129
Abstract
The vegetation of Buna River Protected Landscape is described. The area comprises both the alluvial plain of
the lower course of the Buna river that marks the boundary between Montenegro and Albania, and a carbonatic
range. The vegetation is characterized by a high β-diversity (27 alliances and 46 associations), especially
in wetlands and dry grasslands. In the area it is possible to distinguish 1 dunal and 4 hygro-sequences in the
alluvial plain and 3 xeroseries in the carbonatic range. Associations Clematido viticellae-Punicetum granati and
Periploco graecae-Alnetum glutinosae are described as new.
Key words: aquatic vegetation, Albania, Buna River, alluvial forests, dry grasslands, marshlands, shrublands,
xerothermophilous forests.
Izvleček
V članku je opisana vegetacija v zavarovanega območja ob reki Buna. Območje obsega aluvialno ravnino
ob spodnjem teku reke Buna, ki predstavlja mejo med Črno goro in Albanijo in karbonatnim hribovjem. Za
vegetacijo je značilna visoka β diverziteta (27 zvez in 46 asociacij), še posebej pa mokrišča in suha travišča. V
območju smo izločili 1 sekvenco na sipinah, 4 higro sekvence na aluvialni ravnici in 3 ksero serije na karbonatnem
masivu. Asociaciji Clematido viticellae-Punicetum granati in Periploco graecae-Alnetum glutinosae sta novo
opisani.
Ključne besede: vodna vegetacija, Albanija, reka Buna, poplavni gozdovi, suha travišča, močvirja, grmišča,
kserotermofilni gozdovi.
THE VEGETATION OF THE BUNA RIVER
PROTECTED LANDSCAPE (ALBANIA)
Giuliano FANELLI1, Michele DE SANCTIS2,*, Ermelinda GJETA3,
Alfred MULLAJ4 & Fabio ATTORRE2
1	Department of Biology, Tor Vergata University of Rome, 00133 Roma, Italy.
2	Department of Environmental Biology, Sapienza University of Rome, 00185 Roma, Italy.
3	Department of biology, University “Aleksander Xhuvani", Elbasan, Albania.
4	Flora and Fauna Research Center, Faculty of Nature Science, University of Tirana, Tirana, Albania.
*	Corresponding author, e-mail: michedes@gmail.com
DOI: 10.1515/hacq-2015-0008
HACQUETIA 14/2 • 2015, 129–174
1. INTRODUCTION
The Buna River Protected Landscape extends for
about 20000 ha and is located between the last 15
km of the northern Albanian coast and the Buna
river, which here forms the border with Montenegro
(Figure 1). The Buna river is the outflow of
the Skadar Lake and has the Drin River, the longest
river in Albania, as main tributary. The strong
river flow created one of the most important wetland
systems in Albania providing important
habitats for many animal and plant species. The
sediments of Buna have an important role for the
morphology of the coastline, which is subject to
strong erosion. According to old descriptions
(Reiser & Fuhrer 1896, Kárpáti & Kárpáti 1961,
Kárpáti 1962), the Buna Landscape was an impressive
wilderness area. However, likewise other
Mediterranean wetland areas it has been strongly
transformed during the last decades. Between
1947 and 1980 about 36 km2
of agricultural lands
were reclaimed, extending the only 2 km2
agricultural
land that existed before. In the early ‘70s
a system of hydrovors was installed functional to
the new drainage-irrigation canal system. In the
‘80s three dams and a huge hydropower reservoir
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Hacquetia 14/2 • 2015, 129–174
130
were built along the Drin River. These works have
deeply changed the hydrogeological system and
the land-use of the area: the Buna flooding and
the amount of water greatly decreased and much
of the inner land was converted into agricultural
use. However, as in some other Albanian border
areas, the area was under military protection untill
1991, and due to this status, was effectively
protected. In 1991, the status of the area changed,
and a rapid development of touristic infrastructures
began especially along the coast.
Despite these changes the Buna river area
still hosts a great diversity of important habitats
and species deserving careful conservation policies.
For this purpose the Albanian government
in 2005 decided to establish the Buna River Protected
Landscape for which an integrated and
systemic management plan is in preparation
(Guillet et al. 2012). The analysis of the plant
communities represents important information
for the elaboration of such plan. However, existing
literature about vegetation is rather sparse
and heterogeneous with few phytosociological
studies (Kárpáti & Kárpáti 1961, Kárpáti 1962,
Mullaj 1986, Mullaj et al. 2000). The study of the
vegetation of Albania begun, apart from a few
pioneering works (Markgraaf 1932), in the late
‘80s (e.g. Buzo 1990). However, only recently Albanian
authors adopted the Zürich-Montpellier
method (e.g. Kalajnxhiu et al. 2011) and a first
synthesis based on this approach can be found in
Dring et al. (2002). Nonetheless the phytosociology
of Albania is still imperfectly known and this
study represents a significant contribution to the
knowledge of the vegetation of this country.
2. METHODS
Study area
The Buna River Protected Landscape includes
the area along the Buna river, after the confluence
with the Drin near the Skadar Lake, its delta
on the Adriatic Sea and the area around the Viluni
lagoon (Figure 1).
The protected area includes three main land
unit systems:
1)	 an alluvial plain composed of holocenic loams
deposited by the Buna river (Frasheri et al.
2006) with marshlands, alluvial and riverine
forests, and lagoons;
2)	 a carbonatic outcrop with arid Mediterranean
vegetation. The carbonatic range extends in
SE–NW direction and is composed of upper
Cretaceous-Paleocenic limestones and dolomites;
at the base there are layers of upper Oligocene
flysch with intercalations of clays. The
range reaches 500 m of altitude in the Black
Peak (Maja e Zezë);
3)	 a coastal holocenic dune system, composed of
sands deposited by the Buna river.
The climate is Mediterranean, characterized
by dry-hot summers and mild-wet winters. The
wind of murrlan, which is very characteristic of
the area, makes the winters harsh; whereas in
the summer the wind of Shiroku brings humidity.
Mean annual rainfall and temperature are
1075 mm and 15,3 °C respectively. Precipitation
is concentrated in the period from November to
April (70–80% of total annual). In January mean
temperature is between 5 and 8 °C while in July it
can range between 24 and 26 °C (PHARE 2002).
Figure 1: Location of the Buna River Protected Landscape.
1: Buna River; 2: Domni marsh; 3: Black Peak (Maja e Zezë);
4: Viluni lagoon; 5: Rjolla Beach (Backs Rrjolli); 6: Velipojë
Reserve; a: Black Peak; b: Mount Nikgionih (Mali i Nikgjonit);
c: Campo di Marina (Kampi veror).
Slika 1: Lokacija zavarovanega območja ob reki Buna. 1: reka
Buna; 2: močvirje Domni; 3: Črni vrh (Maja e Zezë); 4: laguna
Viluni; 5: plaža Rjolla Beach (Backs Rrjolli); 6: rezervat
Velipojë; a: Črni vrh; b: gora Nikgionih (Mali i Nikgjonit);
c: Campo di Marina (Kampi veror).
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Giuliano Fanelli et al.: The vegetation of the Buna river protected landscape (Albania)
131
Data collection and analysis
Phytosociological investigation was performed
according to Braun-Blanquet (1964) and Westhoff
& van der Maarel (1978). Data on vegetation
were obtained through 188 relevés (for a total of
389 vascular plant taxa) carried out in June–July
and September 2012–2013. At each site the relevés
were selected in relation to the homogeneity
of physical features and vegetation structure.
Plot sizes ranged from 150 m2
to 1 m2
(Table 4)
depending on the plant community. Cover/abundance
data for all vascular plants were recorded
in the field using a modified Braun-Blanquet
scale (Barkman et al. 1964).
Vascular plants were identified using the ‘‘Flora
e Shqipërisë’’ (Paparisto et al. 1988, Qosja et
al. 1992, 1996, Vangjeli et al. 2000), “Flora ekskursioniste
e Shqipërisë” (Demiri 1983) and “Flora
Europaea” (Tutin et al. 1964–80, 1993), taxon nomenclature
follows the latter, except for Asphodelus
(Diaz-Lifante & Valdés 1996), Bupleurum (Snogerup
& Snogerup 2001), Salicornia (Kadereit et
al. 2007, Kadereit et al. 2012) and Bolboschoenus
(Hroudová et al. 2007).
A database was created in TURBOVEG (Hennekens
& Schaminée 2001) and data were then
exported as percentages for further analyses.
The names of identified syntaxa have been
choosed according the nomenclatural rules codified
in the Third Edition of the International
Code of Phytosociological Nomenclature (Weber
et al. 2000) and the syntaxonomic scheme has
been formulated according the current knowledge
about Albanian vegetation and that of
neighboring regions.
The data collected were used to create a plotsvs-species
matrix. Plant communities of marhslands,
acquatic vegetation and sand dunes which
are characterized by mono-or paucispecific composition
were identified only according to expert
knowledge and scientific literature. Conversely
two classification procedures were perfomed on
ecologically homogeneous aspects such as the
alluvial forests and the vegetation of the carbonatic
outcrop. Using the program Syn-Tax v.
5.0 (Podani 2001) a hierarchical agglomerative
clustering was used, with Euclidean distance as
the similarity coefficient and complete linkage
as the method for grouping the formation. Data
were log transformed (base 10) in order to reduce
skewness and kurtosis (e.g. Johnson & Wichern
2007). Since the syntaxonomy of Albanian vegetation
is still to be completely defined, in order
to support the phytosociological interpretation
of the plant communities of the Buna Landscape
we identified diagnostic species also using the fidelity
coefficient of Tichý & Chytrý (2006) with
a simultaneous calculation of Fisher’s exact test
in the JUICE program with P < 0.001. To avoid
phi coefficient being dependent on the size of the
target site group, group size was standardized to
equal the average size of all groups present in the
entire data set (Tichý & Chytrý 2006).
3. RESULTS AND DISCUSSION
Through the analysis of the relevés 46 plant communities
have been identified and classified in 27
alliances, 23 orders and 17 classes. They are distributed
in 1 dunal and 4 hygro-sequences in the
alluvial plain and 3 xeroseries in the carbonatic
range. Below these communities and the vegetation
sequences are described and discussed in the
same order of the syntaxonomical scheme.
Brackish and halophilous vegetation
Ruppietum cirrhosae Iversen 1934 (Table 1)
= ass. Ruppia maritima Hocquette 1927, Ruppietum
spiralis Hocquette 1927 corr. Iversen 1934
In the Viluni lagoon the vegetation of the
shallow waters is represented mainly by dense
populations of Ruppia cirrhosa together with undetermined
filamentous algae. The association is
abundant close to the mouth of the Lagoon where
there is an inflow of salt water, and rarer on the
northern coast, where waters are less brackish.
The community is widespread in European lagoons
(e.g. Schaminée et al. 1996, Rivas-Martinez
et al. 2001) and has been reported for Albania
by Mullaj (1989) and Ruci et al. (1995).
Salicornietum venetae Pignatti 1966 (Table 2)
= Salicornietum emerici O. de Bolòs ex Brullo &
Furnari 1976
On the shores of the Viluni lagoon, in particular
near the mouth that connects the lagoon to
the sea, there are mud-flats with extensive, almost
monospecific populations of Salicornia. Closer to
the water, where the inundation is prolonged,
the dominant species is the tetraploid Salicornia
procumbens (= S. emerici, S. dolichostachya, S. veneta,
S. stricta). Beyond this belt, where inundation
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Hacquetia 14/2 • 2015, 129–174
132
is shorter, the dominant species is the diploid
Salicornia perennans (= S. patula, S. prostrata). The
two vegetation belts are particularly evident in
October-November, when Salicornia procumbens
becomes brilliant red, whereas Salicornia perennans
remains dirty green.
The vegetation of Salicornia procumbens is
represented by dense populations 20–30 cm tall.
They usually are monospecific (rel. 100), but it is
often possible to observe a transitional belt with
Limonium vulgare subsp. serotinum.
The taxonomy of annual Salicornia is notoriously
difficult, and this complicates the syntaxonomy
of the annual halophilous vegetation,
based largely on the distinction between the
many microspecies (Loidi et al. 1999). Recently,
molecular phylogenetics has greatly clarified the
taxonomical problems of Salicornia (Kadereit et
al. 2007, 2012). Although a few problems remain,
the classification proposed by these authors is
much simpler than earlier attempts (e.g. Ball &
Akeroy 1993, Iberite 1996), recognizing only
one tetraploid taxon, Salicornia procumbens, and
two diploid taxa in Europe, Salicornia europaea
in the Atlantic coasts and Salicornia perennans
in the Mediterranean basin and in East Europe.
The many tetraploid species recognized so far
(Salicornia veneta, Salicornia dolichostachya) are
only growth forms of a single species and, in fact,
they are morphologically and ecologically similar
(Teege et al. 2011). This fact has important consequences
for the syntaxonomy, since it means that
many associations are probably synonyms.
In study area, Salicornietum venetae, considered
endemic of the Venice Lagoon (Géhu et al.
1984b, Géhu & Biondi 1996), is in fact a synonym
of Salicornietum emerici, widespread in the Mediterranean
basin (Bolòs 1962, 1967, Brullo et al.
1988, Lloidi et al. 1999, Frondoni & Iberite 2002).
Salicornia veneta is a luxuriant growth-form of
Salicornia procumbens, but the poor floristic composition
is similar in the two associations. In the
future it will be advisable to mutate the name according
to modern taxonomy, but for the time
being we refrain from proposing a nomen mutatum,
first because Art. 45 dictates that a nomen
mutatum should be proposed only if the name
of taxon on which the association is based is no
longer in use for at least 20 years (Weber et al.
2000), second because adopting the name Salicornia
procumbens for all tetraploid species will
cause Salicornietum venetae to fall in synonymy
with Salicornietum dolichostachyae Géhu & GéhuFranck
1984, widespread on the Atlantic coasts in
intertidal mud flats and therefore with a different
ecology (Ellenberg 1996).
The Association Limonium vulgare + Salicornia
europaea (Buzo 1990), described from Poro, Seman
and Divjaket (Southern Albania) in retrodunal
depressions, is with all probability identical
to our community even if it is not possible to
identify with certainty which annual Salicornia is
reported in the table.
Suaedo spicatae-Salicornietum patulae (Brullo
& Furnari 1976) Géhu & Géhu-Franck 1984.
corr. Alcaraz, Ríos, De la Torre, Delgado &
Inocencio 1998
= Salicornietum patulae Brullo & Furnari 1976 non
Christiansen 1955, Suaedo vulgaris-Salicornietum
patulae (Brullo & Furnari) Géhu & Géhu-Franck
1984; Suaedo maritimae-Salicornietum patulae Brullo
& Furnari ex Géhu & Géhu-Franck 1984 corr.
Rivas-Martínez 1990
Salicornia perennans vegetation is not widespread
in the Velipoje area. It occurs only in the
Viluni lagoon in sites with shorter inundation
than those of Salicornietum venetae and in Campo
di Marina (Kampi veror) lagoon. We did not
carry out relevés of these monospecific stands,
but they can be referred to rather fragmentary aspects
of Suaedo spicatae-Salicornietum patulae.
Puccinellio festuciformi-Aeluropetum litoralis
(Corb 1968) Géhu & Costa 1984 (Table 3)
This community refers to a low scrub vegetation
with grasses and succulent chenopods that
develops extensively on saline muds surrounding
the Viluni lagoon. These perennials occupy a belt
back of the annual Salicornia stands near the water.
The community is absent in Campo di Marina
(Kampi veror), where the zonation of the shore
vegetation is otherwise similar to that of the Viluni
lagoon. Floristically this community is characterized
by the co-occurrence of Aeluropus littoralis,
Halimione portulacoides and Suaeda maritima.
Among the many associations that have been
described in the Mediterranean basin, often on
the base of subtle differences (Brullo et al. 1988,
Rivas-Martinez et al. 2001) and in the Danube
delta and Black Sea (summarized in Tzonev
et al. 2008), our relevés are more similar to the
Puccinellio-Aeluropetum litoralis, although Puccinellia
festuciformis is lacking, and particularly to
the relevés 1–5 in Table 21 in Géhu et al. (1984a)
from Ferrara (Italy).
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Giuliano Fanelli et al.: The vegetation of the Buna river protected landscape (Albania)
133
Juncetum maritimo-acuti Horvatić 1934
­(Table 4)
This association grows on muds on the shores
of the Viluni lagoon and on smaller brackish ponds
not far from the sea. It is characterized by the
dominance of Juncus acutus and Juncus maritimus,
among which succulents grow such as Halimione
portulacoides, Artemisia coerulescens and Limonium
vulgare subsp. serotinum. The floristic composition
matches in some way the Limonio-Artemisietum coerulescentis
Horvatić (1933) 1934 corr. Géhu & Biondi
1996, the only distinction being the presence of
an upper layer dominated by Juncus acutus.
Juncus acutus and Juncus maritimus communities
are widespread in the Mediterranean basin
and along the Southern Atlantic coasts (Molinier
& Tallon 1965), with a rather constant composition.
We can refer without any doubt our association
to Juncetum maritimo-acuti, that has been described
from Dalmatia (Horvatić 1934, 1963), but
also reported with similar aspects in Italy both
along the Adriatic coast (Pignatti 1966 sub Juncetum
maritimi Pignatti 1953, Brullo et al. 1988, Poldini
1989, Pirone 1995a sub Juncetum acuti Molinier
& Tallon 1970) and in the central Tyrrhenian
coast (Frondoni & Iberite 2002). In Albania it has
been already reported for Lalzi Bay near Durres
(Imeri et al. 2010a).
Juncus maritimus and Lippia nodiflora community
(Table 5)
This community develops on the shores south
of the Viluni lagoon, in retrodunal depressions
characterized by moderately brackish waters.
This community is dominated by Juncus acutus
and Juncus maritimus, but the coverage is lower
than that of Juncetum maritimo-acuti and the floristic
composition characterized by species such
as Lippia nodiflora and Inula viscosa. The retrodunal
depressions where this community occurs
have been partly dismantled by erosion, so that
there is an ingression of elements from sanddunes
(Euphorbia paralias, Elymus farctus). Other
halophytes are Crypsis aculeata, Sonchus maritimus,
Artemisia coerulescens and Plantago crassifolia,
suggesting a synclassification in Plantaginion
crassifoliae. This community shows many similarities
with Juncetum acuti Molinier & Tallon 1970
from Camargue in France, (Molinier & Tallon
1970), but there Lippia nodiflora is lacking.
The community is strongly disturbed by erosion
and grazing by cattle, and therefore we refrain
from a formal description.
Psammophilous vegetation
Cakilo-Xanthietum italicae Pignatti 1953
­(Table 6)
= Salsolo kali-Cakiletum maritimae xanthietosum
(Pignatti 1953) Géhu & Scoppola 1984 in Géhu
et al. 1984
This community develops on the foredunes
near the shore in correspondence with the mouth
of the Buna River. Structurally, it is a sparse
grassland, 20–30 cm tall, dominated by succulent
species. The floristic composition is typical,
with Cakile maritima, Salsola kali, Polygonum maritimum,
Xanthium orientale subsp. italicum, and
a few species from Ammophiletalia such as Sporobolus
pungens and Echinophora spinosa.
This association was described for the Venice
Lagoon (Pignatti 1953) and is distinct from
other Cakiletea associations mainly by the presence
of Xanthium orientale subsp. italicum. CakiloXanthietum
is present on the Adriatic and Ionian
coasts of Italy (e.g. Biondi et al. 1989, Corbetta
et al. 1989 sub Salsolo-Cakiletum aegyptiacae Costa
et Manzanet 1981 subass xanthietosum Géhu et
Scoppola 1984) and Greece (Sykora et al. 2003);
it seems to be lacking in France and on the Tyrrhenian
coasts of Italy (Braun-Blanquet et al.
1952). The association is reported from Albania
by Imeri et al. (2010a) with the incorrect name
of Cakilo-Xanthietum strumarii (Beg. 1941) Pign.
1958 and under the name Salsola kali-Xanthium
strumarium association by Buzo (1990).
In our relevés two subspecies of Salsola kali
occur together in equal proportions, Salsola kali
subsp. kali and Salsola kali subsp. tragus. Subspecies
of S. kali are rarely reported in the phytosociological
literature, being rather difficult to distinguish,
but they are probably ecologically quite
distinct.
