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5727
ANALYSIS OF THE FLORISTIC COMPOSITION OF
MOUNTAIN BEECH FORESTS ON THE TERRITORY OF
SERBIA AND MONTENEGRO
Rajko Milosevic1,*
, Milic Curovic2
, Marijana Novakovic-Vukovic1
1
University of Belgrade, Faculty of Forestry, Belgrade, Serbia
2
University of Montenegro, Biotechnical Faculty, Podgorica, Montenegro
ABSTRACT
The mountain beech forest represents a vast
climate-regional belt of vegetation in Serbia, as
well as the most economically important forest
type, and takes up a significant area in Serbia and
Montenegro. This paper focused on a comparison
of the floristic composition of a typical mountain
beech forest in Serbia and Montenegro in order to
determine the differences between the investigated
stands. A total of 12 phytocoenological relevés
were analyzed; 7 on Jastrebac (Serbia) and 5 on
Biogradska Gora (Montenegro). CCA analysis
showed that significant differences are present
among the studied stands. Phytocoenological
relevés from Montenegro correlated with altitude
and assembly, while relevés from Serbia have a
correlation with slopes. Differences in floristic
composition are mostly due to various tree layering.
Stands on Mt. Jastrebac are dispersed, which allow
the penetration of light on the terrestrial flora, leading
to weed growth. The composition of tree layers
is quite dense on Biogradska Gora, thus the shrub
layer is poor, whereas terrestrial flora is mainly
comprised of species that thrive in areas of little
light (sciophytes). In the spectrum of floral elements,
species of Central European areal type dominate
both sites, which expressed extreme mesophilic
characteristics of researched stands. In the
spectrum of life forms on Mt. Jastrebac, phanerophytes
are dominant due to the large number of
species in the tree and shrub layer, while geophytes
are dominant on Biogradska Gora as a result of
dense tree layering, therefore, the ground flora consists
of species that can complete the reproductive
cycle before trees begin to bloom.
KEYWORDS:
beech forest, floristic composition, Serbia, Montenegro
INTRODUCTION
Beech forests cover large areas in South Eastern
Europe and are on different geological substrates,
soils, altitudes, grow in various macro and
micro-climate conditions, thus they are very heterogeneous.
There are many classifications of beech
forests of Southeastern Europe. Horvat [1] first
ranked all beech forests, ranging from the Alps to
Albania and Greece, in alliance Fagion sylvaticae
illyricum. However, within such a wide distribution,
significant floristic differences exist within the
researched stands, thus Soó [2, 3] ranked forests of
Southeastern Serbia, Bulgaria and northern Greece
in alliance Fagion dacicum, which were previously
examined in the Carpathian region. Given that
many diagnostic species of the alliance were not
found in the mentioned areas, the author dubbed
these forests ‘Moesian Beech Forests’ (MoesischeFagionWälder),
marking Moesian beech (Fagus
moesiaca) as a differential type of alliance [4]. All
beech forests of Southeastern Europe have recently
been classified into two alliances [5]: AremonioFagion
and Fagion moesiacae. Pure and mixed
beech forests occupy significant areas in Serbia
because beech (Fagus sylvatica L. ssp. moesiaca) is
the most widespread tree species in Serbia; beech
forests cover 660,400 ha (29.3%) of the total covered
area, of which 64.7% is state-owned [6]. In
syntaxonomic sense, beech forests in Serbia are
classified into class Querco- Fagetea Br.-Bl.
Vlieger & 1937, order Fagetalia silvaticae Pawlowski
and Pawlowski, Sokolowski&Walisch 1928
and alliance Fagion moesiacae Blečić & R. Lakušić
1976. The affiliation of beech forest in the alliance
Fagion moesiacae was confirmed on the area of
Biogradska Gora in Montenegro [7] even though
Illyrian beech forests Fagetum montanum sylvaticae
montenegrinum Blečić 1958 were originally
described [8] on the area of Piva (Montenegro).
