181
Synecology of Cutandia maritima (L.) 
Barbey, a rare psammophytic species 
along the Montenegrin Coast (East 
Adriatic Coast)
Abstract
Cutandia maritima is a circum-Mediterranean species that inhabits sandy dunes 
along the coast line. It is fairly frequent on the western Adriatic coast but fairly 
rare and possibly even non-native in the east. In Croatia, it was discovered in 
1990 in Crnika Bay on the island of Rab, which was considered until 2005 to be 
the only site on the eastern Adriatic coast from the Gulf of Trieste in the north 
to Corfu in the south. In 2009, the species was briefly reported for Velika plaža 
(Long Beach) in Ulcinj (Montenegro) but without details about the habitat 
type and synecology. The aim of this paper is thus to provide a deeper insight 
into the ecology and synecology of C. maritima in the eastern Adriatic part of 
the distribution area. On Velika plaža in Ulcinj, the species was found along the 
whole sea-inland gradient of sand dunes, in various types of vegetation: [1210] 
– annual vegetation of drift lines, [2110] – embryonic shifting dunes, [2120] 
– shifting dunes with Ammophila arenaria (white dunes), [2220] – dunes with 
Euphorbia terracina, [2130*] – fixed coastal dunes with herbaceous vegetation 
(grey dunes), and also [2190] – humid dune slacks.
Izvleček
Cutandia maritima je vrsta, ki uspeva na peščenih sipinah ob morskih obalah ob 
celotnem Mediteranu. Ob zahodnih obalah je precej pogosta, medtem ko je na 
vzhodu redka ali celo tujerodna. Na Hrvaškem so jo odkrili leta 1990 v zalivu 
Crnika na otoku Rabu in to je bilo do leta 2005 edino nahajališče na vzhodni 
Jadranski obali med Tržaškim zalivom na severu in otokom Krfom na jugu. O 
najdbi vrste na Veliki plaži pri Ulcinju so na kratko poročali leta 2009, a brez 
podrobnosti o njenem rastišču in sinekologiji. V članku podajamo natančnejši 
vpogled v ekologijo in sinekologijo vrste C. maritima ob vzhodni Jadranski obali. 
Na Veliki plaži vrsta uspeva na peščenih sipinah na celotnem gradientu od morja 
v notranjost in v različnih vegetacijskih tipih: [1210] – Združbe enoletnic na 
obalnem drobirju, [2110] – Embrionalne premične sipine, [2120] – Premične 
sipine vzdolž obrežij z vrsto Ammophila arenaria (bele dine), [2220] – Sipine z 
vrsto Euphorbia terracina, [2130*] – Utrjene obalne sipine z zeliščno vegetacijo 
(sive dine), in [2190] – Vlažne uleknine na sipinah.
Key words: Cutandia maritima, 
Velika plaža in Ulcinj, sand dunes, 
vegetation.
Ključne besede: Cutandia maritima, 
Velika plaža pri Ulcinju, peščene 
sipine, vegetacija.
