259
Two new mesophilous oriental hornbeam 
communities from the northern Dinaric Alps 
(Bosnia and Herzegovina)
Abstract
The paper describes two new mesophilous communities of oriental hornbeam 
(Carpinus orientalis) coppice from the northern Dinaric Alps in Bosnia and 
Herzegovina (B&H). While oriental hornbeam is mainly considered to be a 
part of thermophilous forests and scrub, numerical analysis of 103 relevés of 
C. orientalis dominated coppice from B&H has shown that two new, rather 
mesophilous, communities thrive on calcareous bedrock of NW B&H. They 
represent secondary successional stages of mesotermic forest vegetation in this 
region. Association Epimedio alpini-Carpinetum orientalis ass. nova hoc loco is 
related to Illyrian oak-hornbeam forests of Erythronio-Carpinion betuli, while 
Asplenio scolopendrii-Carpinetum orientalis ass. nova hoc loco is linked to Balkan 
submediterranean ravine forests of Ostryo-Tilion. Although these two associations 
were recorded only in the NW B&H, their distribution is potentially larger, as 
their source communities are relatively common throughout the Dinaric Alps, 
so the information about their distribution, vertical structure, and syndynamic 
relations could be very useful in a national scale forest management and nature 
conservation.
Iz vleček
V članku opisujemo dve novi panjevski mezofilni združbi kraškega gabra 
(Carpinus orientalis) v severnem delu Dinarskega gorstva v Bosni in Hercegovini 
(B&H). Kraški gaber gradi predvsem termofilne gozdove in grmišča, vendar je 
numerična amaliza 103 popisov panjevcev s prevladujočim kraškim gabrom v 
B&H pokazala, da dve novi mezofilni združbi uspevata na apnenčasti podlagi v 
severozahodni B&H. Predstavljata faze v sekundarni sukcesiji mezotermne gozdne 
vegetacije v tem območju. Asociacija Epimedio alpini-Carpinetum orientalis ass. 
nova hoc loco je povezana z ilirskimi hrastovo-gabrovimi gozdovi zveze Erythronio-
Carpinion betuli, asociacija Asplenio scolopendrii-Carpinetum orientalis ass. nova 
hoc loco pa z balkanskimi submediteranskimi gozdovi plemenitih listavcev zveze 
Ostryo-Tilion. Čeprav sta ti dve asociaciji zabeleženi samo v severozahodnem 
delu B&H je njihova potencialna razširjenost večja, saj so njihove izvorne 
združbe relativno pogoste v celotnem Dinarskem gorstvu. Zato je poznavanje 
njihove razširjenosti, vertikalne strukture in sindinamskih odnosov uporabno za 
gospodarjenje z gozdom in naravovarstvo na nacionalnem nivoju.
Key words: Carpinus orientalis, 
Carpinion orientalis, Ostryo-Tilion, 
Erythronio-Carpinion, Epimedio 
alpini-Carpinetum orientalis, 
Asplenio scolopendrii-Carpinetum 
orientalis, phytosociology, 
syndynamics.
Ključne besede: Carpinus orientalis, 
Carpinion orientalis, Ostryo-
Tilion, Erythronio-Carpinion, 
Epimedio alpini-Carpinetum 
orientalis, Asplenio scolopendrii-
Carpinetum orientalis, fitocenologija, 
sindinamika.
Received: 1. 5. 2020
Revision received: 18. 5. 2020
Accepted: 6. 6. 2020
1 University of Banja Luka, Faculty of Forestry, S. Stepanovića 75A, BA-78000 Banja Luka, Bosnia and Herzegovina. 
* E-mail: vladimir.stupar@sf.unibl.org
Vladimir Stupar1
,
 *

, Jugoslav Brujić1 & Ognjen Lukić1

DOI: 10.2478/hacq-2020-0012 19/2 • 2020, 259–273

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T wo new mesophilous oriental hornbeam communities from the northern 
Dinaric Alps (Bosnia and Herzegovina)
Introduction
Oriental hornbeam (Carpinus orientalis) is a thermophil-
ous and xerophilous tree species distributed in SE Europe 
(southern Italy and Balkan Peninsula) and extending to 
the Asia Minor, Syria, Caucasus region and Crimea (Sik-
kema & Caudullo 2016). Being typical element of Sub-
mediterranean forest vegetation, it can also be found on 
warmer sites of continental regions of its area of distribu-
tion. It is main species of canopy layer in Submediter-
ranean thermophilous oriental-hornbeam forests of the 
Central and Southern Balkans (Syringo-Carpinion orien-
talis) (Mucina et al. 2016) and important understory spe-
cies of many thermophilous deciduous oak-dominated 
forest communities of Carpinion orientalis and Quercion 
frainetto (Blasi et al. 2001, Vukelić 2012, Tomić & Ra-
konjac 2013, Stupar et al. 2016, Tzonev et al. 2019). 
The latter communities are often structurally degraded to 
Carpinus orientalis dominated coppice. Although floristi-
cally they can be very different, such degraded oriental 
hornbeam communities mainly have closed or semi-open 
canopy and, being mainly coppice, all of them share com-
mon prominent synmorphological characteristic, such as 
bushy appearance (related to the fact that after the cut 
back of the oriental hornbeam trees several shoots develop 
simultaneously from one stool).
Regardless of floristic differences, this physiognomical 
similarity led to poor recognition of new vegetation types 
within oriental hornbeam coppices in Western Balkans, 
as was already brought to attention by Šugar & T rinajstić 
(1988) who distinguished continental C. orientalis cop-
pice in Croatia (Cruciato glabrae-Carpinetum orientalis) 
from Mediterranean Querco-Carpinetum orientalis. Ma-
jority of other continental C. orientalis coppices in the 
Western Balkans were treated in the scope of one widely 
comprehended association known under several invalid or 
illegitimate names (Stefanović 1989, Stupar et al. 2015). 
Moreover, until recently, there were only a few associa-
tions of C. orientalis dominated coppice recognized and 
correctly described in the whole Balkan Peninsula (Blečić 
& Lakušić 1967, Šugar & Trinajstić 1988, Bergmeier & 
Dimopoulos 2008, Tzonev 2013).
However, recent research suggests that there is a much 
higher diversity among these communities since they ap-
pear in different regions, reflect different ecological con-
ditions, and are degradations of different types of oak for-
ests (i.e., those of Carpinion orientalis, Quercion confertae 
or Erythronio-Carpinion) (Stupar et al. 2015, Miletić et 
al. 2016). While Stupar et al. (2015) encompassed all of 
the continental communities in Bosnia and Herzegovina 
(B&H) under the association Cruciato glabrae-Carpine-
tum orientalis Šugar et Trinajstić ex Stupar et al. 2015, 
they stated that this association is rather heterogeneous, 
with mesophilous species playing an important role in 
some stands in the NW B&H, and that further research 
is needed to reveal its diversity patterns. Additionally, 
they pointed out that in this transitional zone between 
Central and SE Europe, within the association of Aceri 
obtusati-Quercetum petraeae Stupar et al. 2015 one can 
find together two species of different ecologies, mesophil-
ous Carpinus betulus, and xerothermophilous Carpinus 
orientalis. This suggested that there are possibly some 
more mesophilous communities where C. orientalis plays 
an important role in B&H. 
The study aimed to use the results of the recent field 
research of C. orientalis coppice in B&H to describe two 
new mesophilous communities dominated by oriental 
hornbeam and characterize them by their ecology and 
floristic composition, which would in turn aid to the clas-
sification of this vegetation type in B&H, as well as in the 
wider context.
Methods and materials
Study area
The research took place in the middle part of the Vrbas 
River canyon with its surroundings. The area is situat-
ed in northwestern B&H and it is about a 40 km long 
geomorphological feature that predominantly has south-
north direction in the northern part of the Dinaric Alps 
 (Figure 1). Limestone canyons and gorges are intersected 
by a few rather small valleys in Bočac, Krupa na Vrbasu, 
Rekavice, and Karanovac and surrounded by predomi-
nantly limestone and dolomitic low mountains (up to 
1330 m a.s.l.) and plateaus (400–600 m a.s.l) Čemernica, 
Tisovac, Osmača and Starčevica to the east and Manjača 
Mt. to the west (Mojićević et al. 1976, Marinković & 
Ahac 1979). Vrbas River elevations are between 160 m 
in the north and 270 m a.s.l. in the south. Climate is 
modified continental with the maritime influence from 
the west and Mediterranean from the south (Delijanić 
et al. 1964). The average annual temperature for Banja 
Luka is 11.0 °C and annual rainfall is 1024 mm (Anony-
mous n.d.). Biogeographically, this area lies at the bor-
der between two regions, i.e., Pre-Pannonian (continen-
tal, northern B&H) and Dinaric (mountainous, central 
B&H) (Stefanović et al. 1983). It is also an area that is 
under the strong anthropogenic influence for centuries, 
and large portions of land were either deforested or struc-
turally degraded to coppice or shrub communities.

