Received: 21 February 2021
|
Revised: 10 May 2021
|
Accepted: 14 May 2021
DOI: 10.1111/avsc.12589
Applied Vegetation Science
V E G E TAT I O N S U R V E Y
Classification of forest and shrubland vegetation in
Mediterranean Turkey
Ali Kavgacı1
| Neslihan Balpınar2
| Hafize Handan Öner3
Münevver Arslan4
| Gianmaria Bonari5,6
| Milan Chytrý6
1
Faculty of Forestry, Karabuk University,
Karabuk, Turkey
Faculty of Arts and Sciences, Burdur
Mehmet Akif Ersoy University, Burdur,
Turkey
2
Aegean Forestry Research Institute, İzmir,
Turkey
3
4
|
| Andraž Čarni7,8
Abstract
Questions: What are the main vegetation types of forest and shrubland vegetation
at the alliance level in Mediterranean Turkey? What is their syntaxonomical position?
Can we integrate them into the European vegetation classification system? Which
environmental factors are the main drivers of the floristic differentiation of vegeta-
Research Institute for Forest Soil and
Ecology, Eskişehir, Turkey
tion types?
Faculty of Science and Technology, Free
University of Bozen-Bolzano, BozenBolzano, Italy
Methods: We collected 4,717 vegetation plots of forest and shrubland vegetation
5
Department of Botany and Zoology, Faculty
of Science, Masaryk University, Brno, Czech
Republic
6
7
Research Center of the Slovenian Academy
of Sciences and Arts, Institute of Biology,
Ljubljana, Slovenia
School for Viticulture and Enology,
University of Nova Gorica, Nova Gorica,
Slovenia
8
Correspondence
Ali Kavgacı, Faculty of Forestry, Karabuk
University, Demir Çelik Campus, 78050,
Karabuk, Turkey.
Email: alikavgaci@karabuk.edu.tr
Funding information
AK, NB and MA were funded by TÜBİTAK214O670, GB by the post-doc programme
of the Department of Botany and Zoology,
Masaryk University, and by the Free
University of Bozen-Bolzano through the
CONplant project (TN201H); MC by the
Czech Science Foundation (19-28491X)
and AČ by the Slovenian Research Agency
(ARRS, P1- 0236).
Co-ordinating Editor: Florian Jansen
Location: Southern and western Turkey.
in Mediterranean Turkey and performed an unsupervised classification of this data
set. We described vegetation types based on the classification results, expert knowledge and information from the literature. We defined diagnostic species and prepared distribution maps for each vegetation type. To support the interpretation of
the vegetation types, we determined the most important environmental variables
using canonical correspondence analysis.
Results: The studied vegetation was divided into 21 types related to three vegetation belts: (a) thermo- and meso-mediterranean, comprising coniferous (Pinus brutia,
Pinus pinea) and sclerophyllous forests, as well as macchia, garrigue and phrygana;
(b) supra-mediterranean, comprising Pinus nigra subsp. pallasiana forests, thermophilous deciduous forests dominated by various oak species and Ostrya carpinifolia,
and forests dominated by temperate species such as Fagus orientalis; and (c) oromediterranean, comprising forests and shrublands dominated by Abies cilicica, Cedrus
libani, Juniperus excelsa and Juniperus communis subsp. nana. Elevation was identified
as the main environmental driver of the vegetation pattern. Among climatic variables,
the most important are the mean temperatures (annual and of driest, coldest, and
warmest quarters), minimum temperature of winter, precipitation of warmest and
driest quarters and precipitation seasonality. These factors indicate the decreasing
effect of the Mediterranean climate with increasing elevation.
Conclusions: The vegetation of Mediterranean Turkey is arranged along climatic gradients depending on elevation and the distance from the Mediterranean Sea. Most
This is an open access article under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs License, which permits use and distribution in
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© 2021 The Authors. Applied Vegetation Science published by John Wiley & Sons Ltd on behalf of International Association for Vegetation Science
Appl Veg Sci. 2021;24:e12589.
https://doi.org/10.1111/avsc.12589
wileyonlinelibrary.com/journal/avsc
|
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KAVGACI et Al.
vegetation types in this area correspond to the syntaxa accepted in EuroVegChecklist,
while others were described as new.
KEYWORDS
coniferous forest, deciduous forest, garrigue, macchia, Mediterranean, numerical analysis,
phrygana, plant communities, shrubland, Turkey, vegetation-plot database, vegetation types
1 | I NTRO D U C TI O N
of the integrated database European Vegetation Archive (EVA;
Chytrý et al., 2016) and subsequently of the global database sPlot
The Mediterranean Basin is one of the global biodiversity hotspots
(Bruelheide et al., 2019), which are still growing. One of the national
(Myers et al., 2000). Its vegetation is mainly dominated by broad-
databases developed under these initiatives is the Forest Vegetation
leaved and needle-leaved evergreen tree species, but decidu-
Database of Turkey (GIDV ID: 00-TR- 001), which now includes more
ous trees are also well represented (Gauquelin et al., 2016). The
than 8,500 vegetation plots and aims at collecting all vegetation
knowledge on vegetation diversity varies across this area. The
plots of forest and shrubland vegetation in Turkey (Kavgacı et al.,
floristic and vegetation diversity of the European part of the
2012; Kavgacı & Čarni, 2012).
Mediterranean Basin has been studied extensively (Mucina et al.,
Vegetation plots from the Mediterranean phytogeographical re-
2016), whereas it is still insufficiently explored in the Asian and
gion are the most abundant in the Forest Vegetation Database of
African parts, making it difficult to develop international conser-
Turkey. They represent vegetation across the whole of this region,
vation strategies.
being an invaluable resource for studying and understanding the flo-
The Mediterranean phytogeographical region in Turkey covers
ristic and vegetation diversity of Mediterranean Turkey.
mainly the western (Aegean) and southern parts of Turkey, while
The aims of this study are to: (a) define the main vegetation types
northern Turkey belongs to the Euro-Siberian region and the cen-
of forests and shrublands in Mediterranean Turkey; (b) identify the
tral and eastern part of Turkey to the Irano-Turanian region (Davis,
main environmental variables affecting the diversity of this vegeta-
1971). Southern Mediterranean Turkey with Cyprus is considered
tion; and (c) assess the syntaxonomy of these vegetation types and
as one of ten biodiversity hotspots in the Mediterranean Basin
integrate them into the European vegetation classification system
(Médail & Quézel, 1999). Southern Mediterranean Turkey is the
(Mucina et al., 2016).
richest in endemism among all the Turkish regions (Noroozi et al.,
2019). Like other parts of the Mediterranean Basin, Mediterranean
Turkey comprises human-shaped ecosystems (Blondel, 2006),
2 | M E TH O DS
and it is additionally affected by climate change and related factors, especially the changing fire regime (Viedma et al., 2017).
Our study area is Mediterranean Turkey as defined on the phyto-
Understanding the vegetation diversity of Mediterranean Turkey
geographical map of Davis (1971). However, we also included the
has important implications for nature conservation and ecological
Mediterranean vegetation appearing in inner Anatolia, represent-
restoration.
ing the transition between the Mediterranean and Irano-Turanian
The first vegetation studies based on the Braun-Blanquet ap-
phytogeographical regions. We first checked the Forest Vegetation
proach in Turkey were performed in the 1940s (Ekim & Akman,
Database of Turkey, which was stored in the TURBOVEG (2.149a)
1991). Research intensified during the 1960s and 1970s, especially
database management program (Hennekens & Schaminée, 2001),
in collaboration with the French researchers Pierre Quézel and
and entered missing published and unpublished vegetation plots
Marcel Barbero (Akman et al., 1978, 1979a, 1979b). Since then, the
from forests and shrublands of Mediterranean Turkey. All vegetation
number of phytosociological studies covering different parts of the
plots in the data set were sampled according to the Braun-Blanquet
country has increased and syntaxonomic checklists were prepared
sampling method (Braun-Blanquet, 1964). The species taxonomy
(Quézel et al., 1993; Ketenoğlu et al., 2010). The development of nu-
and nomenclature in the database was unified to follow the Flora of
merical techniques in vegetation science has also positively affected
Turkey (Davis, 1965-1985; Davis et al., 1988; Güner et al., 2000; see
vegetation studies in Turkey. These techniques have been used in-
Appendix S1 for the vegetation data sources).
tensively both in large-scale and regional phytosociological studies
For the analyses, we selected only vegetation plots containing
(Uğurlu et al., 2012; Çoban & Willner, 2019; Karaköse, 2019; Yildirim
a tree or shrub layer with a cover of at least 25%. The selection
et al., 2019; Kenar et al., 2020).
resulted in a total of 4,717 vegetation plots. All records of species
Analysis of vegetation and habitat diversity crucially depends on
represented also by various subspecies and varieties in the data set
the availability of vegetation-plot data (Dengler et al., 2011; Chytrý
were merged to the species level. However, if a species was repre-
et al., 2016, 2020). Therefore, many countries have created their
sented only by a single subspecies or variety in the data set, these
own national vegetation-plot databases over the last decades. The
infraspecific taxa were retained. We removed species identified to
process of aggregation of these databases resulted in the creation
the genus or family level, cryptogams, vegetation plots from azonal
Applied Vegetation Science |
KAVGACI et Al.
vegetation (especially riparian vegetation dominated by Alnus orientalis, Liquidambar orientalis and Platanus orientalis), and vegetation
plots with three or fewer species. Additionally, we merged species
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3 | R E S U LT S
3.1 | Hierarchical classification
records from different layers into a single layer because this information was not consistently recorded among studies. As a result, the
A
classification
dendrogram
of
forests
and
shrublands
in
data set included 4,071 vegetation plots and 2,010 species. These
Mediterranean Turkey is shown in Figure 1. We accepted a division
data were used for the subsequent classification and ordination
into 18 main clusters that correspond to the phytosociological alli-
analyses. We used the JUICE 6.4 program (Tichý, 2002) for data set
ances of forests and shrublands in Mediterranean Turkey. However,
editing.
three of them were further divided into two types due to physiogno-
Before the numerical analyses, square-root transformation
mic (Cluster 2), biogeographical (Cluster 4) and elevational (Cluster
was applied to the percentage values of species cover. We used
7) differences. Consequently, we obtained 21 vegetation types. The
the Sørensen dissimilarity index as a resemblance measure and
classification of Mediterranean forests and shrublands is presented
Beta Flexible clustering (β = −0.25) using the PC- ORD program
in a synoptic table (Table 1 and Appendix S2), in which statistically
(McCune & Mefford, 2006). Beta Flexible clustering is one of the
determined diagnostic species are indicated and ranked by decreas-
most commonly used plot-grouping algorithms to define vegeta-
ing fidelity (phi coefficient). The distribution of each vegetation type
tion types at different hierarchical levels (De Cáceres et al., 2015).
is shown in Figure 2. Photos of typical stands of each vegetation
Vegetation types were based on the clustering results. The num-
type are provided in Figure 3.
ber of clusters representing the vegetation units was determined
Cluster 1 includes low, thorny and chamaephytic commu-
according to the dominant species of clusters conforming to vege-
nities, namely phrygana. This vegetation type is dominated by
tation types at the alliance level in Mediterranean Turkey. In some
Sarcopoterium spinosum. It is characterized by a high number of
cases, we also integrated expert knowledge and information from
diagnostic species. Although these phrygana communities mainly
the literature to divide some clusters into narrower types based on
appear in the Aegean region, they also occur in southern Turkey
the physiognomic, biogeographical, and elevational differences.
(Figure 2-1).
