ПРИЛОЗИ, Одделение за природно-математички и биотехнички науки, МАНУ, том 37, бр. 2, стр. 95–106 (2016)
CONTRIBUTIONS, Section of Natural, Mathematical and Biotechnical Sciences, MASA, Vol. 37, No. 2, pp. 93–104 (2016)
*
The paper was presented as an inaugural lecture by the author on the occasion of his election to be a member of the Macedonian
Academy of Sciences and Arts on 5th
May 2016 in Skopje.
Received: October 4, 2016 ISSN 1857–9027
Accepted: November 6, 2016 e-ISSN 1857–9949
UDC:630*22(497)
DOI: 10.20903/CSNMBS_MASA.2016.37.2.38
Review
VEGETATION OF DECIDUOUS FORESTS IN THE BALKAN PENINSULA*
Andraž Čarni
Institute of Biology, Scientific Research Center of the Slovenian Academy of Sciences and Arts,
Ljubljana, Slovenia
University of Nova Gorica, Nova Gorica, Slovenia
Macedonian Academy of Sciences and Arts, Skopje, Republic of Macedonia
carni@zrc-sazu.si
The article provides an overview of deciduous forests of the Balkan Peninsula. It presents riverine, floodplain
and moor forests, dominated by Alnus incana, A. glutinosa, Fraxinus excelsior, F. angustifolia, Quercus robur, Salix
alba, Ulmus laevis, U. minor; acidophilous forests dominated by Betula pendula, Castanea sativa, Fagus sylvatica,
Quercus petraea; thermophilous forests dominated by Carpinus orientalis, Ostrya carpinifolia, Quercus cerris, Q.
farainetto, Q. petraea, Q. pubescens and mesophilous forests dominated by Carpinus betulus, Fagus sylvatica, F.
orientalis, Fraxinus excelsior, Tilia argentea, T. cordata, T. platyphyllos, Ulmus glabra. At the same time, forests
were classified into the synsystematic framework of the standard Central European system to the level of an alliance.
Keywords: Balkan; forest; vegetation; classification
Nomenclatural source: Euro+Med Plantbase (http://ww2.bgbm.org/EuroPlusMed)
INTRODUCTION
Research into vegetation in southeastern Europe
has a long tradition. The first fundamental
works about vegetation in the region were based on
the so-called physiognomic-ecological approach
and originate from the beginning of the 20th
century,
when Gabriel Beck-Mannagetta published his
work Vegetationsverhältnisse der illyrischen Länder
[1] and Lujo Adamović his work Vegetationsverhälnisse
der Balkanländer [2]. At the beginning
of the 20th
century, a new science – phytosociology
– was established [3]. Soon after its beginning
in Europe, research in the Balkans began
according to this method [4] and the new science
was widely accepted among researchers in the region.
Our presentation is based on the standard
Braun-Blanquet method and its hierarchy; so we
group associations (ending -etum) into alliances
(ending -ion), alliances into orders (ending -etalia)
and those into classes (ending -etea) [5]. As a result
of long and intensive research in the region, during
recent years some important surveys of forest vegetation
of some parts of the region has appeared, e.g.
Slovenia [6], Croatia [7], Serbia [8], Kosovo [9],
Bulgaria [10] and Greece [11, 12]. These and many
other studies have enabled us to make an overview
of forest vegetation of the region.
The large number of publications all over
Europe encouraged a group of authors guided by
Ladislav Mucina to start preparation of an overview
over all vegetation types of the whole of Europe to
the alliance level, from the Canary Islands to the
Urals and from Cyprus to Greenland [13]. It is
based on a list of vegetation classes [14] and on a
preliminary overview of alliances [15]. During the
procedure, literature sources all over Europe have
been checked and many nomenclature corrections
and some revisions of individual syntaxa have been
published, for instance [16, 17].
