultati DCA ordinacije obravnavanih združb: 1, 2 – Do-
rycnio recti-Cladietum marisci; 3 – Junco maritimi-Cladietum marisci; 
4 – Soncho maritimi-Cladietum marisci; 5 – Cladietum marisci (kopna 
varianta); 6 – Cladietum marisci (vodna varianta). Values: MOIST – 
vlažnost; NUTR – hranila; REACT – reakcija tal; LIGHT – svetloba; 
TEMP – toplota; CONT – kontinentalnost.
Differentiation of the associations based on tempera-
ture regime is presented by three groups according to the 
World Map of Köppen-Geiger Climate Classification 
(Peel et. al. 2007): Dorycnio recti-Cladietum marisci in the 
typical hot-summer mediterranean climate type (Csa), 
Junco maritimi-Cladietum marisci and Soncho maritimi-
Cladietum marisci partially represented in the hot-summer 
mediterranean climate, the warm-summer mediterranean 
climate and in the hot-summer humid continental cli-
Figure 9: Differentiation of the associations based on light values. 
Numbers along the abscissa axis correspond to the number of associa-
tions in Figure 8.
Slika 9: Razlike med asociacijami na osnovi vrednosti indeksa svetlobe. 
Številke ob abscisi odgovarjajo številkam asociacij na Sliki 8.
9.2
9.0
8.8
8.6
8.4
8.2
8.0
7.8
7.6
7.4
7.2
7.0
6.8
6.6
6.4
  1 2 3 4 5 6
Light
Group No.

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A Mediterranean element of the vegetation: Junco maritimi-Cladietum marisci – 
a new association for Ukraine
mate types (Csa, Csb, Dfa). The water and terrestrial vari-
ants of Cladietum marisci are mostly found in the temper-
ate oceanic and warm-summer humid continental climate 
types (Cfb, Dfb) (Figure 10). This pattern is also observed 
in the graph with continentality values: Dorycnio recti-
Cladietum marisci tends to the oceanic type, Junco mar-
itimi-Cladietum marisci and Soncho maritimi-Cladietum 
marisci tend to the suboceanic types, the terrestrial variant 
of Cladietum marisci has fairly wide amplitude from the 
oceanic to the suboceanic type, the water variant of Cladi-
etum marisci tends to the suboceanic type. The amplitude 
of the typical Cladietum marisci can be explained by the 
wide distribution of this syntaxon (Figure 11).
The moisture factor demonstrates a great difference 
between hydrophilous and more xerophytic associations 
of Dorycnio recti-Cladietum marisci. Junco maritimi-Cla-
dietum marisci is the most xerophytic of all the studied 
syntaxa. Other associations tend to temporarily flooded 
places (Figure 12).
Figure 10: Differentiation of the associations based on temperature 
values.
Figure 10: Razlike med asociacijami na osnovi vrednosti indeksa toplote.
7.2
7.0
6.8
6.6
6.4
6.2
6.0
5.8
5.6
5.4
5.2
5.0
4.8
4.6
Temperature
  1 2 3 4 5 6
Group No.
Figure 12: Differentiation of the associations based on moisture values.
Figure 12: Razlike med asociacijami na osnovi vrednosti indeksa 
vlažnosti.
Moisture
Group No.
10.2
10.0
9.8
9.6
9.4
9.2
9.0
8.8
8.6
8.4
8.2
8.0
7.8
7.6
7.4
7.2
  1 2 3 4 5 6
Figure 11: Differentiation of the associations based on continentality 
values.
Figure 11: Razlike med asociacijami na osnovi vrednosti indeksa 
kontinentalnosti.
Group No.
5.5
5.0
4.5
4.0
3.5
3.0
2.5
2.0
1.5
Continentality
  1 2 3 4 5 6
Figure 13: Differentiation of the associations based on nutrient values.
Figure 13: Razlike med asociacijami na osnovi vrednosti indeksa hranil.