Received: 2. 12. 2015
Revision received: 31. 1. 2017
Accepted: 2. 2. 2017
1 Faculty of Natural Sciences and Mathematics, Biology Department, Džordža Vašingtona bb, 81000 Podgorica, Montenegro
 *Corresponding author: danijela.stesevic@ac.me
2 Natural History Museum of Montenegro, Trg vojvode Bećir-bega Osmanagića 16, 81000 Podgorica, Montenegro
3 Department of Biology, Biotechnical Faculty, University of Ljubljana, Večna pot 111, SI-1000 Ljubljana, Slovenia
4 Faculty of Hotel Management and Tourism Vrnjačka Banja, University of Kragujevac, Vojvođanska 5A, 36210 Vrnjačka Banja, Serbia
5 ZRC SAZU, Institute of Biology, Novi trg 2, SI-1000 Ljubljana, Slovenia
6 BC Naklo, Strahinj 99, Naklo, Slovenia
Danijela Stešević1,*, Nada Bubanja2, Danka Caković1, Nejc Jogan3,  
Milica Luković4 & Urban Šilc5,6
DOI: 10.1515/hacq-2017-000216/2 • 2017, 181–187
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Synecology of Cutandia maritima (L.) Barbey, a rare psammophytic species along 
the Montenegrin Coast (East Adriatic Coast)
16/2 • 2017, 181–187
182
Introduction
Although Cutandia maritima (L.) Barbey is consid-
ered to be a common species on sandy dunes along the 
Mediterranean coast (van der Maarel & van der Maarel-
Versluys 1996), its record in Montenegro (Velika plaža 
in Ulcinj) dates from 2005 (N. Jogan, unpublished); it 
was published in 2009 (Glasnović et al. 2009). It appears 
frequently on the western Adriatic coast (reported for all 
countries east of Italy (Pignatti 1982), while along the 
eastern coast it is fairly rare (Trinajstić 1994) and may 
even have been recently introduced (Horvat 1939). It 
is considered to be a focal species of 2230 EU habitat – 
Malcolmietalia dune grasslands (Stanisci et al. 2014) and 
occurs in various plant communities of the Ammophiletea, 
Cisto-Micromerietea and Helminthemetea classes (Goday 
& Martinez 1958, Bolos 1996, Costa et al. 2000, Sýkora 
et al. 2003, Zimmitti et al. 2015 etc.). It is a character-
istic species of sabulicolous therophytic plant communi-
ties, which Rivas-Martínez et al. (2002) described as the 
Cutandietalia maritimae order within the class Tuberari-
etaria guttatae (syn. Helianthemetea guttati). This order 
comprises ephemeral vegetation growing on coastal sands 
and dunes with salt spray, as a therophytic vegetation type 
among the perennial vegetation of Ammophiletea, and it is 
distributed over the whole Mediterranean (Díez-Garretas 
et al. 2003). C. maritima is recorded in ca. 30 ephemeral 
plant communities that inhabit coastal sands and sand 
dunes with salt spray, among perennial vegetation of Am-
mophiletea, Crucianelletea and sometimes also Quercetea 
ilicis (Díez-Garretas et al. 2003, Minissale & Sciandrello 
2015). It appears as a dominant species in 7 associations 
and sub-associations: Vulpielo tenuis-Cutandietum mar-
itimae (Rivas-Martinez et al. 2002), Cutandio maritimae-
Parapholidetum marginatae (Bartolo et al. 1988), Sileno 
sericeae-Cutandietum maritimae (Géhu & Biondi 1994), 
Sileno nicaensis-Cutandietum maritimae (Géhu & Biondi 
1994), Matthiolo tricuspidatae-Cutandietum maritimae 
(Géhu et al. 1987), Sileno succulentae-Cutandietum mar-
itimae (Géhu & Géhu-Franck 1986), Sileno ononidetum 
variegatae subass. Cutandietosum maritimae (Minissale & 
Sciandrello 2015), mainly distributed in the western part 
of the Mediterranean. 
Sabulicolous therophytic plant communities in the 
Mediterranean are also classified into the order Malcol-
mietalia. Both orders are mainly dominated by annual 
plants with a short lifecycle growing on non-nitrified deep 
sandy soils. The optimum of Malcolmietalia communities 
is siliceous sands or paleodunes in the western Mediter-
ranean, where they are a subseral stage of Quercus suber 
forests or Juniperus navicularis and J.  broussonetii open 
woodlands. They are assumed to be indicators of conti-
nentality, whereas those of Cutandietalia are indicators 
of less expressed climatic conditions. Good correspond-
ence is also shown along the altitudinal gradient and dis-
tance from the sea (Minissale & Sciandrello 2015). In the 
eastern Mediterranean, C. maritima is characteristic of 
the central fore-dune ridge (Doing 1985).
In other parts of the Mediterranean, the species C. mar-
itima occurs rarely in Cakiletea plant communities (fre-
quency class lower than II), while it is more frequent (fre-
quency class IV) in plant communities of Ammophiletea 
and has the status of differential species (e.g. Limonio 
graeci-Triplachnetum nitentis centaureetosum aegialophilae 
Mayer 1995) (Brullo & Furnari 1970, Brullo et al. 1974, 
Perdigó & Papió 1985, Sykora et al. 2003 etc.). It is also 
a companion species of chamaephytic and hemicrypto-
phytic Crucianelletalia maritimae plant communities (e.g. 