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Data collection and analysis
All 103 relevés used in this study are stored in, and avail-
able from, the Forests Vegetation Database of Bosnia and 
Herzegovina, with the ID EU-BA-001 in the Global In-
dex of Vegetation-Plot Databases (Dengler et al. 2011). 
From the year of 1998, but mainly in the last ten years, we 
recorded 51 relevés of C. orientalis coppice in B&H us-
ing standard Central European phytosociological method 
(Braun-Blanquet 1964). Together with 52 relevés from 
literature (Fabijanić et al. 1963, Stefanović & Manuševa 
1971, Kaćanski 1983, Stefanović 1989, Lakušić & Redžić 
1991, Muratspahić et al. 1991, Bucalo 1999, Miletić et 
al. 2016) they were stored into the Turboveg database 
(Hennekens & Schaminée 2001) and then exported into 
JUICE software (Tichý 2002) for further analysis.
Before numerical analysis was applied, we excluded 
mosses from the dataset, as they were not recorded by all 
authors. For the means of numerical analysis, all layers 
were merged into a single layer to take account of incon-
sistent sampling in the relevés from literature, even though 
we recorded four layers during the field investigation (can-
opy layer: < 10 and > 5 m, understory layer: < 5 and > 1 m, 
shrub layer: < 1 m and herb layer). However, understory 
and shrub layers were combined into a single understory 
layer for use in the text and the preparation of the tables 
with relevés. Records of species determined to the genus 
level were deleted. Taxa occurring in three or fewer rel-
evés were omitted from the analysis to reduce noise (Juvan 
et al. 2013, Stupar et al. 2015). Quercus daleschampi was 
treated as Q. petraea, while Lathyrus vernus and L. venetus 
were combined. Taxa from taxonomically critical groups 
were also combined (Bromus erectus, Carex muricata, Cyt-
isus hirsutus, Festuca pseudovina, and T araxacum officinale).
Figure 1: Locations of relevés.
Slika 1: Lokacije popisov. 


 

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The numerical classification of the data set was per -
formed in PC-ORD (McCune & Mefford 1999) using 
Flexible beta (-0.25) and Relative Sørensen. Consid-
ering that the study aimed to delineate and describe 
mesophytic oriental hornbeam communities outside of 
Carpinion orientalis, after cluster analysis, we selected the 
level of division (four groups of relevés) that best suited 
the aim. Diagnostic species for clusters were determined 
using species fidelity measure (Chytrý et al. 2002) in the 
JUICE software. We also calculated Fischer’s exact test 
and gave a zero fidelity value to a species with P>0.001. 
The threshold phi value for the species to be considered 
as a diagnostic was set at 0.25.
All relevés, together with the unweighted species eco-
logical indicator values (EIVs) for temperature, light, 
moisture, continentality, soil reaction and nutrients (Pig-
natti et al. 2005) were passively projected onto a non-
metric multidimensional scaling (NMDS) plot to relieve 
main ecological factors that affect the variation of the 
floristic composition inside the data set. We used Bray-
Curtis distance measure and square-root transformation 
of cover data. The significance of EIVs correlation with 
the NMDS relevé scores was tested using the modified 
permutation test proposed by Zelený & Schaffers (2012). 
The analysis was done in R software, version 3.4.1 (The 
R Foundation for Statistical Computing 2017) using the 
vegan package (https://github.com/vegandevs/vegan).
Plant nomenclature followed Euro+Med (2006). Syn-
taxonomical concepts and nomenclature of higher syntaxa 
followed Mucina et al. (2016). Descriptions of new syntaxa 
strictly followed the rules of ICPN (Theurillat et al. 2020).
Results
Numerical analysis of Carpinus orientalis dominated cop-
pice from Bosnia and Herzegovina led to the delimita-
tion of four ecologically and floristically distinct groups 
of clusters (Figure 2): xerophilous (clusters 1 and 2) and 
Figure 2: Classification of C. orientalis coppice 
in Bosnia and Herzegovina. Clusters:  
1 – Rusco aculeati-Carpinetum orientalis, 
2 – Cruciato glabrae-Carpinetum orientalis, 
3 – Epimedio alpini-Carpinetum orientalis,  
4 – Asplenio scolopendrii-Carpinetum orientalis.
Slika 2: Klasifikacija panjevcev kraškega gabra 
v Bosni in Hercegovini. Klastri:  
1 – Rusco aculeati-Carpinetum orientalis,  
2 – Cruciato glabrae-Carpinetum orientalis,  
3 – Epimedio alpini-Carpinetum orientalis,  
4 – Asplenio scolopendrii-Carpinetum orientalis.
Figure 3: NDMS spider plot of 103 classified relevés of the C. orien-
talis coppice in B&H with EIVs passively projected. Centroids of 
clusters are indicated by numbers corresponding to Table 1, Figure 2, 
and to cluster numbers used in the text. 
Slika 3: NDMS graf 103 klasificiranih popisov panjevcev kraškega 
gabra v B&H s pasivno projiciranimi EIV vrednostmi. Centroidi 
klastrov so predstavljeni s številkami, ki odgovarjajo klastrom v 
Tabeli 1, Sliki 2 in številkam klastrov v besedilu.
mesophilous (clusters 3 and 4). They are characterized 
by distinctive floristic composition with clearly defined 
diagnostic species (Table 1). This is also supported by the 
NDMS ordination plot, where the main ecological fac-
tors influencing the variation in the floristic composition 
are moisture and nutrients (positively correlated to the 
first axis) and light (negatively correlated to the first axis) 
(Figure 3). Xerophilous clusters (left side of the ordina-
tion and cluster diagrams) belong to already known asso-
ciations, i.e., Rusco aculeati-Carpinetum orientalis Blečić 
et Lakušić 1967 (cluster 1) and Cruciato glabrae-Carpi-
netum orientalis Šugar et Trinajstić ex Stupar et al. 2015 

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(cluster 2), while mesophilous clusters 3 and 4 (right side 
of the ordination and cluster diagrams) are recognized as 
new associations.
These results suggest new syntaxonomical scheme of 
Carpinus orientalis coppice in B&H with four associa-
tions within three alliances and two classes:
Quercetea pubescentis Doing-Kraft ex Scamoni et Passarge 
1959
Quercetalia pubescenti-petraeae Klika 1933
Carpinion orientalis Horvat 1958
Rusco aculeati-Carpinetum orientalis Blečić 
et Lakušić 1967
Cruciato glabrae-Carpinetum orientalis Šugar 
et T rinajstić ex Stupar et al. 2015
Carpino-Fagetea sylvaticae Jakucs ex Passarge 1968
Carpinetalia betuli P . Fukarek 1968
Erythronio-Carpinion (Horvat 1958) Marinček 
in Wallnöfer et al. 1993
Epimedio alpini-Carpinetum orientalis ass. 
nova hoc loco (Table 1 column 3, Table 2)
Aceretalia pseudoplatani Moor 1976 nom. conserv. 
propos.
Ostryo carpinifoliae-Tilion platyphylli (Košir et al. 
2008) Čarni in Willner et al. 2016
Asplenio scolopendrii-Carpinetum orientalis 
ass. nova hoc loco (T able 1 column 4, T able 3)
Description of the new syntaxa
Epimedio alpini-Carpinetum orientalis  
ass. nova hoc loco (Table 1, column 3)
Typus: Table 2, rel. 11 – holotypus hoc loco
Stands of this association are coppices that represent 
secondary succession stages of the association Aceri obtu-
sati-Quercetum petraeae Stupar et al. 2015. They are found 
in NW BIH (Figure 1), on mild to moderately steep 
mainly warmer slopes on calcareous substrates (limestone 
and dolomites) and elevations between 200 and 600 m 
a.s.l. Due to closed canopy, the stands are very shaded 
which results in the poorer species composition than the 
source association, however, the cover of the herb layer 
can sometimes be very high. These stands are coppices 
with a bushy appearance and height between 5 and 10 
m (Figure 4). Due to mild terrain relief and closeness to 
settlements, these stands are continuously being coppiced 
and consequently maintained in the degraded stage.
The dominant species of the canopy layer is Carpinus ori-
entalis, while other diagnostic species include species char-
acteristic for oak-hornbeam forests (Erythronio-Carpinion 
betuli) such as Carpinus betulus, Cyclamen purpurascens, 
Epimedium alpinum, Helleborus odorus, Potentilla micran-
tha, Primula acaulis, Ruscus hypoglossum, T amus communis, 
but also the other species of mesophilous deciduous forests 
Figure 4: Stand of the association Epimedio alpini-Carpinetum orientalis in Starčevica (Drenovača). Photo: Vladimir Stupar.
Slika 4: Sestoj asociacije Epimedio alpini-Carpinetum orientalis pri Starčevici (Drenovača). Foto: Vladimir Stupar.