7.0
6.5
6.0
5.5
5.0
4.5
4.0
3.5
3.0
2.5
2.0
  1 2 3 4 5 6
Nutrients
Group No.

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Anastasia Davydova
A Mediterranean element of the vegetation: Junco maritimi-Cladietum marisci – 
a new association for Ukraine
Differentiation of the associations based on the nutri-
ent content in soils indicates a significant differentiation: 
water Dorycnio recti-Cladietum marisci is marked by the 
lowest trophic level and the typical by a moderate level. 
The broadest amplitude is observed in typical (terrestrial) 
Cladietum marisci and in Soncho maritimi-Cladietum 
marisci. Junco maritimi-Cladietum marisci tends to soils 
that are moderately rich in nutrients (Figure 13).
Differentiation of the associations based on soil reac-
tion showed that the highest alkaline reaction is observed 
in Dorycnio recti-Cladietum marisci associations. The 
communities of Junco maritimi-Cladietum marisci grow 
on slightly alkaline soils. Values of the typical Cladietum 
marisci range from neutral to carbonate soils. Water Cla-
dietum marisci is closer to neutral soils. Values for Soncho 
maritimi-Cladietum marisci also indicate the predomi-
nance of carbonates in soils (Figure 14). 
As a result of this analysis, I propose recognition of 
these syntaxa dominated by Cladium mariscus: Cladietum 
marisci Allorge 1921, Soncho maritimi-Cladietum marisci 
(Br.-Bl. & O. de Bolòs 1957) Cirujano 1980, Dorycnio 
recti-Cladietum marisci Gradstein & Smittenberg 1977 
and Junco maritimi-Cladietum marisci (Br.-Bl. & O. de 
Bolòs 1957) Géhu & Biondi 1988. I believe that Cladio 
marisci-Schoenetum nigricantis sensu Lájer 2006 non Soó 
1930 is a synonym of Soncho maritimi-Cladietum mari-
sci. The most controversial associations Soncho maritimi-
Cladietum marisci and Junco maritimi-Cladietum marisci 
are recognized as separate, based on floristic comparison 
and ecological analysis. The merging of these syntaxa into 
one was proposed recently by Landucci et al. (2020), but 
this was probably due to the small number of relevés with 
Juncus maritimus used in that study. The association Junco 
maritimi-Cladietum marisci is new for Ukraine from the 
Dzharylhach Island. Its special feature is the presence of 
Poacynum russanovii (Mavrodiev et al. 2015) (Table 2), 
an endemic species for this island. The following syntaxo-
nomic scheme for communities with Cladium mariscus 
for “Dzharylhatskyi”  National Nature Park is proposed:
Phragmito-Magnocaricetea Klika in Klika & Novák 1941
Bolboschoenetalia maritimi Hejný in Holub et al. 1967 
Scirpion maritimi Dahl & Hadač 1941
Junco maritimi-Cladietum marisci (Br.-Bl. & O. de 
Bolòs 1957) Géhu & Biondi 1988
Diagnostic species of this association: Carex distans, Cla-
dium mariscus, Juncus maritimus, Mentha aquatica, 
Phragmites australis, Poacynum russanovii, Pulicaria dys-
enterica.
Constant species: Elytrigia elongata.
Dominant species: Cladium mariscus, Phragmites australis.
In relation to the order Bolboschoenetalia maritimi as 
part of Phragmito-Magnocaricetea (Biondi et al. 2014, 
Mucina et al. 2016), the author does not agree with the 
separation of the class Bolboschoenetea maritimi Tx. & 
Vicherek in Tx. & Hülbusch 1971 that is widely ac-
cepted in many Ukrainian papers (Dubyna et al. 2007, 
Dziuba 2008).