Within this alliance, due to a great variety, seven
suballiances and a large number of associations
were isolated where mountain beech forest has the
largest distribution and is economically the most
important type of beech forest, and forests in Serbia
in general [9]. Mountain beech forest (Asperulo
odoratae-Fagetum moesiacae B. Jovanović 1973,
syn. Fagetum moesiacae montanum B. Jovanović
1973) presents a very well defined climate-regional
belt of vegetation in Moesia and transitional Illyrian-Moesia
Provinces. The community is mainly
located on all exposures, at altitudes between 800-
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1200 m, and is observed on most mountains of the
mentioned areas. The aim of this research was to
determine whether there are differences in the floristic
composition of a typical mountain forests of
beech in two localities in Serbia and Montenegro,
given that it amounts to a considerable distance,
and investigated sites with specific characteristics
in terms of severity synergistic environmental impacts
in which these two communities grow.
MATERIALS AND METHODS
Great Jastrebac mountain range stretches between
18 ° 59 'and 19 ° 05' east longitude and 43 °
24 'and 43 ° 28' north latitude. It belongs to the
Rhodope mountain system, and in morphological
terms, belongs to the category of high and medium
mountains with very expressed relief, locatedat
altitudes between 300-1394 m (Figure 1). The average
annual temperature is 9. 7О
С, the average temperature
during the vegetation period is 15.8 О
С.
Geologically speaking, the area of Great Jastrebac
consists of the highest percentage of granodiorite in
its base. Corneits are present on the peaks of the
mountain range, and phyllites at somewhat lower
altitudes ranging from 950-1000 meters. Acidic
brown soils prevail (dystric cambisol), typical acid
brown soil, poor skeletal and typical acid brown
soil of medium depth and skeletal. Podzolic acid
brown soil is less common. Humus-silicate soils are
present at higher altitudes: regosol, colluvial and
podzolic [10]. Biogradska Gora, along with Mt.
Bjelasnica, represent the unique geomorphological
unit in Central Montenegro (Figure 1) with peaks
over 2000 m, where glacier peaks and cirques with
several glacial lakes are present. The average annual
temperature is around 20
C, while the mean air
temperature in the vegetation period is around 60
C
(mountainous areas) to 120
C (lower parts-valley of
Tara). The dominant soil type is acid brown soildystric
cambisol on volcanic surfaces. The geological
foundation consists of various volcanic rocks
that have effused in this area during volcanic activity
in the Middle Triassic and Jurassic period [11].
DATA COLLECTION
For the analysis of floristic composition of the
researched area, 12 phytocoenological relevés made
using the standard method Braun-Blanquet [12]
were done and were taken from literature [10,11]; 7
relevés on Jastrebac and 5 on Biogradska Gora. The
stands at both sites, according to syntaxonomic
origin, were classified as typical mountain beech
forest (Asperulo odoratae-Fagetum moesiacae B.
Jovanović 1973, syn. Fagetum moesiacae montanum
B. Jovanović 1973 subass. typicum).
Statistical analysis. First, the transformation
estimates of abundance and coverage for each species
within the phytocoenological relevés were performed
by the Van Der Maarel method [13]. CCA
analysis of ecological-vegetation data was performed
with the help of statistical software
CANOCO 4.5 [14]. Monte Carlo permutation test
was used to determine the significance of worth
(eigenvalue) with appropriate CCA axes. The study
calculated 499 randomly selected permutation of
the described communities. Syntax names were
given according to [15].
FIGURE 1
Geological location of researched sites
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The spectrum of floral elements were produced
on the basis of systematization of plantgeographic
elements after [16], and the life-form
spectrum after [17]. Indices of diversity and evenness
(Shannon-Wiener diversity index and evenness)
were calculated using the program JUICE 7.0
[18]. The Jaccard index of similarity between the
investigated communities was also calculated [19]
according to formula:
where
UV- is the number of common species, U, Vis
the number of species in the beech and fir forest
on Mt. Jastrebac and Mt. Biogradska Gora, respec-
tively.
RESULTS
Statistical analysis. Stands of typical mountain
beech forests (Asperulo odoratae-Fagetum
moesiacae B. Jovanović 1973, syn. Fagetum
moesiacae montanum subass. typicum) on Jastrebac
(Serbia) appear at altitudes between 700-960 m, on
slopes of 30-35 degrees, and exposure are mostly
hidden- N, NW, NE. The stands at Biogradska Gora
(Montenegro) are at slightly higher altitudes, between
1150 to 1270 m, on slopes of 20-25 degrees
and different exposure-W, NW, S, SW. The graph
(Fig. 2) shows that the relevés from Serbia accumulate
on the right side, while relevés from Montenegro
are on the left side of the chart, which shows
that there are significant differences among them,
although it is the same association and subassocia-
tions.