Freshwater aquatic vegetation
Hydrocharitetum morsus-ranae van Langendonck
1935 (Table 8)
This community occurs in several canals and
ditches near Velipoje with clear, slowly flowing
water. It is characterized by the dominance of
Hydrocharis morsus-ranae, a species classified as
vulnerable in Albania (Vangjeli et al. 1995).
The association is widespread in Europe, in
mesotrophic slowly flowing waters (Šumberová
2011a).
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Hacquetia 14/2 • 2015, 129–174
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Nymphaeetum albae Vollmar 1947 (Table 9)
The community mainly occurs along the drainage
canals in very slow-flowing waters about 1
meter deep. It is dominated by Nymphaea alba and
a few other species. Nymphaeetum albae is characteristic
of mesotrophic waters and has a strong
capacity of filling of the water-bodies due to the
high biomass production (Šumberová 2011b).
This community was probably much more
widespread in the past. The description in Kárpáti
(1962) seems to indicate that Nymphaeion communities
were very frequent on the floodplain of
the Buna. The terrestrialization of the floodplain
may be related to natural phenomena of succession,
but also to the lowering of the water table
due to the artificial drainage.
Trapetum natantis Kárpáti 1963 (Table 10)
This community occurs in a canal of the Domni
marsh. It is characterized by the dominance of
Trapa natans, which covers completely the water,
and is accompanied by a few other species. This
association has already been reported for the Buna
river near the Shkodra Lake and is very rare in
Albania (Schneider-Jacoby et al. 2006).
Potamogetonetum denso-nodosi O. de Bolòs
1957 (Table 11)
= Potametum nodosi (Soó 1960) Segal 1964
This community develops in the Buna River,
close to the shore where the water is 60 cm deep.
From a floristic point of view, as a community
of submerged hydrophytes, it is rather rich with
many Potamogeton species dominated by Potamogeton
nodosus; this richness is probably due to
low turbidity of the waters of the Buna.
The community is typical of rivers and other
habitats with flowing waters, and is generally
widespread but rarer in southern Europe (Bolòs
1962, Šumberová 2011b).
Potamogetonetum pectinati Carstensen ex
­Hilbig 1971 (Table 12)
This community is widespread in the Buna
river. It occurs in relatively deep water (1–3 m),
where it forms extensive populations, but also
close to the shore where the slope of the river
bank is steep. The floristic composition is dominated
by Potamogeton pectinatus, with low frequency
of other Potamogeton (P. crispus, P. nodosus,
P. perfoliatus).
This association has a rather broad ecological
range, but usually occurs in mesotrophic to
eutrophic waters (Testi et al. 2009, Šumberová &
Chytrý 2011).
Elodeetum canadensis Nedelcu 1967 (Table 13)
Elodea canadensis community can be found
along the Buna river, where this species forms
a dense population. The community substitutes
the Potametum denso-nodosi in disturbed areas. In
Albania the association has been also reported
for the Ohrid lake (Imeri et al. 2010b).
Swamp vegetation of fresh and
brackish waters dominated by
graminoids and sedges
Typhetum angustifoliae Pignatti 1953 (Table 14)
This community is one of the most striking
physiognomies of the study area. It occurs in
Domni marsh and in other sites of the alluvial
plain where populations of Typha angustifolia
form dense swards about 2 m tall. The community
is more or less monospecific, and develops
on soils inundated in winter, but that dry out in
summer, leaving a thin veil of water on the damp
ground. The vegetation is subjected to fires set
on by farmers to open the vegetation for grazing,
but this practice does not seem to negatively affect
Typha angustifolia but, on the contrary, to increase
its cover.
The wide diffusion of Typha angustifolia is
probably related to the draining of the marshland.
Typha angustifolia typically grows in deeper
still waters, particularly in lakes and closed ditches
(Pignatti 1953, Šumberová & Chytrý 2011).
Typhetum latifoliae Nowiński 1930 (Table 15)
= Typhetum latifoliae Soó 1927 nom. nudum, Typhetum
latifoliae Lang 1973
This vegetation is characterized by a dense
reed-bed, 150–160 cm tall. It grows on the banks
of the Buna river where the water depth is about
10–20 cm. The community develops in an intermediate
belt between the Phragmites australis and
Bolboschoenus maritimus communities.
The community is widespread in Europe, in
particular, in eutrophic waters in ditches and
along river banks (e.g. Chytrý et al. 2011).
Phragmitetum australis Savič 1926 nom. mut.
prop. (Table 16)
This vegetation occurs along the Buna river
in particular where the flow is slow, but also on
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Giuliano Fanelli et al.: The vegetation of the Buna river protected landscape (Albania)
135
the shores of the Viluni lagoon, where the water
is less brackish; in fact Phragmites australis is only
weakly halotolerant (Pignatti 1952, 1953). The
species needs almost constant inundation and
rather deep waters (50–100 cm).
This community is a dense reed-bed 1.7–3 m
tall, characterized by the dominance of Phragmites
australis, which is accompanied by a few
species (Bolboschoenus maritimus, Iris pseudacorus
and Rumex hydrolapathum). In many stretches,
the community is monospecific.
The community is very widespread (Šumberová
& Chytrý 2011) and in the Mediterranean
basin has been referred to a different association,
Typho angustifoliae-Phragmitetum australis (Tüxen
& Preising 1942) Rivas-Martínez, Báscones, T.E.
Díaz, Fernández-González & Loidi 1991 = ScirpoPhragmitetum
mediterraneum Tüxen & Preising
1942 (Braun-Blanquet et al. 1952, Rivas-Martinez
et al. 2001). However the floristic composition is
so poor that it is difficult to make distinctions,
although ecological differences certainly exist
between water bodies in temperate and Mediterranean
Europe.
Scirpetum lacustris Chouard 1924 (Table 17)
= Schoenoplectetum lacustris Chouard 1924 nom.
mutatum prop.
This community is restricted to the Buna river,
near the banks, in the first helophytic belt at a water
depth of 0.3–0.5 m, where it forms tall, dense
stands. The association is usually mesotrophic to
slightly eutrophic (Šumberová & Chytrý 2011).
The association is widespread, but usually
referred to a facies of Scirpo-Phragmitetum Koch
1926 and therefore its distribution is imperfectly
known.
Glycerio-Sparganietum neglecti Koch 1926
(Table 18)
This plant community is generally found in
the alluvial plain near Velipoje, in clear, slowly
flowing waters, 30–40 cm deep. The community
forms a dense reed-bed 120–130 cm tall. Sparganium
erectum is dominant, but other helophytes
are frequent. The association is floristically interesting,
in particular for the presence of two
species rare in Albania, Butomus umbellatus and
Hydrocharis morsus-ranae (Ruci et al. 2000).
This association is widespread in Europe (e.g.
Schaminée et al. 1996), where is often reported
under the pseudonyms Sparganietum ramosi Roll
1938 or Sparganietum erecti Zutshi 1975 (Dengler
et al. 2004, Šumberová & Chytrý 2011) and has
been reported for Albania in the Ohrid lake (Imeri
et al. 2010b).
Scirpetum maritimo-litoralis (Br.-Bl. in Br.‑Bl.,
Roussine & Nègre 1952) O. de Bolòs 1962
­(Table 19)
= Scirpetum maritimo-litoralis Br.-Bl. 1931 nomen
nudum, Scirpetum compacto-litoralis (Br.-Bl. in
Br.-Bl., Roussine & Nègre 1952) O. de Bolòs 1962
corr. Rivas-Martínez, Costa, Castroviejo & E.
Valdés 1980
This association is a relatively sparse reedbed,
100–170 cm tall, that grows in the brackish
water on the northern side of the Viluni lagoon
and at the mouth of the Buna river. It occupies an
intermediate belt between the Phragmites australis
and the Scirpus litoralis communities in water with
a depth between 15 and 20 cm.
The floristic composition is poor, and is characterized
by the dominance of Bolboschoenus maritimus,
often associated with Phragmites australis.
A number of associations with Bolboschoenus
maritimus have been described around the Mediterranean
basin and in continental Europe (e.g.
Rivas-Martinez et al. 2001). These are usually floristically
poor, and their interpretation is therefore
difficult. Probably different aspects, related
to different ecological conditions occur.
Our relevés are very similar to those from
Provence (Braun-Blanquet et al. 1952), Catalonia
(Bolòs 1962) and the coasts of Croatia (Horvatić
1963). They are instead quite different from those
of the association Scirpetum maritimi Tüxen 1937,
described for the Venice lagoon (Pignatti 1966),
that after Géhu & Biondi (1996) should be referred
to a different association.
This association is reported for Albania (Mullaj
et al. 2000, Imeri et al. 2010a) under the pseudonym
Bolboschoenetum maritimi Eggler 1933 = Bolboschoenetum
yagarae Eggler 1933 corr. Hroudova
et al. 2009, which instead corresponds to a different
freshwater vegetation (Hroudová et al. 2009).
The taxonomy of Bolboschoenus has been recently
reviewed (Hroudová et al. 2009) and Scirpus
compactus has fallen in synonymy with Bolboschoenus
maritimus. This makes superfluos the
correction of Rivas-Martínez et al. (2001) of Scirpetum
maritimo-litoralis (Br.-Bl. in Br.-Bl., Roussine
& Nègre 1952) O. de Bolòs 1962 into Scirpetum
compacto-litoralis (Br.-Bl. in Br.-Bl., Roussine
& Nègre 1952) O. de Bolòs 1962 corr. RivasMartínez,
Costa, Castroviejo & E. Valdés 1980.
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Hacquetia 14/2 • 2015, 129–174
136
Scirpetum litoralis Pignatti 1953 (Table 20)
This community has been found in the north
of the Viluni lagoon, where waters are less saline,
and forms a rather sparse reed-bed dominated by
Scirpus litoralis, in deeper waters than the backstanding
Scirpetum maritimo-litoralis.
The association is reported for the Venice
lagoon, Southern Italy and Tunisia, in slightly
brackish, almost still waters (Pignatti 1966).
Meadows and pastures of moister soils
Schoeno-Erianthetum ravennae Pignatti 1953
(Table 21)
This community grows in depressions north
of the Viluni lagoon, and in fragmentary aspects
in retrodunal depressions near the sea. It
is characterized by large tufts of Erianthus ravennae
that can be up to 1,7 m tall. A lower layer is
dominated by Schoenus nigricans, and these two
dominant species are accompanied by a number
of species of wetlands characterized by a long
drought period (Pulicaria vulgaris, Carex punctata
and Centaurium spicatum). Also a few subruderal
and ruderal subhygrophilous species are present
(Aster squamatus, Lotus tenuis).
The association was described for the Venice
lagoon, and is widespread along the Italian coasts
(Lucchese & Pignatti 1990, Géhu & Biondi 1996).
In Albania it has been reported for the Lalzi bay
(Imeri et al. 2010a).
Paspalo-Agrostidetum Br.-Bl. 1936 (1952)
­(Table 22)
This community has been recorded in the
Domni marshland, in an intermediate position
between Typhetum angustifoliae and Trifolio-Cynodontetum,
in correspondence with conditions almost
as wet as those favouring Typha angustifolia
but in heavily disturbed areas.
The community of Velipoje is poorer in species
than that of Provence where it has been originally
described (Braun-Blanquet et al. 1952), dominated
by Paspalum paspalodes accompanied with
ruderal species such as Plantago major and Rumex
conglomeratus. It is also present in the urban area
of Rome with even poorer, almost monospecific
aspects (Fanelli 2002).
Trifolio fragiferi-Cynodontetum Br.-Bl. & O. de
Bolòs 1957 (Table 23)
In the alluvial plain of the Domni marsh, the
dense reed beds of Typha angustifolia leave place
in some spots to a heavily grazed lawn. This lawn
is 2–4 cm tall and formed by dense mats of the
creeping rhizomatous Trifolium fragiferum and Cynodon
dactylon accompanied by Mentha pulegium
and Plantago lanceolata. The lawn develops where
the soil is relatively dry. This condition is less suitable
for Typha angustifolia, which becomes vulnerable
to disturbance and is replaced by the lawn
species. This community is highly productive and
favoured by cattle, since the nitrogen-fixing Trifolium
allows a high production of proteins.
This association has been described from
Provence, on the floodplain along large rivers
(Braun-Blanquet et al. 1952, Bolòs 1962, 1967), but
has been reported also from the Danube Plain in
Bulgaria (Tzonev 2009). The association is poorly
characterized and probably is more widespread
than the scanty reports in literature may suggest.
VEGETATION OF THE CARBONATIC
OUTCROP
The hierarchical classification identified twelve
groups (Figure 2). They include dry grasslands
and xerothermophilous forests and shrubs described
hereafter. One crevice and one ruderal
Figure 2: Dendrogram of the carbonatic range vegetation.
The capital letters indicate cluster membership.
Slika 2: Dendrogram vegetacije na karbonatnem hribovju.
Velike črke predstavljajo uvrstitev v klaster.
A: Rhamno-Paliuretum; B: Querco-Carpinetum ­orientalis;
C: Seslerio-Ostryetum carpinifoliae; D: Acer monspessulanum
community; E: Salvio-Phlomidetum fruticosae; F: Brachypodio-Cymbopogonetum
hirti; G: Clematido viticellae-Punicetum
granati; H: Quercetum trojanae; I: Seslerio-Putorietum calabricae;
L: Asphodelo-Chrysopogonetum grylli; Asphodelus ramosus
community; N: Stipo-Salvietum officinalis.
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Giuliano Fanelli et al.: The vegetation of the Buna river protected landscape (Albania)
137
community have been also identified and discussed
in their specific section.
Crevice vegetation of limestone
fissures
Seslerio-Putorietum calabricae Horvatić 1963
(Table 7; Figure 2, Cluster I)
This single relevé has been carried out on an
almost vertical slope of rather friable carbonates,
with southern aspect. Putoria calabrica was abundant
nearby, but on the less steep parts of the
slope. The vegetation was rather sparse, with low
cover, characterized by the presence of Sesleria robusta
subsp. skanderbeggii and Moltkia petraea.
This community can be referred to SeslerioPutorietum
calabricae (Horvatić 1963), a coastal
association described from Dubrovnik. Moltkia
petraea has been indicated from a number of associations
under different orders and alliances
(Lakušić 1968, Georgiou et al. 2000).
Perennial grasslands of the
Mediterranean region
Stipo-Salvietum officinalis Horvatić 1958
­(Table 24; Figure 2, Cluster N)
This community occurs on steep slopes (20–
30°) usually with a W–SW aspect. Soil is very
stony (20–80 % rocks and stones) and shallow. It
is a grassland relatively sparse and rather tall (on
average 80 cm) and dominated by Stipa bromoides
and Asphodelus ramosus. These two species are accompanied
by many chamaephytes (Micromeria
juliana, Satureja montana, Teucrium polium subsp.
capitatum). Therophytes are also abundant, in
particular the illyric Bupleurum veronense. Other
therophytes (Avena barbata, Dasypyrum villosum,
Carthamus lanatus) are related to the enrichment
of nutrients probably by sheep and goats.
Relevés 62 and 71 represent a transitional aspect
with Bromo-Chrysopogonetum grylli, since the
floristic composition is similar to Stipo-Salvietum,
but the dominant species is Chrysopogon gryllus.
This aspect occurs on gentler slopes than typical
Stipo-Salvietum, on a northern aspect, but again
in the lower belt. A few species of Festuca-Brometea
are restricted to this aspect (Koeleria splendens,
Leontodon crispus); these species indicate a
tendency toward cooler conditions.
The association is widespread, in an almost
identical aspect, in Dalmatia (Horvatić 1963).
Asphodelo-Chrysopogonetum grylli Horvatić
(1936) 1958 (Table 25; Figure 2, Cluster L)
This community is characterized by the dominance
of Chrysopogon gryllus that is accompanied
by relatively few species of dry grasslands, in
particular Galium lucidum, Helianthemum apenninum,
Stachys recta, Onosma arenaria, Agropyron
elongatum, Scabiosa triniifolia and Satureja montana,
often with low cover or frequency. Salvia
officinalis is sometimes rather abundant, whereas
therophytes are relatively poorly represented.
This community occurs on slopes steeper than
those of Stipo-Salvietum (20–40°), usually with a
S–W aspect. Stones and rocks have on average
a cover of about 30–40%. It occupies an upper
belt with respect to Stipo-Salvietum (see chapter
on zonation).
The association is described from Dalmatia
(Horvatić 1958, 1963), and is reported as widespread
in Albania by Buzo (1990) (under association
Asphodelus ramosus + Chrysopogon gryllus).
Brachypodio-Cymbopogonetum hirti Horvatić
1961 (Table 26; Figure 2, Cluster F)
This association is characterized by the dominance
of Hyparrhenia hirta together with perennial
species of Mediterranean dry grasslands, such
as Convolvulus elegantissimus and Carlina coymbosa.
A few suffruticose species, probably related
to fire, such as Teucrium polium subsp. capitatum,
Micromeria juliana, Petrorhagia saxifraga are in
common with Stipetum, but more abundant here.
This association develops on moderate slopes
(10–15°) on S or W aspects; bare rock has a very
high cover (70–90%) and this community is more
stony than any other community of grasslands or
garrigues in the study area.
The association can be referred to Brachypodio-Cymbopogonetum
hirti, described for the
southern Dalmatia (Korčula, Hvar, Pag, in
Horvatić 1962, 1963). A few associations with
Hyparrhenia hirta have been described for Catalunya
and Italy (Bolòs 1962, Biondi et al. 1988,
Fanelli 2002). The community of Velipoje is actually
easily distinguished by species with a more
or less eastern barycentre of distribution, such as
Convolvulus elegantissimus and Micromeria juliana,
but it is ecologically, structurally and floristically
very closely related to the communities present
in Italy. Buzo (1990) describes for Sarandes an
association with Hyparrhenia hirta + Dasypyrum
villosum that seems different.
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Hacquetia 14/2 • 2015, 129–174
138
Salvio-Phlomidetum fruticosae Barbagallo,
Brullo & Fagotto 1979 (Table 27; Figure 2,
Cluster E)
The floristic composition of this community
is characterized by the dominance of Phlomis fruticosa,
accompanied by a few species of stony dry
grasslands, in particular Stipa bromoides and Melica
ciliata, which also occurr in Stipetum. Other
species are typical of garrigues and probably related
to fire: Micromeria juliana, Petrorhagia saxifraga
and Teucrium polium subsp. capitatum.
This community presents two aspects. The
first one is more floristically interesting, presenting
species such as the subendemic Ballota macedonica,
Satureja calamintha, Geranium pusillum
and Osyris alba together with therophytes such as
Sideritis romana (in a transitional form between
subsp. romana and subsp. purpurea) and Catapodium
rigidum. The other aspect presents many
suffruticose species that are in common with Cymbopogonetum:
Salvia officinalis, Teucrium chamaedrys,
Satureja montana, and a few species of stony
dry grasslands: Orlaya grandiflora, Cephalaria
transylvanica, Allium sphaerocephalon. Stones and
rocks have a higher cover in the latter aspect, that
is identical to Salvio-Phlomidetum fruticosae (Barbagallo
et al. 1989) described for Eastern Sicily
in the surrounding of Siracusa. Phlomis fruticosa
communities are present also in the Apennines
(Pirone 1995b) and in Greece (Raus 1979), but
they are quite distinct, with a more pronounced
nitrophilous character, a higher number of therophytes
and few suffruticose species. Moreover,
they usually grow in deeper and less rocky soils.
The association is widespread in Albania on
dry, calcareous hills along the coasts and on the
southern mountains (Buzo 1990, Hoda & Mersinllari
2000). Buzo (1990) reports a number of associations
(Phlomis fruticosa + Asphodelus ramosus,
Phlomis fruticosa-Psilurus incurvus, Phlomis fruticosa-Urginea
maritima, Phlomis fruticosa-Salvia
officinalis) that are distinguished only by the codominant
species.
Xerothermophilous forests and
scrubs of eastern sub-Mediterranean
regions
Rhamno-Paliuretum Trinajstić 1996 (Table 28,
Figure 2, Cluster A)
=Paliuretum adriaticum Horvatić 1963 nom. illeg.,
Rhamno-Paliuretum (Horvatić) Brullo, Minissale,
Spampinato 1997
Paliurus spina-christi shrubs are widespread in
the carbonatic range, but are usually quite fragmented.