As a result of the studied communities, a distribution 
map of the syntaxa and their types (holotype and neo-
type) is proposed (Figure 15). Two syntaxa – Cladietum 
marisci and Junco maritimi-Cladietum marisci - are thus 
distributed in Ukraine. In the southern part of Ukraine 
Cladium mariscus grows on the territory of the “Danube 
Biosphere Reserve” in the Odesa region and also on the 
“Chornomorsky Biosphere Reserve”. Based on the floris-
tic composition of relevés in the synoptic table from the 
Danube delta (Dubyna et al. 2004), the author consid-
ers them to be Cladietum marisci, although these com-
munities need further investigation. The map shows the 
essential distance (1 358 km) between the locus typicus 
of Junco maritimi-Cladietum marisci and the location of 
the syntaxon indicated by the author. The Mediterranean 
climate partly explains the closeness of Junco maritimi-
Cladietum marisci and Soncho maritimi-Cladietum marisci 
areas, but what is the explanation in this case? Based on 
floristic studies, Illichevskyi (1941) put forward a hypoth-
esis about the origin of Dzharylhach Island as “a kind of 
relic of the former land” and extrapolated the results of 
geological surveys, observing that the flora of the island 
partly consists of rare species with Mediterranean and 
disjunctive areal characteristics typical of the Crimean 
Peninsula, the Balkans, the Caucasus and Asia Minor. Il-
Figure 14: Differentiation of the associations based on soil reaction 
values.
Figure 14: Razlike med asociacijami na osnovi vrednosti indeksa 
reakcije tal.
Group No.
9.5
9.0
8.5
8.0
7.5
7.0
6.5
6.0
5.5
5.0
Soil Reaction
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A Mediterranean element of the vegetation: Junco maritimi-Cladietum marisci – 
a new association for Ukraine
Releve’s № 1 2 3 4 5 6 7 8 9 10 Constancy Phi coeff.
Releve’s area (m²) 25 25 50 50 30 50 50 70 15 50
Number of species 9 7 9 9 8 8 9 4 7 8
Total cover (%) 80 90 90 80 90 100 100 90 90 100
D. s. for the ass. Junco maritimi-Cladietum marisci
Cladium mariscus 3 3 2 2 5 5 5 5 2 5 V 100
Juncus maritimus 2 + + . 2 2 2 . 3 . IV 78.2
Carex distans + . + 2 2 r . . . + III 72.1
Poacynum russanovii . . r 2 . r r . + . III 65.5
D. s. for the cl. Phragmito-Magnocaricetea
Mentha aquatica + 2 2 . + . . + . 2 III 78.2
Phragmites australis 2 2 3 + 2 3 2 2 2 1 V 50.8
Pulicaria dysenterica + . + . 2 . . + . . III 58.3
Althaea officinalis . + . . . . . . . . I 28.7
Calystegia sepium . . . . . . r . . + I 40.8
Other species
Apera maritima . . . . . . . . + . I -
Calamagrostis epigejos  . + + . + . . . . . II -
Carex extensa . . . + . . . . . . I 28.7
Cirsium alatum + . . r . . r . . . III 45.1
Cynanchum acutum 2 . . . . . . . . . I -
Elytrigia elongata + 2 . + . r r . 2 . IV 45.4
Festuca pratensis . . . . + . . . . . I 28.7
Heracleum sibiricum . . . . . . r . . . I 28.7
Inula salicina subsp. aspera . . . + . . r . . 2 II -
Limonium gmelinii . . . . . r . . + . II -
Lythrum virgatum . . . . . . . . . + I -
Plantago cornuti . . . r . . . . . . I -
Plantago maxima . . . . . + . . . . I 28.7
Schoenus nigricans . . . . . . . . . + I
Thalictrum minus . . + . . . . . . + I -
Localities:
1 – N 46° 01' 543", E 32° 94' 92" (09. 07. 2017); 6 – Island Dzharylhach, D. Dubyna (09. 2000); 
2 – N 46° 01' 578", E 32° 93' 535" (09. 07. 2017); 7 – Island Dzharylhach, D. Dubyna (09. 2000); 
3 – N 46° 01' 517", E 32° 94' 677" (09. 07. 2017); 8 – N 46° 01' 217", E 33° 02' 627" (09. 07. 2017);
4 – Island Dzharylhach, D. Dubyna (05. 2002);  9 – N 46° 01' 543", E 32° 94' 92" (09. 07. 2017);
5 – N 46° 01' 472", E 32° 93' 683" (09. 07. 2017); 10 – Island Dzharylhach, D. Dubyna (05. 2001).