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Dinaric Alps (Bosnia and Herzegovina)
(Fagetalia sylvaticae): Ajuga reptans, Aremonia agrimonoi-
des, Asarum europaeum, Brachypodium sylvaticum, Carex 
pilosa, Cruciata glabra, Drymochloa drymeja, Geum urba-
num, Glechoma hirsuta, Hedera helix, Melica uniflora, Poly-
podium vulgare, Pulmonaria officinalis, Sanicula europaea, 
Stellaria holostea, Symphytum tuberosum, etc. Fraxinus ornus 
and Acer obtusatum are usually present in the canopy layer 
with lower cover values. There are only individual plants of 
Quercus petraea in the canopy and understory layers, but 
besides this general lack of Q. petraea, the main difference 
between this association and Aceri obtusati-Quercetum pe-
traeae is lack or low cover and frequency of light-demand-
ing species such as Lathyrus niger, Melittis melissophyllum, 
Tanacetum corymbosum and Aegonychon purpurocaeruleum 
in Epimedio-Carpinetum orientalis, all due to light shortage 
under the closed canopy of C. orientalis coppice. Also, the 
average number of species in Epimedio-Carpinetum orien-
talis is lower by ten when compared to Aceri obtusati-Quer-
cetum petraeae. The stands of this association were formerly 
assigned to Cruciato glabrae-Carpinetum orientalis Šugar et 
Trinajstić ex Stupar et al. 2015 from which they are dif-
ferentiated by many mesophilous species and lack of the 
thermophilous Carpinion orientalis species.
Asplenio scolopendrii-Carpinetum orientalis 
ass. nova hoc loco (Table 1 column 4)
Typus: Table 3, rel. 6 – holotypus hoc loco
This interesting association is found on “microlocali-
ties” in canyons and gorges of the Vrbas River and its 
confluences (Figure 1). It is the secondary succession 
stage of the ravine forests of Ostryo-Tilion. They occupy 
northern, very steep (35–50°) and stony (50–80% of 
the bare rock) slopes on limestone, and the elevations 
between 200 and 400 m a.s.l. These are also coppices 
with a bushy appearance and height between 7 and 10 
m (Figure 5). Although these sites are hardly accessible 
they are still being regularly coppiced and maintained in 
a structurally degraded state.
The main species of the canopy layer is Carpinus ori-
entalis, with sometimes admixed tree species of the po-
tential Ostryo-Tilion community: Fraxinus ornus, Ostrya 
carpinifolia, Tilia tomentosa and Acer monspessulanum. 
Besides oriental hornbeam, other character species are 
mesophilous ferns Polystichum setiferum, Asplenium scolo-
pendrium, Asplenium trichomanes, and Polypodium vul-
gare along with Saxifraga rotundifolia, Veratrum nigrum, 
Figure 5: Stand of the association Asplenio scolopendrii-Carpinetum orientalis in Vrbas River canyon. Photo: Ognjen Lukić.
Slika 5: Sestoj asociacije Asplenio scolopendrii-Carpinetum orientalis v kanjonu reke Vrbas. Foto: Ognjen Lukić.

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Geranium robertianum, Carex digitata, and Asplenium ce-
terach. Among other species characteristic for ravine for-
ests (Ostryo-Tilion) we can find Hedera helix, Helleborus 
odorus, Hepatica nobilis, Cyclamen purpurascens, etc. Spe-
cies characteristic for Carpino-Fagetea are also frequent, 
while of the species characteristic for Quercetea pubescen-
tis there are only Cornus mas and Sesleria autumnalis with 
higher frequency.
Discussion
Our results suggest that Epimedio-Carpinetum orien-
talis is a secondary succession stage of Aceri obtusati-
Quercetum petraeae, which is also found over calcareous 
bedrock in the NW part of B&H. Although Stupar et 
al. (2015) treated Aceri obtusati-Quercetum petraeae as a 
part of thermophilous deciduous forests of Quercetalia 
pubescentis they did not assign it to any of the alliances in 
this order, and stated that this association represents the 
transition from thermophilous forests of Carpinion orien-
talis towards mesophilous forests of Carpinion betuli (i.e. 
Erythronio-Carpinion). Although its understory is made 
of thermophilous species of SE European (Carpinus ori-
entalis, Acer obtusatum, and Fraxinus ornus), and also 
wider distribution (Cornus mas, Euonymus verrucosus, 
and Sorbus torminalis), there are also frequent mesophil-
ous elements, such as Carpinus betulus, Acer campestre 
and Prunus avium. Moreover, the floristic composition 
of its herb layer is mainly mesophilous, with only several 
thermophilous species of SE European distribution. All 
of this is characteristic of Erythronio-Carpinion (Košir et 
al. 2013, Novák et al. 2020), which is probably where 
Aceri obtusati-Quercetum petraeae belongs. Although in 
theory degradation of Aceri obtusati-Quercetum petraeae 
should be more thermophilous, selective logging of oaks 
and regular coppicing of the oriental hornbeam resulted 
in the closed canopy and actually more mesophilous 
 oriental hornbeam coppice of Epimedio-Carpinetum ori-
entalis. 
Although we recorded Epimedio-Carpinetum orienta-
lis only in the NW B&H, there is a mention of orien-
tal hornbeam coppice resulted from the degradation of 
oak-hornbeam forest (Querco-Carpinetum illyricum typi-
cum) from Lepenica in the central B&H (Fabijanić et al. 
1963). Although there are no relevés for this C. orientalis 
coppice, Querco-Carpinetum illyricum typicum is floris-
tically very similar to Aceri obtusati-Quercetum petraeae 
(Carpinus betulus together with C. orientalis and other 
thermophilous species in the understory), so it could be 
expected that the distribution of both, Aceri obtusati-
Quercetum petraeae and its degradation stage Epimedio-
Carpinetum orientalis is potentially larger than recorded. 
It should also be pointed out that Fabijanić et al. (1963) 
argued that Querco-Carpinetum illyricum typicum is typi-
cal oak-hornbeam community (belonging to Carpinion 
betuli illyrico-podolicum) on calcareous bedrock in B&H 
stressing out that it is more thermophilous than original-
ly described climax oak-hornbeam community of con-
tinental Croatia Querco-Carpinetum croaticum Horvat 
1938 (i.e. Epimedio-Carpinetum betuli).
Although there is an evident floristic similarity be-
tween Aceri obtusati-Quercetum petraeae and its second-
ary successional stage Epimedio-Carpinetum orientalis, 
there are also clear differences, which reflect in different 
physiognomy of oriental hornbeam coppice, lack of edi-
fier in the upper layer (Quercus petraea), and the smaller 
overall number of species per relevé. The latter is due to 
the closed canopy of oriental hornbeam coppice, which 
does not allow light to reach the herb layer. Albeit sec-
ondary succession stage, light shortage under the closed 
canopy together with continuous coppicing resulted in 
the formation of the relatively stable degradation stage 
which is hard to convert to high forests of Aceri obtusati-
Quercetum petraeae. This led us to the conclusion that 
secondary succession stages of different oak–hornbeam 
forests, if they are stable, low and dark hornbeam and/
or oriental hornbeam coppices, without upper tree layer, 
and with the low cover or without oaks, could be treated 
as separate associations (or at least subassociations) in-
side the same alliance as their source community. This 
would also relate better with typologies used in forest 
management.
However, there is an opinion that some pure oriental 
hornbeam communities should be classified in its own 
alliance, as is the case with the Submediterranean ther-
mophilous oriental hornbeam forests of the Central and 
Southern Balkans of Syringo-Carpinion (Mucina et al. 
2016). Their source communities with various oaks are 
instead being classified into the Amphiadriatic alliance 
of Carpinion orientalis (Tzonev et al. 2019) although they 
lack Illyrian species which are replaced with eastern-cen-
tral Balkan ones. Following this logic, obviously meso-
philous Arabido turritae-Carpinetum orientalis Tzonev 
2013 (Tzonev 2013) is classified into Syringo-Carpinion 
orientalis (Tzonev et al. 2009), while floristically it be-
longs to Erythronio-Carpinion.
Asplenio scolopendrii-Carpinetum orientalis is a member 
of the Submediterranean ravine forests of Ostryo-Tilion. 
Although this alliance is very well characterized by its 
characteristic species combination it was recognized only 
recently as an independent syntaxon at the level of subal-
liance (Košir et al. 2008), and then later on raised at the 
level of alliance (Willner et al. 2016). Stefanović (1979) 

