ACTA BOT. CROAT. 76 (1), 2017 15
Acta Bot. Croat. 76 (1), 15–26, 2017 CODEN: ABCRA 25
DOI: 10.1515/botcro-2016-0041 ISSN 0365-0588
eISSN 1847-8476
Ecological, ﬂoristic and functional analysis of zonal
forest vegetation in Bosnia and Herzegovina
Vladimir Stupar1*
, Andraž Čarni2, 3
1
University of Banjaluka, Forestry Faculty, Department of Forest Ecology, S.Stepanovića 75a, BA-78000 Banjaluka,
Bosnia and Herzegovina
2
Institute of Biology, Research Centre of the Slovenian Academy of Sciences and Arts, Novi trg 2, SI-1000 Ljubljana,
Slovenia
3
University of Nova Gorica, Vipavska 13, SI-5000 Nova Gorica, Slovenia
Abstract – Zonal vegetation is a large-scale expression of macro-climate and, due to the climatic diversity of
the country, there are seven traditionally recognized zonal forest plant communities in Bosnia and Herzegovina.
Using data from Bosnia and Herzegovina, this study aimed to reveal whether macro-climate is indeed the
most important factor determining the existence of zonal forest plant communities (ZFPC). Detrended correspondence
analysis of 398 relevés of seven ZFPCs revealed that the species turnover along the ﬁrst axis is
strongly related to the macro-climatic gradient (annual mean temperature, mean temperature of the coldest
quarter and precipitation of the warmest quarter). No correlation was detected between this gradient and topographic
factors (slope and aspect) and soil reaction. Floristic analysis revealed clear separation of ZFPCs in
terms of diagnostic species. Functional analysis of all layers showed that competitive ecological strategy has
the highest proportion, while analysis of the herb layer alone expressed a shift of CSR signatures towards the
middle of the C–S axis. Ruderality was overall poorly expressed. Statistically signiﬁcant differences among
communities were discovered along the C–S axis. In terms of life forms, statistically signiﬁcant differences in
the proportions of Phanerophytes, Geophytes and Hemicryptophytes among communities were discovered.
Our study conﬁrms that macro-climatic gradient is the most important determinant of the species turnover
along ZFPCs. CSR signatures show that zonal forest vegetation is represented by productive communities in
a terminal stage of succession. This does not refer to degraded Quercus ilex stands (maquis), which are in the
middle stage of secondary succession.
Keywords: Balkans, climatic gradient, chorotypes, ecological strategies, life forms, ordination, plant functional
types, zonal communities
* Corresponding author, e-mail: vladimir.stupar@sfbl.org
Introduction
Every plant community is a result of a complex interaction
of various ecological factors in a given place and time.
In terms of natural vegetation, there are three types of vegetation
that generally develop in accordance with the biotic,
climatic and soil conditions: zonal, extrazonal and azonal
(Dierschke 1994, Ellenberg 2009, Surina 2014). Zonal vegetation
is a large-scale expression of climate dominating a
particular area, while it is not conﬁned to speciﬁc soil conditions,
i.e., it most precisely reﬂects the macroclimatic conditions
of particular regions (Kovar-Eder and Kvaček 2007).
Zonal vegetation often does not represent a single homogeneous
plant association but rather a number of similar communities,
which can differ to some extent and, consequently, it
is possible to talk of a ‘zonal vegetation group’ (Ellenberg
2009). For example, when different bedrocks and soil series
occur within the same climatic zone or there is a species
turnover due to minor biogeographical differences, several
different, yet similar associations make up the zonal vegetation
group. Furthermore, a mountainous relief of a particular
climatic zone leads to vertical differentiation of the climatic
factors and, accordingly, vertical differentiation of
zonal vegetation communities, which are then often called
‘altitudinal belts’.
Although in the last several thousand years man has deforested
great parts of Europe (Gunia et al. 2012), the potential
natural vegetation and, consequently, the zonal vegetation
of most of temperate Europe is forest (Bohn and
Neuhäusl 2004, Ellenberg 2009). The same is valid for Bosnia
and Herzegovina (B&H) (Horvat et al. 1974, Stefanović
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STUPAR V., ČARNI A.
16 ACTA BOT. CROAT. 76 (1), 2017
et al. 1983). B&H is situated in the western part of the Balkan
Peninsula in SE Europe (Fig. 1). Its climate is very diverse,
since two major climatic zones overlap here: central
European from the north and Mediterranean from the south.
The climate of the transitional zone is highly modiﬁed by
the inﬂuence of mountain massifs (Dinaric Alps), while in
the eastern part of country the inﬂuence of the drier continental
climate can be felt (Milosavljević 1976). While vegetation
studies of forests in the neighboring regions have
produced synthetic overviews (Borhidi et al. 2012, Vukelić
2012, Chytrý 2013, Tomić and Rakonjac 2013), despite the
fairly long tradition of phytosociological studies in B&H
(Horvat 1933, 1941, Horvat and Pawlowski 1939, Tregubov
1941) and the considerable number of relevés, the ecology,
nomenclature and syntaxonomical position of the majority
of B&H forests still remains unsettled (Redžić 2007). With
the exception of thermophilous deciduous forest communities
(Stupar et al. 2015), this also applies to zonal vegetation,
which has only been the subject of a few studies in the
past (Horvat et al. 1974, Stefanović et al. 1983, Beus 1984).
However, following the macro-climatic diversity, these authors
generally agree that seven zonal forest plant communities
(ZFPCs) can be distinguished for the territory of B&H
(Tab. 1, Fig. 1). Four communities are represented by various
oak forests, which occupy the lowlands and hilly region
of B&H, while the other three are altitudinal belts (montane,
altimontane and subalpine) above the oak forests, mainly
built by various types of beech forests (pure and mixed).