FIGURE 2
CCA ordination triplot, fit range for species 20-40%, 44 species
(○-relevé representation, Δ-species representation;(1-7) – phytocoenological relevés in the mountain beech forest on Mt.
Jastrebac (Serbia); (8-12) –phytocoenological relevés of the mountain beech forest on Mt. Biogradska Gora (Montenegro))
Species abbreviations: Ace hel-Acer heldreichii; Fra exc-Fraxinus excelsior; Dap mez-Daphne mezereum; Pru avi-Prunus
avium; Sen nem-Senecio nemorensis; San eur-Sanicula europaea; Fag moe-Fagus moesiaca; Ath fil-Athyrium filix-femina;
Oxa ace-Oxalis acetosella; Ace pse-Acer pseudoplatanus; Lun red-Lunaria rediviva; Pol acy-Polystichum aculeatum; Dry
fil-Dryopteris fili-mas; Sam nig-Sambucus nigra; Hed hel-Hedera helix; Rus hyp-Ruscus hypoglossum; Rub hir-Rubus
hirtus; Sym tub-Symphytum tuberosum; Pol ver-Polygonatum verticillatum; Car bul-Cardamine bulbifera; Abi alb-Abies
alba; Vio syl-Viola sylvestris; Epi mon-Epilobium montanum; Asp odo-Asperula odorata; Lam gal-Lamium galeobdolon;
Pre pur-Prenanthes purpurea; Mer per-Mercurialis perennis; Myc mur-Mycelis muralis; Ane nem-Anemone nemorosa; Cal
off-Calamintha officinalis; Ace pla-Acer platanoides; Ulm gla-Ulmus glabra; Asa eur-Asarum europaeum; (abbreviation
after a species name denotes A- tree layer, B –shrub layer, C-ground flora layer)
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Phytocoenological relevés from Montenegro
correlated with altitude and assembly, while relevés
from Serbia have a correlation with slopes. Exposure
is the variable that has the least impact on the
community, which isn’t a surprise due to the fact
that mountainous beech forests are located in all
aspects within its areal. It is evident that mostly
woody and shrub species are isolated when it comes
to relevés from Serbia: Abies alba, Fraxinus excelsior,
Prunus avium, Ulmus glabra, Fagus moesiaca
and others. Due to more open tree layering (0.5-0.7)
greater light penetration on the ground flora layer
occurred, which led to weed growth and increase
inquantity and coverage of blackberry (Rubus
hirtus), which prevents the development of other
species in this layer. On the other hand, a thick
strand of tree layers in stands in Montenegro hinders
the development of shrub species, however
various sciophyt and semiscyophyte species are
present in the ground flora layer: Asperula odorata,
Mercurialis perennis, Cardamine bulbifera, Mycelis
muralis, Epilobium montanum etc. Common
to both sites is the absolute dominance of the beech
trees in the tree and shrub layer, while the ground
flora layer varies significantly as a result, primarily
to, a different set of tree layers.
Jaccard's similarity index (0.38) shows that
among the study stands there is not a great floristic
similarity, even though it is the same subassocia-
tion.
The spectrum of floral elements. In both researched
sites, the spectrum of floral elements (Fig.
3A) absolutely dominated species belonging to
Central European floral element, which is an indicator
that the community develops in mesophilic
conditions of temperate zone. Other floral elements
have a similar representation, apart from the elements
of northern regions, which in Serbia amounts
to 5%, while in Montenegro there are no records.
The spectrum of life forms. The spectrum of
life forms (Fig 3B) reveals important differences
between the mountain beech stands in Serbia and
Montenegro. On the territory of Serbia, phanerophytes
are significantly more numerous (41%:
25%) which is result of a large number of species of
tree layers and shrubs. On the other hand, geophytes
(early flourishing species) are more numerous
in stands in Montenegro, nearly two fold compared
to Serbia (39%: 18%). This is due to the
dense layer of trees (0.9-1.0), where terrestrial
plants have insufficient sunlight during the growing
season, and therefore, have to complete their reproductive
cycle before trees bloom.