For the separation of these clusters, dominant species responsible
Cluster 2 is represented by garrigue communities dominated by
for the contrasting vegetation structure and subgrouping of clus-
Cistus creticus and Genista acanthoclada. However, Pinus pinea for-
ters for biogeographical and elevational differences were used.
ests were also placed within this cluster due to their floristic simi-
Vegetation plots were moved from one cluster to another only if
larities to garrigue despite an entirely different physiognomy. Since
these two clusters had contrasting vegetation structure (32 veg-
the physiognomic aspect should also be considered in classifica-
etation plots).
tion, in addition to floristic features, we divided Cluster 2 into two
Diagnostic species of vegetation types were defined by calculat-
groups. Cistus creticus and Genista acanthoclada-dominated garrigue
ing the fidelity of each species to each vegetation type (Chytrý et al.,
is a xerophilous dwarf scrub distributed in lowlands throughout
2002) using the phi coefficient of association as fidelity measure.
Mediterranean Turkey (Figure 2-2a). Pinus pinea-dominated forests
The species with a φ-value higher than 0.20 were considered as di-
are scattered throughout Mediterranean Turkey, including both nat-
agnostic. The phi coefficient was calculated for an equalized size of
ural stands and plantations (Figure 2-2b).
clusters (Tichý & Chytrý, 2006). Species with a cover of more than
Cluster 3 corresponds to another garrigue type in Mediterranean
25% in at least 10% of the vegetation plots in each type were con-
Turkey, which is dominated by Erica manipuliflora, one of the typical
sidered as dominant species.
low-shrub species of the region. These garrigues mainly occur in the
We used canonical correspondence analysis (CCA) with a Monte
Carlo test with 999 unrestricted permutations to test the effects
lowlands in the southern Aegean region and western part of southern Turkey (Figure 2-3).
of environmental variables on the floristic composition of vegeta-
Cluster 4 is also divided into two groups. First, Arbutus andrachne,
tion. Elevation and climatic data (from WorldClim 2; Fick & Hijmans,
Quercus coccifera and Juniperus excelsa-dominated macchia and
2017) were used as environmental variables. We also assessed the
sclerophyllous forests occur especially in lowland to submontane
gradients in species composition of the vegetation types using an
belts of the southern Aegean and southern Turkey, but they are also
indirect ordination method, detrended correspondence analysis
found in the northern part of Mediterranean Turkey (Figure 2-4a).
(DCA). Both CCA and DCA are suitable to analyse heterogeneous
Second, Quercus ilex-dominated shrublands and sclerophyllous for-
species-by-plot matrices like ours because they assume unimodal
ests dominated by Quercus ilex occur throughout the precipitation-
response of species to the environment. The most significant en-
richer Aegean region (Figure 2-4b); Arbutus andrachne and Quercus
vironmental variables (according to CCA) were passively projected
coccifera also appear as co-dominant in this vegetation type.
onto the DCA ordination space. Both CCA and DCA were computed
using CANOCO 5 (Šmilauer & Lepš, 2014).
We followed the fourth edition of the International Code of
Cluster 5 corresponds to Olea europaea and Phillyrea latifoliadominated macchia and sclerophyllous forests. Ceratonia siliqua occurs in this vegetation as a diagnostic species. This vegetation type
Phytosociological Nomenclature (ICPN; Theurillat et al., 2021) for
has an extensive distribution in the lowland and low-mountain vege-
nomenclature and typification of syntaxa.
tation belts of southern Turkey (Figure 2-5).
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| Applied Vegetation Science
KAVGACI et Al.
F I G U R E 1 Hierarchical dendrogram of forests and shrublands in Mediterranean Turkey. Dominant species of each cluster are mentioned
in the figure. In addition, Pinus pinea and Quercus ilex (in parentheses) are dominant in subgroups of clusters 2 and 4, respectively. Ceratonia
siliqua is a diagnostic species and an important element for Cluster 5. Red line: thermo- and meso-mediterranean vegetation; yellow line:
supra-mediterranean vegetation; blue line: oro-mediterranean vegetation. Legend: A.: Arbutus, C.: Castanea, F.: Fagus, J.: Juniperus, P.: Pinus,
Q.: Quercus [Colour figure can be viewed at wileyonlinelibrary.com]
Cluster 6 is another Mediterranean evergreen oak-dominated
vegetation type formed by Quercus coccifera. The most charac-
vegetation type. These forests appear in the central and western
Taurus Mountains in southern Turkey (Figure 2-9).
teristic feature of this cluster, in comparison with the previous
Cluster 10 comprises another deciduous oak vegetation type,
Quercus coccifera-dominated vegetation types (4a and 4b), is a lower
which is dominated by Quercus cerris. Styrax officinalis locally dom-
number of macchia shrub and tree species. Additionally, this vegeta-
inates the shrub layer. This vegetation type has a scattered distri-
tion type is more common at higher elevation belts than the previous
bution across a broad geographic range in Mediterranean Turkey
ones. It is distributed throughout Mediterranean Turkey (Figure 2-6)
(Figure 2-10).
and appears across a broad elevational gradient from lowland to
highlands.
Cluster 7 is composed of pure Pinus brutia-dominated forests
throughout Mediterranean Turkey. These forests occur from the
Cluster 11 represents shrublands of the mountain belt dominated by Cistus laurifolius. These shrublands are associated with
Pinus nigra and Quercus cerris-dominated forests and occur mostly in
the inner parts of Mediterranean Turkey (Figure 2-11).
sea level to the montane belt, and contain many garrigue, macchia
Cluster 12 corresponds to Pinus nigra forests. Quercus cerris and
and sclerophyllous forest species, such as Cistus creticus, Phillyrea
Cistus laurifolius are often present in this vegetation, in addition
latifolia, Pistacia terebinthus, Quercus coccifera, Quercus infecto-
to Juniperus oxycedrus. These forests have an extensive distribu-
ria and Styrax officinalis. Pinus brutia forests in Mediterranean
tion range in the inner parts of Mediterranean Turkey throughout
Turkey are represented by two different alliances: thermo- to
the Aegean part and the Taurus Mountains in southern Turkey
meso-mediterranean Pinus brutia forests and meso- to supra-
(Figure 2-12).
mediterranean Pinus brutia forests. Thus, we divided Cluster 7 into
Cluster 13 includes forests dominated by temperate tree spe-
lowland and upland Pinus brutia forests in accordance with the
cies, namely Castanea sativa, Fagus orientalis and Pinus sylvestris.
further subdivision of this cluster in the dendrogram (Figure 2-7a
Pinus nigra is also a dominant tree species for this group. However,
and -7b).
other temperate tree species, such as Abies nordmanniana subsp.
Cluster 8 corresponds to deciduous oak forests in Mediterranean
equi-trojani, Carpinus betulus, Corylus avellana and Quercus petraea
Turkey dominated by Quercus ithaburensis subsp. macrolepis and
appear as diagnostic species for this cluster. These forests mainly
Quercus infectoria. This vegetation type ranges from the lowland
occur in the Aegean region. However, they are also distributed in the
to mountain elevational belt throughout Mediterranean Turkey
Amanos Mountains and nearby areas in the eastern part of southern
(Figure 2-8).
Mediterranean Turkey (Figure 2-13).
Cluster 9 mainly consists of Ostrya carpinifolia-dominated for-
Cluster 14 represents relict extrazonal deciduous Populus trem-
ests. In addition, deciduous oak forests of Quercus trojana and
ula forests. These forests were recorded in two areas, one in the
Quercus vulcanica appear in this cluster. Deciduous trees and
northeastern part of Aegean Turkey, which is in contact with the
shrubs such as Acer hyrcanum, Acer monspessulanum, Cornus san-
Euro-Siberian biogeographical region, and the other in the Amanos
guinea and Fraxinus ornus are among the diagnostic species of this
Mountains in the eastern part of southern Mediterranean Turkey,
Applied Vegetation Science |
KAVGACI et Al.
which is also characterized by a significant occurrence of EuroSiberian flora (Figure 2-14).
5 of 29
The second group includes forests of Ostrya carpinifolia,
Pinus nigra, Populus tremula and Quercus cerris. Additionally, Cistus
Cluster 15 represents shrublands dominated by Juniperus com-
laurifolius-dominated shrublands and forests dominated by temper-
munis subsp. nana. This vegetation type mainly occurs in high moun-
ate trees have similar environmental conditions. This group is related
tains in the inner parts of the Aegean region and the western part of
to high precipitation of the driest month and warmest quarter, in-
southern Mediterranean Turkey (Figure 2-15).
dicating the supra-mediterranean affinity of this vegetation group.
Cluster 16 includes Juniperus excelsa-dominated forests oc-
The third group is represented by the forests dominated by Abies
curring at intermediate and high elevations of the Mediterranean
cilicica, Cedrus libani, Juniperus excelsa and Juniperus communis subsp.
mountains. These forests can reach the timberline in some areas.
nana shrubland. These forests and shrublands occur at high eleva-
Quercus coccifera and Juniperus oxycedrus are often present in these
tions of the Mediterranean mountains (oro-mediterranean belt).
forests, especially at lower elevations. These forests appear over
Forests of Quercus ithaburensis subsp. macrolepis and Quercus in-
large areas in the western and central Taurus Mountains in southern
fectoria are in the centre of the ordination space, corresponding to
Mediterranean Turkey and in the inner part of the southern Aegean
intermediate environmental conditions.
(Figure 2-16).
Cluster 17 consists of mountain and high-mountain forests dominated by Abies cilicica and Cedrus libani occurring in the central and
eastern Taurus Mountains in southern Turkey. These forests cover
extensive areas in the region characterized by the Mediterranean
4 | D I S CU S S I O N
4.1 | Broad vegetation patterns
mountain climate (Figure 2-17). Juniperus excelsa and Juniperus oxycedrus are also common in this vegetation type.
We identified 21 main vegetation types of forests and shrublands
Cluster 18 characterizes mountain and high-mountain for-
in Mediterranean Turkey. Elevation appeared as the most important
ests dominated by Cedrus libani in the western Taurus Mountains
factor affecting the variation in this vegetation in Mediterranean
(Figure 2-18). The absence of Abies cilicica in this group may be
Turkey, followed by macroclimatic factors. In other parts of the
due to wetter site conditions in this western type compared with
Mediterranean Basin, elevation is also an important factor affecting
the previous cluster. Juniperus excelsa is often present in these
climate and, consequently, the development of different vegetation
forests.
types (Blondel & Aronson, 1999; Quézel & Médail, 2003; Gauquelin
et al., 2016; Médail et al., 2019). It shows the correlation between
3.2 | Environmental gradients
elevation, climatic variables and vegetation.
This elevational and macroclimatic variation in Mediterranean
Turkey is reflected in the differentiation of three distinct vegetation
Canonical correspondence analysis (CCA) showed that elevation
belts: (a) thermo- and meso-mediterranean vegetation represented
and all bioclimatic variables are significantly correlated to the floris-
by phrygana, garrigue, macchia, sclerophyllous forests and pine
tic composition of Mediterranean forests and shrublands in Turkey
(Pinus pinea and Pinus brutia) forests; (b) supra-mediterranean pine
(Table 2). Elevation is the most important ecological factor, causing
(Pinus nigra subsp. pallasiana) and deciduous forests (Quercus cerris,
clear elevational differences between vegetation types (Figure 4).
Quercus trojana, Quercus vulcanica, Ostrya carpinifolia and Populus
The elevation is followed by mean temperatures (annual and of dri-
tremula) and also forests dominated by temperate tree species and
est, coldest and warmest quarters), minimum temperature of winter,
Cistus laurifolius shrubland; and (c) oro-mediterranean coniferous
precipitation of warmest and driest quarters and precipitation sea-
forests (Abies cilicica, Cedrus libani and Juniperus excelsa) and shru-
sonality. As expected, this indicates the correlation between climatic
bland (Juniperus communis subsp. nana). This elevational zonation
variables and elevational belts of the Mediterranean region (thermo-
follows the general pattern found in the Mediterranean mountains
and meso-, supra- and oro-mediterranean).
as described by Ozenda (1975).