There are two different conceptual frameworks for the
study of plant communities. The traditional approach to
classiﬁcation is performed at the species level, while a more
recent ‘functional’ approach is based on plant functional
types (Duckworth et al. 2000, Shipley 2010, Škornik et al.
2010). Plant functional types are non-phylogenetic groupings
of species that show close similarities in their response
to environmental and biotic factors and are derived from
plant traits based on species morphology, physiology and/or
life history (Pérez-Harguindeguy et al. 2013). Plant functional
types can aid in the understanding of ecological processes,
such as the assembly and stability of communities
and succession, and facilitate the detection and prediction
of response to environmental change on a range of scales
(Duckworth et al. 2000). With plant functional types, comparisons
between communities of widely differing composition
can be facilitated (Diaz et al. 2004). There are two
main approaches to the classiﬁcation of functional types
Fig. 1. Location of the study area in Bosnia and Herzegovina. The
potential zonal forest plant communities (ZFPCs) are indicated
(Horvat et al. 1974, modiﬁed after Stefanović et al. 1983). Numbers
on the map correspond to community numbers in Tab. 1.
Tab. 1. Zonal forest plant communities in Bosnia and Herzegovina. Community numbers correspond to those used in Tabs. 3–4, and Figs.
1–4. Asterisk (*) denotes provisional invalid names still in use in Bosnia and Herzegovina.
Community
no.
Forest type Related syntaxa
No. of
relevés
No. of
resampled
relevés
1 Quercus ilex Quercion ilicis Br.-Bl. 1931 (1936) 5 5
2
Quercus pubescensCarpinus
orientalis
Querco pubescenti-Carpinetum orientalis Horvatić 1939 (Carpinion orientalis
Horvat 1958)
30 16
3 Quercus frainetto
Quercetum frainetto-cerridis (Rudski 1949) Trinajstić et al. 1996 (Quercion
frainetto Horvat 1954)
38 24
4
Quercus petraeaCarpinus
betulus
Querco-Carpinetum illyricum Horvat et al. 1974* (Erythronio-Carpinion betuli
(Horvat 1938) Marinček in Wallnöfer et al. 1993)
43 26
5
pure Fagus / mixed
Fagus-Abies
Fagetum montanum illyricum Fukarek et Stefanović 1958*, Abieti-Fagetum
dinaricum Tregubov 1957* (Aremonio-Fagion Török et al. ex Marinček et al. 1993)
231 162
6
mixed Picea-
Abies-Fagus
Piceo-Abieti-Fagetum Stefanović et al. 1983* (Abieti-Piceenion Br.-Bl. in Br.-Bl.
et al. 1939)
191 123
7
Subalpine
Fagus sylvatica
Fagetum subalpinum s. lato* (Saxifrago rotundifoliae-Fagenion Marinček et al.
1993)
74 42
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ZONAL FOREST VEGETATION OF BOSNIA AND HERZEGOVINA
ACTA BOT. CROAT. 76 (1), 2017 17
(Shipley 2010). The ﬁrst is used especially by plant geographers,
concentrating on the morphological properties of
plants, e.g., Raunkiaer’s life-forms (Raunkiaer 1934) and
how, on average, such morphologies change as a function
of major climatic variables. A second approach is based on the
notion of ecological ‘strategies’, for which Grime’s model
of CSR triangle (Grime 1974, 1977, 2001) is often used
(Pugnaire and Valladares 2007). This model is a classiﬁcation
based on how plants deal with two groups of external
factors, i.e., stress and disturbance, which results in three
primary plant strategies: competitors (C), stress-tolerators
(S), and ruderals (R) and four secondary, intermediate ones.
The position of each species, as well as each relevé, can be
determined in a CSR triangle. The whole community is
thus given a functional signature, which can be very useful
in comparative studies involving widely differing samples
(Hunt et al. 2004).
While the use of ecological strategies is quite common
in studies of herbaceous vegetation (Hunt et al. 2004, Zelnik
and Čarni 2008, Škornik et al. 2010, Pipenbaher et al.
2013), it has lately also been gaining ground in the study of
forest and woodland communities (Kilinç et al. 2010,
Paušič and Čarni 2012, Juvan et al. 2013, Košir et al. 2013b,
Rozman et al. 2013). Its use is most often related to studies
of changes in communities’ composition as a response to
disturbance, succession, environmental changes or in gradient
analysis.
The aim of this study was to test whether zonal vegetation
is an expression of macro-climatic conditions or whether
there are also other environmental factors involved. The underlying
assumptions were (1) that the macro-climatic gradient
is the most important for the species and structure
turnover along the gradient of ZFPCs in B&H; and (2) that
ZFPCs would demonstrate similarities in Grime’s ecological
strategies, due to the fact that the communities are in
terminal stages of succession, with a low level of degradation,
occupying habitats on moderately fertile soils.
Materials and methods
Data collection and preparation
Seven ZFPCs were identiﬁed for B&H (Tab. 1, Fig. 1,
Stefanović et al. 1983, Beus 1984): (1) eu-Mediterranean
evergreen Quercus ilex forests (a very small area of the
warmest, southernmost part of B&H; represented by
maquis); (2) sub-Mediterranean thermophilous deciduous
Quercus pubescens-Carpinus orientalis forests (a major
part of the lowland and hilly region in southern B&H); (3)
Central Balkans thermophilous Quercus frainetto forests
(relatively narrow lowland and hilly belt of eastern B&H);
(4) Illyrian mesophilous Quercus petraea-Carpinus betulus
forests (major part of northern B&H and marginally in central
B&H); (5) montane mesoneutrophilous pure Fagus or
mixed Fagus-Abies forests (altitudinal belt above oak forests
in all B&H but mainly in the mountainous region of
central B&H (Dinaric Alps); (6) alti-montane, colder and
more acidophilous mixed Picea-Abies-Fagus forests (altitudinal
belt above Community 5, mainly in the Dinaric
mountainous region); and (7) subalpine Fagus sylvatica
forests (uppermost forest altitudinal belt).