Indices of diversity and evenness. The community
of mountain beech on Biogradska Gora
(Table 1) has a slightly higher diversity index
(Shannon-Wiener index) when it comes to ground
flora, compared to Jastrebac.However this value is
not statistically significant. Evenness index (Evenness
index) has approximately the same value at
both sites. Blackberry (Rubus hirtus) can be labeled
as a speciesthat reduces the evenness index value
on Jastrebac, whereas Asperula odorata and Lamium
galeobdolon do the same on Biogradska Gora.
FIGURE 3
Spectrum of floral elements and spectrum of life forms in the mountain beech forests
in Serbia and Montenegro
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TABLE 1
Indices of diversity and evenness in researched
stands
Locality
Average
Shannon-Wiener
index
Average
Evenness
index
Jastrebac (SRB) 1.42 0,54
Biogradska Gora
(MNE)
1,76 0.60
DISCUSSION
At both researched sites (Jastrebac and Biogradska
Gora) fir (Abies alba) was found in the
shrub and ground flora layer. The presence of
young fir is evidence that said community can not
be regarded as a pure beech forest, but as a degraded
deciduous-coniferous forest Abieti-Fagetum. On
both mountain ranges forests of beech and fir were
observed in the vicinity of pure beech forests,
which confirms what was previously said, because
over time man cut more fir than beech, thus beech,
being biologically more resilient, prevailed. The
origin of the secondary forests of mountain beech
on some mountain ranges in Serbia was discussed
in previous research [9, 15, 20, 21], as well as in
Bosnia [22], on the Dinarides [23] and others.
Mountain beech forests occupies large areas
on the Balkans. In the area of Galicica (Macedonia),
the mountain beech forest (Calamintha grandiflorae-Fagetum
(Em 1965) Rizovski& Jack's ex
Matevski et al. Ass. Nova)) is present on altitudes
between 1300-1700 m. This forest is of vast economical
importance and has a lot of common species
with the researched sites in terms of floristic
composition. Dryopteris filix-mas (observed on
Jastrebac and Biogradska Gora), Neottia nidus-avis
(recorded on Biogradska Gora) and Pulmonaria
officinalis (observed on Jastrebac) are all species
labeled as diagnostics [24]. The mountain beech
forest in Macedonia belongs to the alliance Doronico
columnae-Fagion Dzwonki et al.1999, Macedonian
fir (Abies borisii-regis) is a diagnostic type of
alliance.
In the area of Mt. Jadovnik (BiH), the community
Fagetum montanum illyricum Fuk. et Stef
1958. was described, with altitudes ranging from
700m (800m) -1100m (1200) m, on limestones and
dolomites [25]. Cardamine bulbifera, Cardamine
enneaplhyllos, Paris quadrifolia, Polystichum aculeatum,
Geranium robertianum and Oxalis acetosella
are differential types of the subassociation
cardaminetosum (typicum). All of these species
were recorded on the investigated localities Jastrebac
and Biogradska Gora. The author also concludes
that the forests of beech and beech-fir in
Jadovnik have great sociological similarities, 55%
of species are common to both characteristic sets,
spatially very close, grow at the same exposures,
slopes and soils, which supports the conclusion that
the forests of secondary character beech, with anthropological
origins, caused the degradation of
beech-fir forests.
Mountain beech forests (Fagetum montanum
illyricum subass. typicum) in Manjaca (BiH) also
have great floristic similarities with mountain beech
forests in Serbia and Montenegro. It was recorded
at altitudes ranging from 920-1000 m. Fir and
spruce [26] were introduced to this community via
over-seeding. Due to a dense tree layer, ground
flora is impoverished. Typical "fagetal" species
represent a small coverage and quantity: Asperula
odorata, Dryopteris filix-mas, Mercurialis perennis,
Festuca drymeia. European beech (Fagus sylvatica)
is an edificatory for this community.