The results of the DCA ordination of the 21 forest and shrubland
The thermo- and meso-mediterranean belts and the supra-
vegetation types in Mediterranean Turkey are shown in Figure 5. We
mediterranean belt show differences in precipitation in the
found three well-differentiated groups representing the main eleva-
growing season and in the drought period. Although the annual
tional belts in Mediterranean Turkey.
precipitation is lower in the supra-mediterranean than in the ther-
The first group represents vegetation from the thermo- and
mo- and meso-mediterranean belt in Mediterranean Turkey, the
meso-mediterranean belts, such as phrygana, garrigue, macchia,
supra-mediterranean belt has higher precipitation in the driest
sclerophyllous forests and pine forests dominated by Pinus brutia
period (summer; Mayer & Aksoy, 1986). Additionally, winter frost
and Pinus pinea. This group is characterized by high values of the
occurs regularly in the supra-mediterranean area. The mean tem-
annual mean temperature, mean temperature of the coldest month,
perature of the coldest quarter is thus lower in this belt.
mean temperatures of the driest, warmest and coldest quarters
In contrast, the oro-mediterranean belt is characterized by a
and precipitation seasonality, i.e. the characteristic features of the
subhumid mountain climate with cool summers with occasional
Mediterranean climate.
orographic rainfall whereas winters are snowy and cold (Atalay
1
2a
2b
3
4a
4b
5
6
7a
7b
8
9
10
11
12
13
14
15
16
17
18
No. of plots
81
87
32
47
197
66
156
249
649
238
273
63
192
88
655
190
35
44
192
268
269
Pinus pinea
.
1
100
4
.
.
.
.
1
.
.
.
.
.
.
.
.
.
.
.
.
Pinus brutia
2
10
53
49
23
38
9
7
99
100
3
.
8
.
1
.
.
.
.
1
3
Quercus ithaburensis
subsp. macrolepis
2
.
6
.
5
.
1
2
5
.
27
.
4
.
.
.
.
.
.
.
.
Quercus infectoria subsp.
boissieri
.
20
3
32
35
5
7
11
39
29
24
22
31
10
3
1
.
.
3
1
1
Ostrya carpinifolia
.
.
.
.
3
.
.
.
3
.
1
68
1
.
1
2
.
.
.
2
11
Quercus trojana
.
.
.
.
5
.
.
1
1
2
14
44
6
.
1
.
.
.
8
8
.
Quercus vulcanica
.
.
.
.
.
.
.
.
.
.
.
16
.
.
2
9
.
.
1
.
.
Quercus cerris
.
2
.
.
5
9
.
6
13
11
23
35
100
61
41
26
71
.
4
11
1
Pinus nigra subsp.
pallasiana
.
.
.
.
.
.
.
1
2
1
8
.
9
26
100
55
57
2
3
14
4
Pinus sylvestris
.
.
.
.
.
.
.
.
.
.
.
.
.
.
6
37
.
.
.
.
.
Fagus orientalis
.
.
.
.
.
.
.
.
.
.
.
.
.
.
5
47
.
.
.
.
.
Castanea sativa
.
.
6
.
.
.
.
.
1
.
.
.
.
.
4
26
.
.
.
.
.
Populus tremula
.
.
.
.
.
2
.
.
.
.
.
.
.
.
10
27
100
.
.
7
1
Juniperus excelsa
.
.
.
.
36
.
7
12
6
5
13
8
6
1
8
.
.
.
95
35
47
Abies cilicica
.
.
.
.
.
.
.
.
1
.
.
10
4
.
5
1
.
.
9
76
.
Cedrus libani
.
.
.
.
1
.
.
.
1
1
.
8
2
.
8
1
.
.
3
76
92
Sarcopoterium spinosum
100
31
9
34
1
11
13
4
2
7
.
.
.
.
.
.
.
.
.
.
.
Cistus creticus
20
74
78
36
54
45
32
29
49
50
9
.
2
.
2
.
.
.
5
2
1
Genista acanthoclada
4
40
.
47
6
12
19
5
6
4
.
.
.
.
1
.
.
.
.
.
.
Erica manipuliflora
.
21
.
100
4
.
4
.
3
11
.
.
1
.
1
.
.
.
.
.
.
Phillyrea latifolia
4
18
9
68
86
74
59
33
42
55
8
.
10
2
2
1
.
.
9
.
.
Dominant trees
| Applied Vegetation Science
Vegetation type
6 of 29
TA B L E 1 Shortened synoptic table of the percentage frequencies of the dominant and diagnostic species of the vegetation types classified at the alliance level; species are sorted by
decreasing values of the phi coefficient; only species with φ-values >0.20 and >15% frequency are shown and indicated by grey shading; the species that are not diagnostic but have >10%
frequency in the whole table are shown at the bottom of the table. (1) Sarcopoterium spinosum phrygana; (2a) Cistus creticus garrigue; (2b) Pinus pinea forest; (3) Erica manipuliflora garrigue; (4a)
Arbutus andrachne, Quercus coccifera and Juniperus excelsa sclerophyllous forest and shrubland; (4b) Quercus ilex, Arbutus andrachne and Quercus coccifera sclerophyllous forest and shrubland;
(5) Olea europaea sclerophyllous forest and shrubland; (6) Quercus coccifera sclerophyllous forest and shrubland; (7a) Upland Pinus brutia forest; (7b) Lowland Pinus brutia forest; (8) Quercus
ithaburensis and Quercus infectoria forest; (9) Ostrya carpinifolia, Quercus trojana and Quercus vulcanica forest; (10) Quercus cerris forest; (11) Cistus laurifolius shrubland; (12) Pinus nigra forest;
(13) Forests dominated by temperate species; (14) Populus tremula forest; (15) Juniperus communis subsp. nana shrubland; (16) Juniperus excelsa forest; (17) Abies cilicica and Cedrus libani forest;
(18) Cedrus libani forest. Full synoptic table is in Appendix S2
Dominant shrubs
KAVGACI et Al.
(Continues)
(Continued)
Vegetation type
1
2a
2b
3
4a
4b
5
6
7a
7b
8
9
10
11
12
13
14
15
16
17
18
Arbutus andrachne
.
2
.
2
60
91
15
8
8
40
.
5
4
.
1
.
.
.
.
.
1
Quercus coccifera
21
10
56
38
90
71
68
100
68
67
11
2
21
3
3
2
.
.
32
12
24
Quercus ilex
.
.
.
.
1
38
1
.
1
.
.
.
.
.
.
.
.
.
.
.
.
Olea europaea
15
2
19
32
44
52
79
12
12
22
3
.
.
.
.
.
.
.
1
.
.
Cistus laurifolius
.
3
3
.
.
.
.
2
4
.
6
.
26
100
40
19
74
5
.
.
.
Juniperus communis
subsp. nana
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
1
.
100
.
.
.
KAVGACI et Al.
TA B L E 1
Diagnostic species
(1) Sarcopoterium spinosum phrygana
Anagallis arvensis
72
8
.
11
1
.
7
3
2
.
4
.
1
.
1
.
.
.
4
.
.
Hymenocarpos circinnatus
42
.
.
4
3
.
6
8
4
1
1
.
.
.
.
.
.
.
.
.
.
Lysimachia
linum-stellatum
27
1
.
.
1
.
1
.
1
1
1
.
.
.
.
.
.
.
.
.
.
Rumex bucephalophorus
28
2
.
2
1
.
.
1
1
.
7
.
2
.
.
.
.
.
.
.
.
Plantago lagopus
31
.
.
9
1
.
4
3
2
1
1
.
.
.
.
.
.
.
1
.
.
Anthemis arvensis
20
.
.
.
.
.
.
2
1
.
.
.
1
.
.
.
.
.
.
.
.
Sherardia arvensis
46
3
25
.
4
5
17
7
4
1
2
.
1
.
.
.
.
.
1
1
.
Aphanes arvensis
19
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
5
.
.
Pallenis spinosa
17
1
.
.
.
.
2
1
.
.
1
.
.
.
.
.
.
.
.
.
.
Hippocrepis unisiliquosa
17
3
.
.
.
.
.
3
1
.
1
.
.
.
.
.
.
.
.
.
.
.
.
4
.
.
6
5
3
1
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
1
1
2
.
4
.
2
.
1
.
.
.
1
.
.
Plantago cretica
21
7
.
.
.
.
1
7
2
.
.
.
.
.
.
.
.
.
.
.
.
Trifolium stellatum
41
16
9
9
6
8
9
11
11
.
14
.
2
.
1
.
.
.
4
.
.
Medicago orbicularis
22
.
.
2
1
.
3
6
3
1
5
.
.
.
.
.
.
.
.
1
3
Pyrus amygdaliformis
28
1
3
.
1
15
3
11
5
1
7
.
2
1
1
.
.
.
.
.
.
Lagoecia cuminoides
31
.
.
.
5
6
22
13
2
2
17
.
.
.
.
.
.
.
5
2
.
Crepis foetida
38
10
22
.
5
3
25
9
8
5
18
.
1
.
1
1
.
.
6
4
.
Briza maxima
35
26
13
6
3
2
24
6
4
.
9
.
.
.
1
.
.
.
1
.
3
Aira elegantissima
27
5
16
19
1
5
6
4
2
1
2
.
1
2
1
1
.
.
.
.
1
Geranium molle
17
.
.
.
3
.
10
2
2
1
1
.
2
.
1
.
.
.
.
4
.
Tordylium apulum
17
3
.
.
.
.
1
6
7
1
8
.
.
.
.
.
.
.
.
.
.
Scorpiurus muricatus
16
5
.
9
.
.
4
2
1
.
2
.
.
.
.
.
.
.
.
.
.
(Continues)
7 of 29
22
19
Applied Vegetation Science |
Onobrychis caput-galli
Parentucellia latifolia
(Continued)
2a
2b
3
4a
4b
5
6
7a
7b
8
9
10
11
12
13
14
15
16
17
18
Galium brevifolium
17
3
6
.
.
.
15
4
2
.
1
.
.
.
1
.
.
.
.
.
.
Medicago minima
23
13
.
.
4
.
6
17
6
1
15
.
2
.
1
.
.
.
5
1
2
Stipa capensis
4
26
3
.
.
.
1
5
2
.
1
.
.
.
.
.
.
.
.
.
3
Micromeria nervosa
.
18
.
2
6
.
7
2
1
4
.
.
.
.
.
.
.
.
.
.
.
Trifolium angustifolium
17
23
6
.
4
.
3
8
7
3
10
.
2
.
1
.
.
.
.
.
.
Fumana thymifolia
15
20
.
2
4
5
1
9
4
4
1
.
.
.
.
.
.
.
1
.
.
Briza media
.
.
38
.
.
.
.
.
1
.
3
.
1
.
4
3
.
.
.
.
.
Cynosurus cristatus
.
9
34
.
.
.
.
1
.
.
.
.
.
.
1
.
.
.
.
.
.
Bromus erectus
1
2
34
.
1
.
.
.
1
.
.
.
.
.
1
.
.
.
.
.
6
Saxifraga cymbalaria
.
.
22
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
Cynosurus echinatus
14
2
69
30
2
2
5
12
17
4
17
11
8
.
1
.
.
.
14
7
2
Clematis vitalba
.
.
28
.
2
.
.
.
1
.
.
11
1
.
.
.
.
.
.
.
.
Cynosurus effusus
2
2
25
.
1
.
.
1
3
.
.
.
.
.
.
.
.
.
.
.
.
Asplenium
adiantum-nigrum
.
.
19
.
1
.
1
.