Relevés were extracted from the Forest vegetation database
of Bosnia and Herzegovina stored in the Global index
of vegetation-plot databases (Dengler et al. 2011) with the
ID EU-BA-001. This database consists of 2810 published
and available unpublished forest vegetation relevés in
B&H. We compiled all relevés that were assigned to one of
the seven ZFPCs by their authors (Tab. 1), with the exception
of two zonal communities of thermophilous deciduous
forests (Communities 2 and 3), for which we used the results
of formalized classiﬁcation (Stupar et al. 2015). We
did not consider relevés of stands with less than 70% of
canopy cover, nor those with ediﬁer tree species cover value
less than 3 (25%) on the Braun-Blanquet scale, considering
them structurally degraded. Only stands of high, productive
forests were thus taken into account, except in the
case of Community 1 (Quercus ilex stands) because wellestablished
stands do not exist in B&H, and all relevés were
made in structurally degraded maquis. All relevés were
made using the standard Central European phytosociological
method (Braun-Blanquet 1964). Only relevés that could
be georeferenced relatively precisely and those that contained
complete species records were taken into consideration.
In all 612 relevés were compiled in the Turboveg database
(Hennekens and Schaminée 2001) and exported to
JUICE 7 software (Tichý 2002) for further analysis.
Mosses, as well as taxa determined only to the genus
level, were removed from the data set prior to numerical
analysis. All vegetation layers were merged into one layer.
Taxonomy and nomenclature followed Flora Europaea (Tutin
et al. 1968–1993) unless a more modern taxon concept
or circumscription suggested otherwise. These taxa, as well
as those from taxonomically critical groups that were combined
into aggregates (agg.) or species that included several
subspecies that were not always recorded or recognized by
authors and were combined under the abbreviation ‘s.l.’
(sensu lato), were listed in On-line Suppl. Tab. 1. The dubious
taxon Quercus dalechampii was treated as part of Quercus
petraea agg., following Di Pietro et al. (2012). Records
of Fagus moesiaca were treated as F. sylvatica (Marinšek et
al. 2013).
To avoid geographic overrepresentation of some vegetation
types due to oversampling of certain regions, we performed
geographical stratiﬁcation and resampling of the
initial data set (Knollová et al. 2005), a frequent strategy in
recent national and regional level vegetation studies (Chytrý
2013, Košir et al. 2013a, Rodríguez-Rojo et al. 2014).
Stratiﬁcation was performed in a geographical grid with 1
km2
size. If two or more relevés assigned to the same community
fell in the same grid cell, only one of them was selected.
Stratiﬁcation was not performed on Community 1
since it consisted of only ﬁve relevés. The resulting stratiﬁed
data set contained 398 relevés and 669 species.
Data analysis
The data set was then subjected to detrended correspondence
analysis (DCA) in R software, version 2.10.1 (R Development
Core Team 2009) using the vegan package (http://
cc.oulu.ﬁ/~jarioksa/softhelp/vegan.html) on presence-absence
data. To extract the main gradients in species composition,
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STUPAR V., ČARNI A.
18 ACTA BOT. CROAT. 76 (1), 2017
398 relevés, together with the selected ecological variables
were projected onto the two-dimensional ordination space
of DCA. Unweighted average species ecological indicator
values (EIVs) for soil reaction (Pignatti et al. 2005) and selected
climatic variables available from the WorldClim database
(Hijmans et al. 2005) were used as explanatory ecological
variables. The signiﬁcance of EIVs correlation with
the DCA relevé scores was tested using the modiﬁed permutation
test proposed by Zelený and Schaffers (2012).
Climate variables best explaining variation in species composition
were selected through forward selection in canonical
correspondence analysis (CCA) in CANOCO 4.5 software
(Microcomputer Power, Ithaca, NY, US). Other explanatory
variables (altitude, inclination of slope, aspect, chorotypes,
latitude and longitude, life forms and ecological strategies)
were selected and projected based on the strength of correlation
with the ﬁrst and second DCA axis. The signiﬁcances
of correlations between these explanatory variables and
DCA relevé scores were calculated using the Kendall tau
coefﬁcient in Statistica software (StatSoft, Inc.; v 7.0, http://
www.statsoft.com). Chorological spectra of the zonal forest
communities (Gajić 1980, Pignatti et al. 2005) were calculated
for each relevé using presence-absence data. Endemic
Illyrian species were separated from southeast European
species in a broader sense.
In order to reveal the ﬂoristic differences among the
seven ZFPCs, we calculated their diagnostic species in the
resampled data set using phi coefﬁcient in the JUICE 7 program
(Chytrý et al. 2002), after standardization to a relevé
group size equal to a seventh of the total data set size (Tichý
and Chytrý 2006). Fisher’s exact test was calculated giving
a zero ﬁdelity value to species whose phi values were not
statistically signiﬁcant (P>0.001). The threshold phi value
for a species to be considered diagnostic was set at 0.25.