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Vladimir Stupar, Jugoslav Brujić & Ognjen Lukić
T wo new mesophilous oriental hornbeam communities from the northern 
Dinaric Alps (Bosnia and Herzegovina)
wrote about Aceri-Tilietum mixtum in the canyons of the 
Dinaric Alps which fits perfectly into the circumscrip-
tion of the Ostryo-Tilion, although it was not taken into 
consideration by Košir et al. (2008). Asplenio-Carpinetum 
is probably the secondary successional stage of this com-
munity, occurring on the higher parts of canyon slopes 
on colder exposures, whereas closer to the river and more 
humid lower parts are degraded to Ostrya carpinifolia ra-
vine communities.
According to Stefanović (1979), Aceri-Tilietum mixtum 
with the higher cover of Carpinus orientalis is more com-
mon in Neretva and Morača River canyons (Herzegovina 
and Montenegro) so it can be expected that Asplenio-
Carpinetum has a larger distribution in the canyons of 
Dinaric Alps.
Disturbance pattern, along with a high cover of bare 
rock doesn’t allow this community to progress further 
towards the climax community of Ostryo-Tilion. However, 
it still has a huge protective role (antierosion) and presents 
one more authentic forest habitat of B&H canyons.
Acknowledgements
We would like to thank to two anonymous referees for 
very valuable comments and suggestions that have signifi-
cantly improved the first version of the manuscript.
Vladimir Stupar , https://orcid.org/0000-0003-0835-2249
Ognjen Lukić , https://orcid.org/0000-0003-1279-0647
References 
Anonymous: Meteorological station Banja Luka. Report for the period 
1974–2004.
Bergmeier, E. & Dimopoulos, P . 2008: Identifying plant communities 
of thermophilous deciduous forest in Greece: Species composition, dis-
tribution, ecology and syntaxonomy. Plant Biosystems 142: 228–254. 
Blasi, C., Di Pietro, R., Filesi, L. & Fortini, P . 2001: Syntaxonomy, 
chorology and dynamics of Carpinus orientalis communities in Central 
Italy. Phytocoenologia 31: 33–62. 
Blečić, V. & Lakušić, R. 1967: Niederwald und Buschwald der 
orientalischen Hainbuche in Montenegro. Glasnik Botaničkog zavoda i 
bašte Univerziteta u Beogradu 2: 83–94. 
Braun-Blanquet, J. 1964: Pflanzensoziologie, Grundzüge der 
Vegetationskunde. 3rd ed. Springer Verlag, Wien, 865 pp. 
Bucalo, V. 1999: Šumske fitocenoze planine Jadovnik. Šumarski 
fakultet Univerziteta u Banjoj Luci, Beograd, RS, 326 pp. 
Chytrý, M., Tichý, L., Holt, J. & Botta-Dukát, Z. 2002: Determina-
tion of diagnostic species with statistical fidelity measures. Journal of 
Vegetation Science 13: 79–90. 
Delijanić, I., Radičević, D., Sokolović-Ilić, G., Ivanović, D., 
Milošević, S. & Seničar, E. 1964: Klimatska rejonizacija Jugoslavije 
i tabelarni prikaz temperaturnih i pluviometrijskih karakteristika 
klimatskih reona Jugoslavije. Jugoslovenski poljoprivredno šumarski 
centar Beograd: 121–138. 
Dengler, J., Jansen, F ., Glöckler, F ., Peet, R.K., De Cáceres, M., 
Chytrý, M., Ewald, J., Oldeland, J., Lopez-Gonzalez, G., Finckh, M., 
Mucina, L., Rodwell, J.S., Schaminée, J.H.J. & Spencer, N. 2011: The 
Global Index of Vegetation-Plot Databases (GIVD): a new resource for 
vegetation science. Journal of Vegetation Science 22: 582–597. 
Euro+Med 2006: Euro+Med PlantBase – the information resource for 
Euro-Mediterranean plant diversity. Available from: http://ww2.bgbm.
org/EuroPlusMed/query.asp (October 12, 2015) 
Fabijanić, B., Fukarek, P . & Stefanović, V. 1963: Lepenica: Pregled 
osnovnih tipova šumske vegetacije. Naučno društvo SR Bosne i 
Hercegovine, Posebna izdanja 3: 85–129. 
Hennekens, S.M. & Schaminée, J.H.J. 2001: TURBOVEG, a 
comprehensive data base management system for vegetation data. 
Journal of Vegetation Science 12: 589–591.
Juvan, N., Košir, P ., Marinšek, A., Paušič, A. & Čarni, A. 2013: 
Differentiation of the Piceetalia and Athyrio-Piceetalia forests in 
Slovenia. T uexenia 33: 25–48. 
Kaćanski, D. 1983: Dinamika biocenoza ekosistema rijeke Vrbas 
(do Jajca). Biološki institut Univerziteta u Sarajevu, Sarajevo, BA,. 
Elaborat, 66 pp. 
Košir, P ., Čarni, A. & Di Pietro, R. 2008: Classification and 
phytogeographical differentiation of broad-leaved ravine forests in 
southeastern Europe. Journal of Vegetation Science 19: 331–342.
Košir, P ., Casavecchia, S., Čarni, A., Škvorc, Ž., Živković, L. & 
Biondi, E. 2013: Ecological and phytogeographical differentiation of 
oak-hornbeam forests in southeastern Europe. Plant Biosystems 147: 
84–98.
Lakušić, R. & Redžić, S. 1991: Vegetacija refugijalno-reliktnih ekosis-
tema sliva rijeke Une. Bilten Društva ekologa BiH, serija B 6: 25–73. 
Marinković, R. & Ahac, A. 1979: Osnovna geološka karta SFRJ, 
1 : 100 000, list Jajce (sa tumačem). Savezni geološki zavod, Beograd, RS. 
McCune, B. & Mefford, M.J. 1999: PC-ORD. Multivariate Analysis 
of Ecological Data. Version 5.0. MjM Software, Gleneden Beach, 
Oregon, US, 
Miletić, M., Milanović, Đ., Stupar, V. & Brujić, J. 2016: Šumska 
vegetacija T rešnjika kod Banje Luke. Glasnik Šumarskog fakulteta 
Univerziteta u Banjoj Luci 25: 15–40. 
Mojićević, M., Vilovski, S. & Tomić, B. 1976: Osnovna geološka 
karta SFRJ, 1:100 000, list Banja Luka (sa tumačem). Savezni geološki 
zavod, Beograd, RS, 
Mucina, L., Bültmann, H., Dierßen, K., Theurillat, J.-P ., Raus, T., 
Čarni, A., Šumberová, K., Willner, W., Dengler, J., García, R.G., 
Chytrý, M., Hájek, M., Di Pietro, R., Iakushenko, D., Pallas, J., 
Daniëls, F .J.A., Bergmeier, E., Santos Guerra, A., Ermakov, N., 
Valachovič, M., Schaminée, J.H.J., Lysenko, T., Didukh, Y.P ., 
Pignatti, S., Rodwell, J.S., Capelo, J., Weber, H.E., Solomeshch, A., 
Dimopoulos, P ., Aguiar, C., Hennekens, S.M. & Tichý, L. 2016: 
Vegetation of Europe: hierarchical floristic classification system of 
vascular plant, bryophyte, lichen, and algal communities. Applied 
Vegetation Science 19: 3–264.

