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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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
182
Introduction
Although Cutandia maritima (L.) Barbey is considered
to be a common species on sandy dunes along the
Mediterranean coast (van der Maarel & van der MaarelVersluys
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 characteristic
species of sabulicolous therophytic plant communities,
which Rivas-Martínez et al. (2002) described as the
Cutandietalia maritimae order within the class Tuberarietaria
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 Ammophiletea,
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 maritimae
(Rivas-Martinez et al. 2002), Cutandio maritimaeParapholidetum
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 maritimae
(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 Malcolmietalia.
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 Mediterranean,
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 continentality,
whereas those of Cutandietalia are indicators
of less expressed climatic conditions. Good correspondence
is also shown along the altitudinal gradient and distance
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. maritima
occurs rarely in Cakiletea plant communities (frequency
class lower than II), while it is more frequent (frequency
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 hemicryptophytic
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 maritimae
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 considered
to be the longest beach on the eastern Adriatic
coast (ca. 12km), with still well preserved sand dune vegetation.
Since 1968, the beach has been recognized as a
monument of nature (Official Gazette of the Social Republic
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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183
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 studied.
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 system
up to forest vegetation. In each quadrat, the cover
of plant species was visually estimated by the Braun-Blanquet
(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 (McCune&
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, presented
in a single table (Table 1) and classified into different
clusters.
Detrended Correspondence Analysis (DCA) ordination
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 Velika
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-Xanthietum
italici, Cynodon dactylon comm., Euphorbio paraliaeAgropyretum
junceiformis, Medicagini marinae-Ammophiletum
australis, Onobrychis caput-galli comm., Cladietum
marisci, Eriantho-Schoenetum nigricantis, Juncetum maritimo-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), distributed
in a bandwidth from ca. 80 m from the sea to
150 m inland. Its cover value is lower (r, + and 1) in mobile
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 Ulcinj,
C. martima was fairly frequently recorded along
almost the whole gradient of sand dune vegetation, although
never in great abundance. It starts to appear either
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 vegetation
(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) vegetation
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 probably
explained by misidentification and confusion with
robust individuals of Scleropoa rigida (Trinajstić 1997),
although a well conducted study of psammophytic vegetation
of Rab island from the 1930s (Horvat 1939) explicitly
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 Velika
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 natural
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 exposure
to intense sea winds and continual movement of
sand grains of embryonic dunes facing seawards (Minissale
& 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 faciculata,
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 Cakiletea
and Ammophiletea have been reported for the dune
area. C. maritima vegetation with the mentioned species
can certainly not be classified into Ammophilletea.
In the researched transects and numerical classification,
we could not separate communities of Malcolmietalia
-2 14
-410
Cakilo-Xanthietum italici
Juncetum maritimo-acuti
Medicagini marinae-Ammophiletum australis
Euphorbio paraliae-Agropyretum junceiformis
Cynodon dactylon comm.
Holoschoenetum romani
Onobrychis caput-galli comm.
Eriantho-Schoenetum nigricantis
Cladietum marisci
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 synecological
notes on the species, communities of Cutandietalia
maritimi might obviously also be present in Montenegro.
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 (Slovenian
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)
100
102
110
104
106
108
110
104
106
134
136
138
108
110
112
114
82
112
114
116
118
120
122
124
126
128
130
132
134
136
138
146
148
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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