The functional study of zonal forest communities was
performed using data of plant functional traits, i.e., life
forms (Raunkiaer 1934) and ecological strategies (Grime
1977). Plants were classiﬁed according to Grime’s primary
and secondary strategies into seven functional types using
data from the BIOLFLOR database (Klotz et al. 2002). We
thus obtained data for about 80% of species. Ecological
strategies for the remaining species were calculated using
the dichotomous key suggested by Véla (2002). Averages
of each strategy type weighted by cover-percentage were
calculated and standardized for each relevé as suggested by
Hunt et al. (2004). Ecological strategy scores for each
group of relevés were then calculated with the CSR Signature
Calculator 1.2 program (Hunt et al. 2004) and represented
on a CSR ternary plot. We did a separate analysis for
all layers and for the herb layer alone because herb functional
signatures respond more quickly to environmental
changes in the course of late succession (Paušič and Čarni
2012). Plant life forms obtained from Pignatti et al. (2005)
and supplemented by our expert knowledge were used for
the calculation of life forms spectra for each relevé using
presence-absence data, and mean proportions were compared
between ZFPCs. The occurrence of an individual
functional trait in each of the ZFPCs was compared using
the Scheffé post hoc test for normal distributions and Kruskal-Wallis
test by ranks and median for non-normal distributions
using Statistica software. A normality check was
performed using Lilliefors test.
Results
Gradient analysis within ZFPCs in Bosnia and
Herzegovina
Forward selection suggested that the climatic variables
with the highest explanatory value of the variation in species
composition are annual mean temperature (BIO1),
mean temperature of the coldest quarter (BIO11) and precipitation
of the warmest quarter (BIO18). The ﬁrst DCA
axis represents the main gradient in the data set, and all
three climatic variables are signiﬁcantly related to the ﬁrst
DCA axis at P<0.001 (Tab. 2, Fig. 2). It runs from the coldTab.
2. Correlations (Kendall-Tau coefﬁcient) between detrended correspondence analysis (DCA) relevé scores and explanatory variables.
BIO1 – annual mean temperature; BIO11 – mean temperature of the coldest quarter; BIO18 – precipitation of the warmest quarter;
Eur – European and Eurasian; SEur – south and southeast European; EuriMed – Euri-Mediterranean; StMed – Steno-Mediterranean; Illyr
– Endemic Illyrian; Bor – Boreal; SEOro – South European orophytes; Cosm – Widespread species; P – Phanerophytes; NP – Nanophanerophytes;
Ch – Chamaephytes; H – Hemicryptophytes; G – Geophytes; T – Terophytes; C – Competitors; S – Stress-tolerators; R –
Ruderals. Asterisk (*) denotes signiﬁcant correlation at P<0.001.
Ecological factors Geographical factors
Altitude Aspect Slope BIO1 BIO11 BIO18 Latitude Longitude
DCA 1 –0.521* 0.046 0.050 0.569* 0.566* –0.451* –0.200* 0.367*
DCA 2 –0.151* –0.034 –0.103 0.118* 0.103 –0.011 0.028 –0.041
Chorotypes
Eur SEur EuriMed StMed Illyr Bor SEOro Cosm
DCA 1 0.088 0.203* 0.445* 0.309* –0.172* –0.313* –0.510* –0.155*
DCA 2 0.077 –0.034 0.035 –0.004 –0.066 –0.009 –0.138* 0.077
Life forms Ecological strategies
P NP Ch H G T C S R
DCA 1 0.182* 0.020 0.045 0.080 –0.317* 0.109 0.045 –0.048 0.014
DCA 2 0.010 –0.059 0.110 0.057 –0.121* 0.126* –0.049 –0.067 0.109
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ZONAL FOREST VEGETATION OF BOSNIA AND HERZEGOVINA
ACTA BOT. CROAT. 76 (1), 2017 19
est and most mesophilous subalpine beech forests (Community
7) on the left side of the diagram to the most xerothermophilous
Quercus ilex maquis (Community 1) on the
far right side of diagram. It is positively correlated with annual
mean temperature and mean temperature of the coldest
quarter, and negatively with the precipitation of the warmest
quarter (Fig. 2, Tab. 2). These are strong indicators that
the ﬁrst DCA axis represents a macro-climatic gradient that
runs from wet summers, cold winters and lower annual
temperatures to dry summers but warmer winters and higher
overall annual temperatures. There is also a high negative
correlation with altitude, which is again related to macro-climatic
factors. The positive correlation with longitude
and negative correlation with latitude reﬂects the fact that
south and east parts of B&H are warmer and dryer than the
north and west. Correlations between the ﬁrst axis and
slope and aspect are not statistically signiﬁcant (Tab. 2). A
modiﬁed permutation test also showed that the correlation
between DCA 1 and EIVs for soil reaction was not statistically
signiﬁcant (R2
=0.125, P=0.195). In terms of chorotypes,
DCA axis 1 shows a positive correlation with the
proportions of south- and southeast European, Euri-Mediterranean
and Steno-Mediterranean chorotypes, and a negative
correlation with Boreal and south European orophytic
chorotypes, while the correlation with the most abundant
European and Eurasian chorotypes is not statistically signiﬁcant.
Correlations between the ﬁrst axis and life forms
are signiﬁcant only for the proportions of phanerophytes
(positive) and geophytes (negative). There is no signiﬁcant
correlation between ecological strategies and DCA axes
(Tab. 2).
Floristic differentiation
Analysis revealed differences in the ﬂoristic composition
among the seven ZFPCs. Tab. 3 shows a simpliﬁed frequency-ﬁdelity
synoptic table of the ZFPCs of B&H based
on a data set of 398 resampled relevés (full version is given
in On-line Suppl. Tab. 2).
Fig. 2. Detrended correspondence analysis (DCA) spider plot of
398 relevés of zonal forest plant communities with climatic variables,
ecological indicator values (EIVs) for soil reaction and altitude
passively projected. The surface variable is annual mean temperature.
The length of the ﬁrst DCA axis is 7.02 SD units, the
length of the second axis is 2.85 SD units. Centroids of clusters
are indicated by numbers that refer to Tab. 1.
Tab. 3. Frequency-ﬁdelity table of zonal forest plant communities (ZFPCs) in Bosnia and Herzegovina. Frequencies of species are presented
as percentages with phi values multiplied by 100 shown in superscript. Diagnostic species (phi values higher than 0.25) for each
community are shaded (only ten species with the highest phi value for every community are presented). Proportions of chorotypes and
altitudinal ranges of each community are also shown. Community numbers correspond to those used in Tab. 1 and Fig. 1.