Paliurus spina-christi is accompanied by
a few Quercetea ilicis species such as Phillyrea latifolia,
Euphorbia characias and Clematis flammula,
and many species of submediterranean fringes
such as Bupleurum praealtum.
A number of associations with Paliurus spinachristi
have been described for Italy, Catalunya,
Anatolia and the former Yugoslavia (Horvatić
1963, Géhu & Uslu 1989, Biondi 1999, Blasi & Di
Pietro 2001). Our relevés can be referred to the
dalmatic Rhamno-Paliuretum (Horvatić 1963)
Clematido viticellae-Punicetum granati ass.
nova hoc loco (Table 29; Figure 2, Cluster G)
Holotypus Table 34, rel. 27.
This community is widespread in the study
site, in particular in the lower part of the carbonatic
range, where limestone is in contact with the
alluvial plain. It is characterized by the dominance
of Punica granatum, which forms a thick
bush on average 3m tall, with many lianas (Rosa
sempervirens, Clematis flammula, C. viticella) with
Crataegus monogyna and Paliurus spina-christi as
shrubs, and an herb layer not very developed but
rich in relatively mesophilous species typical of
fringes such as Viola odorata and Geum urbanum.
This community is syngenetically very interesting:
Punica granatum is a Tertiary relict, with a
distribution from former Yugoslavia to Afghanistan
(Meusel & Jäger 1992), but other relictual
species are present, in particular Clematis viticella.
The community is well characterized and was
provisionally proposed in Dring et al. (2002). We
therefore proceed to a formal decription.
Querco-Carpinetum orientalis Horvatić 1939
(Table 30; Figure 2, Cluster B)
This community is characterized by the dominance
of Carpinus orientalis. Acer campestre, Celtis
australis, Quercus pubescens and Fraxinus ornus are
particularly important among trees and shrubs.
Rel. 19 presents a few rather mesophilous species:
Tilia cordata, Corylus avellana, Cornus mas. It was
carried out in a ravine, and is probably transitional
to a mesophilous community that is not fully developed
in the study area. The herb layer is poor.
This community develops usually on rather
steep slopes (15–40°); the structure is a thick
bushland 3–4 m tall. It develops mainly on the
southern slope of the carbonatic range, at lower
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Giuliano Fanelli et al.: The vegetation of the Buna river protected landscape (Albania)
139
nity with Acer monspessulanum; nonetheless often
these three associations are in contact and form
a complex mosaic that is difficult to tease apart.
A number of communities with Carpinus orientalis
have been described from former Yugoslavia,
mainly with illegitimate names (Blečić &
Lakušić 1967, Horvat et al. 1974, Poldini 1988) as
well as in central Italy (Blasi et al. 2001). However,
the variability is poorly known mainly because
this vegetation is usually heavily degraded.
This community presents also some affinity with
Phillyreo-Carpinetum (Bergmeier & Dimopoulos
2008), that is more mediterranean.
Acer monspessulanum community (Table 31;
Figure 2, Cluster D)
This community is close to the preceding
one, but is floristically poorer. It is characterized
by the dominance of Acer monspessulanum
with a shrub layer of Paliurus spina-christi. This
community grows on the northern slopes of the
carbonatic range, a few meters higher in altitude
than Carpinetum orientalis. It forms a mosaic with
Rhamno-Paliuretum.
Acer monspessulanum is generally indicated as a
species typical of woodlands dominated by Carpinus
orientalis (Horvat et al. 1974, Bergmeier & Dimopoulos
2008), but in our analysis Acer monspessulanum
community is clearly distinct from them;
the situation is similar in Italy. The herb layer is
nonetheless very poor so as it is very difficult to
characterize this heavily disturbed community
from a phytosociological point of view.
Quercetum trojanae Em 1958 (Table 32; Fig-
ure 2, Cluster H)
Quercus trojana is present in two distinct physiognomic
and ecological aspects in the carbonatic
range. On the steep southern slopes of the range,
toward the sea, it is represented by large, sparse
trees, often centuries old but not taller than 15–
18 m. The shrubland layer is absent, and under
the trees the herb layer is represented by grasses
such as Chrysopogon gryllus and Stipa bromoides
so as to have a savannah-like structure. On the
north-eastern, gentler slopes of the range, trees
are much younger, thinner and smaller, usually
no more than 15 m, but they form a rather dense
degraded forest. In this forest both species of the
evergreen forest (Asparagus acutifolius, Phillyrea
latifolia, Clematis flammula) and species of the
deciduous woodlands (Quercus petraea, Quercus
pubescens, Fraxinus ornus, Brachypodium sylvaticum,
Oenanthe pimpinelloides, Dactylis polygama)
occur. The presence of Quercus petraea, so close
to the sea, is particularly noticeable. This forest is
protected from grazing with fences realized with
the branches of Paliurus spina-christi, which represents
an important source of wood for fire.
The ecology of this eastern Mediterranean oak
species, therefore, is in some way analogous to the
western Mediterranean Quercus suber that occurs
both in “dehesa” or “montado” open formations
and in dense forests together with mesophilous
species (Testi & Lucattini 1994). Both species are
relictual, being evolved probably in the Tertiary
in a climate warmer and moister than today.
Three associations with Quercus trojana have
been described in the Balkans and Italy: Quercetum
trojanae Em 1958 for Macedonia, Euphorbio
apii-Quercetum trojanae Bianco, Brullo, Minissale,
Signorello & Spampinato 1998 and Teucrio siculiQuercetum
trojanae Biondi, Casavecchia, Guerra,
Medagli, Beccarisi & Zuccarello 2004 for Apulia
in Italy (Horvat et al. 1974, Bianco et al. 1998, Biondi
et al. 2004). Euphorbio apii-Quercetum trojanae
seems to be a degradative or ecotonal stage, whereas
Teucrio-Quercetum trojanae is very similar to our
association, in particular for the presence of Carpinus
orientalis, Quercus pubescens s.l. and Oenanthe
pimpinelloides together with species of Quercetea ilicis.
Nonetheless, Quercus petraea is lacking and species
of Quercus cerris woodlands, such as Teucrium
siculum are present. Quercetum trojanae is described
for Macedonia, where it is present in aspects clearly
more mesophile than our relevés. Our community
is therefore intermediate between Quercetum
trojanae and Teucrio-Quercetum trojanae and probably
deserves further investigation. The main diagnostic
character is the co-occurrence of Quercus
pubescens, Quercus petraea and Quercus trojana. We
refrain from a formal description because the area
that we have studied is too small to quantify the
variability of the community. Moreover, albeit
woodlands with Quercus trojana are widespread in
Albania (Proko 1997), but with heavily degraded
aspects, a comprehensive analysis is still lacking.
Seslerio autumnalis-Ostryetum carpinifoliae
Horvat & Horvatić ex Horvat et al. 1974
(Table 33; Figure 2, Cluster C)
On the top of the Black Peak, above about 400
m a.s.l., a woodland occurs that is dominated by
Ostrya carpinifolia, Quercus cerris, Fraxinus ornus
and occasionally by Quercus trojana. The herb
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Hacquetia 14/2 • 2015, 129–174
140
nalis and other species such as Ornithogalum pyrenaicum,
Viola odorata, Geum urbanum and Poa
sylvicola. This woodland is heavily coppiced, but
nemoral species nonetheless predominate. It can
be referred to Seslerio-Ostryetum, a community
widespread in Dalmatia at intermediate altitudes
(400–800 m) near the coast (Horvat et al. 1974).
The species variability of this association is poorly
known; in the north it is rich in Illyrian species
such as Mercurialis ovata and Spiraea chamaedrifolia
that lack in our southern relevés.
Riparian and floodplain forests
Hierachical classification identified four groups
of alluvial forests discussed hereafter (Figure 3).
Junco-Fraxinetum parvifoliae I. Kárpáti &
V. Kárpáti 1961 (Table 34; Figure 3, Cluster D)
This association grows in depressions of the
Velipoje Reserve subjected to long inundation.
Physiognomically, it is characterized by the dominance
of Fraxinus angustifolia subsp. oxy­carpa,
with a herb layer dominated by Juncus maritimus
and other wetland species: Apium nodi­florum, Iris
pseudacorus, Samolus valerandi and others. The
association has been reported for alluvial areas
close to the sea in Northern Albania including
Velipoje by Kárpáti & Kárpáti (1961) and Kárpáti
(1962). These authors emphasize the presence
of species such as Juncus maritimus and Samolus
valerandi that they interpret as halophytes. Conversely
these can be considered as halotolerant
species, that, although preferring moderate levels
of salinity, are generally related to strong fluctuations
of the water level, from completely dried out
to completely inundated. These strong fluctuations
are characteristic of the climate of Albania,
where winter and autumn rainfall are very high,
as already noticed by Kárpáti (1962).
Kárpáti & Kárpáti (1961) and Kárpáti (1962)
also report a different community with dominant
Fraxinus oxycarpa from the Buna river area: Echinodoro-Fraxinetum
parvifoliae I. Kárpáti & V. Kárpáti
1961. The association occupied depressions
where water is present all the year round, and
is characterized by species typical of ponds and
lakes, such as Baldellia ranunculoides, Iris pseudacorus
and Nymphaea alba.
After a careful investigation, we could not
find this association. It has probably disappeared
because of the lowering of the water table after
the drainage of the marshlands. In Horvat et al.
(1974) a picture is reported that shows a physiognomy
no more present in the area, with huge
Fraxinus angustifolia subsp. oxycarpa among abundant
Iris pseudacorus.
Populetum albae Br.-Bl. ex Tchou 1948
(Table 35; Figure 2, Cluster C)
The association is present along the Buna river,
but is heavily disturbed. The floristic composition
is poor, with many species that are transgressive
from other communities (Fraxinus angustifolia
subsp. oxycarpa, Periploca graeca) or indicators of
disturbance (Carduus acanthoides). The community
is highly invaded by Amorpha fruticosa.
The association has been already reported for
the area under the name Populetum albae balcanicum
(Kárpáti 1962), however, it is difficult to distinguish
it from the typical Populetum albae Tchou
1948 described for Southern France (Tchou 1948,
Braun-Blanquet et al. 1952).
Quercus robur and Carpinus betulus community
(Table 36; Figure 2, Cluster A)
= Querco roboris-Carpinetum betuli submediterraneum
Bertović ex Horvat 1974 nom. illeg.
This forest develops locally in small fragments
near the mouth of the Buna river delta where it
grows in the drier areas of the forest.
The species composition is characterized by
the dominance of Carpinus betulus and Quercus
robur, this last being present with an allegedly
Figure 3: Dendrogram of alluvial forest vegetation. The
capital letters indicate the cluster membership of each rele­vé.
Slika 3: Dendrogram poplavnih gozdov. Velike črke predstavljajo
uvrstitev posameznega popisa v klaster.
A: Quercus robur and Carpinus betulus community; B: Periploco
graecae-Alnetum glutinosae; C: Populetum albae;
D: Junco-Fraxinetum parvifoliae.
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Giuliano Fanelli et al.: The vegetation of the Buna river protected landscape (Albania)
141
endemic variety scutariensis. The composition is
typical of mesophile forests: Viola reichenbachiana,
Aristolochia pallida, Arum italicum. Hygrophile
trees such as Fraxinus angustifolia subsp. oxycarpa
and Populus alba are related to catenal contacts.
This community represents a Mediterranean
outpost of planitial forests, widespread in central
Europe, that in the Mediterranean basin retreats
near rivers and occurs relictually in the drier parts
of alluvial plains (Dierschke 1980). In these extrazonal
sites, the floristic composition changes,
showing a transitional character between QuercoFagetea
and Populetea albae, leading to the definition
of a separate alliance, Alno-Quercion roboris
(Horvat 1938, Kárpáti 1962). Many syntaxa have
been referred to this alliance, in particular in the
former Yugoslavia (Raus 1993, Raus & Vukelić
1993), but the alliance is imperfectly known (see
the synthesis in Brullo & Spampinato 1999), and
the classification of our relevés is provisional.
Periploco graecae-Alnetum glutinosae Rudski
in Horvat et al. ex Fanelli, De Sanctis, Gjeta,
Mullaj & Attore ass. nova hoc loco (Table 31;
Figure 2, Cluster B)
= validation of Periploco-Alnetum Rudski in Horvat
et al. 1974 nom. nudum (art. 3b Horvat et al.
1974: p. 174)
Holotypus Table 31, rel. 55
This community develops in the alluvial
plains along the coast, from the mouth of the
Buna river southwards. It is present also in the
floodplain east of the Viluni lagoon, where no
relevé has been carried out. This forest is rather
dense, but the trees are not very tall, up to 12 m.
The floristic composition is characterized by a
large number of lianas: Periploca graeca, Cynanchum
acutum, Humulus lupulus. Beside Alnus glutinosa
the most important trees and shrubs are
Ulmus minor, Cornus mas, Ligustrum vulgare and
Ficus carica, while the herb layer is rather scarce.
Periploca graeca, Cynanchum acutum and Ficus
carica are species with subtropical affinity, and
indicate to the possible relictual character of this
alluvial forest. This hypothesis is also supported
by the abundance of lianas. The presence of Periploca
graeca seems to differentiate this black alder
association from any other.
Kárpáti & Kárpáti (1961) report three relevés
similar to our association from the west bank
of the Shkodra lake under the name Alno-Fraxinetum
parvifoliae Tchou 1949; the first of these
three relevés being the holotype of Aristolochio rotundae-Alnetum
glutinosae (Brullo & Spampinato
1999). The latter name must be corrected since
the Aristolochia is undoubtably Aristolochia pallida
(Aristolochio pallidae-Alnetum glutinosae Brullo &
Spampinato 1999 corr. hoc loco). This association
is different from the black alder forests of the
mouth of the Buna and seems intermediated between
Periploco-Alnetum and Querco-Carpinetum.
It is possible that the relevés are heterogeneous,
including different belts, but, given the accuracy
of the Hungarian authors, it is more probable that
a lowering of the water-table led to a change to
the zonation of vegetation along the Buna River.
Rudski (in Horvat et al. 1974, pag. 174) describes
a Periploco-Alnetum nom. nud. for Macedonia
that is very similar in the composition of
lianas (Hedera helix, Periploca graeca) but quite
different in the herb layer, probably because it
develops on more acidic granites.
The “association” with Alnus glutinosa, Periploca
graeca and Hedera helix described in Mersinllari
& Hoda (1985) from the surroundings of
Durres is identical to this association.
Tamarisk schrublands of moist places
Vitici-Tamaricetum africanae Horvatić 1960
(Table 38)
This community develops on the shores of
the Viluni lagoon and in other brackish ponds
in particular in Campo di Marina (Kampi veror)
and in the Velipoje Reserve. The vegetation is
dominated by Tamarix africana that forms stands
2–5 m tall. Vitex agnus-castus is present when the
vegetation is less dense. Species of the herb layer
are mainly characteristic of Juncetalia maritimae.
The community forms a large belt between
the Juncetum maritimo-acuti and the mud flats
with succulent chenopods near the water.
The association Vitici-Tamaricetum aftricanae
has been described in Dalmatia (Horvatić 1963),
and also reported for Albania (Dring et al. 2002).
Ruderal and sub-ruderal herbaceous
vegetation
Bromo villosi-Haynaldietum villosae Pignatti
1953 (Table 39)
This community occurs on retrodunal sands in
the Rjolla beach (Backs Rrjolli) in disturbed sites
among settlements. It is a rather sparse grassland,
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Hacquetia 14/2 • 2015, 129–174
142
about 50 cm tall. The composition is rich in species
of Brometalia rubenti-tectorum (Dasypyrum villosum,
Bromus madritensis, Avena barbata) Hordeion
(Lepidium graminifolium, Sisymbrium officinale,
Carduus pycnocephalus) and Brachypodietalia distachyae
(Trifolium scabrum, Medicago minima). It
is very close to Laguro-Dasypyretum villosi Fanelli
1998 (Fanelli 1998) widespread along the coasts
of the Italian Peninsula, but is distinguished by
the lack of the characteristic species of the associations
and the presence of a few rather continental
species such as Kochia prostrata. Moreover,
instead of the Western Mediterranean Vicia pseudocracca,
Vicia villosa subsp. eriocarpa is present.
Centaureetum calcitrapae Fanelli & Menegoni
1997 (Table 40)
= Centaureo calcitrapae-Lolietum perennis Blasi et
al. 2009
At the conjuction between the carbonatic
range and the alluvial plain, in abandoned sheep
pens, a vegetation develops that is dominated by
Centaurea calcitrapa or Scolymus hispanicus, spiny
Asteraceae typical of heavily grazed and nitrophile
sites in the Mediterranean basin. The vegetation is
about 90 cm tall, very dense with a relatively rich
floristic composition: Rumex pulcher, Cichorium intybus,
Bromus hordeaceus, Avena barbata and others.
Soils are loamy and compacted by trampling.
This vegetation can be referred to Centaureetum
calcitrapae described from the surroundings
of Rome (Fanelli & Menegoni 1997, Fanelli 2002,
Blasi et al. 2009), although a few continental species
are present such as Anthemis austriaca.
Asphodelus ramosus community
(Table 41; Figure 2; Cluster M)
This community develops on gentle northwestern
slopes of the carbonatic range. It forms a
mosaic with Paliuretum, and is dominated by Asphodelus
ramosus with many ruderal and subruderal
therophytes: Carthamus lanatus, Dasypyrum
villosum, Catapodium rigidum and Nigella arvensis.
It is an open community growing on a stony soil
and subjected to heavy grazing.
The syntaxonomy of this community is difficult
to be defined. It shows similarities with
Carthametalia lanatae described for the ruderal
communities on clays in Sicily (Brullo & Marcenò
1985). The associations described for this order
are rather different, and this is probably related to
the different lithology (clays instead of limestone)
and to the imperfect knowledge of the order.
Carduetum acanthoidis Felföldy 1942
(Table 42)
This community is widespread in the Domni
marshland and in the fields near Velipoje, on raised
terrains of the alluvial plain and shuffled carbonatic
soils. It is characterized by the dominance of tall
thistles such as Carduus acanthoides, Cirsium vulgare
and Cirsium italicum, together with other ruderal
perennial species as Agrostis stolonfiera, Galega officinalis
and Potentilla reptans. The vegetation is tall
(1.7–1.8 m) and presents a high cover.
This association can be referred to Carduetum
acanthoidis, reported for Germany, Slovakia and
Romania. This community is considered a basal
community by Jarolimek et al. (1997), since
Carduus acanthoides occurs with high frequency
in other communities of Dauco-Melilotion and Onopordion
acanthi. In the study area it seems well
characterized.
Vegetation series and
sequences
Dunes
The alluvial plain of the area is fringed seawards
by a wide system of dunes that today are heavily
degradated by strong erosion and the building of
touristic settlements. As a consequence, natural
vegetation is fragmented and localized, in particular
at the mouth of the Buna river.
What remains of the dune vegetation can be
ordered in the following sequence (R1):
•	 on the foredunes near the sea the Cakilo-Xanthietum
italici can be found. This association
is present today only in the Velipoje Reserve,
near the mouth of the Buna river, where is
rather well developed;
•	 back of the foredunes there are strongly eroded
embryonic dunes, with fragments of psammophilous
communities with Euphorbia paralias or
Elytrigia juncea, too degraded to be described.
The embryonic dunes are scarcely developed
and rise only a few dm above the sea;
•	 back of the embryonic dunes, sand-dunes with
Ammophila are lacking and interdunal depressions
develop instead. The vegetation of these
interdunal depressions is heavily affected by
human impact, and is represented by fragments
of populations of Juncus acutus and Vitex
agnus-castus, that derive respectively from
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Giuliano Fanelli et al.: The vegetation of the Buna river protected landscape (Albania)
143
Juncetum maritimo-acuti and the Vitici-Tamaricetum
associations that are well developed
around the lagoons. These fragments, in particular
the populations of Vitex agnus-castus,
often occur directly on the shore, where the
erosion has destroyed the foredunes and embryonic
dunes;
•	 back of the interdunal depressions a few
fragments of retrodunal vegetation still exist,
in particular secondary vegetation with
Dasypyrum villosum and Aegilops triuncialis,
two ruderal species typical of sandy or gravelly
habitats. This vegetation is represented only
by small patches among the recently built
touristic settlements.