19/2 • 2020, 259–273
267
Vladimir Stupar, Jugoslav Brujić & Ognjen Lukić
T wo new mesophilous oriental hornbeam communities from the northern 
Dinaric Alps (Bosnia and Herzegovina)
Muratspahić, D., Redžić, S. & Lakušić, R. 1991: Asocijacija Rusco-
Carpinetum orientalis Bleč. et Lkšić 1966 u dolini rijeke Neretve. 
Glasnik Republičkog zavoda za zaštitu prirode i Prirodnjačkog muzeja 
u Podgorici 24: 7–12.
Novák, P ., Willner, W., Zukal, D., Kollár, J., Roleček, J., Świerkosz, 
K., Ewald, J., Wohlgemuth, T., Csiky, J., Onyshchenko, V. & Chytrý, 
M. 2020: Oak-hornbeam forests of central Europe: a formalized 
classification and syntaxonomic revision. Preslia 92: 1–34.
Pignatti, S., Menegoni, P . & Pietrosanti, S. 2005: Valori di 
bioindicazione delle piante vascolari della flora d’Italia. Braun-
Blanquetia 39: 1–97.
Sikkema, R. & Caudullo, G. 2016: Carpinus orientalis in Europe: 
distribution, habitat, usage and threats. In: San-Miguel-Ayanz, J., de 
Rigo, D., Caudullo, G., Houston Durrant, T., & Mauri, A. (eds.) 
European Atlas of Forest T ree Species. Publ. Off. EU, Luxembourg, 
p. e01bf18+.
Stefanović, V. 1979: Fitocenoza javora i lipa (Aceri-Tilietum mixtum, 
Stef., 1974) u nekim kanjonima Dinarida. Drugi kongres ekologa 
Jugoslavije. Savez društava ekologa Jugoslavije, Zadar-Plitvice, 
pp. 1083–1102. 
Stefanović, V. 1989: Prilog poznavanju fitogeografskih i fitocenoloških 
odnosa bjelograbovih šuma izvan klimazonalnog areala u Bosni. 
Glasnik Zemaljskog muzeja Bosne i Hercegovine, Prirodne nauke, 
Nova serija 28: 103–119. 
Stefanović, V., Beus, V., Burlica, Č., Dizdarević, H. & Vukorep, 
I. 1983: Ekološko-vegetacijska rejonizacija Bosne i Hercegovine. 
Šumarski fakultet u Sarajevu, Posebna izdanja 17: 1–49. 
Stefanović, V. & Manuševa, L. 1971: Šumska vegetacija i zemljišta na 
andezitu i dacitu istočne Bosne. Radovi Šumarskog fakulteta i Instituta 
za šumarstvo u Sarajevu 15: 1–83. 
Stupar, V., Brujić, J., Škvorc, Ž. & Čarni, A. 2016: Vegetation types of 
thermophilous deciduous forests (Quercetea pubescentis) in the Western 
Balkans. Phytocoenologia 46: 49–68.
Stupar, V., Milanović, Đ., Brujić, J. & Čarni, A. 2015: Formalized 
classification and nomenclatural revision of thermophilous deciduous 
forests (Quercetalia pubescentis) of Bosnia and Herzegovina. 
T uexenia 35: 85–130.
Šugar, I. & T rinajstić, I. 1988: Prilog poznavanju bjelograbovih šuma u 
Hrvatskoj. Poljoprivreda i šumarstvo, Titograd 34: 43–51. 
Theurillat, J.-P ., Willner, W., Fernández-González, F ., Bültmann, H., 
Čarni, A., Gigante, D., Mucina, L. & Weber, H. 2020: International 
Code of Phytosociological Nomenclature. 4th edition. Applied 
Vegetation Science. https://doi.org/10.1111/avsc.12491 
Tichý, L. 2002: JUICE, software for vegetation classification. Journal 
of Vegetation Science 13: 451–453.
Tomić, Z. & Rakonjac, L. 2013: Šumske fitocenoze Srbije: Priručnik 
za šumare, ekologe i biologe. Univerzitet Singidunum, Fakultet za 
primenjenu ekologiju Futura i Institut za šumarstvo, Beograd, 177 pp. 
Tzonev, R. 2013: New plant associations from Danubian Plain, 
Bulgaria. Phytologia Balcanica 19: 243–266. 
Tzonev, R., Dimitrov, M., Gussev, C., Pachedjieva, K., Gogushev, 
G., Apostolova-Stoyanova, N., Nikolov, I., Alexandrova, A. & 
Glogov, P . 2019: Phytosociological classification of the thermophilous 
oak forests in Bulgaria: new interpretations and gaps in knowledge. 
Phytocoenologia 49: 369–391. 
Tzonev, R., Dimitrov, M. & Roussakova, V. 2009: Syntaxa according 
to the Braun-Blanquet approach in Bulgaria. Phytologia Balcanica 15: 
209–233. 
Vukelić, J. 2012: Šumska vegetacija Hrvatske. Šumarski fakultet, 
Sveučilište u Zagrebu, DZZP , Zagreb, 403 pp. 
Willner, W., Solomeshch, A., Čarni, A., Bergmeier, E., Ermakov, N. 
& Mucina, L. 2016: Description and validation of some European 
forest syntaxa – a supplement to the EuroVegChecklist. Hacquetia 15: 
15–25.
Zelený, D. & Schaffers, A.P . 2012: Too good to be true: pitfalls of 
using mean Ellenberg indicator values in vegetation analyses. Journal 
of Vegetation Science 23: 419–431.