This paper is intended to give an overview of
deciduous forests of the region on the alliance lev-
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Contributions, Sec. Nat. Math. Biotech. Sci., MASA, 37 (2), 95–106 (2016)
96
el, to synchronize regional classification schemes
with the European one [13], to show the present
state of knowledge and to detect some problems. At
the same time, it will facilitate integration of the
vegetation of deciduous forests into the wider European
framework, into various overviews of forest
vegetation, red lists and similar [18].
DESCRIPTION OF AREA
The Balkan Peninsula has a large altitudinal
diversity, since a great part of the territory has an
altitude of over 1000 m, and provides a large altitudinal
range through a relatively small latitudinal
range of 10o
. At the same time, the Balkan Peninsula
is a highly complex region in terms of geology,
climate and vegetation. The diverse geology is the
most important factor contributing to the diversity
of flora: there are crystalline massifs, such as
Rhodopes and new fold mountains, such as the Dinaric
Alps or the Shar-Pindos mountain range,
composed of limestone [19].
The climatic conditions are parallel its geographical
position, relief and extensive coastline
(Figure 1). Belgrade in the northern part, which is
cut off from the influence of the Mediterranean and
lies open to central and eastern Europe, is dominated
by a continental climate with maximum of precipitation
in May–June. Ljubljana has a mountain
climatic regime, with two peaks of precipitation, in
spring and autumn. Ohrid in the south is also located
in mountains but is influenced by a Mediterranean
climate and an autumn peak can be observed.
Ljubljana has more precipitation but Ohrid and
Belgrade have almost the same amount, although
Belgrade has precipitation during the vegetation
season, that is more propitious for flora and vegetation,
and also for agriculture, whereas in Ohrid the
majority of precipitation falls in the part of the year
in which the vegetation cannot use it.
OBJECT
The potential vegetation of the major part of
the Balkan Peninsula is represented by deciduous
broadleaved forest. Broadleaved deciduous forests
appear in the areas with relatively moist summers
and mild winters. Forests can be divided into two
subgroups: mesic and warm temperate forests. Mesic
temperate forests appear in the humid sub Atlantic
climate and are dominated by mesophilous
deciduous tree species, such as Fagus sylvatica,
Carpinus betulus, Acer pseudoplanatus, Alnus glutinosa
etc. Warm deciduous forests appear in areas
under the influence of continental and Mediterranean
climates, in which summer drought is more pronounced.
These forests are dominated by various
species from the genus Quercus, Ostyra carpinifolia,
Carpinus orientalis etc. [20].
Figure 1. Mean total rainfall (a) and mean monthly
temperatures (b) in the research area. Climatological
information for Ljubljana, Ohrid and Belgrade
(World Meteorological Organization data,
http://www.wmo.int/pages/index_en.html)
CONTACT FORESTS
In the case of increased cooling or increasing
drought, deciduous forests are in contact with coniferous
ones. In cold and humid areas, they are in contact
with forests dominated by Picea abies and Abies
alba, as well as Larix decidua. In drier areas, two
relict coniferous forests appear, Pinus peuce and Pinus
heldreichii, and a hybrid between Abies alba and
Abies cephalonica Abies borisii-regis forests. Deciduous
forest can also be in contact with Pinus syl-
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97
vestris and Pinus nigra but these two species do not
form zonal forests and appear locally in extreme
conditions (e.g. extreme drought or acidity) [21–26].
In areas under the influence of the Mediterranean
climate, deciduous forests are in contact with
evergreen broadleaved forests. A warm climate with
mild winters is characteristic of Mediterranean evergreen
vegetation dominated by evergreen tree species,
such as Quercus ilex, Q. coccifera, and their
substitutes, maquis, garrigues. In some cases, they
can also come in contact with coniferous forests of
Mediterranean mountains [27–29].
HISTORY
One of the main factors determining the present
vegetation is the development of vegetation
during and after the ice ages (Figure 2). This area
was covered with no or relatively little ice during
the last glaciation and the assumed July temperature
was only 5o
cooler than present, which probably
provided a suitable environment for the survival
of temperate tree taxa. Since most water was bound
in ice, the sea level was lower and half of the Adriatic
Sea was part of the mainland. From the point of
view of phytogeography, the land connection to the
Apennines is also important and caused the two
peninsulas to share some common or closely related
taxa and vegetation types. The area of common
(syn)taxa could also be called the Apennine-Dinaric
window in phytogeography [30–32].