.
.
.
.
.
.
1
1
.
.
.
.
.
Bellis perennis
.
.
28
.
.
2
3
1
4
.
.
.
5
.
1
.
.
.
4
1
.
Taraxacum hybernum
.
1
19
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
2
1
.
Capparis spinosa
.
2
22
.
1
.
3
1
1
2
.
.
.
.
.
.
.
.
.
.
.
Aira caryophyllea
.
.
16
.
.
.
.
.
1
.
.
.
.
.
.
.
.
.
.
1
.
Trifolium tomentosum
4
1
28
11
2
3
1
2
2
.
.
.
.
.
.
.
.
.
.
.
.
Nigella arvensis
.
.
19
.
.
.
.
3
.
.
2
.
.
.
1
.
.
.
.
.
1
Globularia alypum
.
.
16
.
.
3
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
Plantago afra
5
.
19
.
1
.
5
.
1
.
.
.
.
.
.
.
.
.
.
.
.
Gagea graeca
2
.
22
9
1
2
5
1
2
.
.
.
.
.
.
.
.
.
.
.
.
Cardamine graeca
.
3
28
.
1
12
6
2
2
1
.
.
2
.
1
3
.
.
1
6
1
Hordeum murinum
4
.
22
.
3
.
1
4
1
.
2
.
1
.
.
1
.
.
.
.
6
Limodorum abortivum
.
.
25
2
.
8
1
.
6
3
1
.
.
.
9
6
.
.
.
1
.
Cerastium illyricum
4
1
19
2
.
.
1
4
3
.
3
.
.
.
1
.
.
.
.
.
.
Hypericum montbretii
1
.
22
.
.
.
.
.
.
.
1
.
6
.
7
7
9
.
.
.
.
Quercus pubescens
.
.
34
.
3
5
.
6
2
.
21
.
24
10
9
3
.
.
5
7
.
(2a) Cistus creticus garrigue
(2b) Pinus pinea forest
(Continues)
KAVGACI et Al.
1
| Applied Vegetation Science
Vegetation type
8 of 29
TA B L E 1
(Continued)
Vegetation type
1
2a
2b
3
4a
4b
5
6
7a
7b
8
9
10
11
12
13
14
15
16
17
18
Psoralea bituminosa
11
6
22
.
4
12
.
1
2
1
.
3
.
.
.
.
.
.
.
.
.
Lotononis genistoides
.
.
19
.
.
.
.
5
4
6
12
.
4
.
2
1
.
.
.
5
.
Hyparrhenia hirta
10
11
25
2
10
.
17
7
2
10
1
.
.
.
.
.
.
.
.
.
12
KAVGACI et Al.
TA B L E 1
(3) Erica manipuliflora garrigue
Cytisopsis dorycniifolia
.
1
.
36
5
.
2
.
1
.
.
.
.
.
.
.
.
.
.
.
.
Hypericum adenotrichum
.
.
.
32
.
.
.
.
1
.
1
.
1
2
2
.
.
9
.
.
.
Stachys byzantina
.
.
.
21
.
.
.
.
.
.
1
.
.
.
.
.
.
.
.
.
.
Scorzonera elata
4
11
13
38
1
.
13
.
2
2
.
.
1
.
1
.
.
.
.
.
.
Lupinus varius
1
.
.
17
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
Linum corymbulosum
4
14
.
32
.
3
6
2
1
.
.
.
.
.
.
.
.
.
.
.
.
Inula heterolepis
1
13
.
32
1
.
13
1
2
1
.
.
.
1
.
.
.
.
.
.
3
Andropogon distachyos
.
3
.
17
.
.
1
.
1
2
.
.
.
.
.
.
.
.
.
.
.
Cytinus hypocistis
.
5
.
23
.
2
.
1
14
3
.
.
.
.
.
.
.
.
.
.
.
Carlina corymbosa
10
.
.
21
3
.
4
1
2
1
.
.
1
.
.
.
.
.
.
.
.
Bupleurum gracile
.
10
.
17
.
.
3
.
1
.
.
.
.
.
.
.
.
.
.
.
1
Fumana arabica
12
14
.
30
6
.
12
4
5
2
2
.
.
.
1
.
.
.
.
.
.
Gladiolus anatolicus
.
1
.
19
2
.
4
.
3
5
.
.
6
.
.
.
.
.
.
.
.
Piptatherum coerulescens
2
18
38
47
9
5
33
18
11
27
2
.
.
.
1
.
.
5
11
4
.
3
9
.
.
1
.
.
.
.
.
6
3
.
(4a) Arbutus andrachne, Quercus coccifera and Juniperus excelsa sclerophyllous forest and shrubland
Phlomis leucophracta
.
.
.
.
20
.
3
.
(4b) Quercus ilex, Arbutus andrachne and Quercus coccifera sclerophyllous forest and shrubland
.
5
.
.
3
42
6
2
2
1
1
.
.
.
1
1
.
.
.
.
.
Tamus communis
1
.
19
2
13
65
12
8
6
3
3
8
7
.
1
2
.
.
3
1
1
Brachypodium retusum
1
5
.
.
.
20
1
.
.
.
.
.
.
.
.
.
.
.
.
.
.
Laurus nobilis
.
.
.
.
7
23
4
1
1
3
.
.
.
.
.
.
.
.
.
.
.
Ruscus aculeatus
.
.
6
13
25
39
7
4
7
10
.
.
2
.
1
.
.
.
.
1
.
Rubia peregrina
2
.
6
9
5
23
.
7
1
.
.
.
.
.
.
.
.
.
.
.
.
Rubia tenuifolia
4
3
.
21
13
39
18
18
6
8
1
5
3
.
1
2
.
.
.
1
4
Piptatherum miliaceum
.
11
.
13
6
30
20
5
4
9
.
.
1
.
.
.
.
.
.
.
.
Salvia fruticosa
.
2
9
.
.
17
4
4
1
.
.
.
2
.
2
.
.
.
.
.
.
Asparagus acutifolius
25
3
19
17
25
50
38
23
33
37
7
3
6
.
.
.
.
.
1
1
1
Ceterach officinarum
.
2
.
.
8
27
18
3
3
3
.
21
2
.
.
.
.
.
10
8
1
9 of 29
(Continues)
Applied Vegetation Science |
Hypericum empetrifolium
(Continued)
Vegetation type
1
2a
2b
3
4a
4b
5
6
7a
7b
8
9
10
11
12
13
14
15
16
17
18
.
3
.
.
1
.
31
2
2
5
.
.
.
.
1
.
.
.
.
.
.
Phagnalon graecum
1
.
.
.
1
.
22
.
1
1
.
.
.
.
.
.
.
.
.
.
.
Ceratonia siliqua
.
.
.
.
7
5
28
2
1
10
.
.
.
.
.
.
.
.
.
.
.
Anthemis chia
10
.
.
.
.
.
22
6
4
.
.
.
.
.
.
.
.
.
3
.
.
Orlaya daucoides
9
.
.
11
1
2
29
5
3
1
8
.
2
.
1
.
.
.
7
1
.
Galium setaceum
12
9
.
.
3
.
23
3
1
1
.
.
.
.
1
.
.
.
.
.
.
Quercus aucheri
10
.
.
.
.
15
24
3
2
3
.
.
.
.
.
.
.
.
.
.
.
Trifolium hirtum
.
.
.
2
3
.
19
2
4
1
3
.
2
5
.
.
.
.
.
.
.
Geranium lucidum
1
.
.
.
1
14
24
6
5
.
1
.
2
.
1
2
.
.
1
2
1
Urospermum picroides
4
5
.
.
3
2
19
4
1
5
.
.
.
.
.
.
.
.
2
.
.
Urginea maritima
19
.
6
.
2
.
24
5
3
8
.
.
.
.
.
.
.
.
.
.
.
Campanula podocarpa
11
7
.
.
.
8
19
.
.
.
.
.
.
.
.
.
.
.
.
.
.
Arabis verna
2
.
.
.
2
3
19
1
6
.
2
.
.
.
.
.
.
.
5
4
3
Origanum onites
19
1
19
2
6
2
31
7
7
13
1
.
.
.
1
.
.
.
.
.
6
Crucianella latifolia
4
10
.
11
18
15
40
15
9
22
9
5
3
3
1
.
.
.
17
3
.
Daphne gnidioides
.
7
13
2
25
.
31
15
11
8
.
.
.
.
1
.
.
.
7
2
.
Euphorbia falcata
10
10
.
.
3
3
19
1
1
4
3
.
1
.
.
.
.
.
.
.
4
| Applied Vegetation Science
(5) Olea europaea sclerophyllous forest and shrubland
Phlomis lycia
10 of 29
TA B L E 1
(6) Quercus coccifera sclerophyllous forest and shrubland
Valerianella vesicaria
7
5
.
.
2
.
.
22
7
.
5
.
2
.
1
1
.
.
9
3
1
Jasminum fruticans
.
1
.
.
26
.
12
30
7
6
13
5
3
2
1
.
.
.
23
5
8
Echinaria capitata
6
.
.
.
.
.
1
18
3
.
15
.
6
.
1
.
.
.
9
2
.
(7a) Upland Pinus brutia forest
Poa annua
.
.
.
.
.
.
.
1
19
1
.
.
.
.
1
.
.
.
.
.
.
Poa trivialis
.
.
.
.
1
.
1
3
17
1
.
.
2
.
2
9
.
.
.
.
.
Lathyrus aphaca
1
2
.
2
3
2
1
6
18
5
3
.
9
.
1
.
.
.
.
2
.
(7b) Lowland Pinus brutia forest
.
6
.
19
24
11
13
2
9
40
.
.
.
.
.
.
.
.
.
.
.
Cotinus coggygria
.
.
.
4
20
3
4
1
8
29
.
6
3
.
.
.
.
.
.
.
.
Lens ervoides
.
.
.
.
2
5
3
1
6
18
1
.
4
.
.
.
.
.
.
1
.
Stipa bromoides
.
16
.
23
21
9
13
10
15
41
2
6
13
.
2
.
.
.
18
8
5
Styrax officinalis
1
.
13
.
37
32
15
21
38
49
13
25
33
.
2
1
.
.
7
7
7
(Continues)
KAVGACI et Al.
Myrtus communis
(Continued)
Vegetation type
1
2a
2b
3
4a
4b
5
6
7a
7b
8
9
10
11
12
13
14
15
16
17
18
Lithodora hispidula
.
6
.
6
14
.
1
.
1
16
.
.
.
.
.
.
.
.
.
.
.
KAVGACI et Al.
TA B L E 1
(8) Quercus ithaburensis and Quercus infectoria forest
Eryngium campestre
6
3
.
.
.
.
.
4
1
1
18
.
6
5
1
1
.
.
3
2
.
Crupina crupinastrum
12
10
.
.
9
5
7
11
7
.
26
.
2
.
1
.
.
.
8
2
.
Ziziphora capitata
2
.
3
.
3
.
.
2
4
1
17
.
3
.
1
2
.
.
5
9
1
(9) Ostrya carpinifolia, Quercus trojana and Quercus vulcanica forest
Opopanax hispidus
.
.
.
.
2
.
1
1
.
.
.
33
1
.
.
.
.
.
3
.
.
Festuca heterophylla
.
.
.
.
.
.
.
3
.
.
1
38
1
1
5
14
.
.
.
4
.
Paeonia mascula
.
.
.
.
.
6
1
4
1
.
.
30
.
.
1
1
.
.
.
2
2
Cornus sanguinea
.
.
.
.
2
.
.
.
1
.
.
22
3
.
.
.
.
.
.
.
.
Geum urbanum
.
.
.
.
1
.
.
1
1
.
1
43
18
1
4
6
.
.
9
7
7
Rubia rotundifolia
.