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Group number 1 2 3 4
No. of relevés 12 50 33 8
Rusco aculeati-Carpinetum orientalis
Paliurus spina-christi 83
87,4
2 . .
Asparagus acutifolius 58
69,8
2 . .
Rubus ulmifolius 58
68,1
4 . .
Petteria ramentacea 50
63,5
2 . .
Brachypodium pinnatum 58
63,3
10 . .
Acer monspessulanum 100
63
14 6 62
Viola alba 67
61,8
10 12 .
Pistacia terebinthus 42
59,1
. . .
Clematis flammula 42
59,1
. . .
Satureja montana 33
49,8
2 . .
Cruciato glabrae-Carpinetum orientalis
Thymus pulegioides ssp. montanus 8 76
71,8
9 .
Euphorbia cyparissias . 62
69,1
6 .
Origanum vulgare . 36
54,5
. .
Galium lucidum 8 52
52,8
9 .
T eucrium chamaedrys 75 88
52,4
9 .
Bromus erectus agg. 8 48
51,6
6 .
Dorycnium pentaphyllum ssp. 
germanicum
. 30
49,3
. .
Stachys recta . 28
47,5
. .
Lotus corniculatus 8 36
45,8
. .
Filipendula vulgaris . 30
45,6
3 .
Sedum acre . 30
45,6
3 .
Cytisus hirsutus . 38
44,4
12 .
Pilosella officinarum 8 34
43,9
. .
Juniperus communis . 48
43,1
27 .
Scabiosa cinerea ssp. cinerea . 22
41,8
. .
Epimedio alpinum-Carpinetum orientalis
Cruciata glabra 8 24 85
64,7
12
Quercus petraea . 26 67
59,5
.
Rosa arvensis . 34 79
59,1
12
Festuca heterophylla 17 8 64
57,9
.
Ajuga reptans 8 6 55
57,1
.
Cyclamen purpurascens . 20 85
54,7
50
Acer obtusatum 8 44 76
54,1
.
Helleborus odorus . 42 100
53
75
Carpinus betulus . 12 45
51,2
.
Acer tataricum . 6 39
51
.
Lathyrus vernus . 4 36
50,4
.
Group number 1 2 3 4
No. of relevés 12 50 33 8
Luzula forsteri . . 30
49,6
.
Sorbus torminalis 8 20 55
48,2
.
Glechoma hirsuta 8 32 79
46,9
38
Drymochloa drymeja . . 27
46,9
.
Sanicula europaea . 2 27
44,2
.
Stellaria holostea . 6 42
43,7
12
Aremonia agrimonoides . 12 55
43,5
25
Hedera helix 50 24 94
42,4
62
Brachypodium sylvaticum 25 14 73
42,3
38
Epimedium alpinum . 2 36
41
12
Asplenio scolopendrium-Carpinetum orientalis
Saxifraga rotundifolia . . 6 100
96,1
Veratrum nigrum . 10 . 100
93,7
Polystichum setiferum . 2 27 100
83,6
Asplenium scolopendrium . 2 15 88
80,6
Carex digitata . 10 36 100
76
Hepatica nobilis . 8 33 88
68,3
Geranium robertianum 17 4 24 88
66,7
Valeriana officinalis . 2 . 50
63,5
Asplenium trichomanes 25 22 39 100
61,8
Peltaria alliacea . 4 15 62
60,3
Lamium galeobdolon . 8 12 62
59,7
Mercurialis perennis . 2 18 62
59,6
Sambucus nigra . . . 38
55,7
Galium schultesii . . 12 50
54,9
Galium sylvaticum . . 12 50
54,9
Polypodium vulgare . 2 61
27,5
88
59,6
Other species with high frequency
Carpinus orientalis 100 100 100 100
Sesleria autumnalis 100 76 24 75
Crataegus monogyna 58 54 76 38
Fraxinus ornus 75 94 91 88
Cornus mas 25 60 79 100
Asplenium ceterach 50 30 30 88
Ruscus aculeatus 75 16 48 25
Table 1: Frequency-fidelity table of Carpinus orientalis coppice communities in B&H (fidelity value in superscript multiplied by 
100). Diagnostic species (phi values higher than 0.25) for each community are shaded (only species with the phi value higher than 
0.4 are presented). Cluster numbers correspond to those used throughout the text.
Tabela 1: Tabela frekvenc in navezanosti panjevski združb vrste Carpinus orientalis v B&H (nadpisane vrednosti navezanosti vrst so 
pomnožene s 100). Diagnostične vrste (fi vrednosti večje od) vsake združbe so zasenčene (prikazane so samo vrste s fi vrednostjo, 
večjo od 0,4 ). Številke klastrov so enake kot v besedilu.

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Relevé number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
Relevé area (m2) 100 100 100 225 400 400 400 400 400 400 400 400 400 400 225
Altitude (m) 340 390 477 310 370 340 308 543 552 523 230 323 258 270 280
Aspect SE SE S W E W SW SE E S E SE SE SE W
Slope (degrees) 5 25 15 35 25 25 15 5 5 3 15 3 7 10 35
Hight of the canopy layer (m) 6 9 8 8 10 9 4 6 6 6 7 8 9 8 13
Cover total (%) 100 90 100 80 70 95 95 80 90 80 90 90 70 80 100
Cover A (%) 100 90 100 80 100 90 0 80 90 80 90 90 70 80 60
Cover B (%) 20 50 50 25 25 20 90 20 10 10 20 20 50 40 15
Cover C (%) 5 60 20 50 90 60 20 30 20 40 30 20 90 90 70
Cover bare rock (%) 50 0 10 0 0 20 0 10 10 40 5 0 20 20 0
Characteristic species of the association
Carpinus orientalis A 5 4 5 4 4 4 . 4 4 4 5 5 4 4 2
Carpinus orientalis B 1 . . 1 1 1 4 2 2 2 1 2 1 1 3
Acer obtusatum A . 1 1 . 1 2 . + . + + . . . 1
Acer obtusatum B + . + + 2 + . + . 2 2 + + + .
Potentilla micrantha C + . + + + . + 1 + 1 + 1 + + .
Brachypodium sylvaticum C 1 3 + 1 . . 1 1 1 1 + 1 2 1 +
Lathyrus vernus+venetus C . 1 + . + + . + + . r + 1 + +
Glechoma hirsuta C + . . + . . 1 1 1 1 1 + 1 1 .
Pulmonaria officinalis C r . . + . . + + 1 . + r + + +
Festuca heterophylla C . + 2 . . . 1 1 r + + 1 1 1 .
Asarum europaeum C . + . . + . + 1 r . + + 1 1 +
Polypodium vulgare C . + . + + . + r r + . . + . +
Erythronio-Carpinion
Carpinus betulus A . . . . . + . 1 1 . + + 1 1 .
Carpinus betulus B . + . . . . 2 2 . . 1 1 . . .
Ruscus aculeatus B . 3 + 2 2 1 . . . . + . . . 2
Ruscus hypoglossum B . + . + + 1 . . . . . . . . +
Helleborus odorus C + + + + 1 1 1 1 1 1 + 1 1 1 +
Cyclamen purpurascens C . 1 + + + 1 . + 2 1 + . 1 1 +
Primula acaulis C . + . + . . 1 + + . + + + + .
Epimedium alpinum C . 3 . r + . 1 . . . + . . . +
T amus communis C + 2 1 + 1 . . . . . . . . . 1
Carpino-Fagetea
Crataegus monogyna A + . . . . . . . . + 1 1 . . .
Crataegus monogyna B + 2 + + . . 2 + 1 2 2 2 1 . .
Quercus petraea A 1 + 1 . 2 1 . . . . . . . . .
Quercus petraea B 1 . 1 . + + . + 2 . + + . 1 +
Acer campestre A . . . . . 1 . . . 1 + . . . .
Table 2: Epimedio alpini-Carpinetum orientalis ass. nov. hoc loco, holotypus: relevé 11 (A – canopy layer (> 5 m),  
B – understory layer (< 5 m), C – herb layer).
Tabela 2: Epimedio alpini-Carpinetum orientalis ass. nov. hoc loco, holotip: popis 11 (A – drevesna plast (> 5 m),  
B – grmovna plast (< 5 m), C – zeliščna plast).

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Relevé number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
Acer campestre B 1 . + + + 2 2 . . . 2 + + . 1
Rosa arvensis B 1 + + + + . . + + + + + + + +
Hedera helix B 1 2 1 3 1 1 . 1 1 2 + . 1 2 3
Ligustrum vulgare B . 1 . + . 1 1 + + 1 1 + + + .
Clematis vitalba B r + . . + . + r 1 . . + r . .
Rubus hirtus B . . . . . + + + + . + . . . .
Corylus avellana B . . . . . + 1 . . . r . . . .
Cruciata glabra C . 3 + + + . + + + + 1 1 + + +
Aremonia agrimonoides C + . . . . . 1 1 1 + . + + + .
Symphytum tuberosum C . + + . + + . r . . + + + r .
Ajuga reptans C + 1 . + . . 1 . . . r . + + r
Dactylis glomerata C + . + + . + . + . + . . . . +
Carex sylvatica C + . . . . . + + + . . + 2 1 .
Luzula forsteri C . . + . . . . r + + . + + 1 .
Carex digitata C . . . + + 1 . . . . + . 1 2 r
Sanicula europaea C . + . . . . . + . . + + . + .
Drymochloa drymeia C . . . + 5 1 . + . + . . . . .
Hepatica nobilis C . . . + . + . . . . + . + 1 .
Euphorbia amygdaloides C . . . r . . . . . . + . + + +
Stellaria holostea C . + . . . . . . + 1 . . 1 . .
Bromus benekenii C . . r . . . . . . r + . . . 2
Daphne laureola B . . . + . . . + + . . . . . +
Polystichum setiferum C . . . + . . . r + . . . . . +
Geum urbanum C . . . + . . . . r + . . + . .
Viola reichenbachiana C . . . . . . + . + . + + . . .
Melica uniflora C . + + . . . . . r . . . . . .
Veronica chamaedrys C . . + + . r . . . . . . . . .
Galium schultesii C . . r r . . . . . . . . . + .
Carex pilosa C . . . . . 3 . . . . . . + 3 .
Quercetea pubescentis
Fraxinus ornus A 2 2 1 2 3 2 . . + 1 + + 1 1 .
Fraxinus ornus B 2 . 1 + + 2 . . 2 2 2 2 1 1 .
Euonymus verrucosus B . 2 . + 1 r . . . + + . + 1 .
Cornus mas A 1 . . . 1 1 . . . . . . . 1 .
Cornus mas B + . . 2 . 1 2 . 2 . 1 . 1 2 2
Sorbus torminalis A 1 . 2 . . . . . . . . . . . .
Sorbus torminalis B 1 . + + . . . . . . 2 1 1 . 1
Cotinus coggygria A 1 . . . . . . . . . . . . . .
Cotinus coggygria B 1 . . . + . . . . . . . + . +
Viburnum lantana B + 1 . . 2 . + . . . + . 1 . +
Tilia tomentosa B . . . . + 2 . . . . . . 1 . .
Carex flacca C . . . + . . 1 . . . 2 1 . r .
Viola hirta C + . + + + . . . . . . + . . .
Campanula persicifolia C . . + + + + . + . . . . . . .