Before the retrodunal depression there is a
large plantation of Pinus pinea, which probably
replaces a belt with Ammophila.
Buzo (1990) and Mersinllari & Hoda (1985)
describe a similar zonation respectively for southern
Albania and Durres.
Brackish waters
Brackish waters are extensively present in Velipoje.
The most important water body is the Viluni
lagoon, but many smaller, sometimes temporary
ponds are also occurring.
The vegetation of this unit is diverse and complex.
Nonetheless, patterns are highly repetitive
and recurrent, and it is possible to reconstruct a
sequence, with a variant, along an environmental
gradient determined by increasing time of inundation
and salinity, more or less complete or
truncated.
Lagoons
Hereafter a few case studies are described.
Campo di Marina (Kampi veror)
This is a large temporary pond, with saline
waters, near the Buna river. The zonation of the
vegetation is in form of concentric belts. Outside
the pond, Alnus glutinosa woody patches extensively
develop. The first outer belt of the pond is
represented by rather sparse treelets of Tamarix
dalamatica. Then a vegetation belt dominated by
Juncus acutus, Juncus maritimus, with many halophytes
such as Limonium vulgare subsp. serotinum
and Inula crithmoides appears, often interrupted
because of the grazing by cattle. In the centre of
the pond there is a large population of Salicornia
perennans.
Small ponds in the Velipoje Reserve
The brackish ponds in the Velipoje Reserve
are important for wildlife, especially migrating
birds. Close to the Buna river there is a large belt
of Phragmitetum. This is followed by a narrower
belt of Scirpetum compacto-litoralis, and then by
an interrupted belt of Tamarix; mixed with Tamarix,
but extending further inward, there are large
populations of Juncus acutus with Amorpha fruticosa.
In this case the water is only slightly brackish,
and the halophile character increases in the
drier areas, due to intense evaporation that concentrates
the salts.
Viluni lagoon
The Viluni lagoon is rather large, and tends to
be more brackish near the sea, where there is an inflow
of sea water, and less brackish inwards, where
there are many channels bringing fresh­water.
Table 43: Vegetation sequences of the lagoons.
Tabela 43: Vegetacijske sekvence v laguni.
Series H1 H1a
I Fraxinus angustifolia subsp. oxycarpa Alnus glutinosa
↓
Salinity
Inundation
II Phragmites australis Phragmites australis
III Typha latifolia Scirpus litoralis (inflow of freshwater)
IV Bolboschoenus maritimus
V Juncus maritimus-Juncus acutus
VI Tamarix africana-Vitex agnus-castus
VII Halimione portulacoides-Aeluropus litoralis
VIII Salicornia perennans
IX Salicornia procumbens
X Ruppia cirrhosa
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Hacquetia 14/2 • 2015, 129–174
144
In the part near the sea, the shores are characterized
by mud colonized by halophytes: close
to the shore there is a vegetation belt with locally
large stands of Salicornia procumbens, followed by
a vegetation dominated by Salicornia perennans.
Inward a belt with Halimione portulacoides and
Aeluropus littoralis can be found preceding stands
of Juncus maritimus.
In the drier parts, seaward, there are also populations
of Euphorbia paralias or Elytrigia juncea
and other dunal species that indicate the presence
of embryo sand dunes.
The water body is colonized by a dense population
of submerged Ruppia cirrhosa, together
with many filamentous algae.
In the intermediate part, between the mud-flats
and the water there is a belt with Tamarix africana.
In the inner part of the lagoon the zonation is
rather different. The mud-flats are lacking, and
from the shores to the water it is possible to observe
three belts: first Phragmitetum, than Scirpetum
maritimo-litoralis, and finally Scirpetum litoralis.
Behind the lagoon, in the north, there is a plain
with damp soils where extensive populations of
Juncus maritimus develop, fringed by woodlands
with dominant Alnus glutinosa. In areas with more
water, Phragmites australis appears again.
Part of this plain has been reclaimed and destined
to agriculture, while in the few areas where
there is still natural vegetation, interesting examples
of Schoeno-Erianthetum ravennae are present.
Freshwaters
The zonation of freshwaters is more complex
than that of brackish waters. It is possible, in fact,
to distinguish three sequences and a variant, according
to a gradient from still to flowing waters.
Domni marsh
The Domni marsh is characterized by a tall
reed-bed dominated by Typha angustifolia and
Phragmites australis forming monospecific large
stands. This dominant vegetation types is probably
the result of the drainage of a natural marsh.
In fact, where pits have been excavated, the vegetation
is dominated by Nymphaea alba and Potamogeton
spp. that probably represent relics of formerly
deeper still waters.
Where Typha angustifolia is not present, mainly
because it has been destroyed for providing pasture
for cattle, secondary vegetation appears with
Paspalum paspalodes in moister sites and TrifolioCynodontetum
in dryer sites. On artificial risings
of the ground level along unpaved roads, a rich
ruderal vegetation dominated by Carduus acanthoides
grows.
Buna River
The vegetation along the water course of the Buna
river is still rather well preserved. It is possible to
observe the typical sequence of river banks (H2),
that begins with two belts of floating submerged
macrophytes, an outer belt on deep waters with
dominant Potamogeton pectinatus and an inner
belt in less deep waters with dominant Potamogeton
nodosus, and locally, in disturbed sites, Elodea
canadensis. More close to river bank four belts of
helophytes develop: in deeper waters the rather
sparse Scirpus lacustris, then Iris pseudacorus, Typha
angustifolia and finally Phragmites australis.
Table 44: Vegetation sequences of the freshwaters.
Tabela 44: Vegetacijske sekvence v celinski vodi.
Series H2 H3 H4 H4b
Flowing waters Slowly flowing channels Still water Disturbed
I Carduus acanthoides
↓
Waterdepth
II Populus alba Quercus robur-Carpinus
betulus
Trifolium fragiferumCynodon
dactylon
III Phragmites australis Paspalum distichum
IV Typha angustifolia Typha angustifolia Typha angustifolia
V Iris pseudacorus Sparganium erectum
VI Schoenoplectus lacustris
VII Potamogeton nodosuscrispus
(Elodea)
Hydrocharis morsus-
ranae
Nymphaea alba
VIII Potamogeton pectinatus (Trapa natans)
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Giuliano Fanelli et al.: The vegetation of the Buna river protected landscape (Albania)
145
These belts are often lacking or intermixed, due
to the changing profile of the river bank; when
the profile is steep, the sequence is less developed
with respect to areas with a gentler profile. In
some areas populations of Bolboschoenus maritimus
are present, probably indicating some mixing
with seawaters.
Beyond the helophytic belts, the remnant of
an alluvial forest vegetation can be found, that
has attracted the attention of botanists since a
long time (Kárpáti & Kárpáti 1961, Kárpáti 1962).
Woodland vegetation of the Buna river
alluvial plain
The zonation of the vegetation of the alluvial
forest is rather simple, with the different associations
ordered along a single gradient of increasing
moisture from the Querco-Carpinetum, that
grows in relatively dry soils, to the Alnetum that
is influenced by the high water table but is rarely
inundated, to the Populetum that develops close
to the river and is often subjected to flooding, to
Fraxinetum that grows in areas with permanent
or prolonged inundation. A similar zonation is
described by Mersinllari & Hoda (1985) for areas
near Durres. The spatial distribution of these
associations is highly fragmented with frequent
ecotonal belts due to the complex dynamism of
the delta of the Buna river in the Holocene. This
in turn determined a complex morphology, with
depressions and risings, and a variable water table
level creating the “catenas” indicated in the
rows of Table 6.
Kárpáti & Kárpáti (1961) describe the succession
at the mouth of the Buna river as leading
from Nymphaeion albae to Echinodoro-Fraxinetum
up to Alno-Fraxinetum. We could not find examples
of the latter two associations. This probably
means that the water table has been lowered, with
the disappearance of an entire sequence.
Spatial distribution of the sequences
of the alluvial plain
When the sequences of the alluvial plain are
mapped, an interesting coincidence with the
geological map appears (Figure 4). Series H1 of
brackish waters is related to the “marshland deposits”
near the sea; series H1a of less brackish
waters to “marshland deposits” farther from the
sea, series H3 of slowly flowing waters is related
with “lagoon deposits”, and series H4 of still waters
with “alluvial-marshland deposits”. These
four belts are parallel to the sea, and probably
are related the progression of the delta of the Buna
river in the Holocene: belts farther from the
sea are older while belts near the sea are younger.
The advancement of the delta of the Buna created
a mosaic of environments (brackish and freshwaters,
still and flowing waters) that enhances the
great complexity of this landscape.
Carbonatic range
Forest vegetation
It is difficult to reconstruct the zonation of the
forest and shrub vegetation, given the present state
of heavy degradation. In the vegetation below 400
m a.s.l., at the base of the carbonatic range, there
are three physiognomic units: 1) a lower belt of
Punica granatum; 2) on the steep southern slopes
Figure 4: Map of the vegetation series and sequences of the
Buna river Protected Landscape. H1: series of brackish and
halophilous vegetation related to the “marshland deposits”;
H1a: series of less brackish waters farther from the sea; H3:
series of slowly flowing waters related with “lagoon deposits”;
H4: series of still waters related with “alluvial-marshland deposits”;
X1: moister carbonatic series of Carpinetum orientalis
at the base of the south-western slope; X2: dryer carbonatic
series on north-east slopes; X3: carbonatic series of the upper,
steep slopes of the Black Peack (Maja e Zezë).
Slika 4: Karta vegetacijskih serij in sekvenc v zavarovanem
območju ob reki Buna. H1: serija brakične in halofilne vegetacije
na “močvirnih naplavinah”; H1a: serija manj brakičnih
voda oddaljenih od morja; H3: serija počasi tekočih voda na
“naplavinah lagune”; H4: serija stoječih voda na “aluvialnomočvirnih
naplavinah”; X1: vlažna karbonatna serija Carpinetum
orientalis ob vznožju jugozahodnega pobočja; X2: suha
karbonatna serija na severovzhodnem pobočju; X3: karbonatna
serija na zgornjem, strmem pobočju Črnega vrha (Maja
e Zezë).
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Hacquetia 14/2 • 2015, 129–174
146
a sparse savanoid woodland with Quercus trojana,
that at lower altitudes and on gentler slopes presents
an heavily grazed undergrowth of Carpinus
orientalis; 3) on north-eastern slopes a mixed,
heavily degraded woodland with Carpinus orientalis,
Quercus pubescens, Celtis australis, Pistacia
terebinthus, Paliurus spina-christi, Phillyrea latifolia,
and, locally, Quercus trojana and Quercus petraea.
These physiognomic units are nonetheless the
result of a long history of disturbance, in particular
grazing, coppicing and fire. The areas with
more developed vegetation are protected from
grazing with fences made with branches of Paliurus
spina-christi, probably to preserve a source
of wood for charcoal; in the other sites pigs and
goats are allowed to graze everywhere.
The phytosociological analysis of the table
of relevés reveals a different situation, with three
woodland and two shrub associations. Such a
high β-diversity suggests that more than one series
of vegetation is present, today homogenized and
partly destroyed by the human impacts. Probably,
two series were present: a moister series (X1), in
particular at the base of the south-western slope,
and a dryer series (X2) on north-east slopes:
Series X1: Carpinetum orientalis → Punicetum granatae
→ Asphodelus ramosus
Series X2: Quercetum trojanae → Acer monspessulanum
→ Paliuretum → Salvio-Phlomidetum →
Stipo-Salvietum officinalis.
On the upper, steep slopes of the Black Peak
there is a different woodland type, Seslerio-Ostryetum
carpinifoliae, belonging to a third series:
Series X3: Seslerio-Ostryetum carpinifoliae → Asphodelo-Chrysopogonetum
grylli.
In order to support such an interpretation few
hints can be presented:
1)	 in Carpinetum orientalis a few mesophile species
occur: Tilia cordata, Acer campestre;
2)	 a few species of Quercetalia pubscentis occur
both in Carpinetum orientalis and Punicetum
but not in both Quercetum trojanae and in the
community with Acer monspessulanum: Crataegus
monogyna Lithospermum purpureo-coeruleum,
Viola odorata;
3)	 spiny species such as Asparagus acutifolius, Paliurus
spina-crhisti are more frequent in Acer
monspessulanum and Quercetum trojanae than
in Carpinetum orientalis.
The distinction between the two series is probably
related to differences in the soils. The legend
of the geologic map (Halili 1985) shows that both
dolomites and limestones are present in the carbonatic
range. However, there is no information
on the distribution of these two lithotypes, but our
direct observations seem to show that limestones
are prevalent at the base of the range, whereas dolomites
are prevalent at higher altitudes, above 90–
100 m on the northern slope, above 150 m on the
southernslope.Thisissuggestedbytheratherbrisk
break of slope at these altitudes. Significantly, in
correspondence with this slope break Carpinus orientalis
disappears or becomes less abundant. This
observation is particularly evident on the southern
slopes of the Black Peak near Blacs-Rjolli. The associations
of series X1 develop at lower altitudes
than the associations of the series X2, so it is probable
that the former develops on limestones, the latter
on dolomites. Limestone is calcium carbonate,
whereas dolomite contains also magnesium, and
this leads to different chemical and pedological
properties of soils. This topic is little studied from
a vegetational and ecophysiological point of view,
but it is well known. For instance, many Mediterranean
pines grow mainly on dolomites (Barbero
et al. 1998), whereas a few grassland types are restricted
to this rock, for instance in the Causses,
in Southern France, where on dolomites develops
even an endemic alliance, Armerion junceae Br.-Bl.
in Br.-Bl., Roussine & Nègre 1952 (Braun-Blanquet
et al. 1952). It is possible that dolomites induce a
sort of xerophytism in the vegetation that develops
on soils derived from this rock.
Grasslands
The carbonatic range is 561 m in altitude with
rather steep slopes on the southwest side, and
gentler on the eastern side. There is a clear zonation
of the grassland vegetation related to altitude
and variation in steepness. As an example, the
south-western slope of the Black Peak is rather
gentle at the beginning, than becomes steeper up
to the top where the steepness can be up to 50°.
The flora is structured in four belts (Figure 5).
At the base, where the slopes are about 20°, there
is a belt dominated by Stipa bromoides; other species
are Linum nodiflorum, Lolium rigidum subsp.
lepturoides, Stachys recta, Aurinia petraea and Campanula
lingulata. Higher up, there is a transitional
belt, and thereafter, where the slopes are about
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Giuliano Fanelli et al.: The vegetation of the Buna river protected landscape (Albania)
147
30–40°, a belt where Chrysopogon gryllus becomes
dominant. Here a few, relatively rare species are
abundant: Centaurea salonitana, Onosma arenaria
and Scabiosa triniifolia. Near the top the slope is
very steep, and also small vertical cliffs are present.
Here the dominant species is Achnatherum
calamagrostis, usually in monospecific stands,
and on the slopes Putoria calabrica, Moltkia petraea
and Sesleria robusta subsp. skanderbegii occur.
Northwards, on the southern slopes of Mount
Nikgionih, the slope is very steep at the base
(about 40°) and gentler higher up (10–25°). On
the steeper belt the dominance of Hyparrhenia
hirta can be observed, whereas on gentler slopes it
is possible to observe the dominance of Stipa bromoides.
The northern slope is gentle (15–30°) and
a transitional belt with the dominance of Chrysopogon
gryllus occurs, but with dominated species
similar to Stipo-Salvietum officinalis (Figure 6).
In summary it is possible to observe the following
sequence according to an increasing steepness:
Asphodelus microcarpus → Stipo-Salvietum officinalis
→ Stipo-Salvietum officinalis with Chrysopogon
gryllus → Asphodelo-Chrysopogonetum gryllus →
Achantherum calamagrostis → Brachypodio-Cymbopogonetum
hirti→ Seslerio-Putorietum calabricae
Steepness is the distal ecological factor; it covaries
with soil thickness and rockiness, and indirectly
with water and nutrient availability. In
fact, there is an interaction with aspect. Typical
Stipo-Salvietum occurs at the base of the southern
slope, whereas higher up Chrysopogonetum develops;
the transitional facies of Stipo-Salvietum
with dominant Chrysopogon gryllus occurs at the
Figure 5: Vegetation belts of Black Peak (Maja e Zezë).
Slika 5: Vegetacijski pasovi na Črnem vrhu (Maja e Zezë).
Figure 6: Vegetation belts of Mt. Nikgionih (Mali i Nikgjonit).
Slika 6: Vegetacijski pasovi na gori Nikgionih (Mali i Nikg-
jonit).
base of the moister, cooler north-eastern slopes.
In Croatia, the two associations are strongly differentiated:
Stipo-Salvietum grows on thin soils
on stony hills; Chrysopogonetum grows in plain on
deep soils (Horvat 1962, Horvat et al. 1974).
Moreover, it is possible that the break of slope
is related also to lithotype, since both dolomites
and limestones are present in the carbonatic
range. Dolomite is less erodible, and can form
very steep slopes, like in the eponym range of
Dolomites, whereas limestones are more erodible
and tend to form gentler slopes. Unfortunately,
information on the detailed distribution of lithotypes
is lacking.
4. CONCLUSIONS
Although strongly modified by human activities,
the Buna River Protected Landscape is still characterized
by a high diversity of community types
classified in 46 associations, 27 alliances, 23 orders
and 17 classes.
This diversity is related not to the richness of
the flora, that is not exceptional for a Mediterranean
area, but to a complex interaction between
morphological and geological features. In turn,
this complexity determined the presence of 1
dunal and 4 hygro-sequences, related to the age
of deposits of Buna River, in relationship to the
advancement of the delta in the Holocene, and
of 3 xeroseries in the carbonatic range, related to
lithological differences and altitudinal gradient.
The vegetation is very interesting, with many
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Hacquetia 14/2 • 2015, 129–174
148
communities with a probably relictual character,
as shown by species such as Periploca graeca,
Clematis viticella, Punica granatum, Ficus carica,
Laurus nobilis.
Despite the heavy environmental modifications
caused by the land reclamation works of the
second half of the last century, the conservation
status of the remnant vegetation is still relatively
good. Nonetheless, the area is subjected to a process
of increasing human impact that is particularly
evident when comparing the present alluvial
vegetation with the description made by Kárpáti
& Kárpáti (1961) and Kárpáti (1962); a few types
have disappeared or are nowadays rare, in particular
those related to permanent freshwaters.
More recently, the touristic development is
threatening the coastal environments. The Rjolla
beach (Backs Rrjolli) presented, probably not
far ago, interesting examples of dunal vegetation
that have today almost completely disappeared.
5. ACKNOWLEDGMENTS
We would like to thank the personnel of the Forest
District Service Eduard Gajtani, Tonin Macaj,
Elvin Shala, Zequir Ujca and Edoardo Scepi,
Francesconi Fabio for their support during the
field work. A special thank to Nadia Abdelahd,
Emanuela Congestri, Emanuela Viaggiu for the
determination of algae, to Andraž Čarni, Mauro
Iberite, Gianfranco Pirone and Roberto Venanzoni
for providing valuable scientific references
and to Sandro Pignatti for making available his
phytosociological library.
This study was carried out within the framework
of the IUCN Project “Institutional Support
to the Albanian Ministry of Environment, Forest and
Water Administration (MoEFWA) for Sustainable
Biodiversity Conservation and Use in Protected Areas
and the Management of Waste” funded by the
DGCS (Italian Development Conservation).
Syntaxonomic list
RUPPIETEA MARITIMAE J. Tüxen. ex Den Hartog & Segal 1964
RUPPIETALIA J. Tüxen ex Den Hartog & Segal 1964
Ruppion maritimae Br.-Bl. ex Westhoff in Bennema, Sissingh & Westhoff 1943
Ruppietum cirrhosae Iversen 1934
THERO-SALICORNIETEA STRICTAE Tüxen in Tüxen & Oberdorfer 1958
THERO-SALICORNIETALIA STRICTAE Pignatti 1952
Thero-Salicornion strictae Br.-Bl. 1933
Salicornietum venetae Pignatti 1966
Suaedo spicatae-Salicornietum patulae (Brullo & Furnari 1976) Géhu & Géhu-Franck 1984. corr.