.
.
.
.
.
.
.
.
.
.
17
.
.
.
1
.
.
.
.
.
Serratula grandifolia
.
.
.
.
.
.
.
.
.
.
.
17
.
.
1
.
.
.
.
.
.
Myrrhoides nodosa
.
.
.
.
.
.
.
.
.
.
3
22
5
.
.
.
.
.
.
.
.
Eragrostis cilianensis
.
.
.
.
.
.
.
.
.
1
.
16
.
.
.
.
.
.
.
.
.
Fraxinus ornus
.
.
.
.
2
11
2
.
4
.
.
40
13
.
4
1
.
.
1
5
15
Epipactis condensata
.
.
.
.
.
3
.
3
1
.
1
24
.
.
4
.
.
.
3
3
.
Laser trilobum
.
.
.
.
.
.
.
1
.
.
2
22
1
.
4
7
.
.
.
1
.
.
.
3
.
12
5
.
6
7
8
2
56
31
5
16
25
.
7
3
26
2
.
.
.
.
1
.
1
1
1
.
1
25
1
.
2
1
3
.
.
12
3
Acer monspessulanum
.
.
.
.
1
.
.
3
1
.
6
25
2
.
1
.
.
.
6
2
8
Acer hyrcanum
.
.
.
.
.
.
.
.
1
.
.
25
.
.
1
2
.
.
4
9
13
Erysimum goniocaulon
.
.
.
.
1
.
.
1
1
.
1
16
.
.
1
.
.
.
.
2
2
Coronilla emerus subsp.
emeroides
.
.
.
.
19
9
.
7
7
5
3
38
18
.
2
1
.
.
1
4
3
Physospermum
cornubiense
.
.
.
.
.
2
.
.
2
.
1
22
3
.
5
7
.
.
.
3
.
Lathyrus aureus
.
.
.
.
.
.
.
.
.
.
.
16
1
.
.
8
.
.
.
.
.
Galium lucidum
.
.
.
.
.
.
.
.
2
.
.
17
.
9
1
.
.
.
.
.
1
Lapsana communis
.
.
.
.
2
.
2
12
3
.
4
29
13
.
11
17
3
2
3
16
1
11 of 29
(Continues)
Applied Vegetation Science |
Clinopodium vulgare
Elymus panormitanus
(Continued)
Vegetation type
2a
2b
3
4a
4b
5
6
7a
7b
8
9
10
11
12
13
14
15
16
17
18
Agrimonia eupatoria
.
.
.
.
1
.
.
1
1
.
1
.
18
.
1
2
.
.
.
1
2
Falcaria vulgaris
.
.
.
.
1
.
.
3
2
1
8
13
31
3
1
.
.
.
8
10
3
Prunus spinosa
2
.
.
.
.
.
.
.
1
.
5
.
16
.
1
1
.
.
.
.
.
Bunium ferulaceum
.
.
.
.
.
.
.
.
3
1
.
.
16
.
2
.
.
.
.
.
9
Galium verum
2
1
.
.
2
.
.
15
1
.
12
.
30
5
9
6
.
2
14
5
1
Cephalorrhynchus
tuberosus
.
.
.
.
1
.
2
.
1
1
.
8
19
.
3
1
.
.
1
13
3
Crataegus monogyna
2
.
3
.
8
3
1
22
15
12
19
8
38
3
11
15
.
.
5
3
1
Festuca valesiaca
.
.
.
.
2
.
1
4
2
4
3
.
24
9
7
4
.
.
6
3
13
Lathyrus digitatus
.
.
.
.
.
.
.
8
3
.
6
17
27
9
8
2
.
.
6
12
.
Trifolium physodes
.
.
.
.
2
.
.
1
5
8
4
5
20
.
2
1
.
.
3
10
1
Coronilla varia
.
.
.
.
.
.
1
7
4
2
10
22
28
2
8
9
17
.
6
8
.
(10) Quercus cerris forest
| Applied Vegetation Science
1
12 of 29
TA B L E 1
(11) Cistus laurifolius shrubland
.
.
.
.
2
2
.
1
1
.
1
.
.
40
1
.
.
.
1
1
.
Linaria simplex
.
2
.
.
.
.
.
1
2
1
5
.
.
32
1
.
.
.
2
2
.
Herniaria glabra
.
.
.
.
.
.
.
1
.
.
.
.
.
19
.
.
.
.
.
.
.
Silene supina subsp.
pruinosa
.
.
.
.
.
.
.
4
.
.
2
.
.
23
1
.
.
.
1
1
.
Logfia arvensis
1
.
.
.
.
.
.
10
3
1
16
.
4
45
.
1
17
.
15
2
.
Verbascum insulare
.
.
.
.
.
.
.
.
.
.
.
.
.
16
1
.
.
.
.
.
.
Alyssum desertorum var.
desertorum
.
.
.
.
1
.
1
2
4
.
.
.
1
20
1
.
.
.
.
1
.
Ziziphora taurica
1
7
6
.
.
.
1
5
2
.
5
.
.
32
1
.
.
7
3
2
.
Bromus squarrosus
4
14
.
.
1
.
.
6
2
1
15
3
1
43
2
.
29
5
9
1
5
Arabis nova
.
.
.
.
.
.
.
2
1
.
4
.
2
25
1
.
.
.
4
6
9
Sedum pallidum
.
.
.
.
1
.
.
2
1
.
1
.
2
28
5
6
.
11
14
4
1
Phleum exaratum
.
.
.
.
.
.
1
14
1
.
9
.
4
25
1
.
.
.
9
.
1
Salvia tomentosa
.
.
.
.
25
.
5
12
12
24
5
30
35
58
18
8
.
2
21
27
9
Holesteum umbellatum
.
.
.
.
.
.
.
2
2
.
.
.
.
16
1
.
.
2
3
.
4
Bromus tectorum
23
14
34
2
3
.
1
11
2
2
16
.
7
47
9
1
29
5
13
18
3
Juniperus oxycedrus
.
6
3
.
32
2
.
28
27
16
46
49
57
68
38
14
3
.
34
38
21
Vulpia ciliata
2
.
3
2
1
.
1
4
2
1
13
.
.
18
1
1
.
.
.
.
2
(Continues)
KAVGACI et Al.
Galium floribundum
(Continued)
Vegetation type
1
2a
2b
3
4a
4b
5
6
7a
7b
8
9
10
11
12
13
14
15
16
17
18
Minuartia mesogitana
4
.
.
.
4
.
5
1
2
1
5
3
.
23
1
.
17
.
10
1
1
Trifolium arvense
4
24
.
6
4
5
4
6
10
6
14
.
10
30
5
1
.
.
13
2
1
KAVGACI et Al.
TA B L E 1
(13) Forest dominated by temperate species
Fragaria vesca
.
.
.
.
.
2
.
.
1
.
1
.
6
1
14
43
.
2
.
1
.
Asperula involucrata
.
.
.
.
.
.
.
.
.
.
6
.
15
.
12
41
.
2
.
1
.
Carpinus betulus
.
.
.
.
.
.
.
.
.
.
.
.
.
.
1
18
.
.
.
.
.
Cirsium hypoleucum
.
.
.
.
.
.
.
.
.
.
.
.
.
.
2
19
.
.
.
.
.
Galium paschale
.
.
.
.
.
.
.
.
.
.
.
.
2
.
9
25
.
.
.
.
.
Veronica officinalis
.
.
.
.
.
.
.
.
.
.
.
.
.
.
1
17
.
.
.
.
.
Quercus petraea
.
.
.
.
.
.
.
.
.
.
3
.
.
.
9
30
9
.
.
3
.
Corylus avellana
.
.
.
.
.
.
.
1
.
.
.
.
.
.
1
16
.
.
.
.
.
Stellaria holostea
.
.
.
.
.
.
.
.
1
.
.
.
2
.
3
19
.
.
.
.
.
Galium rotundifolium
.
.
.
.
1
.
.
.
.
.
.
.
.
.
4
19
.
.
.
2
.
Veronica chamaedrys
.
.
.
.
.
.
1
.
1
.
.
.
5
.
15
24
.
.
.
.
1
Rubus caesius
.
.
.
.
.
.
.
.
1
1
.
13
2
.
2
17
.
.
.
.
.
Sorbus torminalis
.
.
.
.
1
.
.
.
2
.
.
17
6
.
9
24
.
.
.
7
3
Viola sieheana
.
.
.
.
2
.
.
.
1
.
1
6
2
.
9
18
.
.
.
3
.
Cytisus pygmaeus
.
.
.
.
.
.
.
.
.
.
1
.
2
5
11
20
.
.
.
.
3
Trifolium alpestre
.
.
.
.
.
.
.
.
.
.
.
.
8
10
11
18
.
.
.
.
.
Pilosella piloselloides
.
.
3
.
.
.
.
2
3
.
10
.
8
13
14
26
6
.
2
4
3
(14) Populus tremula forest
.
.
.
1
3
.
.
1
.
.
.
5
1
3
7
86
.
.
.
.
.
.
.
.
.
.
.
.
.
.
2
.
1
13
1
.
89
5
.
1
3
Saponaria glutinosa
.
.
.
.
.
.
.
.
.
.
1
.
.
.
1
1
69
.
.
.
5
Myosotis discolor
.
.
.
.
.
.
.
.
.
.
.
.
.
23
.
.
80
.
.
.
.
Silene compacta
.
.
.
.
.
.
.
1
.
.
.
3
2
3
6
4
80
.
.
6
1
Verbascum × splendidum
.
.
.
.
.
.
.
1
.
.
.
.
.
.
.
.
57
.
.
.
.
Arabis sagittata
.
.
.
.
.
.
.
.
1
1
.
5
3
.
2
1
63
.
.
.
.
Petrorhagia alpina
.
.
.
.
.
.
.
.
.
.
.
.
1
5
8
11
54
2
.
7
.
Micropyrum tenellum
.
.
6
.
.
.
.
1
.
.
.
.
.
14
1
1
46
.
.
.
.
Poa nemoralis
.
.
.
.
.
3
.
3
1
.
2
30
35
2
13
23
86
.
15
18
1
Linaria genistifolia subsp.
genistifolia
.
.
.
.
.
.
.
.
.
.
.
.
.
1
.
.
23
.
.
.
.
(Continues)
13 of 29
.
Erysimum smyrnaeum
Applied Vegetation Science |
Cytisus hirsutus
(Continued)
Vegetation type
1
2a
2b
3
4a
4b
5
6
7a
7b
8
9
10
11
12
13
14
15
16
17
18
.
.
.
.
1
3
3
4
2
.
5
11
4
.
5
3
46
.
.
4
1
Thlaspi violascens
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
23
.
.
2
.
Fibigia eriocarpa
.
.
.
.
1
.
1
5
1
.
1
.
2
.
1
.
40
.
12
6
6
Anthemis tinctoria
.
.
.
.
2
.
.
14
3
.
27
32
21
26
17
13
80
.
21
18
1
Veronica dillenii
.
.
.
.
.
.
.
.
.
.
.
.
.
11
1
1
29
.
.
.
.
Stachys lavandulifolia
.
.
.
.
.
.
.
2
1
.
1
.
1
.
1
.
29
.
11
4
.
Cephalanthera rubra
.
.
.
.
.
.
.
5
5
1
4
2
5
3
14
10
40
.
2
9
1
Campanula lyrata
6
2
38
.
5
.
7
12
10
.
9
43
5
22
22
8
69
.
2
7
9
Trifolium medium
.
.
.
.
.
.
.
.
.
.
1
.
2
2
9
3
23
.
.
.
.
Hypericum perforatum
.
1
.
.
4
.
2
6
1
4
1
.
1
5
4
7
34
2
5
11
1
Silene italica
1
.
31
.