Figure 2. Vegetation cover of the Balkan during ice age
(Würm) (after [33]).
The area was covered by forest, tundra and
steppe and it was one of the refugial areas for tree
species in Europe. Even now, the highest diversity
of tree species in Europe is found in the southern
part of Macedonia, where there are over 130 woody
species [34].
]
Figure 3. Refugial areas of beech (after [35]).
There were macro refugia of mesophilous
forest in the northwestern part of the region (Figure
3). The major refugia of beech forests were in the
northwestern part of the Balkans and beech spread
out from these refugia practically all over Europe.
These refugia were at an altitude between 600 and
800 m, since in the lowlands it was too dry, and at
higher altitudes it was too cold. So called Illyricoid
flora (e.g. Lamium orvala, Omphalodes verna etc.)
(sensu Trinajstić) also survived here, i. e. mesophilous
endemic species that survived in these
mesophilous forests. A distinction has thus been
made between Illyric species – endemics surviving
on extreme habitats and Illyrioid species – mesophilous
endemics [35, 36].
The situation was different with thermophilous
forests; their primary refugia were in the south
but in warmer periods they expanded towards the
north and could sometimes survive colder periods
in those areas, in so called secondary refugia (Fig.
4). Evidence exists that in the southern Balkans
there were macro refugia of warm temperate forests
and also cryptic (smaller) refugia of mesic temperate
forests. The macro refugia were more species
rich but cryptic refugia also had their floristic individuality
[20, 37, 38].
Most research of the history of forests has
been done by people from outside the region and
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Contributions, Sec. Nat. Math. Biotech. Sci., MASA, 37 (2), 95–106 (2016)
98
the focus has been on the location of refugia and
the time needed for temperate trees to migrate from
these areas to northern Europe [39]. The vegetation
history of the Balkans has much greater significance
for the development of forests than this. If
the temperate trees were present in the region during
the last glacial period, then it is an important
region in which we can study them in terms of understanding
forest development without the arrival
of any immigrant taxa. If migration was not a significant
factor in the postglacial vegetation succession
of the region, other factors need to be considered
to explain the type and rates of postglacial
vegetation change. This can include external forces
such as climate change and internal factors such as
soil development and forest dynamics [40, 41].
Figure 4. Refugial areas of oak species (after [37]).
The species that were present in the region
could adapt to climatic and other conditions and
could even genetically change in a fairly short time
during the Holocene. Here there can be found, for
instance, two subspecies of the species Cistus incanus:
C. incanus subsp. incanus and C. incanus
subsp. creticus. These subspecies can be explained
by a process of adaptive radiation within Cistus
incanus agg. Specimens found in the continental
parts of the Balkans have larger leaves, a straight
leaf margin and do not have glandules, whereas
specimens in the coastal areas have glandules and
undulated leaf margins. These differences are treated
taxonomically on the level of subspecies; specimens
in the continent are treated as the typical subspecies,
whereas those in the coastal regions as the
subspecies creticus. It should still be tested whether
these differences are genetically fixed; what would
happen if plants from the coastal region were planted
in continental conditions and vice-versa [42, 43].
BIOGEOGRAPHICAL DIVISION
OF THE REGION
The following biogeographical regions can
be found in the area: the Mediterranean, which is
linked to the coastal area; the mountain area is classified
within the alpine region, while the majority
of the area is connected to the continental region
and there is a small part that can be attached to the
Pannonian region; there is also a Black Sea region
along the Black Sea coast. (European Environmental
Agency: www.eea.eu.int).
This is the commonly accepted solution of the
biogeographical division. However, other divisions
can also be found, including that of Oberdorfer, who
drew a distinction between the Mediterranean area
and the Eurosibirian region in the northern part of
the Balkans. This distinction is based on two factors.