Community number 1 2 3 4 5 6 7
No. of relevés 5 16 24 26 162 123 42
Altitudinal range (m) 0–150 100–650 150–700 200–900 700–1400 800–1600 1400–1800
Chorotype (% of all species in a community)
Widespread species 3 1 3 5 7 7 4
European and Eurasian 13 29 56 67 59 55 49
south and southeast European 21 30 21 16 13 10 12
Eurimediterranean 29 29 13 5 1 1 0
Stenomediterranean 33 7 0 0 0 0 0
Endemic Illyrian 1 3 1 0 1 2 3
Boreal 0 1 6 6 10 13 13
South European orophytes 0 0 0 1 9 12 19
ZFPC 1
Quercus ilex 10096.5
6–
. . . . .
Arbutus unedo 8088
. . . . . .
Teucrium polium ssp. capitatum 8084.1
6–
. . . . .
Centaurium erythraea 8083.2
. . 8–
. . .
Pistacia terebinthus 10082.6
3818.3
. . . . .
Juniperus phoenicea 6075
. . . . . .
Pistacia lentiscus 6075
. . . . . .
Juniperus oxycedrus ssp. macrocarpa 4060.3
. . . . . .
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STUPAR V., ČARNI A.
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Community number 1 2 3 4 5 6 7
No. of relevés 5 16 24 26 162 123 42
Altitudinal range (m) 0–150 100–650 150–700 200–900 700–1400 800–1600 1400–1800
Crepis sancta 4060.3
. . . . . .
Cistus salvifolius 4060.3
. . . . . .
ZFPC 2
Quercus pubescens 20–
10084.5
12–
. . . .
Cornus mas . 8883.6
8–
8–
. . .
Acer monspessulanum . 6276.7
. . . . .
Sesleria autumnalis 20–
8176.5
. . 1–
1–
.
Frangula rupestris . 5672.4
. . . . .
Viola hirta . 7569.4
17–
12–
. . .
Rubus ulmifolius . 5067.9
. . . . .
Carex halleriana 20–
6264.2
. . . . .
Brachypodium sylvaticum . 7561.1
12–
19–
5–
6–
7–
Juniperus oxycedrus 40–
6960
. . . . .
ZFPC 3
Quercus frainetto . 19–
10090.3
. . . .
Chamaecytisus hirsutus agg. . . 5870.4
4–
. 1–
.
Quercus cerris . 5023
10069.8
27–
1–
1–
.
Thymus pulegioides . . 5465.7
. 4–
1–
2–
Lychnis coronaria . . 4664.8
. . . .
Lathyrus niger . 12–
5863
4–
. . .
Euphorbia cyparissias . 19–
5861.7
. . . .
Dianthus armeria . . 4261.6
. . . .
Carex caryophyllea . 19–
5857.8
8–
. . .
Silene viridiﬂora . 12–
4655.4
. . . .
ZFPC 4
Carpinus betulus . . 4219.6
10077.3
11–
. .
Cruciata glabra . . 4–
6967.5
11–
8–
.
Prunus avium . . 25–
6560.7
6–
1–
.
Luzula luzuloides . . . 5859.9
1–
8–
12–
Pteridium aquilinum . . 38–
7757.7
12–
15–
.
Stellaria holostea . . 8–
5056.6
1–
. 7–
Melampyrum pratense . . 17–
5054.5
. 1–
2–
Erythronium dens-canis . . . 2749
. . .
Crataegus monogyna . 44–
29–
6945.3
15–
2–
.
Hieracium racemosum . . . 2345.2
. . .
ZFPC 5
Galium odoratum . . . 12–
7646.5
59–
36–
Acer pseudoplatanus . . . 12–
8144.6
69–
52–
Cardamine bulbifera . . . 12–
6742.3
30–
55–
Lonicera xylosteum . . . . 3138.9
15–
2–
Cardamine enneaphyllos . . . . 6238.1
35–
62–
Daphne mezereum . . . . 5135.1
37–
38–
Salvia glutinosa . . 4–
4–
3134.9
14–
2–
Glechoma hirsuta . . 17–
19–
5034.6
26–
12–
Rubus hirtus . . 12–
42–
5733.1
36–
14–
Cardamine trifolia . . . . 2832.9
20–
2–
ZFPC 6
Picea abies . . . . 51–
10059.7
69–
Tab. 3. – continued
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ZONAL FOREST VEGETATION OF BOSNIA AND HERZEGOVINA
ACTA BOT. CROAT. 76 (1), 2017 21
Analysis of functional traits
In the standard CSR triangle plot, communities are positioned
in the upper right part of the diagram (Fig. 3a). This
means that zonal forest plant communities are dominated by
species with considerable competitive capacity. However,
there is apparent divergence of Communities 1, 2 and 4 from
the main cluster along the C–S axis, whereby statistically
signiﬁcant differences were discovered (Tab. 4). The highest
ratio of stress tolerant-species occurs in Community 1,
much less in Community 2, while the most competitive ability
is expressed by Community 4. CSR signatures of the herb
layer are shifted to the middle of the C–S axis (Fig. 3b).
The most signiﬁcant differences in functional traits of a
life form were detected for Phanerophytes and Geophytes
between Communities 1–4 and Communities 5–7, and for
Hemicryptophytes between Communities 1 and 5 and the
rest of the data set (Tab. 4). The highest ratios of PhaneroFig.
3. CSR triangle of the ordination of zonal forest plant communities
in Bosnia and Herzegovina according to ecological strategy
scores (proportion of competitors – C, stress-tolerators – S
and ruderals – R components in each community). Community
numbers correspond to those used in Tab. 1 and Fig. 1. (a) All layers
together; (b) Herb layer.