Alcaraz, Ríos, De la Torre, Delgado & Inocencio 1998
JUNCETEA MARITIMI Br.-Bl. in Br.-Bl., Roussine & Nègre 1952
JUNCETALIA MARITIMI Br.-Bl. ex Horvatić 1934
Agropyro-Artemision coerulescentis Pignatti 1953
Puccinellio festuciformis-Aeluropetum litoralis (Corb. 1968) Géhu & Costa 1984
Juncion maritimi Br.-Bl. ex Horvatić 1934
Juncetum maritimo-acuti Horvatić 1934
Plantaginion crassifoliae Br.-Bl. in Br.-Bl., Roussine & Nègre 1952
Juncus maritimus and Lippia nodiflora community
CAKILETEA MARITIMAE Tüxen & Presing ex Br.-Bl. & Tüxen 1952
EUPHORBIETALIA PEPLIDIS Tüxen ex Oberdorfer 1949
Euphorbion peplis Tüxen ex Oberdorfer 1952
Cakilo-Xanthietum italicae Pignatti 1953
LEMNETEA O. de Bolòs & Masclans 1955
LEMNETALIA MINORIS Tüxen ex O. de Bolòs & Masclans 1955
Hydrocharition morsus-ranae (Passarge 1964) Westhoff & den Held 1969 = Lemno minorisHydrocharition
morsus-ranae Rivas-Martínez, Fernández-González & Loidi 1999, [Hydrocharition
morsus-ranae Passarge 1996, non Rübel 1933]
Hydrocharitetum morsus-ranae van Langendonck 1935
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Giuliano Fanelli et al.: The vegetation of the Buna river protected landscape (Albania)
149
POTAMOGETONETEA Klika in Klika & Novak 1941
POTAMOGETONETALIA W. Koch 1926
Nymphaeion albae Oberdorfer 1957
Nymphaeetum albae Vollmar 1947
Trapetum natantis Kárpáti 1963.
Potamogetonion Miljan 1933 = Magnopotamion (Vollmar 1947) den Hartog & Segal 1964,
Parvopotamion (Vollmar 1947) den Hartog & Segal 1964; Potamion pectinati (Koch 1926)
Görs 1977
Potamogetonetum denso-nodosi O. de Bolòs 1957
Potamogetonetum pectinati Carstensen ex Hilbig 1971
Elodeetum canadensis Nedelcu 1967
PHRAGMITI-MAGNOCARICETEA Klika in Klika & Novak 1941
PHRAGMITETALIA COMMUNIS W. Koch 1926
Phragmition communis W. Koch 1926
Typhetum angustifoliae Pignatti 1953
Typhetum latifoliae Nowiński 1930
Phragmitetum australis Savič 1926 nom. mut. prop.
Scirpetum lacustris Chouard 1924
Glycerio-Sparganietum neglecti Koch 1926
SCIRPETALIA MARITIMI Heijný in Holub, Heijný, Moravec & Neuhäusl 1967
Scirpion maritimi Dahl & Hadač 1941
Scirpetum maritimo-litoralis (Br.-Bl. in Br.-Bl., Roussine & Nègre 1952) O. de Bolòs 1962
Scirpetum litoralis Pignatti 1953
MOLINIO-ARRHENATHERETEA Tüxen 1937
HOLOSCHOENETALIA Br.-Bl. ex Tchou 1948
Molinio-Holoschoenion Br.-Bl. ex Tchou 1948
Schoeno-Erianthetum ravennae Pignatti 1953
PASPALO-HELEOCHLOETALIA Br.-Bl. in Br.-Bl. et al. 1952
Paspalo-Agrostidion Br.-Bl. in Br.-Bl. et al. 1952
Paspalo-Agrostidetum Br.-Bl. 1936 (1952)
Trifolio-Cynodontion Br.-Bl. & O. de Bolòs 1957
Trifolio fragiferi-Cynodontetum Br.-Bl. & O. de Bolòs 1957
ASPLENIETEA TRICHOMANIS (Br.-Bl. in Meier & Br.-Bl. 1934) Oberdorfer 1977
MOLTKIETALIA PETREAE Lakušić 1968
Seslerion tenuifoliae Horvat 1930
Seslerio-Putorietum calabricae Horvatić 1963
FESTUCO-BROMETEA Br.-Bl. & Tüxen ex Klika & Hadeč 1944
SCORZONERO-CHRYSOPOGONETALIA Horvatić & Horvat in Horvatić 1963
Chrysopogono-Saturejion Horvat & Horvatić in Horvatić 1934
Stipo-Salvietum officinalis Horvatić 1958
Asphodelo-Chrysopogonetum grylli Horvatić (1936) 1958
THERO-BRACHYPODIETEA RAMOSI Br.-Bl. ex A. et O. de Bolòs 1950 (= Lygeo-Stipetea tenacissimae
Rivas-Mart. 1978 nom cons. prop.)
CYMBOPOGONO HIRTI-BRACHYPODIETALIA RAMOSI Horvatić 1958
Cymbopogono hirti-Brachypodion ramosi Horvatić 1958
Brachypodio-Cymbopogonetum hirti Horvatić 1961
CISTO-MICROMERIETEA JULIANAE Oberdorfer 1954
CISTO-ERICETALIA Horvatić 1958
Cisto-Ericion manipuliflorae Horvatić 1958
Salvio-Phlomidetum fruticosae Barbagallo, Brullo & Fagotto 1979
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QUERCETEA PUBESCENTIS Doing-Kraft ex Scamoni et Passarge 1959
FRAXINO ORNI-COTINETALIA Jakucs 1960
Paliuro-Carpinion orientalis Crisurean & Ţeculescu 1968
Rhamno-Paliuretum Trinajstić 1996
Clematido viticellae-Punicetum granati ass. nova hoc loco
QUERCETALIA PUBESCENTI-PETREAE Klika 1933
Carpinion orientalis Horvat 1954
Querco-Carpinetum orientalis Horvatić 1939
Acer monspessulanum community
Quercetum trojanae Em 1958
Seslerio autumnalis-Ostryetum carpinifoliae Horvat & Horvatić ex Horvat et al. 1974
POPULETEA ALBAE Br.-Bl. 1962
FRAXINETALIA Scamoni & Passarge 1959
Lauro-Fraxinion angustifoliae I. Kárpáti et V. Kárpáti 1961 = Fraxinion angustifoliae Pedrotti
1970 nom. prov.
Junco-Fraxinetum parvifoliae I. Kárpáti & V. Kárpáti 1961
POPULETALIA ALBAE Br.-Bl. ex Tchou Yen-Tcheng 1948
Populion albae Br.-Bl. ex Tchou Yen-Tcheng 1948
Populetum albae Br.-Bl. ex Tchou 1948
Alno-Quercion roboris Horvat 1938
Quercus robur and Carpinus betulus community
Periploco graecae-Alnetum glutinosae Rudski in Horvat et al. ex Fanelli, De Sanctis, Gjeta, Mullaj,
Attore ass. nova hoc loco
NERIO-TAMARICETEA Br.-Bl. & O. de Bolòs 1958
TAMARICETALIA AFRICANAE Br.-Bl. & O. de Bolòs 1958
Tamaricion africanae Br.-Bl. & O. de Bolòs 1958
Vitici-Tamaricetum africanae Horvatić 1960
STELLARIETEA MEDIAE Tüxen Lohmeyer et Preising in Tüxen ex von Rochow 1951
BROMETALIA RUBENTI-TECTORUM Rivas Mart. & Izco 1977 = Thero-Brometalia annuae (Rivas Goday
& Rivas Mart. ex Esteve 1973) O. de Bolòs 1975
Laguro ovati-Bromion rigidi Géhu & Géhu-Franck 1985 nom. nud.	
Bromo villosi-Haynaldietum villosae Pignatti 1953
SISYMBRIETALIA OFFICINALIS J. Tüxen ex W. Matuszkiewicz 1962
Hordeion murini Br.-Bl. in Br.-Bl., Gajewski, Wraber & Walas 1936	
Centaureetum calcitrapae Fanelli & Menegoni 1997
CARTHAMETALIA LANATAE Brullo in Brullo & Marcenò 1985
Asphodelus ramosus community
ARTEMISIETEA VULGARIS Lohmeyer, Preising & Tüxen ex von Rochow 1951
ONOPORDETALIA ACANTHII Br.-Bl. & Tüxen ex Klika & Hadač 1944
Dauco-Melilotion Görs 1966 ex Rostański & Gutte 1971
Carduetum acanthoidis Felföldy 1942
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7. Appendix
Coordinates and date of vegetation relevés
Rel.	 01:	 N 4635488.67,	 E 366422.98,	 26/06/2012;
rel.	 02:	 N 4635473.54,	 E 366393.02,	 26/06/2012;
rel.	 03:	 N 4634077.49,	 E 365346.05,	 26/06/2012;
rel.	 06:	 N 4642898.92,	 E 370341.97,	 27/06/2012;
rel.	 07:	 N 4642893.37,	 E 370344.94,	 27/06/2012;
rel.	 08:	 N 4642902.13,	 E 370359.14,	 27/06/2012;
rel.	 09:	 N 4642895.72,	 E 370365.44,	 27/06/2012;
rel.	 10:	 N 4642808.34,	 E 370130.74,	 27/06/2012;
rel.	 11:	 N 4642816.46,	 E 370126.72,	 27/06/2012;
rel.	 12:	 N 4642666.76,	 E 369951.83,	 27/06/2012;
rel.	 13:	 N 4642673.23,	 E 369972.34,	 27/06/2012;
rel.	 14:	 N 4642882.30,	 E 370238.04,	 27/06/2012;
rel.	 15:	 N 4641417.10,	 E 370660.76,	 27/06/2012;
rel.	 16:	 N 4641529.20,	 E 370802.94,	 27/06/2012;
rel.	 17:	 N 4641107.57,	 E 370661.49,	 27/06/2012;
rel.	 18:	 N 4641126.37,	 E 370654.22,	 27/06/2012;
rel.	 19:	 N 4640378.36,	 E 370214.78,	 27/06/2012;
rel.	 20:	 N 4639576.28,	 E 370903.21,	 27/06/2012;
rel.	 21:	 N 4639597.16,	 E 370910.26,	 27/06/2012;
rel.	 22:	 N 4639155.30,	 E 371246.68,	 27/06/2012;
rel.	 23:	 N 4635139.35,	 E 373019.20,	 27/06/2012;
rel.	 24:	 N 4642309.22,	 E 366473.43,	 27/06/2012;
rel.	 25:	 N 4642305.37,	 E 366448.69,	 27/06/2012;
rel.	 26:	 N 4642259.85,	 E 366543.43,	 27/06/2012;
rel.	 27:	 N 4642278.14,	 E 366134.45,	 27/06/2012;
rel.	 28:	 N 4642220.81,	 E 366056.24,	 27/06/2012;
rel.	 29:	 N 4635920.60,	 E 373983.00,	 28/05/2012;
rel.	 30:	 N 4636193.31,	 E 373833.74,	 28/05/2012;
rel.	 31:	 N 4636378.04,	 E 373827.81,	 28/05/2012;
rel.	 32:	 N 4636472.10,	 E 373541.50,	 28/05/2012;
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Hacquetia 14/2 • 2015, 129–174
156
rel.	 33:	 N 4636464.87,	 E 373994.27,	 28/05/2012;
rel.	 34:	 N 4636370.87,	 E 373807.33,	 28/05/2012;
rel.	 35:	 -	 -	 29/06/2012;
rel.	 36:	 N 4636366.50,	 E 371284.85,	 29/06/2012;
rel.	 37:	 N 4636366.50,	 E 371284.85,	 29/06/2012;
rel.	 38:	 N 4636347.03,	 E 371271.28,	 29/06/2012;
rel.	 39:	 N 4636454.70,	 E 370814.40,	 29/06/2012;
rel.	 40:	 N 4636440.40,	 E 370811.10,	 29/06/2012;
rel.	 41:	 N 4636426.80,	 E 370800.80,	 29/06/2012;
rel.	 42:	 N 4636388.18,	 E 370778.28,	 29/06/2012;
rel.	 43:	 N 4637838.35,	 E 370047.42,	 29/06/2012;
rel.	 44:	 N 4637826.00,	 E 370037.00,	 29/06/2012;
rel.	 45:	 N 4636311.00,	 E 371991.00,	 29/06/2012;
rel.	 49:	 N 4635697.75,	 E 370295.07,	 29/06/2012;
rel.	 50:	 -	 -	 29/06/2012;
rel.	 51:	 N 4637191.54,	 E 366339.67,	 30/06/2012;
rel.	 52:	 N 4637042.60,	 E 365242.73,	 30/06/2012;
rel.	 53:	 N 4637115.69,	 E 365235.47,	 30/06/2012;
rel.	 54:	 N 4636741.52,	 E 365263.08,	 30/06/2012;
rel.	 55:	 N 4636702.67,	 E 365200.76,	 30/06/2012;
rel.	 56:	 N 4636666.45,	 E 365258.20,	 30/06/2012;
rel.	 57:	 N 4636381.74,	 E 365371.10,	 30/06/2012;
rel.	 58:	 N 4641317.03,	 E 368621.14,	 30/06/2012;
rel.	 59:	 N 4641311.14,	 E 368626.88,	 30/06/2012;
rel.	 60:	 N 4640129.00,	 E 370502.37,	 30/06/2012;
rel.	 61:	 N 4640149.92,	 E 370534.19,	 30/06/2012;
rel.	 62:	 N 4639444.30,	 E 372834.80,	 30/06/2012;
rel.	 63:	 N 4640791.94,	 E 371404.37,	 30/06/2012;
rel.	 64:	 N 4640810.13,	 E 371380.06,	 30/06/2012;
rel.	 65:	 N 4639599.13,	 E 370860.37,	 01/06/2012;
rel.	 66:	 N 4639497.74,	 E 371020.20,	 01/06/2012;
rel.	 67:	 N 4639475.42,	 E 371135.57,	 01/06/2012;
rel.	 68:	 N 4639405.50,	 E 371252.20,	 01/06/2012;
rel.	 69:	 N 4639431.82,	 E 371247.81,	 01/06/2012;
rel.	 70:	 N 4639460.15,	 E 371283.49,	 01/06/2012;
rel.	 71:	 N 4639462.22,	 E 371278.88,	 01/06/2012;
rel.	 72:	 N 4639084.33,	 E 371483.04,	 01/06/2012;
rel.	 73:	 N 4638900.54,	 E 371552.59,	 01/06/2012;
rel.	 74:	 N 4636001.00,	 E 372136.00,	 29/06/2012;
rel.	 75:	 N 4634582.60,	 E 365044.77,	 03/07/2012;
rel.	 76:	 N 4634823.80,	 E 365055.90,	 03/07/2012;
rel.	 77:	 N 4634823.90,	 E 365055.80,	 03/07/2012;
rel.	 78:	 N 4635090.30,	 E 365064.80,	 03/07/2012;
rel.	 79:	 N 4635090.30,	 E 365064.80,	 03/07/2012;
rel.	 80:	 N 4635753.79,	 E 365204.34,	 03/07/2012;
rel.	 81:	 N 4635751.08,	 E 365203.80,	 03/07/2012;
rel.	 83:	 N 4636633.65,	 E 365198.34,	 03/07/2012;
rel.	 84:	 N 4637057.70,	 E 365220.82,	 03/07/2012;
rel.	 85:	 N 4637300.67,	 E 365313.44,	 03/07/2012;
rel.	 86:	 N 4637606.19,	 E 365489.48,	 03/07/2012;
rel.	 87:	 N 4638060.75,	 E 365774.91,	 03/07/2012;
rel.	 88:	 N 4638220.60,	 E 365793.40,	 03/07/2012;
rel.	 89:	 N 4638220.66,	 E 365793.48,	 03/07/2012;
rel.	 90:	 N 4634938.15,	 E 372709.73,	 18/09/2012;
rel.	 91:	 N 4634994.58,	 E 372619.28,	 18/09/2012;
rel.	 93:	 N 4635036.88,	 E 372420.82,	 18/09/2012;
rel.	 94:	 N 4635160.30,	 E 4635160.70,	 18/09/2012;
rel.	 95:	 N 4635088.09,	 E 371968.39,	 18/09/2012;
rel.	 96:	 N 4635217.47,	 E 371931.44,	 18/09/2012;
rel.	 97:	 N 4634755.69,	 E 373358.84,	 18/09/2012;
rel.	 99:	 N 4634457.28,	 E 374230.51,	 18/09/2012;
rel.	100:	 N 4635735.99,	 E 372005.49,	 18/09/2012;
rel.	 101:	 N 4635734.31,	 E 372003.98,	 18/09/2012;
rel.	 102:	 N 4640124.05,	 E 369212.97,	 19/09/2012;
rel.	 103:	 N 4637992.86,	 E 369830.38,	 19/09/2012;
rel.	 104:	 N 4637962.73,	 E 369814.57,	 19/09/2012;
rel.	 105:	 N 4637932.77,	 E 369878.97,	 19/09/2012;
rel.	 106:	 N 4637932.70,	 E 369878.97,	 19/09/2012;
rel.	 107:	 N 4637569.69,	 E 369610.53,	 19/09/2012;
rel.	 108:	 N 4637055.47,	 E 369950.47,	 19/09/2012;
rel.	 109:	 N 4635581.45,	 E 366086.18,	 22/09/2012;
rel.	 110:	 N 4635436.49,	 E 365921.77,	 22/09/2012;
rel.	 111:	 N 4636351.31,	 E 365334.50,	 22/09/2012;
rel.	 112:	 N 4635436.40,	 E 365921.70,	 22/09/2012;
rel.	 113:	 N 4636534.83,	 E 365453.40,	 22/09/2012;
rel.	 151:	 N 4636118.90,	 E 374899.70,	 28/06/2012;
rel.	 152:	 N 4636389.70,	 E 374431.30,	 28/06/2012;
rel.	 168:	 -	 -	 31/05/2013;
rel.	 169:	 -	 -	 29/05/2013;
rel.	 170:	 -	 -	 29/05/2013;
rel.	 191:	 N 4637196.00,	 E 366401.00,	 16/06/2013;
rel.	 192:	 N 4637125.00,	 E 366290.00,	 16/06/2013;
rel.	 193:	 N 4637185.00,	 E 366258.00,	 16/06/2013;
rel.	 194:	 N 4637212.00,	 E 366183.00,	 16/06/2013;
rel.	 195:	 N 4637170.00,	 E 366522.00,	 16/06/2013;
rel.	 196:	 N 4637200.00,	 E 366427.00,	 16/06/2013;
rel.	 197:	 N 4637239.00,	 E 366490.00,	 16/06/2013;
rel.	199:	 N 4637231.00,	 E 365283.40,	 16/06/2013;
rel.	200:	 N 4636863.00,	 E 365232.00,	 16/06/2013;
rel.	 201:	 N 4636750.00,	 E 365237.00,	 16/06/2013;
rel.	202:	 N 4636667.00,	 E 365256.00,	 16/06/2013;
rel.	203:	 N 4635455.00,	 E 365165.00,	 16/06/2013;
rel.	204:	 N 4635451.00,	 E 365191.00,	 16/06/2013;
rel.	 207:	 N 4637031.00,	 E 373434.00,	 18/06/2013;
rel.	208:	 N 4637136.00,	 E 373704.00,	 18/06/2013;
rel.	209:	 N 4636777.00,	 E 374273.00,	 18/06/2013;
rel.	 210:	 N 4636755.00,	 E 373781.00,	 18/06/2013;
rel.	 211:	 N 4635451.00,	 E 365191.00,	 16/06/2013;
rel.	212:	 N 4641725.00,	 E 370694.00,	 17/06/2013;
rel.	 213:	 N 4641725.00,	 E 370696.00,	 17/06/2013;
rel.	 214:	 N 4642016.00,	 E 370738.00,	 17/06/2013;
rel.	 215:	 N 4643138.00,	 E 370706.00,	 17/06/2013;
rel.	 216:	 N 4643038.00,	 E 370529.00,	 17/06/2013;
rel.	 217:	 N 4642873.50,	 E 370264.00,	 17/06/2013;
rel.	 218:	 N 4642890.00,	 E 370336.00,	 17/06/2013;
rel.	 219:	 N 4642890.00,	 E 370336.00,	 17/06/2013;
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Giuliano Fanelli et al.: The vegetation of the Buna river protected landscape (Albania)
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rel.	220:	 N 4642843.00,	 E 370219.00,	 17/06/2013;
rel.	221:	 N 4642843.00,	 E 370014.00,	 17/06/2013;
rel.	223:	 N 4642634.00,	 E 369849.00,	 17/06/2013;
rel.	224:	 N 4642388.00,	 E 369512.00,	 17/06/2013;
rel.	225:	 N 4642388.00,	 E 369512.00,	 17/06/2013;
rel.	 227:	 N 4642987.00,	 E 369511.00,	 17/06/2013;
rel.	228:	 N 4642349.00,	 E 369460.00,	 17/06/2013;
rel.	230:	 N 4639113.00,	 E 371126.00,	 17/06/2013;
rel.	 231:	 N 4639151.00,	 E 371092.00,	 17/06/2013;
rel.	232:	 N 4637372.00,	 E 371818.00,	 17/06/2013;
rel.	233:	 N 4635759.00,	 E 366776.00,	 19/06/2013;
rel.	234:	 N 4635897.00,	 E 366712.00,	 19/06/2013;
rel.	235:	 N 4635867.20,	 E 366741.60,	 19/06/2013;
rel.	236:	 N 4635783.00,	 E 366207.00,	 19/06/2013;
rel.	 237:	 N 4635783.00,	 E 366207.00,	 19/06/2013;
rel.	238:	 N 4635616.00,	 E 365893.00,	 19/06/2013;
rel.	240:	 N 4635609.00,	 E 365858.00,	 19/06/2013;
rel.	 241:	 N 4635244.00,	 E 365210.00,	 19/06/2013;
rel.	242:	 N 4634222.00,	 E 365468.00,	 19/06/2013;
rel.	243:	 N 4634182.00,	 E 365390.00,	 19/06/2013;
rel.	244:	 N 4634184.00,	 E 365359.00,	 19/06/2013;
rel.	245:	 N 4634183.40,	 E 365365.90,	 19/06/2013;
rel.	246:	 N 4634177.00,	 E 365366.00,	 19/06/2013;
rel.	 479:	 -	 -	 19/06/2013;
rel.	480:	 -	 -	 19/06/2013;
rel.	481:	 -	 -	 19/06/2013;
rel.	482:	 -	 -	 19/06/2013;
rel.	485:	 -	 -	 29/06/2013;
rel.	486:	 -	 -	 29/06/2013;
rel.	 487:	 -	 -	 30/06/2013;
rel.	488:	 -	 -	 17/06/2013;
rel.	489:	 -	 -	 17/06/2013;
rel.	490:	 -	 -	 17/06/2013;
rel.	 491:	 N 4641529.20,	 E 370802.94,	 27/06/2013;
rel.	492:	 -	 -	 27/06/2013;
rel.	 493:	 -	 -	 01/06/2013.