Evergreen vegetation can be found only in the
southernmost part of Greece and there is quite a
large area of submediterranean thermophilous deciduous
forest there, unlike the situation in the western
Mediterranean. The other factor is that the transition
between evergreen and deciduous vegetation is fairly
continuous; there is maquis consisting only of evergreen
species, then pseudomaquis consisting of a
mixture of evergreen species and deciduous species
and šibljak consisting only of deciduous species. On
the other hand, the division between mesophilous
deciduous and thermophilous deciduous forest is
fairly obvious [27, 45].
OVERVIEW OF FORESTS
Riverine, floodplain and moor forest
These forests appear in areas that are under
the influence of a watercourse or underground water.
The boundary of these habitats in floodplains is
the line that is reached by the highest floods. The
time when the highest water level is attained depends
on the water regime; rivers that have their
source in the mountains have the highest level during
late spring, when the snow melts, while others
have their maximum during high precipitation, and
floods can happen unexpectedly due to sudden high
precipitation, often causing destruction. When the
water level is low, these forests can often be dry.
Species need to be able to adapt to floods, to survive
the submerged period, to have flexible branches
not be broken by a water course etc. and also to
survive the lack of humidity during low water lev-
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els. On the other hand, these habitats are very rich
in nutrients, which the water flow brings and deposits
and this makes the habitats one of the richest
in the landscape. At the same time, these deposits
can be carried out by the next floods and so the best
position for growth is near the edge of average
floods, where the fine deposits are left and are not
too often carried off. In these stands, the development
of a soil profile can already be found. The river
brings deposits and gravel with flow, which are deposited
in its bed. The river climbs above the landscape
in this way and moor vegetation can be found
in the hinterland. These moor forests are often dominated
by Alnus glutinosa, since this species well
supports the high ground water over the whole year.
These stands can be occasionally flooded. This vegetation
seems to be to a large extent azonal and does
not show a special geographic pattern. In lowlands,
such stands are often in contact with zonal oakhornbeam
forest in the north and with thermophilous
deciduous forest in the south; pine forests can sometimes
be found on old gravel deposits [46–50].
The majority of these forests are classified
into Alno-Populetea albae, which encompass floodplain
and riverine forests of the Eurosibirian and
Mediterranean regions. The class is divided into
two orders, which reflect macroclimatic conditions:
Populetalia albae in the Mediterranean region and
Alno-Fraxinetalia in the temperate region. Within
the Mediterranean part, Platanus dominated communities
can be distinguished, classified as Platanion
orientalis and Lauro nobilis-Fraxinion angustifoliae
encompassing other floodplain and riparian
forests [51]. Forests of the temperate order AlnoFraxinetalia
can be divided into two alliances; Alnion
incanae encompassing forests appearing along
the upper stream of rivers and dominated to a large
extent by alders (Alnus glutinosa, A. incana) and
Alno-Quercion, which appear downstream and are
dominated by Fraxinus, Ulmus and Quercus. Alnion
incanae is in close contact with oak-hornbeam
forests and, in some cases, these forests share many
common species. Some authors do not recognize
the individuality of the two alliances: Alnion
incanae and Alno-Quercion [7, 52]. Riparian willow
forests, appearing mainly on lowland alluvia,
are classified within Salicetea purpureae, Salicetalia
purpuraea and Salicion albae. Mesotrophic forests
appearing in waterlogged sites are classified
into Alnetea glutinosae, Alnetalia glutionsae and
Alnion glutinosae [46–52].
Acidophilous forests
These forests appear on less productive soils,
where more demanding species such as hornbeam,
maple and others cannot thrive. Deciduous acidophilous
forests are dominated by Fagus sylvatica,
Quercus petraea and Betula pendula. It has
been established that Quercus robur is rarely a
dominant in such forests in the region. Fagus sylvatica
dominated forests are classified into the
group of beech forests (Carpino-Fagetea). Oak and
birch forests are also classified into two different
classes (see further). Mesic-acidophilous oak dominated
forests appear only in the western part of the
region, while in the eastern part acidothermophilous
oak forests belong to the group of
thermophilous deciduous forests and are classified
within the class Quercetea pubescentis. Birch forests
can be found all over the region. These forests
are to a large extent the result of human activities
over centuries. With the abandonment of exploitation,
succession is liberated and they can convert to
mesic forests [7, 8, 53, 54].