Community number 1 2 3 4 5 6 7
No. of relevés 5 16 24 26 162 123 42
Altitudinal range (m) 0–150 100–650 150–700 200–900 700–1400 800–1600 1400–1800
Abies alba . . . 8–
73–
9852.5
76–
Athyrium ﬁlix-femina . . . 8–
38–
6349
14–
Galium rotundifolium . . . . 9–
4146.5
12–
Oxalis acetosella . . . . 64–
8346.1
69–
Lonicera nigra . . . . 19–
4643.9
17–
Sorbus aucuparia . . . . 33–
6342.1
50–
Senecio nemorensis s.l. . . . . 38–
5940.8
38–
Prenanthes purpurea . . . 4–
46–
6838.9
67–
Lamiastrum galeobdolon . . . 27–
56–
7338.8
52–
ZFPC 7
Saxifraga rotundifolia . . . . 12–
24–
7666.7
Luzula sylvatica . . . . 3–
20–
6765.6
Adenostyles alliariae . . . . 5–
23–
6461.8
Valeriana montana . . . . 4–
8–
4555.3
Cicerbita alpina . . . . 1–
10–
4353.9
Veronica urticifolia . . . . 4–
23–
5252.8
Ranunculus platanifolius . . . . 2–
5–
3852.4
Astrantia major . . . . . 1–
3151.8
Homogyne alpina . . . . . . 2950.5
Veratrum lobelianum . . . . . . 2446
Species diagnostic for more than one community
Phillyrea latifolia 10078.2
5028.4
. . . . .
Clematis ﬂammula 8064.5
5033
. . . . .
Asparagus acutifolius 6040.5
8162.1
. . . . .
Carpinus orientalis . 8868.3
5434.5
. . . .
Fraxinus ornus 60–
10050.4
8336.5
31–
2–
1–
.
Genista tinctoria . . 5450.6
3526.7
1–
. .
Quercus petraea . . 7143.7
10071.3
1–
1–
.
Acer campestre . 12–
4227.5
5845.1
4–
. .
Fagus sylvatica . . 12–
46–
10039.8
10039.8
100–
Vaccinium myrtillus . . . 4–
12–
5740.3
5740.6
Tab. 3. – continued
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STUPAR V., ČARNI A.
22 ACTA BOT. CROAT. 76 (1), 2017
phytes and Nanophanerophytes are found in the ﬁrst two
communities, with a decline in Phanerophytes towards
Community 7 (Fig. 4). Beech-dominated forests (Communities
5–7) have twice the ratio of Geophytes compared to
oak-dominated forests (Communities 1–4). There is the
highest proportion of Chamaephytes in Community 1,
while the biggest share of Therophytes is found in Communities
1 and 3.
Discussion
Our study strongly suggests macro-climatically based
differentiation of ZFPCs in B&H. While gradient analysis
revealed a major inﬂuence of climatic factors on the species
turnover, there is little or no impact of topographic (slope
and aspect) or edaphic conditions (Fig. 2, Tab. 2), which is
congruent with the traditional understanding of zonal vegetation
(Dierschke 1994, Ellenberg 2009, Surina 2014).
Macro-climatic gradient is supported by the signiﬁcant correlation
with the geographic factors (altitude, longitude and
latitude), which are all determinants of climate on a larger
scale (Ellenberg 2009, Adams 2010). A recent study of zonal
forests of Korean Peninsula similarly showed that the
main factor discriminating individual forest types was temperature
(annual mean temperature, as well as temperature
extremes) followed by precipitation, altitude and aridity
(Černý et al. 2015). While chorotypes indicate units that describe
distribution patterns shared by several species, explaining
origins and history of development of particular
ﬂoras (Pignatti and Pignatti 2014, Passalacqua 2015), some
studies suggest that there is a correlation between chorotypes
and climatic conditions (Ferrer-Castán and Vetaas
2003, Abbate et al. 2012). In the case of our study, the proportion
of ‘warmer’ (Euri-Mediterranean and Steno-Mediterranean)
chorotypes increases in a direction of the main
gradient, while the proportion of ‘colder’ (Boreal and Orophytic)
chorotypes decreases (Tab. 3), which again suggests
the macro-climatic gradient.
Floristic analysis showed a clear separation of seven
ZFPCs in B&H. Community 1 (Quercus ilex maquis, Tab.
3, col. 1) occupies only a small area (about 20 km2
) in the
extreme south of B&H. It is represented by the secondary
succession stage of the eastern Adriatic eu-Mediterranean
zonal association Fraxino orni-Quercetum ilicis (Kutleša
and Lakušić 1964). Mean annual temperatures are above 15
°C (Fig. 2) and elevations 0–150 m. Diagnostic species are
mainly of Steno-Mediterranean, Euri-Mediterranean and
south and southeast European chorotypes. Diagnostic herb
species are mainly indicators of rocky habitats due to structural
degradation. Although in the last 50 years this community
has expanded its distribution area in B&H, due to
the abandonment of coppicing, burning and goat breeding,
there are still no high stands (Drešković et al. 2011). The
situation is similar, though a little better, in neighboring
Croatia (Vukelić 2012). Community 2 (Quercus pubesFig.
4. Proportions of life forms in zonal forest plant communities
in Bosnia and Herzegovina. Community numbers correspond to
those used in Tab. 1 and Fig. 1. P – Phanerophytes; NP – Nanophaneorphytes;
Ch – Chamaephytes; H – Hemicryptophytes; G –
Geophytes; T – Therophytes.