Table 1 (Tabela 1): Ruppietum cirrhosae Iversen 1934
Relevé number 35 45
Relevé area (m2
) 20 5
Cover total (%) 20 100
Water depth (cm) 50 100
Char. And diff. species of the ass.
Ruppia cirrhosa 2b 5
Table 2 (Tabela 2): Salicornietum venetae Pignatti 1966
Relevé number 36 37 74 94 100
Relevé area (m2
) 5 3 5 300 80
Cover herb layer (%) 80 80 60 100 90
Char. and diff. species of the ass.
Salicornia procumbens 2b 4 3 5 5
Other species
Halimione portulacoides 4 + . . .
Juncus maritimus + + . + .
Aeluropus littoralis 2a + . . .
Limonium vulgare subsp. serotinum + + 2a + .
Salsola soda . . . + .
Table 3 (Tabela 3): Puccinellio festucaeformi-Aeluropetum
litoralis (Corb 1968) Géhu & Costa 1984
Relevé number 39 111 485 486
Relevé area (m2
) 3 20 5 7
Cover herb layer (%) 100 75 80 80
Char. and diff. species of the ass.
Aeluropus littoralis 5 2m 3 3
Other species
Halimione portulacoides + . + .
Salicornia patula . 3 1 .
Suaeda maritima . + + .
Limonium vulgare subsp. serotinum . + . .
Salsola soda . . . +
Artemisia caerulescens . . . +
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Hacquetia 14/2 • 2015, 129–174
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Relevé number 2 91 93 97 99
Relevé area (m2
) 60 80 30 35 45
Cover herb layer (%) 90 90 100 60 85
Char. and diff. species of the ass.
Juncus maritimus . 2a 3 . 3
Lippia nodiflora . + . + +
Inula viscosa . 2b . + +
Other species
Potentilla reptans + . . . .
Amorpha fruticosa 2a + . . .
Periploca graeca 2a . . . .
Cynanchum acutum 1 . . . .
Juncus acutus 4 4 3 3 2b
Agrostis stolonifera + . . . .
Phragmites australis + . . . .
Elymus athericus + . . . .
Table 4 (Tabela 4): Juncetum maritimo-acuti Horvatić 1934
Relevé number 38 41 42 57 90 96 101 103 108 113
Relevé area (m2
) 25 30 60 15 120 30 25 40 25 20
Cover herb layer (%) 90 100 90 90 100 95 100 100 0 100
Char. and diff. species of the ass.
Juncus acutus . 4 3 . + . 4 . + 5
Juncus maritimus 5 . + 4 5 5 2b 5 3 +
Other species
Halimione portulacoides 1 + . . . + + . . .
Limonium vulgare subsp. serotinum 1 3 3 2b 2b 1 + 1 5 2b
Asparagus maritimus . r + . . . . . . .
Phragmites australis . + . . . . . + . +
Aeluropus littoralis . + . + . . . . . .
Elymus athericus . + 2b . . . . . . .
Aster tripolium . . + + . . . 2m . .
Scirpus maritimus . . + . . . . . . .
Juncus gerardi . . + . . . . . . .
Lotus tenuis . . + . . . . . . .
Bolboschoenus maritimus . . . 1 . . . + . .
Salicornia patula . . . . + . . . . +
Artemisia caerulescens . . . . + . + . + .
Salicornia procumbens . . . . . 2b 2b . . .
Salsola soda . . . . . + + . . .
Atriplex latifolia . . . . . . + + + +
Elymus farctus . . . . . . + + + 1
Sonchus maritimus . . . . . . . . + .
Juncus subulatus . . . . . . . . . +
Crypsis aculeata . . . . . . . . . +
Relevé number 2 91 93 97 99
Euphorbia paralias . + . + .
Plantago crassifolia . + . . .
Sonchus maritimus . + + . .
Aeluropus littoralis . + . . .
Cynodon dactylon . + . . .
Artemisia caerulescens . + 2m . .
Elymus farctus . + + . +
Arthrocnemum perenne . . + . .
Inula crithmoides . . + . .
Limonium vulgare subsp. serotinum . . 2m . .
Bolboschoenus maritimus . . + . .
Polypogon monspeliensis . . . + .
Crypsis aculeata . . . + 2m
Dittrichia graveolens . . . + .
Scirpoides holoschoenus . . . + .
Table 5: Juncus maritimus and Lippia nodiflora community
Tabela 5: Združba z vrstama Juncus maritimus in Lippia nodiflora
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Table 6 (Tabela 6): Cakilo-Xanthietum italicae Pignatti
1953
Relevé number 3 242 243 246
Relevé area (m2
) 30 20 5 5
Aspect (degrees) 160 - - Slope
(degrees) 4 - - Cover
herb layer (%) 45 40 40 10
Char. and diff. species of the ass.
Cakile maritima + . . .
Xanthium strumarium 2b 3 1 2a
Polygonum maritimum 1 + . +
Salsola kali subsp. kali 2a . . .
Other species
Echinophora spinosa 1 1 2a +
Sporobolus pungens + . . .
Atriplex prostrata subsp. latifolia + . . .
Salsola kali subsp. tragus + . . .
Eryngium maritimum . + . .
Catapodium marinum . + 1 .
Euphorbia paralias . . + .
Pseudorlaya pumila . . + .
Hedypnois cretica . . + .
Plantago cynops . . + +
Elymus farctus subsp. farctus . . 2a .
Euphorbia peplus . . . +
Stachys maritima . . . 1
Lagurus ovatus . . . +
Table 7 (Tabela 7): Seslerio-Putorietum calabricae
Horvatić 1963
Relevé number 34
Relevé area (m2
) 15
Altitude (m a.s.l.) 482
Aspect (degrees) 180
Slope (degrees) 80
Cover herb layer (%) 5
Cover bare rock (%) 100
Char. and diff. species of the ass.
Moltkia petraea 1
Sesleria robusta subsp. skanderbeggii +
Other species
Ficus carica +
Pistacia terebinthus +
Punica granatum +
Satureja montana +
Achnatherum calamagrostis 1
Table 8 (Tabela 8): Hydrocharitetum morsus-ranae van
Langendonck 1935
Relevé number 58 219 224 225 234
Relevé area (m2
) 18 6 5 7 15
Cover herb layer (%) 95 90 90 85 70
Water depth (cm) 60 100 110 85 70
Char. and diff. species of the ass.
Hydrocharis morsus-ranae 4 4 5 . 5
Other species
Myriophyllum spicatum 3 2a + . .
Potamogeton natans 2b . . . .
Potamogeton crispus . 3 . . .
Phragmites australis . + . . .
Lemna minor . . + . .
Apium nodiflorum . . . + .
Alisma plantago-aquatica . . . + .
Sparganium erectum . . . 5 .
Typha angustifolia . . . + .
Typha latifolia . . . . +
Table 10 (Tabela 10): Trapetum natantis Kárpáti 1963
Relevé number 11 223
Relevé area (m2
) 7 12
Cover herb layer (%) 90 90
Water depth (cm) 160 200
Char. and diff. species of the ass.
Trapa natans 5 5
Other species
Phragmites australis + .
Lemna minor 2b +
Sparganium erectum + .
Typha angustifolia + .
Table 9 (Tabela 9): Nymphaeetum albae Vollmar 1947
Relevé number 6 216 217 218 220 233 240
Relevé area (m2
) 30 4 6 6 7 10 8
Cover herb layer (%) 80 60 60 100 70 80 90
Water depth (cm) 100 90 75 90 120 100 110
Char. and diff. species of the ass.
Nymphaea alba 3 3 4 5 4 5 5
Other species
Myriophyllum spicatum 2b . . 4 . . .
Potamogeton perfoliatus 1 . . . . . .
Typha angustifolia 1 . . . + . .
Elodea canadensis . + . . . . .
Potamogeton pectinatus . + 1 . + + .
Lemna minor . + + . 2a . +
Paspalum paspalodes . + + . . . .
Typha latifolia . . . . . + .
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Hacquetia 14/2 • 2015, 129–174
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Table 11 (Tabela 11): Potamogetonetum denso-nodosi
O. de Bolòs 1957
Relevé number 84 488 489
Relevé area (m2
) 4 8 6
Cover herb layer (%) 100 100 100
Water depth (cm) 60 80 100
Char. and diff. species of the ass.
Potamogeton nodosus 5 4 4
Other species
Potamogeton perfoliatus 1 + .
Potamogeton crispus + . .
Typha latifolia + . .
Potamogeton natans . . +
Table 12 (Tabela 12): Potamogetonetum pectinati
Carstensen ex Hilbig 1971
Relevé number 53 78 80 83 88 89 199
Relevé area (m2
) 10 15 4 5 7 30 10
Cover herb layer (%) 100 100 100 100 100 100 100
Water depth (cm) 100 150 200 100 300 200 150
Char. and diff. species of the ass.
Potamogeton pectinatus 5 3 5 5 5 5 4
Other species
Potamogeton nodosus r . . . . . .
Myriophyllum spicatum . + . . . 1 .
Potamogeton perfoliatus + r 2a . . . .
Potamogeton crispus . + . + . . .
Elodea canadensis . . . + . . 2b
Table 13 (Tabela 13): Elodeetum canadensis Nedelcu
1967
Relevé number 87 487
Relevé area (m2
) 10 8
Cover herb layer (%) 100 100
Water depth (cm) 100 80
Char. and diff. species of the ass.
Elodea canadensis 5 5
Other species
Potamogeton crispus 1 +
Potamogeton natans . +
Table 14 (Tabela 14): Typhetum angustifoliae Pignatti
1953
Relevé number 8 13 221 479 480
Relevé area (m2
) 30 40 5 10 15
Cover herb layer (%) 100 100 100 100 100
Char. and diff. species of the ass.
Typha angustifolia 5 5 5 5 5
Other species
Alisma plantago-aquatica + 1 + . +
Paspalum paspalodes 3 . . . .
Sparganium erectum . 1 2a . .
Phragmites australis . . + . .
Butomus umbellatus . . . + .
Table 15 (Tabela 15): Typhetum latifoliae Nowinski
1930
Relevé number 105 109 235 244
Relevé area (m2
) 30 25 4 13
Cover herb layer (%) 70 100 85 80
Char. and diff. species of the ass.
Typha latifolia 3 5 5 5
Other species
Phragmites australis + + + .
Bolboschoenus maritimus . . . +
Scrophularia nodosa . . + .
Baldellia ranunculoides . . + .
Alisma plantago-aquatica . . + .
Myriophyllum spicatum . . . +
Elymus farctus subsp. farctus + . . .
Tamarix africana + . . .
Atriplex prostrata subsp. latifolia 2m . . .
Table 16 (Tabela 16): Phragmitetum australis Savic
1926 nom. mut. prop.
Relevé number 75 236 245 85 227 110
Relevé area (m2
) 15 10 10 5 6 20
Cover herb layer (%) 100 100 100 95 95 100
Water depth (cm) 100 50 70 40 30 80
Char. and diff. species of the ass.
Phragmites australis 5 5 5 1 5 5
Other species
Iris pseudacorus . . . 5 . .
Scirpus lacustris . . . . . 1
Bolboschoenus maritimus . . . . . +
Rumex hydrolapathum . . + . . .
Eupatorium cannabinum . . + . . .
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Table 17 (Tabela 17): Scirpetum lacustris Chouard 1924
Relevé number 79 81 86
Relevé area (m2
) 15 6 5
Cover herb layer (%) 100 100 100
Water depth (cm) 50 50 30
Char. and diff. species of the ass.
Scirpus lacustris 5 5 1
Other species
Typha latifolia . + .
Bolboschoenus maritimus . + +
Iris pseudacorus . . 5
Table 18 (Tabela 18): Glycerio-Sparganietum neglecti
Koch 1926
Relevé number 59 215 228
Relevé area (m2
) 10 2 6
Cover herb layer (%) 100 90 95
Water depth (cm) 35 40 25
Char. and diff. species of the ass.
Sparganium erectum 5 5 4
Other species
Alisma plantago-aquatica . + +
Phragmites australis . + +
Butomus umbellatus + . .
Hydrocharis morsus-ranae + . .
Potamogeton natans + . .
Typha angustifolia + . .
Scirpus lacustris + . .
Oenanthe fistulosa . + .
Pulicaria odora . + .
Lemna minor . + .
Apium nodiflorum . . 2b
Table 19 (Tabela 19): Scirpetum maritimo-litoralis (Br.Bl.
in Br.-Bl., Roussine & Nègre 1952) O. Bolòs 1962
Relevé number 1 43 237 106
Relevé area (m2
) 3 0 15 10
Cover herb layer (%) 90 80 100 100
Water depth (cm) 20 15 25 20
Char. and diff. species of the ass.
Bolboschoenus maritimus 4 4 5 5
Other species
Phragmites australis + 2b . +
Typha angustifolia 2a . + .
Scirpus lacustris + . . .
Paspalum paspalodes + . . .
Alisma plantago-aquatica . . + .
Agrostis stolonifera + . . .
Table 20 (Tabela 20): Scirpetum litoralis Pignatti 1953
Relevé number 481 482 44
Relevé area (m2
) 15 7 10
Cover herb layer (%) 80 75 70
Char. and diff. species of the ass.
Scirpus litoralis 4 5 4
Other species
Phragmites australis + . .
Scirpus lacustris . + .
Table 21 (Tabela 21): Schoeno-Erianthetum ravennae
Pignatti 1953
Relevé number 102 107 95
Relevé area (m2
) 40 30 30
Cover herb layer (%) 90 100 100
Char. and diff. species of the ass.
Erianthus ravennae 5 3 3
Schoenus nigricans + + .
Char. species of upper syntaxa
Scirpus holoschoenus 1 . 1
Sonchus maritimus . + .
Aster squamatus + + +
Lotus tenuis + + .
Other species
Lythrum salicaria + + +
Carex punctata + 3 1
Inula viscosa + 1 +
Centaurium spicatum . + +
Pulicaria vulgaris + . +
Phragmites australis + + +
Juncus acutus 2m + .
Amorpha fruticosa + . .
Bromus hordeaceus + . .
Clematis flammula . . +
Cynanchum acutum . + .
Equisetum arvense + . .
Euphorbia paralias . . +
Juncus maritimus . + .
Mentha aquatica + . +
Petroselinum segetum . + +
Populus alba + . .
Rubus ulmifolius . . +
Samolus valerandi . + .
Sporobolus pungens . . +
Trifolium repens + . .
Vulpia membranacea . . +
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Table 22 (Tabela 22): Paspalo-Agrostidetum Br.-Bl.
1936 (1952)
Relevé number 9 12 214
Relevé area (m2
) 5 5 5
Cover herb layer (%) 100 100 90
Char. and diff. species of the ass.
Paspalum paspalodes 5 5 5
Other species
Plantago major . . +
Rumex conglomeratus + . .
Alisma plantago-aquatica + . .
Trifolium repens . . +
Eleocharis palustris . . +
Juncus sp. . . +
Table 23 (Tabela 23): Trifolio fragiferi-Cynodontetum
Br.-Bl. et O. de Bolòs 1957
Relevé number 7 212 213
Relevé area (m2
) 2 2 2
Cover herb layer (%) 98 100 100
Char. and diff. species of the ass.
Trifolium fragiferum 3 2a 3
Cynodon dactylon 3 5 3
Other species
Mentha pulegium + + 2a
Plantago lanceolata . + 1
Oenanthe fistulosa + . .
Matricaria chamomilla + . .
Paspalum paspalodes 2a . .
Plantago macrorhiza . + .
Setaria viridis . + .
Juncus species . + .
Rumex conglomeratus . . +
Potentilla reptans . . +
Cirsium vulgare . . +
Table 24: Stipo-Salvietum officinalis Horvatić 1958. Species
with phi-value above 0.50 (only species with a statistically
significant affinity to a cluster according to Fisher’s
exact test with P <0.001 are included) are in bold.
Tabela 24: Stipo-Salvietum officinalis Horvatić 1958.
Krepko so označene vrste z phi>0,50 (vključene so samo
vrste, ki so statistično značilno navezane na klaster
glede na Fisherjev eksaktni test, p <0,001).
Relevé number 29 73 72 62 71
Relevé area (m2
) 60 10 15 10 4
Altitude (m a.s.l.) 144 152 138 83 125
Aspect (degrees) 110 90 100 330 140
Slope (degrees) 20 32 20 15 18
Cover herb layer (%) 80 75 80 100 100
Cover bare rock (%) 0 5 20 90 70
Cover stones (%) 80 45 30 0 0
Char. and diff. species of the ass.
Bupleurum veronense + + 1 + +
Asperula aristata subsp. scabra + . + + +
Satureja montana + . + 2a +
Salvia officinalis 2b 2a + . .
Convolvulus elegantissimus + + + . .
Koeleria splendens . . . 2a +
Leontodon crispus . . . + +
Other species . . . . .
Dasypyrum villosum . . + r 2m
Asphodelus ramosus . 3 + 2b +
Stipa bromoides + 3 4 . 2a
Chrysopogon gryllus + . + 3 5
Micromeria juliana 1 . 1 + +
Teucrium polium subsp. capitatum + . + 2a 1
Avena barbata . + . + +
Brachypodium distachyon 2m . . + +
Petrorhagia saxifraga subsp. saxifraga + . + . +
Teucrium chamaedrys + + + . .