Centaureo-Oninidetum ramosissimae) usually confined to 
dunes further inland with little movement (Zimmitti et 
al. 2015), as well as Malcolmietalia Rivas Goday 1958 
communities (e.g. Violo henriquesii-Silenetim littoreae 
Izco & Guitian 1988; Costa et al. 2000). 
In previous studies of sand dune vegetation of Velika 
plaža in Ulcinj, only communities of Cakiletea mariti-
mae and Ammophiletea were mentioned (Trinajstić 1989, 
Mijović 1994, Mijović et al. 2006, 2012, Šilc et al. 2015), 
while sabulicolous therophytic communities were not 
taken into account. The aim of this paper is to provide 
a deeper insight into the ecology and synecology of the 
species C. maritima in Montenegro and to initiate more 
detailed investigation of annual sand dune vegetation on 
Velika plaža. 
Materials and methods
Study area
Velika plaža in Ulcinj (Velika plaža, Montenegro) is con-
sidered to be the longest beach on the eastern Adriatic 
coast (ca. 12km), with still well preserved sand dune veg-
etation. Since 1968, the beach has been recognized as a 
monument of nature (Official Gazette of the Social Re-
public of Montenegro, SRCG 30/68). According to the 
Spatial Plan of Special Purpose for the Coastal Zone, one 
spatial unit on the east side of Velika plaža is recognized 
as a new protected nature reserve, with forests, marshes 
and meadows. The area is protected from trampling or 
grazing with a wooden fence (JPMD 2015, Šilc et al. 
2017). However, the rest of the beach, which represents 
more than 75 % of natural coast and 97 % of whole 
sand dune system, is heavily impacted by tourism, illegal 
dumping, sand exploitation and urbanization (Petrović 
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16/2 • 2017, 181–187
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D. Stešević, N. Bubanja, D. Caković, N. Jogan, M. Luković & U. Šilc
Synecology of Cutandia maritima (L.) Barbey, a rare psammophytic species along 
the Montenegrin Coast (East Adriatic Coast)
& Karaman 2009). Eleven NATURA 2000 habitats are 
reported for the beach and its hinterland (Petrović et 
al. 2012) and the site is included in the list of IPA areas 
(Petrović & Karaman 2009).
Data collection and analysis
During 2015, the floristic composition and zonation of 
plant communities on Velika plaža in Ulcinj were stud-
ied. Twenty transects were set perpendicular to the coast, 
at a distance from each other of cca. 500 m. Contiguous 
quadrats (2 × 2 m) were laid out in the form of a belt 
transect (Kent 2012), starting from the area with the first 
colonizing plants towards the end of the sand dune sys-
tem up to forest vegetation. In each quadrat, the cover 
of plant species was visually estimated by the Braun-Bl-
anquet (1964) scale. Plants were determined following 
Pignatti (1982) and Tutin et al. (1964-80). All gathered 
relevés were entered into the TURBOVEG (Hennekens 
& Schaminée 2001) database. 
Multivariate analysis (Ward method and Euclidean 
distance) of the plots was performed in PC-ORD (Mc-
Cune& Grace 2002) to recognize plant communities on 
sand dunes. Clusters were identified by their diagnostic 
species, using the JUICE program (Tichý 2002). Three 
species with the highest fidelity were selected for each 
cluster. Relevés with C. maritima were selected, present-
ed in a single table (Table 1) and classified into different 
clusters.
Detrended Correspondence Analysis (DCA) ordina-
tion was used to visualize the floristic similarity of plots. 
Species cover values (mean percentage values) were 
square-root transformed. DCA was done in CANOCO 5 
program (ter Braak & Šmilauer 2012).
The nomenclature of taxa is according to Tutin et al. 
(1964-80).