Tab. 4. Comparison of differences in functional traits occurrences between each pair of zonal forest plant communities (ZFPCs). Statistical
signiﬁcance established using Kruskal-Wallis test by ranks and median, except for R, H and G, for which the Scheffé post hoc test for
a normal distribution was used. Community numbers correspond to those used in Tab. 1 and Fig. 1. FT – Functional type; C – Competitors;
S – Stress tolerators; R – Ruderals; P – Phanerophytes; NP – Nanophanerophytes; Ch – Chamaephytes; H – Hemicryptophytes; G –
Geophytes; T – Therophytes. *** p<0.001; ** p<0.01; * p<0.05.
FT
Compared pairs of ZFPCs
1–2
1–3
1–4
1–5
1–6
1–7
2–3
2–4
2–5
2–6
2–7
3–4
3–5
3–6
3–7
4–5
4–6
4–7
5–6
5–7
6–7
C *** * ** * ** *** *
S ** *** ** * ** *** *** * *** ** ***
R ** *** **
P * *** * ** *** *** ** *** *** *** ***
NP *** *** *** * ***
Ch ** * ** *** * * **
H *** *** ** ** *** ** *** * *** *** *** ***
G *** ** *** *** *** *** *** *** *** *** *** *** *** *
T * ** *** ** ***
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ZONAL FOREST VEGETATION OF BOSNIA AND HERZEGOVINA
ACTA BOT. CROAT. 76 (1), 2017 23
cens-Carpinus orientalis forests, Tab. 3, col. 2) is represented
by high, not degraded stands of zonal thermophilous
deciduous forest of the association Querco pubescenti-Carpinetum
orientalis (Stupar et al. 2015) found in the lowlands
and hilly area of sub-Mediterranean B&H. However,
due to the negative human impact, they have mainly been
replaced by the secondary scrub community Rusco aculeati-Carpinetum
orientalis (Muratspahić et al. 1991),
while their present distribution area is restricted to small
patches of mainly private old groves (Stupar et al. 2015).
Mean annual temperatures are between 12 and 15 °C (Fig.
2), occupying elevations between 100 and 650 m. Diagnostic
species are thermophilous and xerophilous plants of
mainly S/SE European or Euri-Mediterranean distribution,
although some widespread nemoral herbs and shrubs appear
with high frequency, e.g.: Brachypodium sylvaticum,
Dactylis glomerata, Veronica chamaedrys, Crataegus monogyna,
Hedera helix etc., indicating more mesophilous
microclimatic conditions under the closed canopy. Community
3 (Quercus frainetto forests, Tab. 3, col. 3) is found
in the eastern parts of B&H, where the inﬂuence of the continental
drier climate increases. This is a zonal community
of Central Balkans lowlands and hilly areas (Horvat et al.
1974) and is represented by the fairly heterogeneous association
Quercetum frainetto-cerridis (Tomić and Rakonjac
2013, Stupar et al. 2015). Although we included in the analysis
only stands with 70% and more canopy cover, these
forests are mainly found in the proximity of human settlements,
so they are frequently degraded by grazing and
browsing, as well as occasional ﬁres. Mean annual temperatures
are between 10 and 12 °C (Fig. 2), and elevations are
between 150 and 700 m. They are dominated by a mixture
of thermophilous, acid tolerant and widespread nemoral
species, as well as some more light-demanding herbs, indicating
wood pasture and cutting. Community 4 is represented
by sessile oak-common hornbeam forests, which are
considered a zonal forest vegetation community for the area
of NW Balkans (Quercus petraea-Carpinus betulus forests,
Tab. 3, col. 4). In B&H they are found mainly in the hilly
area of northern parts of the country but can also penetrate
deeper into the south (e.g., Central Bosnian basin, Fig. 1).
Habitats of these forests have been under intensive anthropogenic
inﬂuence since the Neolithic (Horvat et al. 1974),
so it is hard to ﬁnd stands of high, productive forests with
Quercus petraea cover value in the tree layer more than 3
on the Braun-Blanquet scale (25–50% of cover) and more
than 70% of overall canopy cover. They are often, due to
bad management, structurally degraded to Carpinus betulus
coppice. Mean annual temperatures are between 9 and 10
°C (Fig. 2) and they occupy elevations between 200 and
900 m. Several types of these forests occur in B&H but neither
the syntaxonomy nor nomenclature has been settled
(Lakušić et al. 1978, Redžić 2007). In addition, there is disagreement
about the syntaxonomical position of these forests
on the regional level. While Croatian and Slovenian
authors assign these forest to the Illyrian alliance Erythronio-Carpinion
betuli (Vukelić 2012, Košir et al. 2013a),
some other authors question this view, arguing that the
group is weakly characterized by diagnostic species (Willner
and Grabherr 2007, Borhidi et al. 2012). This is sustained
by our results, in the sense that this ZFPC is the only
one in B&H that completely lacks endemic Illyrian species
(Tab. 3). Diagnostic species are mainly represented by mesophilous
European and Eurasian elements. Community 5
(pure Fagus / mixed Fagus-Abies forests, Tab. 3, col. 5)
represents the ﬁrst altitudinal belt above oak forests and is
made up of mesoneutrophilous montane pure beech, as well
as mixed ﬁr-beech forests. The syntaxonomy and nomenclature
of these forests have not yet been the subject of
thorough analysis but two main types, or widely understood
associations, are included: Fagetum montanum illyricum
and Abieti-Fagetum dinaricum (Beus 1984). They belong
to the alliance Aremonio-Fagion (Marinšek et al. 2013).