Relevé number 29 73 72 62 71
Bromus erectus + + . . .
Briza maxima . + . + .
Carlina corymbosa . + 2b . .
Catapodium rigidum + + . . .
Eryngium amethystinum . + . 1 .
Melica ciliata . + + . .
Quercus trojana + . + . .
Aegilops geniculata . . . + .
Aethionema saxatile . + . . .
Allium guttatum subsp. guttatum . . + . .
Bituminaria bituminosa + . . . .
Carthamus lanatus . . . . +
Centaurea alba subsp. deusta . . . r .
Cynosurus echinatus . + . . .
Dactylis glomerata subsp. glomerata . . + . .
Dactylis glomerata subsp. hispanica . + . . .
Dorycnium hirsutum . . + . .
Echium plantagineum 2m . . . .
Erysimum repandum . + . . .
Euphorbia spinosa . . . + .
Festuca rupicola . . . . +
Helichrysum italicum subsp. italicum . . . . +
Hyparrhenia hirta . . + . .
Hypericum humifusum . + . . .
Hypericum perforatum subsp. veronense . . . + .
Inula viscosa + . . . .
Linum trigynum . . . + .
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Table 25: Asphodelo-Chrysopogonetum grylli Horvatić
1957. Species with phi-value above 0.50 (only species
with a statistically significant affinity to a cluster
according to Fisher’s exact test with P <0.001 are included)
are in bold.
Tabela 25: Asphodelo-Chrysopogonetum grylli Horvatić
1957. Krepko so označene vrste z phi>0,50 (vključene
so samo vrste, ki so statistično značilno navezane na
klaster glede na Fisherjev eksaktni test, p <0,001).
Relevé number 31 30 32 33
Relevé area (m2
) 25 30 8 20
Altitude (m a.s.l.) 353 277 483 484
Aspect (degrees) 200 120 150 140
Slope (degrees) 23 40 25 25
Cover herb layer (%) 80 60 95 95
Cover bare rock (%) 0 30 0 35
Cover stones (%) 20 40 30 40
Char. and diff. species of the ass.
Chrysopogon gryllus 3 3 5 4
Galium lucidum + . + +
Linum trigynum + + . +
Helianthemum apenninum . . + +
Coronilla scorpioides + . . +
Other species
Teucrium polium subsp. capitatum + . . +
Stachys recta subsp. labiosa + . . .
Anthemis austriaca + + . +
Onosma arenaria . + . .
Salvia officinalis . 2a + +
Bituminaria bituminosa + . + .
Brachypodium distachyon 2b + . .
Convolvulus elegantissimus + . . +
Dactylis glomerata subsp. glomerata + . + .
Satureja montana . 1 . +
Sideritis romana + . . +
Relevé number 31 30 32 33
Stipa bromoides 1 . + .
Aethionema saxatile + . . .
Elymus elongatus . + . .
Althaea hirsuta . . . +
Bromus erectus + . . .
Carlina corymbosa . . . +
Catapodium rigidum . + . .
Dorycnium herbaceum . + . .
Eryngium amethystinum . + . .
Linum nodiflorum . + . .
Pistacia terebinthus . + . .
Quercus trojana . . + .
Micromeria juliana 1 . . .
Scabiosa triniifolia . . + .
Teucrium montanum . + . .
Verbascum phlomoides + . . .
Table 26: Brachypodio-Cymbopogonetum hirti Horvatić
1961. Species with phi-value above 0.50 (only species
with a statistically significant affinity to a cluster according
to Fisher’s exact test with P <0.001 are included)
are in bold.
Tabela 26: Brachypodio-Cymbopogonetum hirti Horvatić
1961. Krepko so označene vrste z phi>0,50 (vključene
so samo vrste, ki so statistično značilno navezane na
klaster glede na Fisherjev eksaktni test, p <0,001).
Relevé number 67 493 69
Relevé area (m2
) 5 15 3
Altitude (m a.s.l.) 70 90 102
Aspect (degrees) 50 50 180
Slope (degrees) 15 15 10
Cover herb layer (%) 50 70 95
Cover bare rock (%) 70 70 90
Cover stones (%) 30 25 15
Char. and diff. species of the ass.
Hyparrhenia hirta 2b 3 4
Aethionema saxatile + + +
Crupina vulgaris + +
Sherardia arvensis + +
Hypericum perforatum subsp. veronense + +
Other species
Convolvulus elegantissimus + +
Carlina corymbosa + +
Teucrium polium subsp. capitatum 2b + 1
Petrorhagia saxifraga subsp. saxifraga + + +
Micromeria juliana + + 2a
Cynosurus echinatus + +
Arenaria leptoclados + +
Brachypodium distachyon 2a +
Anthemis austriaca +
Asphodelus ramosus 1
Avena barbata + + +
Bromus intermedius +
Relevé number 29 73 72 62 71
Orlaya grandiflora . . . + .
Paliurus spina-christi + . . . .
Pistacia terebinthus + . . . .
Potentilla aurea . . . . +
Potentilla recta . . . + .
Prunus spinosa + . . . .
Sanguisorba minor subsp. muricata . . . + .
Securigera cretica . . + . .
Silene vulgaris subsp. vulgaris . + . . .
Teucrium montanum 2m . . . .
Thesium divaricatum . . . + .
Tragopogon pratensis . . . + .
Trifolium angustifolium . . + . .
Trifolium campestre . . + . .
Urospermum picroides . . . . +
Verbascum phlomoides + . . . .
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Hacquetia 14/2 • 2015, 129–174
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Relevé number 67 493 69
Bupleurum veronense +
Chrysopogon gryllus +
Dasypyrum villosum +
Euphorbia spinosa +
Phlomis fruticosa +
Quercus trojana +
Satureja montana +
Sedum rupestre +
Sideritis romana +
Stachys recta subsp. recta +
Thesium divaricatum +
Table 27: Salvio-Phlomidetum fruticosae Barbagallo, Brullo &
Fagotto 1979. Species with phi-value above 0.50 (only species
with a statistically significant affinity to a cluster according
to Fisher’s exact test with P <0.001 are included) are in bold.
Table 27: Salvio-Phlomidetum fruticosae Barbagallo, Brullo &
Fagotto 1979. Krepko so označene vrste z phi>0,50 (vključene
so samo vrste, ki so statistično značilno navezane na klaster
glede na Fisherjev eksaktni test, p <0,001).
Relevé number 68 65 21 66 60 61
Relevé area (m2
) 20 50 50 20 70 30
Altitude (m a.s.l.) 91 - - 44 Aspect
(degrees) 120 150 135 130 160 140
Slope (degrees) 25 15 25 20 15 40
Cover herb layer (%) 70 70 75 95 60 80
Cover bare rock (%) 10 95 0 10 70 100
Cover stones (%) 0 0 70 75 50 0
Char. and diff. species of the ass.
Phlomis fruticosa 4 4 3 4 2b 4
Stipa bromoides + + 2b + + 1
Melica ciliata + . 1 . + +
Micromeria juliana + . + + + .
Petrorhagia saxifraga subsp.
saxifraga . + + + + .
Other species
Teucrium polium subsp. capitatum . . 1 + + .
Ballota macedonica + 1 . . . .
Satureja calamintha . 1 . . . .
Geranium pusillum + . . . . .
Sideritis romana . . + 1 . .
Osyris alba . . + + . .
Catapodium rigidum . . + + . .
Salvia officinalis . . . + + +
Teucrium chamaedrys . . + + + +
Satureja montana . . + . 2a +
Orlaya grandiflora . . . . + +
Cephalaria transylvanica . . . . + .
Allium sphaerocephalon . . . . . +
Brachypodium distachyon + . + + + .
Aethionema saxatile . . . + + .
Anthemis austriaca . . . . + +
Relevé number 68 65 21 66 60 61
Briza maxima . . + . + .
Dactylis glomerata subsp. glomerata
+ . + . . .
Phillyrea latifolia . . 1 . . +
Punica granatum + . . + . .
Rhus coriaria . . . . 2b +
Arenaria leptoclados . . . . + .
Asparagus acutifolius . . . . . +
Asperula aristata subsp. scabra . . . . . +
Asphodelus ramosus . + . . . .
Avena barbata . . . . + .
Bituminaria bituminosa . . . . . +
Carex hallerana . . . . + .
Carlina corymbosa + . . . . .
Carthamus lanatus . . + . . .
Convolvulus cantabrica . . . . . +
Convolvulus elegantissimus . . . + . .
Cynosurus echinatus . . + . . .
Filago germanica . . . + . .
Fraxinus ornus . . . . + +
Linum trigynum . . . . . +
Origanum vulgare . . . . . 2a
Paliurus spina-christi . . + . . .
Phillyrea latifolia . . . + . .
Pistacia terebinthus . . . . + 1
Sedum rupestre . . . . + .
Sherardia arvensis . . + . . .
Urospermum picroides + . . . . .
Table 28: Rhamno-Paliuretum Trinajstić 1996. Species
with phi-value above 0.50 (only species with a statistically
significant affinity to a cluster according to Fisher’s
exact test with P <0.001 are included) are in bold.
Tabela 28: Rhamno-Paliuretum Trinajstić 1996. Krepko
so označene vrste z phi>0,50 (vključene so samo vrste,
ki so statistično značilno navezane na klaster glede na
Fisherjev eksaktni test, p <0,001).
Relevé number 16 491 169
Relevé area (m2
) 50 15 20
Altitude (m a.s.l.) 50 32 25
Aspect (degrees) 20 30 Slope
(degrees) 10 5 Cover
shrub layer (%) 100 100 100
Cover herb layer (%) 5 5 35
Height (highest) shrubs (m) 3 3 4
Height lowest shrubs (m) 1 1 1,5
Char. and diff. species of the ass.
Paliurus spina-christi 5 4 3
Phillyrea latifolia + 3 5
Euphorbia characias + + +
Clematis flammula + + +
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Giuliano Fanelli et al.: The vegetation of the Buna river protected landscape (Albania)
165
Table 29: Clematido viticellae-Punicetum granatae ass. nova
hoc loco. Species with phi-value above 0.50 (only species
with a statistically significant affinity to a cluster according
to Fisher’s exact test with P <0.001 are included) are in bold.
Tabela 29: Clematido viticellae-Punicetum granatae ass.
nova hoc loco. Krepko so označene vrste z phi>0,50
(vključene so samo vrste, ki so statistično značilno
navezane na klaster glede na Fisherjev eksaktni test, p
<0,001).
Relevé number 20 27* 28 22
Relevé area (m2
) 80 60 50 60
Altitude (m a.s.l.) 13 - 25 18
Aspect (degrees) 180 - - 120
Slope (degrees) 15 - - 15
Cover shrub layer (%) 90 100 100 100
Cover herb layer (%) 10 20 5 60
Height (highest) shrubs (m) 17 35 30 40
Cover stones (%) 0 0 0 10
Char. and diff. species of the ass.
Punica granatum 5 5 4 5
Clematis viticella + + + .
Rosa sempervirens 1 + + .
Asplenium onopteris . + + +
Geum urbanum + + + .
Other species
Paliurus spina-christi + 1 2b .
Ruscus aculeatus . 1 1 .
Viola odorata . 2a . 2b
Clematis flammula + + . .
Carex distachya + . . +
Crataegus monogyna . + 1 .
Carpinus orientalis . . 2a .
Catapodium rigidum + . . .
Consolida ajacis . . . +
Dactylis glomerata subsp. glomerata . . . 2b
Relevé number 16 491 169
Bupleurum praealtum + + .
Other species
Asparagus acutifolius + + +
Pistacia terebinthus . 2a 2b
Carthamus lanatus + + .
Rubus ulmifolius 2b 1 .
Rhus coriaria . 1 +
Ruscus aculeatus + . 3
Scolymus hispanicus + + .
Dactylis glomerata subsp. glomerata + . +
Carpinus orientalis 1 . .
Crataegus monogyna . . +
Oenanthe pimpinelloides + . .
Quercus trojana . . +
Relevé number 20 27* 28 22
Hedera helix . . . +
Lathyrus aphaca . . . +
Buglossoides purpurocaerulea . 2a . .
Phillyrea latifolia 1 . . .
Phlomis fruticosa + . . .
Stipa bromoides + . . .
Torilis arvensis subsp. arvensis . . . +
Torilis nodosa + . . .
Brachypodium retusum + . . .
Brachypodium sylvaticum . . . 3
Bupleurum praealtum . + . .
Table 30: Carpinetum orientalis Horvat 1939. Species with
phi-value above 0.50 (only species with a statistically significant
affinity to a cluster according to Fisher’s exact test
with P <0.001 are included) are in bold.
Tabela 30: Carpinetum orientalis Horvat 1939. Krepko so
označene vrste z phi>0,50 (vključene so samo vrste, ki so
statistično značilno navezane na klaster glede na Fisherjev
eksaktni test, p <0,001).
Relevé number 18 19 192
Relevé area (m2
) 80 50 100
Altitude (m a.s.l.) 24 - Aspect
(degrees) - 180 Slope
(degrees) - 15 Cover
tree layer (%) 80 100 80
Cover shrub layer (%) 2 90 2
Cover herb layer (%) 3 10 10
Cover bare rock (%) 5 0 0
Height (highest) trees (m) 6 8 7
Height lowest trees (m) 3 4 6
Height (highest) shrubs (m) 2 2 3
Char. and diff. species of the ass.
Carpinus orientalis 4 5 4
Ruscus aculeatus + 2b 2b
Fraxinus ornus + 2a +
Quercus pubescens + + .
Crataegus monogyna + + .
Acer campestre 2b . 2a
Celtis australis + + .
Other species
Hedera helix . + +
Asparagus acutifolius . + +
Clematis vitalba + . .
Vitis vinifera subsp. sylvestris + . .
Phillyrea latifolia + . .
Buglossoides purpurocaerulea . + .
Asplenium onopteris . + .
Corylus avellana . + .
Clematis flammula . + .
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166
Table 31: Acer monspessulanum community.
Tabela 31: Združba z vrsto Acer monspessulanum.
Relevé number 26 17
Relevé area (m2
) 80 50
Altitude (m a.s.l.) 12 16
Cover tree layer (%) 50 30
Cover shrub layer (%) 15 50
Cover herb layer (%) 20 10
Cover bare rock (%) 5 50
Height (highest) trees (m) 6 5
Height (highest) shrubs (m) 2,5 3
Char. and diff. species of the ass.
Acer monspessulanum 3 2b
Paliurus spina-christi 2a 2b
Phillyrea latifolia + 2b
Asparagus acutifolius + +
Other species
Quercus pubescens . 2a
Carpinus orientalis 2a .
Rosa sempervirens 1 .
Celtis australis + .
Clematis flammula + .
Cruciata glabra + .
Dactylis glomerata subsp. glomerata . 2b
Fraxinus ornus . 2a
Punica granatum . +
Ruscus aculeatus 2b +
Salix caprea 1 .
Table 32: Quercetum trojanae Em 1958. Species with
phi-value above 0.50 (only species with a statistically
significant affinity to a cluster according to Fisher’s exact
test with P <0.001 are included) are in bold.
Tabela 32: Quercetum trojanae Em 1958. Krepko so
označene vrste z phi>0,50 (vključene so samo vrste,
ki so statistično značilno navezane na klaster glede na
Fisherjev eksaktni test, p <0,001).
Relevé number 63 64 170 208
Relevé area (m2
) 80 20 50 90
Altitude (m a.s.l.) 53 55 100 362
Cover tree layer (%) 100 100 100 100
Cover shrub layer (%) 3 50 0 0
Cover herb layer (%) 15 60 50 70
Height (highest) trees (m) 6 10 12 13
Height lowest trees (m) 0 6 10 5
Height (highest) shrubs (m) 4 2 3 2
Height lowest shrubs (m) 1 1 0,5 0
Char. and diff. species of the ass.
Quercus trojana 3 4 2a 2b
Quercus petraea . 2b 4 2a
Dactylis glomerata subsp. aschersoniana + 2b . +
Cornus sanguinea + . + +
Other species
Carpinus orientalis 4 2b 5 2b
Asparagus acutifolius + 3 + .
Clematis flammula + 1 + .
Ruscus aculeatus 2b + 3 .
Fraxinus ornus . . + 3
Hedera helix . 2b 2a .
Paliurus spina-christi + 1 . .
Quercus pubescens + 1 . .
Phillyrea latifolia . 1 . .
Brachypodium sylvaticum + . . .
Oenanthe pimpinelloides . + . .
Dactylis glomerata subsp. glomerata . . . +
Acer monspessulanum . . . +
Aremonia agrimonoides . . . +
Aristolochia pallida . . . +
Aristolochia rotunda . . . +
Buglossoides purpurocaerulea . . . 2b
Festuca heterophylla . . . +
Lathyrus montanus . . . +
Ligustrum vulgare . . 1 .
Milium effusum . . . +
Origanum vulgare . . . +
Pistacia terebinthus . + . .
Potentilla species . . . +
Quercus cerris . . 2a .
Rosa sempervirens . . + .
Sesleria autumnalis . . . 3
Stachys scardica . . . +
Relevé number 18 19 192
Acer campestre . 2a .
Paliurus spina-christi . + .
Tilia cordata . + .
Punica granatum . + .
Cornus mas . + .
Fraxinus ornus . 4 .
Dactylis polygama . + .
Carpinus orientalis . . +
Ulmus minor . . +
Aristolochia pallida . . +
Anemone apennina . . +
Ranunculus lanuginosus . . +
Rosa canina . . +
Crataegus monogyna . . +
Vicia tetrasperma . . +
Geranium purpureum . . +
Viola odorata . . 1
Cornus mas . . 2b
Ligustrum vulgare . . 1
Tamus communis . . +
Luzula forsteri . . +
Chaerophyllum temulentum . . +
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Giuliano Fanelli et al.: The vegetation of the Buna river protected landscape (Albania)
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Relevé number 63 64 170 208
Teucrium chamaedrys . . . +
Thymus pulegioides . . . +
Vincetoxicum hirundinaria . . . +
Viola odorata . . . 2a
Table 33: Seslerio-Ostryetum carpinifoliae Horvat & Horvatić 1950. Species with phi-value above 0.50 (only species
with a statistically significant affinity to a cluster according to Fisher’s exact test with P <0.001 are included) are in bold.
Tabela 33: Seslerio-Ostryetum carpinifoliae Horvat & Horvatić 1950. Krepko so označene vrste z phi>0,50 (vključene
so samo vrste, ki so statistično značilno navezane na klaster glede na Fisherjev eksaktni test, p <0,001).
Relevé number 151 152 207 209 210
Relevé area (m2
) 100 80 100 90 90
Altitude (m a.s.l.) 514 520 432 483 468
Aspect (degrees) - - - 30 30
Slope (degrees) - - - 4 4
Cover tree layer (%) 90 80 60 90 90
Cover shrub layer (%) 30 25 35 5 5
Cover herb layer (%) 10 60 50 80 60
Cover bare rock (%) 0 35 0 0 90
Height (highest) trees (m) 9 12 12 10 12
Height lowest trees (m) 8 8 8 6 10
Height (highest) shrubs (m) 2 2 4 2 4
Height lowest shrubs (m) 1 0,5 2 0,5 1
Char. and diff. species of the ass.
Ostrya carpinifolia 5 2b 2a 5 5
Sesleria autumnalis . 2a 1 4 2b
Viola odorata 1 2b 1 + +
Ornithogalum pyrenaicum + + . + +
Geum urbanum 1 2a . . +
Poa trivialis subsp. sylvicola + . + . .
Other species
Dactylis glomerata subsp. glomerata + 2a + + +
Fraxinus ornus 2a 3 2b 2a 2b
Geranium purpureum + . + + +
Quercus trojana + 3 2b . +
Acer monspessulanum + + . . .
Cornus sanguinea . . + . +
Fragaria vesca . . 2a . +
Hieracium piloselloides . . . + +
Melica arrecta . + + . .
Milium effusum . . + . +
Mycelis muralis . . + . +
Poa nemoralis + 2a . . .
Teucrium chamaedrys . . + + .
Quercus cerris + . . . .
Agrimonia eupatoria . + . . .