Table 2: Phytocoenological table of assotiation Junco maritimi-Cladietum marisci.
Tabela 2: Fitocenološka tabela acociacije Junco maritmi-Cladietum marisci.
lichevskyi concludes that Dzharylhach was originally part 
of a larger territory connected with the Crimea, Balkans 
and Asia Minor (Illichevskyi 1941). This idea was also 
confirmed by later research by geologists (Ryan et al. 
1997). Barbarych (1962) believed that the distribution of 
the genus Cladium in the past tended to the sea coasts, so 
its growth in the Eastern and Central Europe is also due 
to the presence of sea basins. The age of origin of southern 
Ukrainian localites is Pleistocene and western Ukrainian 
– Miocene (Barbarych 1962). 
Cladium mariscus therefore has a disjunct habitat. The 
existence of individual localities of C. mariscus with halo-
philic elements is due to regional features – soil type, 
 salinization and flooding.
Cladium mariscus s.l. is a key species in the habitats 
from Habitat Directive “7210 Calcareous fens with  Cla-
dium mariscus and species of the Caricion davallianae” (In-
terpretation Manual of European Union Habitats 2013) 
and in “D 5.2 Large thickets without free standing water” 
of Resolution No. 4 of the Bern Convention (2016) 

19/2 • 2020, 275–291
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A Mediterranean element of the vegetation: Junco maritimi-Cladietum marisci – 
a new association for Ukraine
 (Perrino et al. 2013, Detailed final conclusions … 2016, 
Habitats Naturels supports de la biodiversité 2018, 
Habitat Italia 2018). We propose distinguishing a new 
sub-type of habitat D 5.2 “Large thickets without free 
standing water” (“D 5.2.1. Halophytic wetlands and wet 
depressions with Cladium mariscus”). 
Conclusions
The isolated position of plant communities with Cladium 
mariscus s.l. on Dzharylhach Island was thus confirmed 
on the basis of the results of the analysis of geobotanical 
relevés. A new subtype of habitat (“D 5.2.1. Halophytic 
wetlands and wet depressions with Cladium mariscus”) 
is proposed. The name “ Cladio marisci-Schoenetum nig-
ricantis sensu Lájer 2006 non Soó 1930” is a synonym 
of Soncho maritimi-Cladietum marisci. The floristic and 
ecological independence of Soncho maritimi-Cladietum 
marisci (Br.-Bl. & Bolòs 1957) Cirujano 1980 is sub-
stantiated. A new association Junco maritimi-Cladietum 
marisci (Br.-Bl. & Bolòs 1957) Géhu & Biondi 1988 for 
vegetation of Ukraine was confirmed. Further research 
of Cladium mariscus communities is needed, particularly 
a comparison with data from the Caucasus and Western 
Asia. 
Acknowledgements
The autor are very grateful to editor in chief Urban Šilc 
and two anonymous reviewers for comments and recom-
mendations that greatly improved the manuscript. Special 
thanks are due to Denis Davydov and Martin Cregeen for 
English proofreading, to Dmytro Dubyna for providing 
the relevés and to Anna Kuzemko for providing images.
Anastasia Davydova , https://orcid.org/0000-0001-7839-962X
Figure 15: Map of syntaxa based on the results of this study.
Slika 15: Zemljevid sintaksonov na osnovi rezultatov te raziskave.