Results 
Within the part of the study concerning the floristic 
composition and zonation of plant communities on Ve-
lika plaža in Ulcinj, 20 transects were set perpendicularly 
from the sea towards inland and 876 relèves were done. 
In further analysis, they were classified into 9 clusters 
corresponding to 9 plant communities (Cakilo-Xanthie-
tum italici, Cynodon dactylon comm., Euphorbio paraliae-
Agropyretum junceiformis, Medicagini marinae-Ammophi-
letum australis, Onobrychis caput-galli comm., Cladietum 
marisci, Eriantho-Schoenetum nigricantis, Juncetum mar-
itimo-acuti, Holoschoenetum romani) or to 7 NATURA 
2000 habitats (Figure 1). The species C. maritima was 
recorded in 4 transects and 33 quadrats (Table 1), dis-
tributed in a bandwidth from ca. 80 m from the sea to 
150 m inland. Its cover value is lower (r, + and 1) in mo-
bile parts of dunes and higher (1) in more stabilized parts 
of dunes covered with Onobrychis caput-galli comm. and 
Holoschoenetum romani (Figure 1).
During the floristic survey along Velika plaža in Ul-
cinj, C. martima was fairly frequently recorded along 
almost the whole gradient of sand dune vegetation, al-
though never in great abundance. It starts to appear ei-
ther in the zone of annual vegetation of drift lines (EU 
habitat 1210) or in the transitional zone among annual 
vegetation of drift line sand embryonic shifting dunes 
(EU habitat 2110), which is characterised by intensive 
impact of wind and salt spray and fairly low vegetation 
cover. It also penetrates deeper into the vegetation of 
shifting dunes with Ammophila arenaria (white dunes, 
EU habitat 2120), dunes with Euphorbia terracina (EU 
habitat 2220), fixed coastal dunes with herbaceous veg-
etation (grey dunes, EU habitat 2130*) and humid dune 
slacks (EU habitat 2190). 
Discussion
The first record of C. maritima in Montenegro dates 
back to 2005 (N. Jogan unpublished) and it was later 
published in Glasović et al. (2009) since it had not been 
mentioned in previous (Trinajstić 1989, Mijović 1994, 
Mijović et al. 2006) and recent (Mijović et al. 2012) veg-
etation studies of the dune vegetation of Velika plaža. 
In neighbouring Croatia, the species was also discovered 
fairly late, in 1990 in Crnika Bay on the island of Rab, 
so up to 2005 it was considered to be the only site on 
the eastern Adriatic coast from the Gulf of Trieste in the 
north to Corfu in the south. Its late record was prob-
ably explained by misidentification and confusion with 
robust individuals of Scleropoa rigida (Trinajstić 1997), 
although a well conducted study of psammophytic veg-
etation of Rab island from the 1930s (Horvat 1939) ex-
plicitly rejected the presence of the mentioned species, 
despite the fact that it could be expected in that type of 
vegetation. In the case of Montenegro, it is important 
to note that the species has not been found in Albanian 
coastal dunes, which are a natural prolongation of Ve-
lika plaža (Fanelli et al. 2015). It is thus possible that the 
species has colonized Velika plaža recently, as a result of 
touristic impact. On the other hand, scattered localities 
may be the result of quite old introduction, or even natu-
ral occurrence, which cannot be completely excluded, 
especially since the whole lowland Montenegro has been 
much less explored than the mountainous inland. 
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D. Stešević, N. Bubanja, D. Caković, N. Jogan, M. Luković & U. Šilc
Synecology of Cutandia maritima (L.) Barbey, a rare psammophytic species along 
the Montenegrin Coast (East Adriatic Coast)
16/2 • 2017, 181–187
184
C. maritima is known to be capable of tolerating ex-
posure to intense sea winds and continual movement of 
sand grains of embryonic dunes facing seawards (Minis-
sale & Sciandrello 2015). This explains its high frequency 
within the habitat types: annual vegetation of drift lines 
(Cakilo-Xanthietum italici (Beg. 1941) Pign. 1953) and 
embryonic dunes (Echinophoro spinosae-Elymetum farcti 
Géhu 1988) (Figure 1). It can thus be considered to be 
one the first pioneer species or colonizer of dunes.