Aremonia agrimonoides . . + . .
Aristolochia pallida . . 2a . .
Brachypodium sylvaticum . . 1 . .
Campanula rapunculus + . . . .
Carex spicata + . . . .
Carpinus orientalis . . 2a . .
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Hacquetia 14/2 • 2015, 129–174
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Table 34: Junco-Fraxinetum angustifoliae Kárpáti I. & Kárpáti V. 1961. Species with phi-value above 0.50 (only species
with a statistically significant affinity to a cluster according to Fisher’s exact test with P <0.001 are included) are in bold.
Table 34: Junco-Fraxinetum angustifoliae Kárpáti I. & Kárpáti V. 1961. Krepko so označene vrste z phi>0,50
(vključene so samo vrste, ki so statistično značilno navezane na klaster glede na Fisherjev eksaktni test, p <0,001).
Relevé number 76 77 168 193 197 204 211 238
Relevé area (m2
) 80 70 80 90 90 110 80 100
Cover tree layer (%) 90 95 75 95 95 80 90 95
Cover shrub layer (%) 15 3 0 0 0 5 5 0
Cover herb layer (%) 70 15 40 10 10 60 60 95
Height (highest) trees (m) 14 17 10 11 11 15 15 13
Height lowest trees (m) 12 13 4 5 5 4 13 4
Height (highest) shrubs (m) 2 2 0 0 0 3 6 2
Height lowest shrubs (m) 1 1 0 0 0 1 4 1
Char. and diff. species of the ass.
Fraxinus angustifolia subsp. oxycarpa 4 5 5 5 5 5 5 5
Juncus acutus + + 2b . + + + 4
Iris pseudacorus + + . + + 1 1 +
Carex remota 2b + . 1 + . . .
Oenanthe fistulosa . . . . . + + +
Other species
Agrostis stolonifera + + . . . 3 3 2b
Carex otrubae . + 2b + + . . +
Mentha aquatica . + + . . + + .
Samolus valerandi 1 + . . . + + .
Alnus glutinosa 1 . . . . + + .
Oenanthe pimpinelloides + . + . . . . +
Ulmus minor . + + . . . . +
Aegopodium podagraria . . . . . 2a 2a .
Relevé number 151 152 207 209 210
Carya species . . + . .
Clematis vitalba + . . . .
Clinopodium vulgare . 2a . . .
Cornus mas . + . . .
Cornus sanguinea . . . . +
Geranium columbinum . . + . .
Hedera helix . . . . +
Hieracium murorum . . . . +
Platanthera bifolia . . . . +
Potentilla recta . + . . .
Quercus petraea . . . + .
Ranunculus bulbosus . . + . .
Rubus hirtus + . . . .
Satureja montana . + . . .
Stachys officinalis . + . . .
Stachys scardica . . . + .
Thalictrum aquilegifolium . . . + .
Torilis arvensis subsp. arvensis . . + . .
Trifolium ochroleucon . + . . .
Trifolium pignantii . . . + .
Veronica spicata . . 2a . .
Vincetoxicum hirundinaria + . . . .
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Relevé number 76 77 168 193 197 204 211 238
Callitriche stagnalis . . . . . + + .
Circaea lutetiana . . . + + . . .
Cornus sanguinea . . . 2a 2b . . .
Crataegus monogyna . . . 2a 2a . . .
Cynanchum acutum . . . + + . . .
Galium species . . . . . + + .
Lycopus europaeus . . + . . . . +
Lysimachia nummularia . . . + + . . .
Periploca graeca . . . + + . . .
Polygonum hydropiper . . . + + . . .
Populus alba + . . + . . . .
Potentilla reptans . . . + + . . .
Ranunculus flammula . . . . . + + .
Rubus ulmifolius 2b + . . . . . .
Ulmus minor . . . . . + + .
Veronica anagallis-aquatica . . . . . + + .
Acer campestre . . . . 1 . . .
Ajuga reptans + . . . . . . .
Amorpha fruticosa . . . . . . . +
Apium nodiflorum + . . . . . . .
Berberis vulgaris . . . + . . . .
Bidens tripartita . + . . . . . .
Blackstonia perfoliata + . . . . . . .
Carex distans . . . . . . . +
Carex sylvatica . . . . + . . .
Carpinus betulus . . . . + . . .
Cornus sanguinea + . . + . . . .
Crataegus monogyna . . . . + . . .
Eupatorium cannabinum + . . . . . . .
Fraxinus ornus . . . . . + . .
Galium elongatum . . . . . . . +
Galium rotundifolium . + . . . . . .
Inula viscosa . . . . . . . +
Juncus bufonius + . . . . . . .
Ligustrum vulgare . . . . + . . .
Limonium vulgare subsp. serotinum . . + . . . . .
Lotus species . . . . . . . +
Periploca graeca . + . . . . . .
Polygonum hydropiper + . . . . . . .
Plantago major . . . . . . . +
Prunella vulgaris . + . . . . . .
Prunus spinosa . . . + . . . .
Pulicaria dysenterica . . . . . . . +
Ranunculus arvensis . . + . . . . .
Rubus ulmifolius . . + . . + 1 .
Sparganium erectum . . . + . . . .
Tamarix africana . . 3 . . . . .
Tamus communis . . . . . . . +
Ulmus minor 2b 1 + . . . . .
Vincetoxicum hirundinaria . . . . . . . +
Viola reichenbachiana . . . . + . . .
Vitex agnus-castus . . + . . . . .
Alisma plantago-aquatica . + + . . 2a 2a +
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Hacquetia 14/2 • 2015, 129–174
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Table 35: Populetum albae balcanicum Kárpáti I. & Kárpáti
V. 1961 nom. illeg. Species with phi-value above
0.50 (only species with a statistically significant affinity
to a cluster according to Fisher’s exact test with P <0.001
are included) are in bold.
Tabela 35: Populetum albae balcanicum Kárpáti I. &
Kárpáti V. 1961 nom. illeg. Krepko so označene vrste
z phi>0,50 (vključene so samo vrste, ki so statistično
značilno navezane na klaster glede na Fisherjev eksaktni
test, p <0,001).
Relevé number 56 194 202 203
Relevé area (m2
) 100 90 90 95
Cover tree layer (%) 100 100 80 80
Cover shrub layer (%) 3 0 10 10
Cover herb layer (%) 7 25 15 20
Height (highest) trees (m) 22 35 22 25
Height lowest trees (m) 0 7 12 6
Height (highest) shrubs (m) 25 0 4 3
Height lowest shrubs (m) 1 1,5 1 1
Char. and diff. species of the ass.
Populus alba 5 5 5 5
Pulicaria odora . + 1 +
Aegopodium podagraria 1 . + 2a
Other species
Fraxinus angustifolia subsp. oxycarpa 4 + + +
Ulmus minor + + + .
Rubus ulmifolius + + 1 2a
Hedera helix + + 2a .
Ficus carica + . + .
Arum italicum + + . .
Humulus lupulus 1 . + .
Agrostis stolonifera . . 2a 1
Alnus glutinosa . . + 2b
Carex otrubae . . + 2a
Periploca graeca + . . .
Carduus acanthoides 2m . . +
Amorpha fruticosa + + . .
Fallopia convolvulus . + + .
Ligustrum vulgare + + . .
Rosa sempervirens + . . +
Samolus valerandi . . + +
Viola odorata . + . 1
Acer campestre . + . .
Alisma plantago-aquatica . . + .
Bidens tripartita . . . +
Carex sylvatica . + . .
Carpinus betulus . + . .
Carpinus orientalis . + . .
Cornus mas . + . .
Crataegus monogyna 1 . . .
Cynanchum acutum . + . .
Eupatorium cannabinum . . + .
Geum urbanum . + . .
Relevé number 56 194 202 203
Humulus lupulus 1 . . .
Oenanthe pimpinelloides . . . +
Primula vulgaris . + . .
Punica granatum + . . .
Quercus robur . + . .
Rubus caesius . + . .
Rumex sanguineus . . . +
Ruscus aculeatus . 2a . .
Viola reichenbachiana . + . .
Table 36: Querco roboris-Carpinetum betuli submediterraneum
Bertović ex Horvat 1974 nom. illeg. Species
with phi-value above 0.50 (only species with a statistically
significant affinity to a cluster according to Fisher’s
exact test with P <0.001 are included) are in bold.
Tabela 36: Querco roboris-Carpinetum betuli submediterraneum
Bertović ex Horvat 1974 nom. illeg. Krepko
so označene vrste z phi>0,50 (vključene so samo vrste,
ki so statistično značilno navezane na klaster glede na
Fisherjev eksaktni test, p <0,001).
Relevé number 51 191 195 196
Relevé area (m2
) 150 60 120 110
Cover tree layer (%) 80 100 100 100
Cover shrub layer (%) 10 20 2 2
Cover herb layer (%) 60 75 70 70
Height (highest) trees (m) 20 18 18 30
Height lowest trees (m) 12 5 8 6
Height (highest) shrubs (m) 3 2 3,5 4
Char. and diff. species of the ass.
Quercus robur 4 5 3 5
Viola reichenbachiana + + + 1
Carpinus betulus 1 . 3 2a
Arum italicum + + + +
Aristolochia pallida . . + +
Other species
Clematis viticella . + + .
Carpinus orientalis 4 2a 3 3
Hedera helix + + 2a 2b
Ruscus aculeatus 4 3 3 3
Acer campestre + . + +
Cornus mas + 2b . +
Crataegus monogyna 1 + + .
Ligustrum vulgare . 1 + +
Primula vulgaris . + 1 +
Viola odorata + . + +
Ajuga reptans . . + +
Asparagus acutifolius . + + .
Carex sylvatica . + . +
Fallopia convolvulus . + . +
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Giuliano Fanelli et al.: The vegetation of the Buna river protected landscape (Albania)
171
Relevé number 51 191 195 196
Rubus ulmifolius . + . +
Acer monspessulanum . + . .
Fraxinus angustifolia subsp. oxycarpa + . . .
Populus alba 2b . . .
Ulmus minor . . . 1
Aristolochia lutea + . . .
Brachypodium sylvaticum . . + .
Relevé number 51 191 195 196
Carex depauperata . . + .
Carex distans + . . .
Carpinus orientalis 4 . . .
Clematis vitalba . . . +
Dactylis glomerata . . + .
Geranium purpureum . . . +
Tamus communis . . . +
Table 37: Periploco graecae-Alnetum glutinosae Rudski in Horvat et al. ex Fanelli, De Sanctis, Gjeta, Mullaj, Attore
ass. nova hoc loco. Species with phi-value above 0.50 (only species with a statistically significant affinity to a cluster
according to Fisher’s exact test with P <0.001 are included) are in bold.
Tabela 37: Periploco graecae-Alnetum glutinosae Rudski in Horvat et al. ex Fanelli, De Sanctis, Gjeta, Mullaj, Attore
ass. nova hoc loco. Krepko so označene vrste z phi>0,50 (vključene so samo vrste, ki so statistično značilno
navezane na klaster glede na Fisherjev eksaktni test, p <0,001).
Relevé number 52 54 55* 200 201 230 231 232 241
Relevé area (m2
) 50 50 60 105 110 150 90 120 95
Cover tree layer (%) 100 95 98 85 95 95 95 95 95
Cover shrub layer (%) 15 10 15 60 40 10 10 2 2
Cover herb layer (%) 1 35 5 5 15 50 35 3 15
Height (highest) trees (m) 8 12 12 12 10 12 12 13 13
Height lowest trees (m) 4 3 6 10 8 9 9 8 10
Height (highest) shrubs (m) 3 3 2 3 5 2 2 0 0
Height lowest shrubs (m) 1,5 2 1 1 1 1,5 1,5 0,8 1,3
Char. and diff. species of the ass.
Alnus glutinosa 5 5 5 5 5 5 5 5 5
Periploca graeca + + + + 2m + 1 . .
Circaea lutetiana . + . + . + + . .
Humulus lupulus 2m 2m + + + . . . +
Cornus sanguinea . 2a 2m + 2a + + . .
Ficus carica . 1 + + + . . . 2a
Cynanchum acutum + + . . + . . . +
Other species
Rubus ulmifolius 2b 2a + 3 . 2a 2a + +
Fraxinus angustifolia subsp. oxycarpa . + + + . + 2a 2a .
Ulmus minor + + + . + . . + 2a
Bidens tripartita + . . + 1 1 2a . .
Hedera helix . 2a 1 + 2a . . . +
Ligustrum vulgare . + + 2a 2a . . . +
Iris pseudacorus . . . + . + + + .
Crataegus monogyna . 2a + + 2a . . . .
Lycopus europaeus . + . . + 1 + . .
Quercus robur . + + . + . . . .
Mentha aquatica . . . . . + + . +
Oenanthe pimpinelloides . . . . + + 2a . .
Rumex sanguineus . . . . . + + + .
Carex remota . . . . . . . + 2a
Agrostis stolonifera . . . . 1 . + . .
Alisma plantago-aquatica . . . . . + . + .
Amorpha fruticosa 1 . . + . . . . .
Apium nodiflorum . . . . . + 1 . .
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Hacquetia 14/2 • 2015, 129–174
172
Relevé number 52 54 55* 200 201 230 231 232 241
Carex spicata . . . . . + + . .
Carex distans . + . . + . . . .
Carex otrubae . . . . . 3 . + .
Carex sp. . . . + . . + . .
Fallopia convolvulus . . . + + . . . .
Galium elongatum . . . . . + + . .
Polygonum hydropiper . . . . . + + . .
Salix alba . + . + . . . . .
Vitis vinifera . . . . . + + . .
Arum italicum . . . . + . . . .
Carduus acanthoides . . + . . . . . .
Carex sylvatica . . . . . . . . +
Clematis vitalba . + . . . . . . .
Clematis viticella . . . . 1 . . . .
Cornus mas . . . . . . . . +
Cynanchum acutum . . . . + . . . .
Galium debile . . . . + . . . .
Geum urbanum . 2a . . . . . . .
Buglossoides purpurocaerulea . + . . . . . . .
Lysimachia nummularia . . . . . + . . .
Poa nemoralis . . . . . + . . .
Potentilla reptans . . . . . . + . .
Prunella vulgaris . . . . . + . . .
Rosa sempervirens . . . . + . . . .
Samolus valerandi . . . . . . . + .
Vincetoxicum canescens . . . . . + . . .
Vincetoxicum hirundinaria . . . . . . + . .
Viola reichenbachiana . + . . . . . . .
Table 38 (Tabela 38): Vitici-Tamaricetum africanae
Horvatić 1960.
Relevé number 23 40 104 112
Relevé area (m2
) 80 15 25 30
Cover shrub layer (%) 75 95 90 75
Cover herb layer (%) 75 2 0 45
Height (highest) shrubs (m) 5 3 5 3
Height lowest shrubs (m) 2 0 0 2
Char. and diff. species of the ass.
Tamarix africana 4 5 5 4
Vitex agnus-chastus + . . +
Atriplex prostrata subsp. latifolia + + . +
Other species
Aster tripolium + . . .
Juncus acutus 4 . . +
Phragmites australis + . + .
Cladonia portentosa + . . .
Elymus athericus + . . .
Limonium vulgare subsp. serotinum . + . 3
Salsola soda . . . +
Polypogon monspeliensis . . . +
Table 39 (Tabela 39): Bromo villosi-Haynaldietum
­villosae Pignatti 1953.
Relevé number 49 50
Relevé area (m2
) 8 3
Altitude (m a.s.l.) 330 Slope
(degrees) 5 Cover
herb layer (%) 60 80
Char. and diff. species of the ass.
Dasypyrum villosum 1 4
Bromus madritensis + +
Vulpia membranacea + +
Vulpia myuros 2a +
Kochia prostrata + +
Aegilops triuncialis 1 +
Relevé number 23 40 104 112
Crypsis aculeata . . . 1
Suaeda species . . . +
Elymus farctus subsp. farctus . . . +
Bolboschoenus maritimus . . . +
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Giuliano Fanelli et al.: The vegetation of the Buna river protected landscape (Albania)
173
Table 40 (Tabela 40): Centaureetum calcitrapae Fanelli
& Menegoni 1997.
Relevé number 24 25 490
Relevé area (m2
) 80 0 10
Altitude (m a.s.l.) - - Cover
herb layer (%) 100 100 90
Char. and diff. species of the ass.
Centaurea calcitrapa 5 . 4
Scolymus hispanicus 2a 5 1
Rumex pulcher + . +
Other species
Bromus hordeaceus + + .
Avena barbata . + +
Anthemis austriaca 2m . +
Filago germanica 2m . +
Matricaria disciformis 2m . +
Vicia villosa subsp. eriocarpa . + +
Cichorium intybus . + .
Carduus acanthoides + . .
Vulpia ciliata + . +
Hordeum hystrix + . .
Cynodon dactylon + . .
Trifolium campestre . . +
Trifolium scabrum . . +
Agrostis stolonifera . . +
Relevé number 49 50
Other species
Medicago minima 1 2a
Lagurus ovatus + 1
Lepidium graminifolium . +
Sisymbrium officinale . +
Carduus pycnocephalus . +
Vicia villosa subsp. eriocarpa + .
Avena barbata + .
Trifolium scabrum 1 1
Daucus carota 2b .
Lolium perenne + +
Briza maxima + +
Bromus rigidus + .
Bromus tectorum + .
Salsola soda . +
Verbascum banaticum . +
Scolymus hispanicus . +
Bromus hordeaceus . +
Cynodon dactylon . +
Table 41: Asphodelus ramosus community.
Tabela 41: Združba z vrsto Asphodelus ramosus.
Relevé number 15 70 492
Relevé area (m2
) 5 2 8
Altitude (m a.s.l.) - 125 50
Aspect (degrees) 40 60 50
Slope (degrees) 3 10 5
Cover herb layer (%) 90 70 60
Cover bare rock (%) 0 35 0
Cover stones (%) 40 70 40
Char. and diff. species of the ass.
Asphodelus ramosus 4 3 3
Carthamus lanatus + + .
Catapodium rigidum + + +
Nigella arvensis . + +
Other species
Teucrium polium subsp. capitatum 2b 2b 2a
Sideritis romana + + +
Petrorhagia saxifraga subsp. saxifraga 1 + +
Crepis zacintha + + .
Arenaria leptoclados 2m . 1
Sedum hispanicum + . +
Bupleurum veronense 1 + .
Medicago praecox + . +
Trifolium scabrum 1 . +
Daucus aureus 1 . +
Centaurea solstitialis 1 . 1
Crepis foetida + . +
Aegilops geniculata + . .
Avena barbata + . +
Filago gallica + . +
Melica ciliata . + +
Brachypodium distachyon . + +
Dasypyrum villosum . 1 +
Carex distachya . + +
Micromeria juliana . + +
Asperula aristata subsp. condensata 1 . .
Anthemis austriaca + . .
Carlina corymbosa + . .
Rostraria cristata + . .
Bothriochloa ischaemum + . .
Medicago minima . + .
Trifolium campestre . + .
Eryngium amethystinum . + .
Satureja montana . + .
Helichrysum italicum subsp. italicum . + .
Dactylis glomerata subsp. glomerata . + .
Gastridium ventricosum . + .
Chrysopogon gryllus . + .
Sanguisorba minor subsp. muricata . + .
Plantago bellardii . + .
Carya cordiformis . . +
Sanguisorba minor subsp. muricata . . +
Trifolium campestre . . +
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Hacquetia 14/2 • 2015, 129–174
174
Table 42 (Tabela 42): Carduetum acanthoidis
Felföldy 1942.
Relevé number 10 14
Relevé area (m2
) 15 20
Cover herb layer (%) 100 100
Char. and diff. species of the ass.
Carduus acanthoides 5 4
Cirsium italicum + 2b
Cirsium vulgare + +
Other species
Cirsium vulgare + +
Galega officinalis + +
Medicago lupulina + +
Rumex conglomeratus + +
Convolvulus arvensis + +
Agrostis stolonifera + +
Potentilla reptans + .
Euphorbia platyphyllos + .
Daucus carota + .
Galium elongatum + .
Scolymus hispanicus + .
Phragmites australis + .
Satureja calamintha + .
Paspalum paspalodes + .
Atriplex patula . +
Prunella vulgaris . +
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