The most common, and the most clearly defined,
Illyrian beech forest in Croatia is the community
Lamia orvalae-Fagetum (I. Horvat 1938)
Borhidi 1963 classified in alliance Aremon-Fagion
(I. Horvat 1938) Török, Podani et Borhidi 1989. It
is found in the Dinaric area, between 400 -800 m
above sea level, below the beech-fir belt [27]. By
floristic composition, this community differs significantly
from the mountain beech forests in the
Moesia province, because it‘s floristic composition
is comprised of a vast number of Illyrian species:
Lamium orvala, Rhamnus fallax, Cardamine enneaphyllos,
Hacquetia epipactis, but also a lot of species
that occur in the mountainous beech forest in
the Moesia Province: Polystichum aculeatum, Lonicera
nigra, Prenanthes purpurea, Senecio
nemorensis, Lilium martagon and others. The
community Lamia orvalae-Fagetum in Slovenia
represents the zonal vegetation mountainous belt at
altitudes between 600-900 m [28]. It is mainly distributed
in limestones and dolomites. In Slovenia,
this community is of similar floristic composition
compared to the mountain forests of beech in the
Moesia Province, as can be seen from the combination
of diagnostic species: Acer pseudoplatanus,
Ulmus glabra, Acer platanoides, Euonymus latifolia,
Sambucus nigra, Athyrium filix-femina, Dentaria
bulbifera, Dryopteris filix-mas Oxalis acetosella,
Paris quadrifolia, etc.
CONCLUSION
Mountain beech forests make up an ecocoenological
and spatially differentiated whole in
terms of developed and formed climate-regional
belt. Typical forest community of mountain beech
(Fagetum moesiacae montanum subass. typicum) is
most widely spread and the most common within
this belt.
There are significant differences in the floristic
composition of the plant community between the
forest communities typical mountain beech
(Fagetum moesiacae montanum subass. typicum)
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5732
on Veliki Jastrebac (Serbia) and Biogradska Gora
(Montenegro), although according to the classification
they belong to the same association and sub-
associations.
Jaccard's similarity index (0.38) shows that
among the studied stands there is great floristic
similarity, even though it is the same subassocia-
tions.
It is evident that within the records from Serbia,
mostly woody and shrub species are isolated:
Abies alba, Fraxinus excelsior, Prunus avium,
Ulmus glabra, Fagus moesiaca etc., while in the
stands in Montenegro on the ground flora many
sciophytes and semi-sciophyte species are present:
Asperula odorata, Mercurialis perennis, Cardamine
bulbifera, Mycelis muralis, Epilobium montanum,
etc. Common to both sites is the absolute
dominance of beech trees in the tree layer.
In the spectrum of floral elements on both
studied sites, dominant species belong to Central
European floral element, which is an indicator that
the community develops in mesophilic conditions
of temperate zone. Other floral elements have a
similar representation, except for the elements of
the northern regions, which in Serbia accounted for
5%, while in Montenegro they are not recorded.
In the spectrum of life forms significant differences
between the mountain beech stands in Serbia
and Montenegro were observed. On the territory
of Serbia, phanerophytes are significantly more
numerous (41%: 25%) as a result of a large number
of species of trees and shrubs. On the other hand, in
stands in Montenegro, geophytes are more numerous,
early flourishing species, which is more than
doubled compared to Serbia (39%: 18%).
The community of mountain beech on Biogradska
Gora has a slightly higher diversity index
(Shannon-Wiener index) of ground flora layer
compared to Jastrebac, but this value is not statistically
significant. Evenness index also has approximately
the same value at both sites. Species that
reduce the value of the evenness index in the area
of in Biogradska Gora are Asperula odorata and
Lamium galeobdolon, whereas in Jastrebac, blackberry
(Rubus hirtus) is known to reduce the evenness
index value.
ACKNOWLEDGEMENTS
This paper was done as a part of scientific project:
Sustainable management of the total forest
potential in Republic of Serbia” EVBR 37008, Ministry
of education and science of Republic of Ser-
bia.
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of Faculty of forestry. Belgrade. 98, 89-106
(in Serbian).
[27]Vukelić, J., Baričević, D. (2002) Recent phytocoenological
findings of beech forests in Croatia.
Sumar list. 9-10, 439-57 (in Croatian).
[28]Marinček, L., Čarni, A. (2002) Commentary to
the vegetation map of forest communities of
Slovenia in a scale of 1:400 000. Biološki inštitut
Jovana Hadžija ZRC SAZU. Ljubljana.
120p.
Received: 31.10.2017
Accepted: 01.05.2019
CORRESPONDING AUTHOR
Rajko Milosevic
Faculty of Forestry,
University of Belgrade,
Kneza Višeslava 1,
11 000 Belgrade – Serbia
e-mail: rajko.milosevic@sfb.bg.ac.rs
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