Although the species was recorded in only 4 transects 
located in the western part of the beach, it is sparsely 
distributed over almost the entire beach. It should be 
noted that the 2 m wide transects were set systematically 
at a distance of 500 m, so some occurrences may have 
been overlooked. Furthermore, the cover of C. martitima 
never exceeded the value 1.
As shown in Table 1, C. maritima frequently occurs 
in the studied area in annual grasslands with Vulpia fac-
iculata, Lagurus ovatus and Onobrychis caput-galli. This 
vegetation type has not been studied in detail on Velika 
plaža in Ulcinj. To date, only plant communities of Cak-
iletea and Ammophiletea have been reported for the dune 
area. C. maritima vegetation with the mentioned spe-
cies can certainly not be classified into Ammophilletea. 
In the researched transects and numerical classification, 
we could not separate communities of Malcolmietalia 
 






	
		
	



	


Figure 1: Detrended Correspondence Analysis (DCA) ordination of relevés from 20 transects classified into 9 plant communities (clusters). Size of 
red circles indicates cover value of Cutandia maritima.
Slika 1: DCA ordinacija popisov iz 20 transektov klasificiranih v 9 rastlinkih združb (klastrov). Velikost rdečih krogov nakazuje pokrovnost vrste 
Cutandia maritima.
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16/2 • 2017, 181–187
185
D. Stešević, N. Bubanja, D. Caković, N. Jogan, M. Luković & U. Šilc
Synecology of Cutandia maritima (L.) Barbey, a rare psammophytic species along 
the Montenegrin Coast (East Adriatic Coast)
and Crucinelletalia orders. More detailed investigations 
are needed to complete knowledge about the synecology 
of this species, as well as sabulicolous therophytic plant 
communities on Velika plaža in Ulcinj.
Conclusion
In previous studies of the sand dune vegetation of Velika 
plaža in Ulcinj, only communities of Cakiletea maritimae 
and Ammophiletea have so far been recognized, while 
sabulicolous therophytic communities have certainly not 
been taken into account. After the record of C. maritima 
on Velika plaža in Ulcinj, and some ecological and synec-
ological notes on the species, communities of Cutandieta-
lia maritimi might obviously also be present in Montene-
gro. However, more detailed phytocoenological research 
is needed.
Acknowledgements
The research was partly financed through the Rufford 
project 14048-1 and research core funding P1-0236 (Slo-
venian Research Agency).
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16/2 • 2017, 181–187
187
D. Stešević, N. Bubanja, D. Caković, N. Jogan, M. Luković & U. Šilc
Synecology of Cutandia maritima (L.) Barbey, a rare psammophytic species along 
the Montenegrin Coast (East Adriatic Coast)
Table number 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
Cover herb layer (%) 1 1 2 6 12 35 45 1 3 25 25 35 35 45 50 40 15 60 65 85 95 80 75 70 75 75 70 70 70 70 60 50 40
Distance from the sea  
(m) 10
0
10
2
11
0
10
4
10
6
10
8
11
0
10
4
10
6
13
4
13
6
13
8
10
8
11
0
11
2
11
4
82 11
2
11
4
11
6
11
8
12
0
12
2
12
4
12
6
12
8
13
0
13
2
13
4
13
6
13
8
14
6
14
8
Habitat type 1210 2110 2130*
Cutandia maritima r r r + 1 1 1 + + + + + + + + 1 1 1 1 1 1 1 + 1 1 1 + + + + + + +
Cakiletea maritimae
Xanthium italicum . . . + 1 1 . + . 2 1 1 . . . . 1 . + . . . . . . . . . . . . . .
Salsola kali . . r . . . . . r . . . . . r . . . . . . . . . . . . . . . . . .
Ammophiletea
Cyperus capitatus . . . . . . . . . + 1 1 . + 1 + 2 2 2 3 3 2 2 2 1 1 2 2 2 1 1 . .