Mean annual temperatures are between 6 and 8 °C (Fig. 2)
and they are mainly found at elevations between 700 and
1400 m. Floristically and ecologically similar to this type is
Community 6 (mixed Picea-Abies-Fagus forests Tab. 3,
col. 6), which also has a considerable amount of Boreal and
south European orophytic elements, while annual mean
temperatures are between 5 and 7 °C (Fig. 2). In the view of
Beus (1984), which is supported by our results, the main
difference between Community 6 and the previous community
is that Community 6 has a higher proportion of spruce
(100% frequency) and acidophilous Vaccinio-Piceetea species,
while the proportion of mesoneutrophilous AremonioFagion
species is remarkably lower. In addition, it comes as
an altitudinal belt above Community 5 (elevations between
800 and 1600 m) and can be found exclusively in the central
range of Dinaric Alps, while Community 5 extends
more to the north and south (Fig. 1). This is explained by
the fact that the hot dry summers of the northern and southern
chains of Dinaric Alps do not favor spruce (Beus 1984).
The syntaxonomy and nomenclature of Community 6 have
also not been settled, so in B&H this complex type is identiﬁed
under the invalid provisional name Piceo-Abieti-Fagetum.
Community 7 is the highest altitudinal belt, represented
by subalpine Fagus sylvatica forests (Tab. 3, col. 7).
This is yet another group the syntaxonomy and nomenclature
of which in B&H has not been analyzed and it is known
by the generic name of Fagetum subalpinum s. lato. These
forests belong to the suballiance Saxifrago rotundifoliaeFagenion
of the alliance Aremonio-Fagion (Marinšek et al.
2013). These are mainly mesoneutrophilous pure beech forests
found mainly on the mountains of the central and
southern chains of the Dinaric Alps (Fig.1). Mean annual
temperatures are between 5 and 6 °C (Fig. 2), with elevations
between 1400 and 1800 m. This community harbors
the highest proportion (32%) of Boreal and south European
orophytic elements.
Functional analysis of ecological strategies showed
small overall differences among ZFPCs. Based on the communities’
locations on the CSR triangle, competitive strategy
(C) is dominant in all seven communities, followed by
CSR and SC strategies (Fig. 3). Considering the course of
succession (Fig. 3a), site production is high (Grime 1974),
while the importance of ruderality is insigniﬁcant, which
indicates late stages of succession, as pointed out by Prévosto
et al. (2011). The terminal stage of zonal communiBrought
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STUPAR V., ČARNI A.
24 ACTA BOT. CROAT. 76 (1), 2017
ties is supported by the position of the communities on the
CSR plot when only the herb layer was analyzed (Fig. 3b).
In this plot, stress tolerance becomes more important since
shading and lack of nutrients in the herb layer coincide with
the development of large long-lived forest trees (Grime
1977, 1988). Only Community 1 (Quercus ilex maquis),
which has the largest share of ruderals, expresses conditions
of moderate productivity and middle to late stage of
secondary succession (maquis). The increased proportion
of stress-tolerators in Community 1 and also, to a much
lesser extent, in Community 2 is related to drought stress
during the summer season in the Mediterranean limestone
region of B&H. On the other hand, Community 4 (Quercus
petraea-Carpinus betulus forests) has an increased proportion
of competitors, which indicates higher productive capacities.
Also, this community lies in the middle of the
macro-climatic gradient (Fig. 2), which altogether suggests
the most favorable conditions for forest vegetation development
in B&H.
Changes in the life form proportions of different communities
are expressed in the greater share of woody species
in the more thermophilous vegetation types, which corresponds
with the statement that ‘the Phanerophyte is the
plant type that belongs to warm regions’ (Raunkiaer 1934).
Similar results were obtained in the study of the gradient
from warm to mesic temperate forests on the Galičica
mountain range in Macedonia (Čarni et al. 2016). The proportion
of Geophytes is higher in Communities 5–7 (beech
dominated forests), which have spring ephemerals (mainly
bulbous Geophytes), also abundant in oak communities
(Popović et al. 2016), in addition to harboring a considerable
number of non-ephemeral rhizomatous ferns and species
from the family Liliaceae. This is congruent with the
notion that rhizomatous Geophytes are adapted to life in regions
with a severe unfavorable period (e.g., hard winters),
but have at the same time a long period of vegetation
(Raunkiaer 1934). The proportion of Therophytes is insigniﬁcant
except in Communities 1 and 3, which can be explained
by water shortage in the conditions of harsher Mediterranean
and continental climates (Kavgaci et al. 2012,
Raju et al. 2014). The same can be said for the proportion
of Chamaephytes in Community 1. On the other hand, according
to Bloch-Petersen et al. (2006) plant life forms differentiate
not only due to climatic variations, but seem also
to relate to humane disturbance and management. Prasad
(1995) studied the effects of grazing on the plant species
and life form composition and found that the percentage of
Therophytes was higher on grazed areas than on protected
areas, which reﬂects the disturbance through grazing in
Communities 1 and 3.
To conclude, the results of our analysis support the hypothesis
that zonal forest vegetation in B&H is an expression
of macro-climatic conditions and that there are no remarkable
differences in CSR plant strategies among the
ZFPCs in B&H (excluding Community 1). However, having
in mind that the currently protected forest area in B&H
is very small and covers only a few types of ZFPCs (Stupar
2011), in order to facilitate further research into the ecological,
ﬂoristic and functional characteristics of ZFPCs there is
a need for establishment of a national network of protected
areas which would cover all ZFPC types, thus preserving
the representative stands in as natural conditions as possible
(Milanović et al. 2015). Particular emphasis should be given
to thermophilous forests, especially to Community 1
(stands of Quercus ilex) which should be converted from
maquis to high forests.
Acknowledgements
The research was partly carried out by the BH-SI bilateral
project »Vegetation of thermophilous oak forests of the
Pre-Pannonian area of the Western Balkans« no. BI-BA/
12-13-015. AČ also acknowledges the ﬁnancial support of
the Slovenian Research Agency (P1-0236). We would like
to express our gratitude to the Institute of Biology, Research
Center of the Slovenian Academy of Sciences and Arts
(ZRC SAZU) and Forestry Faculty, University of Banja
Luka for logistic support.
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