2
14
.
13
9
.
7
43
24
6
25
16
57
9
11
34
6
Colutea cilicica
.
13
19
15
4
14
.
11
7
1
1
8
10
.
1
1
34
.
4
1
5
Bromus sterilis
11
1
.
11
3
3
31
11
7
3
20
5
9
1
3
.
34
.
17
7
3
| Applied Vegetation Science
Galium aparine
14 of 29
TA B L E 1
(15) Juniperus communis subsp. nana shrubland
.
.
.
.
.
.
1
.
.
.
.
.
.
.
1
.
.
39
3
1
1
Hypericum linarioides
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
23
.
.
.
Bromus tomentellus
.
.
.
.
.
.
.
3
1
.
10
.
1
9
6
.
.
48
21
7
1
Thymus longicaulis
.
.
.
.
.
.
.
.
.
.
1
.
4
8
6
11
.
39
2
1
.
Poa alpina
.
.
.
.
.
.
.
1
1
.
.
.
.
.
.
.
.
25
.
.
7
Minuartia juressi
.
.
.
.
.
.
.
.
1
.
.
.
.
.
3
.
.
20
.
.
.
Daphne oleoides
.
.
.
.
1
.
1
4
1
.
3
.
.
.
6
.
.
39
7
14
5
Astragalus brachypterus
.
.
.
.
.
.
.
.
.
1
.
.
.
.
.
.
.
16
.
.
.
Rosa pulverulenta
.
.
.
.
.
3
.
.
1
.
.
.
3
.
3
9
.
30
.
4
12
Acantholimon ulicinum
subsp. ulicinum
.
.
.
.
.
.
2
.
.
.
1
.
.
.
1
.
.
25
7
4
8
Minuartia anatolica
.
.
.
.
.
.
.
2
.
.
3
.
2
.
1
.
.
18
2
.
.
Pilosella hoppeana
.
.
.
.
.
.
.
4
1
1
4
.
8
15
20
22
.
39
1
3
1
Thymus zygioides
.
.
.
.
.
.
.
7
1
.
10
.
4
6
4
1
.
30
9
3
.
Astragalus angustifolius
.
.
.
.
.
.
.
16
4
1
16
.
8
5
5
.
.
41
28
12
3
Euphorbia herniariifolia
.
.
.
.
.
.
.
.
.
.
1
.
.
1
3
.
.
18
2
3
8
Cerastium dichotomum
4
.
.
.
.
.
.
2
1
.
8
6
4
.
3
2
.
18
6
5
2
Digitalis ferruginea
.
.
.
.
.
.
.
1
.
.
3
.
3
10
17
22
.
23
2
6
3
(Continues)
KAVGACI et Al.
Marrubium astracanicum
(Continued)
Vegetation type
1
2a
2b
3
4a
4b
5
6
7a
7b
8
9
10
11
12
13
14
15
16
17
18
Minuartia hamata
.
.
.
.
.
.
.
14
.
.
17
.
2
.
5
.
.
.
37
6
6
Ziziphora tenuior
.
.
.
.
1
.
2
6
1
.
10
.
.
.
3
.
.
.
24
4
.
Aubrieta canescens subsp.
canescens
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
16
.
10
Arenaria serpyllifolia
.
.
.
.
1
.
.
7
1
.
4
.
3
.
3
.
.
.
20
4
.
Minuartia multinervis
.
.
.
.
.
.
.
4
1
.
1
.
.
.
2
.
.
.
16
6
.
Phlomis grandiflora
.
.
.
.
8
.
1
3
13
2
.
.
2
.
.
.
.
.
22
7
3
Phlomis armeniaca
.
.
.
.
.
.
.
4
1
.
3
.
.
.
1
.
.
9
17
2
3
Berberis crataegina
.
.
.
.
.
.
.
7
3
3
21
22
18
3
10
.
.
18
40
16
20
Geranium tuberosum
.
.
.
.
1
.
1
11
2
.
5
5
4
.
4
1
.
2
23
20
6
Briza humilis
.
.
.
.
2
3
1
26
10
1
17
27
19
16
5
.
.
2
36
16
6
(16) Juniperus excelsa forest
KAVGACI et Al.
TA B L E 1
(17) Abies cilicica and Cedrus libani forest
Cyclamen cilicicum
.
.
.
.
2
.
.
.
1
1
.
.
.
.
1
.
.
2
6
23
.
Corydalis solida
.
.
.
.
.
.
.
1
2
.
.
.
.
.
4
2
.
5
9
27
.
Veronica cuneifolia
.
.
.
.
.
.
.
.
1
1
.
.
4
.
3
.
.
.
1
18
11
Anemone blanda
.
.
.
.
1
.
.
.
5
.
1
8
1
.
2
1
.
.
4
21
8
Juniperus drupacea
.
.
.
.
.
.
.
1
1
.
.
14
2
.
2
.
.
.
13
21
1
Thlaspi perfoliatum
.
.
.
.
1
5
.
4
4
.
6
21
7
.
7
2
.
.
17
26
12
Myosotis alpestris
.
.
.
.
.
.
.
.
.
.
3
2
7
.
5
3
3
5
6
17
7
Anthemis rosea
.
.
.
.
1
.
2
2
1
.
.
.
.
.
.
.
.
.
3
3
35
Lonicera nummulariifolia
.
.
.
.
.
.
.
.
1
.
2
13
3
.
2
.
.
.
3
15
41
Vicia villosa
.
.
.
.
1
.
1
1
2
1
.
.
2
.
2
.
.
.
.
.
24
Astragalus macrourus
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
17
Lolium temulentum var.
temulentum
.
.
.
.
.
.
.
.
1
.
.
.
.
.
.
.
.
.
.
.
16
Bromus lanceolatus
.
.
.
.
.
.
.
.
1
.
.
.
.
.
.
.
.
.
.
.
16
Ranunculus argyreus
.
.
.
.
.
.
.
.
1
1
.
.
1
3
.
.
.
.
2
1
19
Crepis macropus
.
.
.
.
.
.
.
1
.
.
1
.
2
.
.
.
.
.
4
.
16
Verbascum sinuatum
.
.
.
.
1
.
.
1
2
1
.
.
2
.
1
.
.
.
.
.
16
Alliaria petiolata
.
.
.
.
.
.
.
.
.
.
.
13
5
.
1
3
.
.
.
2
20
Orchis palustris
.
.
.
.
.
6
2
.
1
.
.
.
.
.
1
.
.
.
.
7
17
(18) Cedrus libani forest
Applied Vegetation Science |
15 of 29
(Continues)
(Continued)
Vegetation type
1
2a
2b
3
4a
4b
5
6
7a
7b
8
9
10
11
12
13
14
15
16
17
18
Asphodelus aestivus
52
3
9
47
.
9
27
11
6
3
14
.
.
.
.
.
.
.
4
.
.
Galium murale
28
.
22
.
2
2
6
4
3
.
.
.
.
.
.
.
.
.
.
1
.
Theligonum cynocrambe
21
.
16
.
1
.
3
1
1
1
.
.
.
.
1
.
.
.
1
.
.
Valantia hispida
25
3
.
.
5
.
19
4
1
1
.
.
.
.
1
.
.
.
4
.
.
Trifolium campestre
59
23
56
30
10
15
22
29
26
20
13
.
11
15
7
1
.
.
9
1
1
Avena barbata
35
9
50
9
9
5
17
7
2
.
9
.
.
.
1
.
.
.
5
2
.
Trachynia distachya
31
28
.
9
9
3
37
10
5
7
3
.
.
.
2
2
.
.
5
1
1
Cistus salviifolius
5
45
13
47
14
24
19
6
4
21
.
.
.
.
1
.
.
.
.
.
.
Lavandula stoechas
6
34
91
34
3
2
12
1
4
4
.
.
.
.
.
.
.
.
.
.
.
Coridothymus capitatus
10
29
31
.
7
5
11
2
1
1
.
.
.
.
.
.
.
.
.
.
.
Anthyllis hermanniae
1
6
28
.
1
21
.
1
1
1
.
.
.
.
.
.
.
.
.
.
.
Euphorbia characias
subsp. wulfenii
.
.
34
23
13
3
41
8
5
14
.
.
.
.
.
.
.
.
.
.
.
Helichrysum stoechas
1
5
19
.
2
20
6
.
1
.
.
.
.
.
.
.
.
.
.
.
.
Pteridium aquilinum
.
.
22
.
.
3
.
.
3
1
.
3
4
.
11
22
.
.
.
.
.
Aetheorhiza bulbosa
5
1
3
21
2
2
20
1
5
6
.
.
.
.
1
.
.
.
.
.
.
Teucrium divaricatum
.
18
19
32
3
39
15
6
4
3
.
.
.
.
.
.
.
.
.
.
.
.
.
.
23
5
18
35
4
5
3
.
.
.
.
.
.
.
.
1
.
.
2
8
6
47
80
52
35
49
41
58
23
2
17
5
2
1
.
.
11
2
4
Fontanesia phillyreoides
.
.
.
.
33
2
1
.
19
28
.
.
12
.
.
.
.
.
.
.
.
Eryngium falcatum
.
2
.
.
28
.
4
5
14
33
.
.
5
.
1
.
.
.
1
.
1
Smilax aspera
1
.
13
21
38
56
35
5
11
35
.
.
.
.
1
.
.
.
.
.
.
Melica minuta
.
.
.
.
2
24
21
2
3
3
.
.
.
.
1
.
.
.
.
.
1
Pistacia lentiscus
4
11
.
2
9
2.9
29
3
2
7
.
.
.
.
.
.
.
.
.
.
.
Arisarum vulgare
1
1
.
.
3
24
33
1
5
3
.
.
.
.
.
.
.
.
.
.
.
Selaginella denticulata
.
6
.
4
.
17
16
1
2
1
.
.
.
.
.
.
.
.
.
.
.
Cerastium fragillimum
.
.
.
.
.
.
.
4
2
2
13
29
6
3
6
.
.
.
28
24
4
Papaver apokrinomenon
.
.
.
.
.
.
.
1
1
.
1
14
.
.
1
.
.
18
1
1
.
Carex divulsa
.
.
.
.
.
.
.
1
2
3
2
22
31
2
5
5
.
.
1
7
.
Lathyrus laxiflorus
.
.
.
.
.
.
.
7
4
3
11
22
43
1
36
47
.
7
5
30
.
Brachypodium pinnatum
.
.
.
.
3
.
.
.
1
5
.
.
22
.
11
24
.
.
.
.
1
(Continues)
KAVGACI et Al.
Clematis cirrhosa
Pistacia terebinthus
| Applied Vegetation Science
Diagnostic species for more than one alliance
16 of 29
TA B L E 1
KAVGACI et Al.
TA B L E 1
(Continued)
Vegetation type
1
2a
2b
3
4a
4b
5
6
7a
7b
8
9
10
11
12
13
14
15
16
17
18
Veronica grisebachii
.
.
.
.
.
.
.
2
.
.
1
.
.
30
1
.
57
.
.
.
.
Hypericum origanifolium
.
.
.
.
.
.
.
.
1
.
.
.
.
22
1
.
34
.
3
2
.
Allium flavum
.
.
.
.
1
.
6
2
2
1
1
.
.
18
1
.
17
.
1
2
2
Acinos rotundifolius
.
1
.
.
1
.
6
13
4
1
10
13
6
40
4
.
51
.
35
11
18
Alyssum strigosum
4
.
.
.
5
.
9
34
8
7
23
.
5
38
4
.
.
2
42
22
1
Hypericum olympicum
.
.
.
.
.
.
.
.
1
.
.
.
.
14
2
4
.
16
.
.
.
Viscum album
.
.
.
.
.
.
.
.
1
.
1
.
2
.
17
14
.