Castanea sativa dominated forests need further
research. Castanea sativa originates from the
Quercion frainetto zone and is probably non-native
in the northwestern part of the Balkans. It grows
not only on acidic soils but can also appear on other
substrata, so these forests can be classified into various
groups: acidophilous, mesophilous, thermophilous
deciduous and ravine forests [12, 55–59].
Acidophilous oak dominated forests are classified
into the group of acidophilous oak and birchoak
forests on nutrient poor soils, Quercetea roboris-petraeae
and Quercetalia roboris. Two alliances
appear; one from temperate regions, Agrostio-Quercion
petraeae and the other from submediterranean
regions, Castaneo-Quercion. The division
between these two alliances in the area should be
reconsidered in the future. In recent years, proposals
have been made to separate birch forests
from oak forests on the class level and to classify
them within the group of hemiboral pine and birchpine
herb rich open forests on fertile soils and relict
birch-poplar forest Brachypodio pinnati-Betuletea
pendulae and further into relict extrazonal temperate
deciduous birch-poplar woods on mineral soil
Fragario vescae-Populetalia tremulae and
Fragario vescae-Populion tremulae [17, 60, 61].
Thermophilous deciduous forests
The most widespread forests of the region
are thermophilous deciduous forests. They can be
divided into two larger groups. (1) The first is dominated
by various oak species. They form zonal forests
in the southern-eastern part of the region and
extrazonal forests on loess substratum on the edges
of the Pannonian basin, as well as forests on south-
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ern slopes on carbonate bedrock in the northwestern
part. Thermophilous-acidophilous oak forests
appearing in the central and eastern part of the region
are also attached to this group. (2) The second
group is dominated by Ostrya carpinifoila, Quercus
pubescens and Carpinus orientalis. Such vegetation
can be found along the coast in the submediterranean
area, on steep slopes with shallow soils on carbonate
mountain chains that separate the coast and
the continent, such as the Dinaric Alps or Shar–
Pindos mountain range and on dry calcareous sites
in the hinterland [62–65].
These forests are classified within Quercetea
pubescentis and Quercetalia pubescenti-petraeae, a
group of forests that are dominated by various oak
species (Q. pubescens, Q. cerris, Q. petraea, Q.
frainetto, etc.), oriental hornbeam (Carpinus orientalis),
hop hornbeam (Ostrya carpinifolia), as well
as Mediterranean relict fir species (A. cephalonica).
This vegetation is fairly diverse and can be divided
into several subunits. Within this framework can be
distinguished submediterranean forests dominated
by Quercus pubescens and Carpinus orientalis
Carpinion orientalis, meso-thermophilous supraMediterranean
and relic forests dominated by Ostyra
carpinifolia Fraxino orni-Ostryion and continental
thermophilous forests dominated by Carpinus
orientalis Syringo-Carpinion orientalis, thermophilous
continental forests of deep, neutral to slightly
acidic soils dominated by Quercus frainetto and
Quercus cerris Quercion confertae and those under
maritime influence Melitto albidae-Quercion, dry
acido-thermophilous (sub)montane continental forests
dominated by Quercus petraeae and Quercus
cerris Quercion petraeo-cerridis, acido-thermophilous
northern-dinaric and southern-pannonian
Quercus petraea dominated forests Quercion pubescenti-petraeae,
and thermophilous oak forests
on deep soils in the forest-steppe zone in the northern
part of the region Aceri tatarici-Quercion [12,
66, 67].
Ravine forests
Broadleaved ravine forests grow especially
on restricted sites with specific soil conditions.