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Relevé number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
Lathyrus niger C r . + . + . . . . . . . + . .
Melittis melissophyllum C . + . . + . . . . . . . . . +
Aegonychon purpurocaeruleum C 1 . + + . . . . . . . . . . .
Asplenium adiantum-nigrum C . . . + . + . . . . . . . . +
Asplenium ceterach C . . . + . . . . r + . . . . .
Allium carinatum C . . . . . . . r + r . . . . .
Silene nutans C . . . . . . . . . 1 . . + + .
Companions
Acer tataricum B r . r + + . . . . . + . 1 1 .
Juniperus communis B + + . . . . + r . . . + + . .
Fragaria moschata C . . + . . . . . . . . . + + .
Asplenium trichomanes C . . . + . . . . r . . . . . +
In one or two relevés:
A: Sorbus torminalis 1: 1, 3: 2; Cotinus coggygria 1: 1; Quercus cerris 2: +; Tilia tomentosa 5: 1, 6: 1; Ostrya carpinifolia 9: +, 10: +; Quercus 
pubescens 1: 1; Prunus avium 9: +; Fagus sylvatica 12: +; Fraxinus excelsior 15: 1;
B: Cornus sanguinea 2: +; Quercus cerris 7: 2; Pyrus communis ssp. pyraster 1: +, 12: +; Fagus sylvatica 3: +, 7: 2; Staphylea pinnata 4: +, 5: 2; 
Euonymus latifolius 4: +, 5: +; Euonymus europaeus 7: +, 11: +; Ostrya carpinifolia 9: 2, 10: 2; Prunus avium 9: 2, 11: 2; Quercus pubescens 1: 
1; Prunus spinosa 1: r; Rhamnus cathartica 2: 1; Cytisus hirsutus 2: 1; Ulmus minor 8: +; Tilia platyphyllos 8: r; Quercus robur 13: 2; Tilia 
cordata 15: +; Fraxinus excelsior 15: +;
C: Sesleria autumnalis 8: +, 14: +; Cephalanthera longifolia 2: +, 7: r; Mercurialis perennis 4: +, 15: +; Iris graminea 4: 1; Lilium martagon 5: +; 
Pteridium aquilinum 7: +; Vincetoxicum hirundinaria 1: r, 4: +; Melica nutans 2: 1, 4: +; Melampyrum pratense 2: +, 4: +; Aristolochia lutea 
2: +, 7: 1; Galium sylvaticum 2: +, 11: +; Fragaria vesca 2: +, 15: +; Campanula trachelium 3: +, 4: +; Polygonatum multiflorum 3: +, 9: +; 
Hylotelephium maximum 3: +, 11: r; Pseudoturritis turrita 4: +, 9: +; Asplenium scolopendrium 4: +, 15: +; Cardamine bulbifera 6: +, 9: +; 
Galanthus nivalis 6: r, 9: +; Taraxacum officinale 8: r, 9: r; Peltaria alliacea 9: +, 10: +; Arum maculatum 9: +, 11: +; Geranium robertianum 
9: r, 10: +; Aposeris foetida 11: 1, 12: 1; Melampyrum nemorosum 11: +, 14: +; Galium verum 2: 2; Thymus pulegioides 2: +; Poa angustifolia 
2: +; Luzula campestris 2: +; Asplenium ruta-muraria 2: +; Chaerophyllum hirsutum 3: r; Platanthera bifolia 3: r; Holcus lanatus 3: r; Lamium 
galeobdolon 4: +; Peucedanum austriacum 4: +; Piptatherum virescens 4: +; Silene viridiflora 4: +; Polygonatum odoratum 4: r; Anemone 
ranunculoides 6: +; Erythronium dens-canis 6: +; Orchis pallens 6: r; Orchis simia 7: +; Campanula bononiensis 7: r; Filipendula vulgaris 7: r; 
Corydalis cava 9: +; Neottia nidus-avis 9: +; Crocus vernus 9: +; Scilla bifolia 9: r; Galium mollugo 10: +; Orchis purpurea 11: 1; Anemone 
nemorosa 11: +; Smyrnium perfoliatum 13: +; Ornithogalum pyrenaicum 13: +; Peucedanum carvifolia 13: r; Pastinaca sativa 14: r.
Details of relevés (indicated in the following order: relevé number, GIVD EU-BA-001 relevé number, date (year/month/day), description 
of locality, longitude, latitude, bedrock): 1) 309, 2014/06/24, Ćićina kosa, Gradina (Bočac), 17.147586, 44.545781, limestone; 2) 1073, 
1998/04/28, Cer (Starčevica), 17.212744, 44.7504, dolomite; 3) 3026, 2017/05/02, Jagare (Starčevica), 17.207547, 44.712043, limestone; 
4) 3579, 2014/08/19, Magareći potok (Starčevica), 17.184836, 44.745235, dolomite; 5) 3580, 2014/07/31, T rešnjik (Starčevica), 17.17307, 
44.73713, dolomite; 6) 3632, 2020/04/03, Drenovača (Ponir), 17.221471, 44.747957, limestone; 7) 3636, 2020/04/05, Donja Kola (Manjača), 
17.090543, 44.749972, dolomite; 8) 3638, 2020/04/09, above Vrbas canyon (Ljubačevo), 17.18372, 44.683289, limestone; 9) 3639, 
2020/04/09, above Vrbas canyon (Ljubačevo), 17.184923, 44.682865, limestone; 10) 3640, 2020/04/09, Grabež Mala (Ljubačevo), 17.187266, 
44.680652, limestone; 11) 3642, 2020/04/09, Savići (Rekavice), 17.13603, 44.674709, limestone; 12) 3643, 2020/04/09, Rekavice, 17.131842, 
44.679936, marl limestone; 13) 3644, 2020/04/10, Krupa na Vrbasu, 17.135722, 44.617552, limestone; 14) 3645, 2020/04/10, Krupa na 
Vrbasu, 17.137349, 44.618658, limestone; 15) 3578, 2014/07/30, Banj brdo (Starčevica), 17.174481, 44.746997, dolomite.