.
.
2
.
Luzula forsteri
.
.
25
.
.
.
.
1
5
1
2
.
10
.
27
40
.
.
.
.
.
Rubus canescens
.
.
.
.
.
5
.
2
2
.
3
2
16
15
28
40
.
2
.
3
.
Turritis laxa
.
.
.
.
.
.
.
.
1
.
1
6
1
2
21
25
29
.
1
19
.
Euphorbia kotschyana
.
.
.
.
.
.
2
7
3
5
11
24
8
.
8
.
.
23
48
46
7
Juniperus foetidissima
.
.
.
.
1
.
.
2
1
.
10
.
1
3
2
.
.
.
28
13
26
Lamium garganicum
.
.
.
.
1
.
1
4
1
.
2
14
8
.
9
7
.
2
17
31
33
Bunium microcarpum
.
.
.
.
1
3
.
2
3
1
1
5
2
.
7
.
.
.
21
24
23
Calicotome villosa
6
48
25
49
28
26
47
7
7
16
2
.
1
.
.
.
.
.
3
.
.
Arabis caucasica
.
.
.
.
.
.
.
.
1
.
2
49
1
.
4
2
40
5
13
35
11
Galium peplidifolium
1
.
.
.
2
.
.
9
2
1
20
44
15
.
12
.
69
.
41
47
10
Non-diagnostic species with more than 10% frequency in the whole data set
38
24
41
40
28
20
25
39
38
34
47
24
52
39
34
24
26
11
23
51
20
31
25
19
38
5
12
24
45
42
2
42
24
24
53
43
11
57
25
64
43
7
Teucrium chamaedrys
.
3
.
.
15
5
14
22
13
20
27
32
35
33
18
12
11
14
44
24
4
Teucrium polium
7
47
16
36
21
.
14
31
12
24
33
13
5
14
3
.
3
5
33
11
5
Vicia cracca
1
.
3
11
1
3
.
20
10
5
21
21
32
1
18
12
34
.
9
22
21
Doronicum orientale
.
.
6
.
1
6
.
8
6
2
3
13
16
.
28
11
.
11
7
25
19
Dorycnium pentaphyllum
.
.
.
.
3
2
1
14
14
9
11
10
15
5
19
28
.
5
4
11
.
Micromeria myrtifolia
14
29
31
30
24
15
41
18
9
22
3
.
.
.
1
.
.
.
21
4
.
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Dactylis glomerata
Poa bulbosa
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F I G U R E 2 Distribution of vegetation types in Mediterranean Turkey based on the analysis of a data set of 4,071 vegetation plots. Red
points: thermo- and meso-mediterranean vegetation; yellow points: supra-mediterranean vegetation; blue points: oro-mediterranean
vegetation [Colour figure can be viewed at wileyonlinelibrary.com]
et al., 2014). Especially the Taurus Mountains in southern Turkey,
were classified as the suballiance Cisto salviifolii-Lavandulenion
running parallel to the coastline, receive humid air from the
stoechadis. The invalidity of the alliance name makes invalid also
Mediterranean Sea, resulting in summer fog and rainfall (Atalay
the sub-alliances (ICPN, Art. 4). Moreover, the authors failed to
et al., 2018). This situation supports the productive Abies cilicica
select a validly described association as a holotype (ICPN, Art. 5).
and Cedrus libani forests, which occur in the Taurus Mountains
Therefore, we decided to describe the new alliance Cisto salviifolii-
but are absent in western Turkey. This can be due to the per-
Lavandulion stoechadis to include vegetation on non- carbonate
pendicular orientation of the mountain ranges in the western
bedrock. This new alliance (Appendix S2-1) is grouped under the
part to the coastline, which supports the penetration of the
order Lavandulo stoechadis-Hypericetalia olympici representing
Mediterranean climate further inland and broader distribution of
all phrygana vegetation on acidic siliceous and ultramafic sub-
Mediterranean vegetation than in the south (Akman, 1995; Mayer
strates from the eastern Mediterranean within Cisto-Lavanduletea
& Aksoy, 1986).
stoechadis (Mucina et al., 2016).
4.2 | Syntaxonomy of Turkish Mediterranean
forests and shrublands
4.2.2 | Garrigue
Garrigue is an open Mediterranean low scrub formation of browsed
4.2.1 | Phrygana
evergreen trees and shrubs, sub-shrubs and herbs resulting from
Based on the studies describing phrygana and garrigue communities
a successional stage and is maintained by grazing, fire, and summer
in Turkey (Ayaşlıgil, 1987; Brullo et al., 1997; Kavgaci et al., 2017),
aridity (Kavgacı et al., 2017).
long-term grazing, cutting and burning (Bergmeier et al., 2010). It is
the vegetation of Clusters 1 to 3 has been traditionally classified
Barbero & Quézel (1989) divided garrigue vegetation of the
within the class Cisto-Micromerietea julianae and the order Cisto-
eastern Mediterranean based on geographical distribution into two
Micromerietalia julianae. The class Cisto-Micromerietea julianae and
alliances: Hyperico empetrifolii-Micromerion graecae appearing in
the order Cisto-Micromerietalia julianae were merged with the class
more humid areas in the western part of the eastern Mediterranean
Ononido-Rosmarinetea by Mucina et al. (2016).
and Helichryso sanguinei- Origanion syriaci (recte Origano syriaci-
Phrygana is a vegetation type formed of low, thorny, chamaeph-
Hypericion thymifolii sensu Mucina et al., 2016) in the drier eastern
ytic (dwarf shrub) species. These communities are mostly the result
parts. Cistus creticus and Genista acanthoclada-dominated garrigues
of (over-)grazing, fire, and land abandonment. They usually repre-
appearing on calcareous bedrock are therefore grouped under the
sent a stage of secondary succession (Oberdorfer, 1954; Barbero &
alliance Origano syriaci-Hypericion thymifolii. This is also supported by
Quézel, 1989; Bergmeier, 2002).
Kavgacı et al. (2017; Appendix S2-2a).
Cluster 1 includes Sarcopterium spinosum-dominated communi-
The current syntaxonomic scheme of garrigue vegetation high-
ties that are common throughout the eastern Mediterranean region.
lights the importance of bedrock. Some syntaxonomic changes
Barbero & Quézel (1989) stated that Sarcopterium spinosum creates
should therefore be implemented to meet this criterion. Erica-
well-recognizable low phrygana communities, but it is difficult to
dominated communities in the eastern Mediterranean occurring on
identify the diagnostic species of higher syntaxa. This is because
non-calcareous substrates are classified under the alliance Helichryso
Sarcopoterium spinosum appears on formerly cultivated land and cre-
barrelieri-Phagnalion graeci (Mucina et al., 2009). Since the diagnostic
ates dense vegetation, which lasts for about 15 years in the succes-
species of this alliance (Mucina et al., 2016; Erica manipuliflora, Cistus
sional series (Barbero & Quézel, 1989).
salviifolius, Genista acanthoclada, Helichrysum stoechas subsp. barre-
Sarcopoterium spinosum- dominated communities are rather
difficult to classify. They need a broad-scale assessment in the
future. However, Barbero & Quézel (1989) classified these com-
lieri) are also diagnostic of our group, it has been classified under this
alliance.
The alliance Helichryso barrelieri-Phagnalion graeci encompasses
munities within the alliance Helichryso sanguinei- Origanion syriaci,
communities appearing in the western part of Turkey, which is more
which was suggested as nomen dubium by Mucina et al. (2016).
humid. Therefore, succession is faster approaching the next succes-
Therefore, a new name, Origano syriaci-Hypericion thymifolii,
sional stage of macchia, i.e., taller shrubland. Many elements of mac-
was proposed to encompass phryganas over calcareous bedrock
chia can be found in these communities, such as Myrtus communis,
(Mucina et al., 2016). Barbero & Quézel (1989) divided their al-
Phillyrea latifolia and Pistacia terebinthus (Appendix S2-3).
liance into three sub-alliances according to bedrock differences.
Other garrigue communities can be found at the upper eleva-
In this classification, Sarcopoterium spinosum-dominated com-
tional limit of Mediterranean vegetation (supra-mediterranean belt).
munities thriving over green rocks, serpentinites and gabbro
They are stages of degradation of Pinus nigra forests and share many
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F I G U R E 3 Typical stands of each vegetation type: (1) Sarcopoterium spinosum phrygana (Manavgat, Antalya, 200 m a.s.l.); (2a) Cistus
creticus garrigue (Serik, Antalya, 160 m a.s.l.); (2b) Pinus pinea forest (Bergama, İzmir, 550 m a.s.l.); (3) Erica manipuliflora garrigue (Termessos,
Antalya, 350 m a.s.l.); (4a) Arbutus andrachne, Quercus coccifera and Juniperus excelsa sclerophyllous forest and shrubland (Döşemealtı,
Antalya, 550 m a.s.l.); (4b) Quercus ilex, Arbutus andrachne, Quercus coccifera sclerophyllous forest and shrubland (Davutlar, Aydın, 50 m a.s.l.);
(5) Olea europaea sclerophyllous forest and shrubland (Kaş, Antalya, 200 m a.s.l.), (6) Quercus coccifera sclerophyllous forest and shrubland
(Gelendost, Isparta, 1,100 m a.s.l.); (7a) Upland Pinus brutia forest (Korkuteli, Antalya, 900 m a.s.l.); (7b) Lowland Pinus brutia forest (Kemer,
Antalya, 220 m a.s.l.); (8) Quercus ithaburensis and Quercus infectoria forest (Termesos, Antalya, 750 m a.s.l.); (9) Ostrya carpinifolia, Quercus
trojana and Quercus vulcanica forest (Bayatbademler; Antalya; 950 m a.s.l.); (10) Quercus cerris forest (Aksu, Isparta, 1,200 m a.s.l.); (11)
Cistus laurifolius shrubland (Simav, Kütahya, 1,050 m a.s.l.); (12) Pinus nigra forest (Beyağaç, Denizli, 1,300 m a.s.l.); (13) Forests dominated by
temperate species (Dursunbey, Balıkesir 1,600 m a.s.l.); (14) Populus tremula forest (Çameli, Denizli, 1,750 m a.s.l.); (15) Juniperus communis
subsp. nana shrubland (Simav, Kütahya; 1,950 m a.s.l.), (16) Juniperus excelsa forest (Sütçüler, Isparta, 1,450 m a.s.l.); (17) Abies cilicica and
Cedrus libani forest (Alanya, Antalya, 1,750 m a.s.l.); (18) Cedrus libani forest (Elmalı, Antalya, 1,650 a.s.l.). Photo credits: M. Arslan (1, 11, 13,
15), A. Kavgacı (2a, 3, 4a, 5, 6, 7a, 7b, 8, 9, 10, 12, 16, 17, 18), E. Örtel (2b), Ü. Akkemik (4b), Y.S. Bostancı (14) [Colour figure can be viewed at
wileyonlinelibrary.com]
TA B L E 2 Effects of environmental
variables with the percentage of the total
variance (PTV) of species data explained
by canonical correspondence analysis
(CCA) analysis
Variable
PTV
P-value
F-statistic
Elevation
0.55
0.001
33.58
Mean temperature of driest quarter (BIO9)
0.46
0.001
28.06
Annual mean temperature (BIO1)
0.46
0.001
28.01
Mean temperature of coldest quarter (BIO11)
0.46
0.001
27.96
Mean temperature of warmest quarter (BIO10)
0.45
0.001
27.87
Min temperature of coldest month (BIO6)
0.45
0.001
27.56
Precipitation of warmest quarter (BIO18)
0.45
0.001
27.41
Precipitation seasonality (BIO15)
0.44
0.001
27.19
Precipitation of driest month (BIO14)
0.44
0.001
27.08
Max temperature of warmest month (BIO5)
0.43
0.001
26.46
Mean temperature of wettest quarter (BIO8)
0.43
0.001
26.42
Precipitation of driest quarter (BIO17)
0.43
0.001
26.39
Precipitation of wettest quarter (BIO16)
0.38
0.001
23.11
Precipitation of coldest quarter (BIO19)
0.38
0.001
23.03
Precipitation of wettest month (BIO13)
0.37
0.001
22.88
Temperature seasonality (BIO4)
0.33
0.001
20.48
Isothermality (BIO3)
0.32
0.001
19.80
Annual precipitation (BIO12)
0.30
0.001
18.60
Temperature annual range (BIO7)
0.26
0.001
15.66
Mean diurnal range (BIO2)
0.19
0.001
11.56
common species with them (Atalay & Efe, 2010). In phytosociolog-
4.2.3 | Macchia and sclerophyllous forests
ical studies describing Cistus laurifolius shrubland in Turkey, these
communities are often identified as an association and are related
Macchia is evergreen sclerophyllous shrubland with a more or less
to relict Pinus nigra forests (Hamzaoğlu & Duran, 2004; Ture et al.,
closed canopy structure. It is a stage of vegetation succession to-
2005) or thermophilous deciduous forests (Sağlam, 2013) in terms of
wards the forest, replacement stage of climax forests, or even per-
floristic similarity. Since the contemporary vegetation classification
manent communities on xeric sites. These stages are maintained
integrates, in addition to floristic criteria, the physiognomy of the
by grazing, forest clearing and fires. Unless the succession is inter-
vegetation (Pignatti et al., 1995; Mucina et al., 2016; Bonari et al.,
rupted, macchia develops into a sclerophyllous forest, but the spe-
2021), Cistus laurifolius-dominated phrygana must be separated from
cies composition is nearly the same (Kavgacı et al., 2010). Macchia
forest vegetation. A vicariant alliance called Cistion laurifolii exists in
and sclerophyllous forests are therefore often treated together
the western Mediterranean (Escudero et al., 1996; de Foucault et al.,
in vegetation classification (e.g., Čarni et al., 2011, 2018; Kavgacı
2012). Because of the floristic and macroecological differences in
et al., 2017).