They occur on slopes, at the foot of slopes, in sinkholes,
gorges and hollows with colluvial, skeletal
and primarily unstable soils, which allow broadleaved
trees such as Acer platanoides, Acer pseudoplantus,
Fraxinus excelsior, Tilia sp. div. to settle
and replace competitively stronger species,
mainly Fagus sylvatica. These species are called
noble trees, because their litter decays quickly and
ameliorates the soils, and so many high stalk plants
can be found in these forests, such as Lunaria rediviva,
Urtica dioica and many others. Within the
area of the southern Alps and southeastern Europe,
these forests can be classified within two large
groups: thermophilous scree and ravine forests
dominated by Tilia sp. div. and Ostrya carpinifolia
and mesophilous scree and ravine forests dominated
by Acer pseudoplatanus, Fraxinus excelsior and
Ulmus glabra [68].
These forests are classified within mesic deciduous
and mixed forests Carpino-Fagetea, as well
as scree and ravine forests Aceretalia pseudoplatani.
In the region, this group can be divided into mesophilous
ash-maple forests Fraxino-Acerion and
thermophilous lime-hop hornbeam forests Ostryo
carpinifoliae-Tilion platyphylli [17, 69].
Oak-hornbeam forests
Oak-hornbeam forests appear as zonal vegetation
of the plains and hilly region of the northwestern
part of the Balkan Peninsula, whereas in
the southeastern part hornbeam forests can be
found only extrazonally in valleys, along water
courses and other habitats that offer humidity and
mesic site conditions. In the southeastern part of the
region, oak-hornbeam forests are replaced by thermophilous
deciduous forests as zonal lowland forests.
In the northwestern part of the region, oakhornbeam
forests are in contact with beech forests,
which form the submontane vegetation belt. In the
lowlands, oak and hornbeam outcompete beech because
they better support summer drought and spring
frosts. In the transitional zone, beech forests are often
converted into oak-hornbeam coppiced forests.
In natural stands, oak can be found in the upper and
hornbeam in the lower tree layer. However, this vertical
zonation in a stand can often be changed by
forest management; pure hornbeam or oak forests
can be found. Oak-hornbeam forests also appear on
more humid sites. Since hornbeam is more susceptible
to underground and surface water, it gradually
disappears from stands and those forests are classified
into floodplain forests. [57, 70–72]
Oak-hornbeam forests are classified within
the class of mesic deciduous and mixed forests
Carpino-Fagetea; specifically they belong to oakhornbeam
and mesic oak forests on deep nutrient
rich soils Carpinetalia betuli. Oak-hornbeam forests
of the Balkans are classified into ErythronioCarpinion
betuli; within this framework two suballiances
can be distinguished, as moesian oakhornbeam
forests Aceri tartarici-Carpinenion and
illyric oak-hornbeam forests Lonicero caprifolii-
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Carpinenion. Hornbeam forests in the southeastern
part of the Balkans can be classified into CastaneoCarpinion
orientalis, which has its center of distribution
in northern Anatolia [49, 73].
Beech forests
Beech forests cover a large proportion of the
European forest area. It has to be considered that
beech gained the dominant role in European forests
in the subatlantic period about 2500 years ago and
in about 30 generations has formed well established
forest communities. The population originating
from the northwestern Balkans is the most vital,
since it has settled nearly the whole continent.
Beech can be found in intermediate sites that are
not too wet and not too dry, not too nutrient rich
nor too poor, not too warm or too cold. It has its
optimum habitats in the montane belt, where it
builds monodominant stands that are sometimes
mixed with fir (Abies alba, A. borisii-regis). In the
northwestern part of the region, beech forests occupy
various sites and have a wide altitudinal range,
while in the southeastern part of the region and on
the edge of the Pannonian plain, in their border
range, they can only be found in humid mountain
areas. [11, 35, 74, 75].
During elaboration of beech forests, we treated
only the taxa Fagus sylvatica subsp. sylvatica and
Fagus sylvatica subsp. orientalis. The intermediate
taxon Fagus sylvatica subsp. moesiaca was attached
to Fagus sylvatica subsp. sylvatica [75].