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A Mediterranean element of the vegetation: Junco maritimi-Cladietum marisci – 
a new association for Ukraine
Syntaxa Country Year № of table/releve
Cladietum marisci Spain 1957 tab. 15, rel. 1–5
Great Britain 1980 tab. 3, rel. 2
Ireland 1990 tab. 2, rels. 16–29
Netherlands 1992 tab. 4, rels. 1–19
Poland 2005 tab. 1, rels. 10–18
Croatia 2010 tab. 3, rels. 24–34
Ukraine 2013 tab. 31, rels. 1–10 
Ukraine 2013 p. 10–11
Italy 2017 tab. 8, rels.1–8
Cladietum marisci var. Carex elata Poland 2005 tab. 1, rels. 1–9
Soncho maritimi-Cladietum marisci Italy 1980 tab. 4, rels. 1–3
Spain 1980 tab. 7, rels. 1–5
Italy 1991 tab. 7, rels. 1–2
Soncho maritimi-Cladietum marisci myrtetosum communis Italy 2006 tab. 4, rels. 4–6
Soncho maritimi-Cladietum marisci schoenetosum nigricantis Spain 1980 tab. 7, rels. 10, 12, 14, 17, 19
Junco maritimi-Cladietum marisci Italy 1988 rels. 1–5
Italy 1994 tab. 70 p. 120
Italy 2011 tab. 15, rels. 3–4
Ukraine 2000–2018 in abstract
Dorycnio-Cladietum marisci typicum Crete 1977 tab. 6, rels. 12–17
Greece 2006 tab. 2, rels. 16–27
Dorycnio-Cladietum marisci inops Crete 1977 tab. 6, rels. 18–25
Cladio marisci-Schoenetum nigricantis Hungary 2006 tab. 2, rels. 16–19
Schoeno nigricantis-Erianthetum ravennae cladietosum marisci Italy 2006 tab. 7, rels.1–4
Table 3: Data about the geobotanical relevés included in the database.
Tabela 3: Podatki o vegetacijskh popisih vključenih v podatkovno bazo.
Table 4: Synoptic table with percentage frequency and modified fidelity index phi coefficient.
Table 4: Sinoptična tabela s frekvencami v odstotkih in modificiranim fi koeficientom navezanosti.
Group Number 1 2 3 4 5 6
Number of relevés 4 8 21 29 66 32
Phragmites australis . . 67 39 62 91
39.7
Cladium mariscus 100 100 100 100 100 100
Carex lasiocarpa . . . . 2 62
73.1
Lythrum salicaria . . 5 28 26 59
44.8
Carex elata . . . . 2 53
68.4
Galium palustre . . 5 . 5 50
60.1
Lysimachia vulgaris . . . 3 33
27.3
41
37
Peucedanum palustre . . . . 3 41
57.4
Comarum palustre . . . . 2 38
56.2
Utricularia vulgaris . . . 24
20.4
. 38
40.1
Calliergonella cuspidata . . . . 2 34
53.5
Scorpidium scorpioides . . . . . 34
Utricularia minor . . . . 2 31
50.8
Menyanthes trifoliata . . . . . 28
49.6
Mentha aquatica 50 38 28 35 25
Juncus subnodulosus . . . 21 5 25
26.8
Nymphaea alba . . . . . 25
46.6
Equisetum fluviatile . . . . 9 25
37.3
Calliergon cuspidatus . . . . 2 25
44.8

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Group Number 1 2 3 4 5 6
Number of relevés 4 8 21 29 66 32
Carex buxbaumii . . . . 2 22
41.5
Campylium stellatum . . . . . 19
40.2
Lycopus europaeus . . . 10 14 19
Lemna minor . . . . 2 19
38.1
Salix cinerea . . . . 3 19
36.2
Lysimachia thyrsiflora . . . . 3 16
32.2
Potentilla palustris . . . . 2 16
34.3
Epilobium palustre . . . . 6 16
28.8
Drepanocladus revolvens . . . . . 16
36.6
Stachys palustris . . . . 2 16
34.3
Lemna gibba . . . . . 12
32.6
Thelocarpon pallidum . . . . 3 12
27.9
Chara species . . . . 3 12
27.9
Lathyrus palustris . . . . . 12
32.6
Galium uliginosum . . . . . 12
32.6
Spirodela polyrhiza . . . . . 12
32.6
Poa palustris . . . . . 12
32.6
Carex rostrata . . . . 2 12
30.1
Eupatorium cannabinum . . . . 23
37.3
6
Solanum dulcamara . . . . 15
27.9
6
Typha angustifolia . . 5 . 18
27.7
6
Schoenus nigricans . . 14 72
69.6
2 6
Agrostis stolonifera . . . 17
27.9
3 6
Typha angustata 25 . . . 2 3
Hydrocotyle vulgaris 25 . . . 3 3
Iris pseudacorus . 50
62.4
. . 3 3
Carex panicea . . . . 11
24.9
3
Holoschoenus romanus . 12 . 3 . .