Echinophora spinosa . . . + 2 2 2 . . + + . r r + . + 1 1 + + 1 1 + 2 1 1 . . . . . .
Euphorbia terracina . . . . . 1 1 . . . . . . . . . . . + 2 2 2 2 2 2 2 1 2 2 1 2 2 3
Alkanna tinctoria . . . . . . . . . . . + . . . . . . . 1 + . 1 + + + + 1 + 2 2 . +
Medicago marina . . . . . . . + 1 . . . . + + + . . . . . 1 1 . 2 + . . 1 1 1 . .
Pancratium maritimum . . . . . . + . + r . . . . . . . . . . . . 1 1 2 + + . . r + . .
Eryngium maritimum . . . . . 1 . . . . . . . . . . . + . . . . 1 + . . r 1 + . + r .
Elymus farctus . . . 1 . 2 2 . . . . . . . . . . + 2 2 1 . 1 . . . . . . . . . .
Scolymus hispanicus . . . . . . . . . . . . . . . . . r . . . r + + + . + + . . . . .
Lagurus ovatus . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . 2 2 2 1 1
Pseudorlaya pumila . . . . . . . . . . . . . . . . 1 . . . . . . . . . . . . . . . .
Sedo-Scleranthetea
Hypochaeris radicata . . . . 1 + . . . . . . . . . . . 1 + + + 1 1 1 1 2 2 2 2 1 1 1 1
Arenaria serpyllifolia agg. . . . . . . . . . . . . . . . . . . . . . 2 2 2 2 2 2 2 2 1 1 1 1
Tunica saxifraga . . . . . . . . . . . . . . . . . . . . . . . + . 1 1 1 1 1 1 + 1
Trifolium campestre . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . .
Helianthemetea guttati
Vulpia fasciculata . r . . 1 . 2 1 1 1 1 2 1 1 1 + . 2 2 2 2 2 2 3 2 2 2 2 2 2 2 2 2
Catapodium rigidum . r . . . . . + + . . + + + + 1 . . . . . . . . . . . . . + 1 + 1
Artemisietea vulgaris
Bromus diandrus . . . . . . + . . 1 1 + . . . . . 1 . 1 1 + + . 1 . . . + 1 1 1 1
Verbascum sinuatum . . . . . . . . . . + r . . . . . . . . . . . . . . . . . + + . .
Dasypyrum villosum . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . .
Juncetea maritimi
Imperata cylindrica . . . . . . . . . 2 2 2 . . . . . . . . . . . . . . 1 1 1 . . . .
Holoschoenus romanus . . . . . . . . . . . . . . . . . . . . . . . . . 3 2 . . . . . .
Parapholis incurva + . . . . . . r + . . . . . . . . . . . . . . . . . . . . . . . .
Others
Medicago littoralis . + 1 1 1 1 1 + 1 1 . . 3 3 3 3 1 1 1 1 1 . . . . . . 1 1 1 + 1 1
Onobrychis caput-galli . . . . . 1 2 r . . . 1 . . + + . 2 2 2 2 2 2 2 3 1 2 2 2 2 2 2 2
Crepis foetida . . . 1 1 1 . . . 1 1 1 . r . r 1 1 1 1 1 . . 1 1 1 1 + . 1 + . 1
Cerastium sp. . . . . . . . . . . . . . . . . . . . + . . 1 1 1 1 + . + . . . .
Chondrilla juncea . . . . . . . . . 1 1 1 . . . . . r . . . . . . . r r . . . . . .
Cynodon dactylon . . . . . . 1 . . . . . . + 1 . . . . . . . . . . . . . . . . 2 1
Conyza albida . . . . . . . + . . . . . . . . . . . . . . . . . . . . r . . . .
Phleum arenarium . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . .
Oenothera spp. . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . .
Pinus sp. . . . . . . . . . . . r . . . . . . . . . . . . . . . . . . . . .
Rubus ulmifolius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . .
Table 1: Cutandia maritima in the vegetation of Velika plaža (Montenegro).
Tabela 1: Vrsta Cutandia maritima v rastlinskih združbah na Veliki plaži (Črna Gora).
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