Classification of beech forests has been taking
place for nearly a century and a common consensus
could not be reached. Three main gradients can be
observed with beech forests: soil pH, temperature
and geography, of which the last includes macroclimatic
gradients and, at the same time, also a history
of the spread of beech forests. A consensus has been
achieved about acidophilous beech forests and beech
forests on acid soils in temperate Europe are classified
within a separate group. However, basiphilous
beech forests have given rise to major discussion
among researchers. It is not obvious which factor is
more important, temperature or geography. If temperature
is more important, units would be ecologically
defined and subunits geographically, and viceversa.
A consensus has recently been achieved that
both approaches can be considered equally relevant.
With the European classification system, it has been
accepted that the mostly well-defined alliances
should be retained. [13, 76]
Acidophilous beech forests, illyric beech forests,
moesian beech forests and south Balkan beech
forests form individual groups in the region. All
beech forests from Europe have been classified into
the class of mesic deciduous and mixed forests
Carpino-Fagetea. Acidophilous beech forests are
classified into Luzulo-Fagetalia sylvaticae and Luzulo-Fagion.
Basiphilous beech forests are classified
into Fagetalia sylvaticae. These forests from the illyric
floristic province are further classified into
Aremonio-Fagion. Moesian beech forests are not
well defined floristically and it was decided to attach
them partly to Central European beech forests
Fagion sylvaticae and partly to south Balkan beech
forests. South Balkan beech forests are grouped into
Geranio striati-Fagion. The latter group also comprises
some acidophilous communities. [11, 13, 76]
Oriental beech forests
Fagus orientalis appears in Europe in the extreme
southeastern part, in Turkey, Bulgaria and
Greece. The distribution center of these forests is in
the Euxinian province, which is situated in the central
part of the Turkish Black Sea region. On the basis
of analysis, these forests were divided into seven
groups (alliances). Only two groups can be found in
Europe: submontane and ravine beech forests. Within
the European classification system, only one alliance
is recognized, containing all oriental beech forests
of Europe. Oriental beech forests are classified
within the class of mesic deciduous and mixed forests
Carpino-Fagetea and within oriental beech forests
Rhododendro pontici-Fagetalia orientalis. Oriental
beech forests appearing in Europe are classified
within Fagion orientalis [13, 76, 77].
CONCLUSIONS
The presentation provides a brief overview
of deciduous forests of the Balkan Peninsula on the
alliance level. An attempt is made to put all forests
of the region into a logical system that is integral
within the region and corresponds to the European
classification scheme. However, other classification
concepts still exist, each with its own shortcomings
and inconsistencies, and much effort will be needed
to attain the optimal solution.
Acknowledgement. I owe thanks to Academician
Vlado Matevski, who stimulated the preparation of this
contribution. I also thank Iztok Sajko, who kindly prepared
the illustrations. The preparation of the article was
funded by the Slovenian Research Agency (P1-0236).
Andraž Čarni
Contributions, Sec. Nat. Math. Biotech. Sci., MASA, 37 (2), 95–106 (2016)
102
Appendix
Synsystematic scheme (after [13]):
Alno glutinosae-Populetea albae P. Fukarek et Fabijanić 1968
Populetalia albae Br.-Bl. ex Tchou 1949 nom. conserv. propos.
Platanion orientalis I. Kárpáti et V. Kárpáti 1961
Lauro nobilis-Fraxinion angustifoliae I. Kárpáti et V. Kárpáti 1961
Alno-Fraxinetalia excelsioris Passarge 1968
Alnion incanae Pawłowski et al. 1928
Alno-Quercion roboris Horvat 1950
Salicetea purpureae Moor 1958
Salicetalia purpureae Moor 1958
Salicion albae Soó 1951
Alnetea glutinosae Br.-Bl. et Tx. ex Westhoff et al. 1946
Alnetalia glutinosae Tx. 1937
Alnion glutinosae Malcuit 1929
Quercetea robori-petraeae Br.-Bl. et Tx. ex Oberd. 1957
Quercetalia roboris Tx. 1931
Agrostio-Quercion petraeae Scamoni et Passarge 1959
Castaneo-Quercion petraeae Soó 1964
Brachypodio pinnati-Betuletea pendulae Ermakov et al. 1991
Fragario vescae-Populetalia tremulae Willner et Mucina in Willner et al. 2016
Fragario vescae-Populion tremulae Willner et Mucina ined.