Carex hispida 25 25 . 21
12.4
. .
Linum maritimum . . . 14
34.3
. .
Saccharum ravennae . . . 17
38.5
. .
Equisetum ramosissimum . 12 . 3 . .
Rubus ulmifolius . . . 17
38.5
. .
Carex distans . 25 29
29.1
3 . .
Pulmonaria dacica 25
46.6
. . . . .
Arundo donax 75
84.5
. . . . .
Polypogon monspeliensis 25
46.6
. . . . .
Galium constrictum 25 12 5 . . .
Nerium oleander                                     . 12 . . . .
Anagallis tenella . 12 . . . .
Sonchus glaucescens . 12 . . . .
Panicum repens . 25
46.6
. . . .
Rubus sanctus . 38
57.7
. . . .
Oenanthe pimpinelloides 25 62
60.
. . . .
Poa trivialis s. sylvicola . 25
46.6
. . . .
Juncus heldreichianus 25 38
39.6
. . . .
Rumex conglomeratus . 25
46.6
. . . .
Agrostis semiverticillata 25
34.6
12 . . . .
Lythrum junceum 50 25
16.9
. . . .
Imperata cylindrica 25
50.7
25
27.0
. . . .
Cynodon dactylon . 12 . . . .

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a new association for Ukraine
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Group Number 1 2 3 4 5 6
Number of relevés 4 8 21 29 66 32
Inula viscosa . 25 . 28
29.8
. .
Ipomoea sagittata . . 5 31
47.3
. .
Myrtus communis . . . 24
45.8
. .
Phillyrea media . . . 14
34.3
. .
Sonchus maritimus . . 10 45
55.7
. .
Oenanthe lachenalii                                 . . 5 34
50.5
. .
Juncus acutus . . 14
26.1
7 . .
Juncus articulatus . 12 . . . .
Gaudinia fragilis . 12 . . . .
Rubia peregrina . . . 10
29.6
. .
Juncus maritimus . . 76
80.7
7 . .
Orchis laxiflora subsp. palustris 25 25
22.2
. 10 . .
Cirsium creticum . 38
46.6
. 10 3 .
Samolus valerandi 25 12 . 48
42.8
2 .
Apium nodiflorum 50
59.1
. 5 . 6 .
Dorycnium rectum 25 75
66.4
. 3 5 .
Calamagrostis epigejos . . 14
28.6
. 5 .
Calystegia sepium . . 38
32.7
31
23.4
11 .
Carex flava . . . . 11
30.0
.
Sparganium erectum . . . . 11
30.0
.
Pulicaria dysenterica 25 29
24.7
10 3 .
Cirsium alatum . . 14
34.9
. . .
Inula salicina subsp. aspera . . 14
34.9
. . .
Poacynum russanovii . . 24
45.5
. . .
Elytrigia elongata . . 29
50.0
. . .
Carex extensa . . 19
40.5
. . .
Inula crithmoides . . 14
34.9
. . .
Festuca arundinacea . 12 . . 2 .

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