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272
Vladimir Stupar, Jugoslav Brujić & Ognjen Lukić
T wo new mesophilous oriental hornbeam communities from the northern 
Dinaric Alps (Bosnia and Herzegovina)
Relevé number 1 2 3 4 5 6 7 8
Relevé area (m2) 400 100 400 100 400 400 225 225
Altitude (m) 313 330 350 215 317 406 220 230
Aspect NE NE N NW E N NW NE
Slope (degrees) 45 45 50 45 35 40 35 50
Hight of the canopy layer (m) 8 8 7 10 10 8 8 10
Cover total (%) 100 100 100 100 80 100 100 85
Cover A (%) 100 100 100 90 80 100 45 80
Cover B (%) 20 10 0 30 50 30 80 25
Cover C (%) 60 50 50 50 50 80 40 50
Cover bare rock (%) 80 50 70 90 75 60 50 80
Characteristic species of the association
Carpinus orientalis A 5 3 4 5 4 5 1 5
Carpinus orientalis B 2 + 2 2 2 2 3 .
Polystichum setiferum C 4 + 3 2 1 1 2 3
Asplenium scolopendrium C 3 2 2 2 2 1 2 2
Asplenium trichomanes C 1 1 + + 1 + 1 1
Asplenium ceterach C + + . + 1 + + +
Polypodium vulgare C 1 . 2 1 3 2 1 1
Saxifraga rotundifolia C + + 1 + + 1 + +
Veratrum nigrum C 2 1 2 + . + r +
Geranium robertianum C 1 + + + 1 . + +
Carex digitata C + + + + . + + +
Ostryo-Tilion
Fraxinus ornus A 1 2 3 . 2 1 1 1
Fraxinus ornus B 2 1 2 . 2 2 2 +
Ostrya carpinifolia A . 2 . 1 1 . 2 1
Ostrya carpinifolia B . . . . . + 2 .
Tilia tomentosa A 1 . 1 . . . . .
Tilia tomentosa B 1 . + . . 1 . .
Acer monspessulanum A + 1 1 1 . . . .
Hedera helix B + . + + 2 . + +
Euonymus verrucosus B . + + + . . + 1
Staphylea pinnata A + 2 . . . . . .
Staphylea pinnata B + 1 . . . . 2 .
Sambucus nigra B . . . + . . 2 +
Tilia platyphyllos B . . . + . . + +
Helleborus odorus C 1 . + + + 1 + +
Hepatica nobilis C + 1 + + . r + +
Cyclamen purpurascens C r . + + 2 . . .
Lathyrus vernus+venetus C + . r + . 1 + +
Peltaria alliacea C + . + + . r . .
Arum maculatum C r . . . + . . +
Fagetalia
Corylus avellana A . 1 . . . . . .
Corylus avellana B . 1 . . + . 1 .
Euonymus europaeus B . + . + + . . +
Table 3: Asplenio scolopendrii-Carpinetum orientalis ass. nov. hoc loco, holotypus: relevé 6 (A – canopy layer (> 5 m),  
B – understory layer (< 5 m), C – herb layer). 
Tabela 3: Asplenio scolopendrii-Carpinetum orientalis ass. nov. hoc loco, holotip: popis 6 (A – drevesna plast (> 5 m),  
B – grmovna plast (< 5 m), C – zeliščna plast).

19/2 • 2020, 259–273
273
Vladimir Stupar, Jugoslav Brujić & Ognjen Lukić
T wo new mesophilous oriental hornbeam communities from the northern 
Dinaric Alps (Bosnia and Herzegovina)
Relevé number 1 2 3 4 5 6 7 8
Dactylis glomerata C + + 1 + + 2 . +
Mercurialis perennis C + 1 r + + . . .
Lamium galeobdolon C + + + + . . + .
Veronica chamaedrys C r . r . + r . +
Galium sylvaticum C . + . + . . + +
Glechoma hirsuta C . . . . 1 + + +
Symphytum tuberosum C + + + . . + . .
Brachypodium sylvaticum C + + . + + . . .
Cardamine bulbifera C + . 1 . + . . .
Campanula trachelium C r . r . . + + .
Geum urbanum C . . . r + + . +
Galium schultesii C + . + . . + . .
Polygonatum multiflorum C + + + . . . . .
Asarum europaeum C + + . . . r . .
Silene dioica C + . 1 . . . . +
Galanthus nivalis C + . + . + . . .
Melica uniflora C + . . + . . . 1
Ruscus hypoglossum B + . . . . . . +
Lactuca muralis C . + . r + . . .
Cardamine impatiens C r . r . . . . +
Quercetea pubescentis
Cornus mas A 1 3 . 1 1 . . .
Cornus mas B 1 1 2 2 2 2 2 2
Sesleria autumnalis C + 1 1 + . 2 . .
Campanula persicifolia C + . r . . + . .
Potentilla micrantha C + . 1 + . . . .
Companions
Pseudoturritis turrita C + . . . + + . .
Valeriana officinalis C r + r . . . . .
Verbascum nigrum C r . r r . . . .
In one or two relevés:  
A: Staphylea pinnata 1: +, 2: 2; Crataegus monogyna 2: 1, 8: +; Corylus avellana 2: 1, 9: +; Acer obtusatum 5: 1, 9: 1; Carpinus betulus 5: 1, 9: +; 
Acer hyrcanum ssp. intermedium 2: 1; Tilia cordata 2: 1; Sorbus aria 3: 1; Tilia platyphyllos 7: 1; Acer campestre 7: 1; Ulmus glabra 5: 1; 
Ulmus minor 9: +;
B: Cotinus coggygria 2: 1, 6: +; Ostrya carpinifolia 6: +, 7: 2; Ruscus aculeatus 1: +, 6: 1; Acer campestre 1: r, 3: r; Crataegus monogyna 2: 1, 4: +; 
Acer monspessulanum 2: 1, 4: +; Viburnum lantana 2: +, 4: r; Ulmus minor 4: +, 7: +; Acer obtusatum 5: 2, 9: 1; Hippocrepis emerus ssp. 
emeroides 6: 1; Ligustrum vulgare 1: +; Rosa arvensis 1: +; Tilia cordata 2: +; Sorbus aria 3: +; Daphne laureola 5: 1; Lonicera xylosteum 7: +; 
Quercus robur 7: r; Clematis vitalba 8: +; Crataegus laevigata 9: +;
C: Viola hirta 1: +, 3: r; Hylotelephium maximum 4: +, 6: +; Moehringia trinervia 1: +, 3: +; Tamus communis 1: +, 6: r; Smyrnium perfolia-
tum 1: r, 6: +; Arabidopsis arenosa 4: +, 8: +; Poa angustifolia 6: +, 8: +; Fragaria moschata 6: r; Stellaria holostea 1: +; Carex sylvatica 1: +; 
Primula acaulis 1: +; Alliaria petiolata 1: +; Epimedium alpinum 2: 1; Clematis recta 2: +; Convallaria majalis 2: +; Laserpitium krapfii ssp. 
krapfii 2: +; Asplenium ruta-muraria 2: +; Euphorbia amygdaloides 2: +; Vincetoxicum hirundinaria 2: +; Symphyandra hofmannii 2: r; Lilium 
martagon 2: r; Tanacetum corymbosum 2: r; Urtica dioica 2: r; Adoxa moschatellina 3: r; Pulmonaria officinalis 4: +; Scutellaria altissima 4: r; 
Carex divulsa ssp. leersii 5: +; Galium lucidum 5: +; Arum italicum 5: +; Poa nemoralis 5: +; Anemone ranunculoides 5: +; Cardamine hir-
suta 5: r; Galium aparine 5: r; Digitalis grandiflora 6: +; Cruciata glabra 6: +; Isopyrum thalictroides 6: +; Geranium lucidum 6: r; Lunaria 
rediviva 7: +; Viola reichenbachiana 7: +; Inula conyza 7: +; Parietaria officinalis 7: +; Pseudofumaria alba ssp. leiosperma 8: +.
Details of relevés (indicated in the following order: relevé number, GIVD EU-BA-001 relevé number, date (year/ month/day), description of 
locality, longitude, latitude):  
1) 2936, 2015/05/14, Ispod Greben grada (Krupa na Vrbasu), 17.13748, 44.61054; 2) 3109, 2018/06/21, Kanjon Crne rijeke (desna obala), 
17.165446, 44.463635; 3) 2935, 2015/05/14, Ispod Greben grada (Krupa na Vrbasu), 17.137663, 44.610092; 4) 3102, 2018/06/20, Kanjon 
Vrbasa (desna obala, Tijesno), 17.197794, 44.68809; 5) 3646, 2020/04/15, Čelinski potok (Starčevica), 17.267045, 44.726266; 6) 2933, 
2015/05/14, Ispod Greben grada (Krupa na Vrbasu), 17.137304, 44.609612; 7) 3613, 2018/06/20, Kanjon Vrbasa (desna obala, Tijesno), 
17.197921, 44.687727; 8) 3610, 2018/06/20, Kanjon Vrbasa (desna obala, Tijesno), 17.19911, 44.68801.