Quercetea pubescentis Doing-Kraft ex Scamoni et Passarge 1959
Quercetalia pubescenti-petraeae Klika 1933
Carpinion orientalis Horvat 1958
Fraxino orni-Ostryion Tomažič 1940
Syringo-Carpinion orientalis Jakucs (1959) 1960
Quercion confertae Horvat 1958
Melitto albidae-Quercion Barbero et Quézel 1976
Quercion petraeo-cerridis Lakušić et B. Jovanović in B. Jovanović et al. ex Čarni et Mucina 2015
Quercion pubescenti-petraeae Br.-Bl. 1932 nom. mut.
Aceri tatarici-Quercion Zólyomi 1957
Carpino-Fagetea sylvaticae Jakucs ex Passarge 1968
Aceretalia pseudoplatani Moor 1976 nom. conserv. propos.
Fraxino excelsioris-Acerion pseudoplatani P. Fukarek 1969
Ostryo carpinifoliae-Tilion platyphylli (Košir et al. 2008) Čarni in Willner et al. 2016
Carpinetalia betuli P. Fukarek 1968
Erythronio-Carpinion (Horvat 1958) Marinček in Wallnöfer et al. 1993
Castaneo sativae-Carpinion orientalis Quézel, Barbero et Akman ex Quézel et al. 1993
Luzulo-Fagetalia sylvaticae Scamoni et Passarge 1959
Luzulo-Fagion sylvaticae Lohmeyer et Tx. in Tx. 1954
Fagetalia sylvaticae Pawłowski 1928
Aremonio-Fagion (Horvat 1950) Borhidi in Török et al. 1989
Fagion sylvaticae Luquet 1926
Geranio striati-Fagion Gentile 1970
Rhododendro pontici-Fagetalia orientalis Passarge 1981
Fagion orientalis Soó 1964
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ВЕГЕТАЦИЈА НА ЛИСТОПАДНИ ШУМИ НА БАЛКАНСКИОТ ПОЛУОСТРОВ
Андраж Чарни
Институт за биологија, Научно истражувачки центар на Словенска академија на науките и
уметностите, Љубљана, Словенија
Универзитет на Нова Горица, Нова Горица, Словенија
Македонска академија на науките и уметностите, Скопје, Република Македонија
Во трудот се дава преглед на листопадните шуми на Балканскиот Полуостров. Претставени се речните,
заливните и мочуришните шуми, во кои доминираат Alnus incana, А. glutinosa, Fraxinus excelsior, F. angustifolia,
Quercus robur, Salix alba, Ulmus laevis, U. minor; ацидофилни шуми во кои доминираат Betula pendula, Castanea
sativa, Fagus sylvatica, Quercus petraea; термофилни шуми во кои доминираат Carpinus orientalis, Ostrya
carpinifolia, Quercus cerris, Q. farinetto, Q. petraea, Q. pubescens и мезофилни шуми во кои доминираат
Carpinus betulus, Fagus sylvatica, F. orientalis, Fraxinus excelsior, Tilia argentea, T. cordata, T. platyphyllos,
Ulmus glabra. Во исто време, шумите се класифицирани во синсистематски стандрадни рамки според
Системот на Централна Европа до ниво на сојуз.
Клучни зборови: Балкан; шума; вегетација; класификација
The author has requested enhancement of the downloaded file. All in-text references underlined in blue are linked to publications on ResearchGate.The author has requested enhancement of the downloaded file. All in-text references underlined in blue are linked to publications on ResearchGate.