the eastern Mediterranean, Cistus laurifolius-dominated shrublands
Macchia and sclerophyllous forests used to be grouped with Pinus
in Mediterranean Turkey have been identified as a new alliance
brutia and Pinus pinea forests and classified within the class Quercetea
called Galio floribundi-Cistion laurifolii (Appendix S2-11).
ilicis in Turkey (Quézel et al., 1993; Ketenoğlu et al., 2010). Recently,
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F I G U R E 4 Forest and shrubland
types in Mediterranean Turkey along an
elevational gradient. See Table 1 and the
text for the explanation of the vegetation
type codes [Colour figure can be viewed
at wileyonlinelibrary.com]
Pinus brutia and Pinus pinea forests were separated within the new
siliquae-Pistacion lentisci represents thermo-mediterranean sclero-
class Pinetea halepensis (Bonari et al., 2021). Our results show that
phyllous xerophilous macchia and forests dominated by Olea euro-
macchia and sclerophyllous forests in Turkey can be divided into four
paea of the eastern Mediterranean (Mucina et al., 2016; Gianguzzi
groups corresponding to four alliances classified within three orders.
& Bazan, 2019; Şekerciler & Ketenoğlu, 2019; Appendix S2-5). This
The first order, Quercetalia ilicis, occurs in more humid parts of the
vegetation has been traditionally classified within Oleo-Ceratonion si-
area. It includes two alliances, Arbuto andrachnes- Quercion cocciferae
liquae in Turkey (Ayaşlıgil, 1987; Akman, 1995; Ketenoğlu et al., 2010).
and Cyclamini cretici- Quercion ilicis. The first alliance includes Arbutus
The third order, Quercetalia cocciferae, includes macchia and
andrachne, Quercus coccifera and Juniperus excelsa-dominated mac-
sclerophyllous forests dominated by Quercus coccifera from higher
chia and sclerophyllous forests of southern Mediterranean Turkey
elevations of the meso-mediterranean to the supra-mediterranean
(Kavgacı et al., 2017). It was described as evergreen basiphilous
belt. Quercus coccifera has an extensive distribution in Mediterranean
mesic kermes oak forests of the Eastern Mediterranean by Mucina
Turkey and occurs in many different vegetation types. However,
et al. (2016; Appendix S2-4a). However, especially in the more humid
these types at higher elevations differ from the others in their floris-
western Turkey, Quercus ilex also appears as a co-dominant species
tic composition, especially in the absence or low abundance of other
of vegetation with Arbutus andrachne and Quercus coccifera. These
macchia shrubs and sclerophyllous trees.
communities are classified within Cyclamini cretici- Quercion ilicis,
There are various taxonomic concepts of Quercus coccifera agg.
encompasing mesic macchia and sclerophyllous forests (Appendix
It is possible to understand it as two separate species, Quercus cal-
S2-4b). In the literature, Quercus ilex-dominated vegetation from
liprinos and Quercus coccifera (Pignatti, 2017-2019), two subspecies
western, more humid parts of the Mediterranean Turkey has always
(nominal and subsp. calliprinos; Schwarz, 1936a), or a single species
been placed in a different alliance (Quercion ilicis) than the southern
Quercus coccifera (Greuter et al., 1986; Jasprica et al., 2016). Like
Mediterranean Arbutus andrachne, Quercus coccifera and Juniperus
the Flora of Turkey (Davis, 1965-1985, Vol. 7, p. 681), the check-
excelsa-dominated vegetation (Arbuto andrachnes- Quercion coccif-
list of Flora of Turkey (Güner et al., 2012, p. 507 and Euro+Med
erae; Akman, 1995; Ketenoğlu et al., 2010). However, the alliance
PlantBase (Euro+Med, 2006), we have accepted the last solution and
Quercion ilicis was identified as an alliance of holm oak forests of the
treated Quercus calliprinos as a younger synonym of Quercus coccif-
western Mediterranean (Mucina et al., 2016).
era. Consequently, in accordance with Art. 44 of ICPN (Theurillat
The second order, Pistacio lentisci-Rhamnetalia alaterni, in-
et al., 2021), we consider the names Quercion calliprini Zohary 1955
cludes the thermo-mediterranean calcicolous macchia communities
(Zohary, 1955, p. 352) and Quercetalia calliprini Zohary 1955 (Zohary,
from the whole Mediterranean area. Within this order, Ceratonio
1955, p. 323, 338) as inadequate names (nom. inept.; ICPN, Art. 44)
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Pinus brutia is a tree species with the largest distribution in
Turkey, dominating forest stands across a total area of 5 million hectares (Boydak et al., 2006; Bonari et al., 2020). Like Pinus halepensis,
it is a typical eastern Mediterranean pine which is mainly distributed
in the western Mediterranean (Mauri et al., 2016). Many phytosociological studies have been carried out in Turkish Pinus brutia forests
and have classified these forests in several different alliances of the
Quercetalia ilicis (Quézel et al., 1993; Akman, 1995; Ketenoğlu et al.,
2010). However, Pesaresi et al. (2017) emphasized that classification
into Quercetalia ilicis does not discriminate Mediterranean needleleaved forests from the evergreen sclerophyllous and deciduous
Mediterranean vegetation and described the independent order
Pinetalia halepensis. Bonari et al. (2021) included this order in the
new class Pinetea halepensis. Pinus brutia forests in Mediterranean
Turkey were classified in two alliances with different elevational
distributions (Bonari et al., 2021): Styraco officinalis-Pinion brutiae
(Appendix S2-7a) in the highlands and Pinion brutiae (Appendix S27b) at lower elevations.
F I G U R E 5 Detrended correspondence analysis (DCA)
ordination of forest and shrubland vegetation types in
Mediterranean Turkey. Elevation and BioClim data were passively
projected on the ordination diagram. Only the eight most important
BioClim variables in the canonical correspondence analysis (CCA)
analysis are shown. For an explanation of vegetation type codes
and BioClim data, see Tables 1 and 2, respectively [Colour figure
can be viewed at wileyonlinelibrary.com]
In Turkish phytosociological studies, forests dominated by
Pinus nigra subsp. pallasiana, the only subspecies of black pine in
Turkey (Gülsoy et al., 2014), have been grouped under two different alliances: Cisto laurifolii-Pinion pallasianae in the Black Sea
region and central Turkey and Adenocarpo complicati-Pinion pallasianae in Mediterranean Turkey (Akman, 1995; Ketenoğlu et al.,
2010). The first alliance was also named Pino- Cistion laurifolii
without any indication of association, making it invalid (Quézel
and correct them to Quercion cocciferae Zohary 1955 nom. corr. and
et al., 1978). Bergmeier et al. (2018) were of the same opinion
Quercetalia cocciferae Zohary 1955 nom. corr. (Appendix S2-6).
and they classified Pinus nigra- dominated forests in the Aegean
and southern Mediterranean parts of Turkey under Adenocarpo
complicati-Pinion pallasianae. Mediterranean Pinus nigra subsp.
4.2.4 | High-mountain scrub
pallasiana forests are therefore grouped within this alliance
(Appendix S2-12). However, some extrazonal occurrences of Cisto
Subalpine and supra-montane chionophobous calcicolous dry low
laurifolii-Pinion pallasianae in Mediterranean Turkey were also
juniper shrubland of the central and southern Apennines, south-
found (Kavgacı et al. 2012, 2013) and are described below. The
central Balkans, Hellenic mainland and Turkey are grouped in the
lack of unique diagnostic species of Adenocarpo complicati-Pinion
alliance Daphno oleoidis-Juniperion alpinae (Mucina et al., 2016;
pallasianae is most likely related to the broad distribution range of
Bergmeier et al., 2018). We agreed with this classification and
the alliance and of its wide elevation range. Therefore, a revision
grouped Juniperus communis subsp. nana-dominated shrubland
of these forests is needed.
under Daphno oleoidis-Juniperion alpinae by including western Turkey
within the geographical range of the alliance (Appendix S2-15).
Juniperus excelsa has a wide distribution in the Mediterranean
and Euro-Siberian phytogeographical regions of Turkey. These communities have been intensively studied. Many communities were described under different names (Ketenoğlu et al., 2010). In addition
4.2.5 | Coniferous forests
to the appearance of Juniperus excelsa in macchia and sclerophyllous forests in Mediterranean Turkey, it dominates juniper forests
Pinus pinea forests show a high floristic similarity to Cistus creticus
in the Mediterranean high mountains, often forming the timberline.
and Genista acanthoclada garrigues. Because of these similari-
Bergmeier et al. (2018) grouped the montane tall juniper forests of
ties, these forests were previously included in Cisto-Lavanduletea
western Turkey under the alliance Juniperion excelso-foetidissimae,
(Brullo et al., 2002). However, the physiognomic difference (forest
like those in the south-central Balkans and Greece (Matevski et al.,
vs shrubland) suggests a classification of these forests within the
2010; Mucina et al., 2016). We also agree with the classification of
alliance Pinion pineae under Pinetalia halepensis of the class Pinetea
Bergmeier et al. (2018) on Juniperus excelsa-dominated forests in
halepensis (Feinbrun, 1959; Mucina et al., 2016; Sarmati et al., 2019;
Mediterranean Turkey (Appendix S2-16).
Bonari et al., 2021). These communities are also distinguished from
Cedrus libani covers an area of almost 400,000 hectares in the
garrigues by the presence of other tree species, such as Pinus brutia,
Taurus Mountains in southern Turkey (Kavgacı & Čarni, 2012), in addi-
Quercus coccifera and Quercus pubescens (Appendix S2-2b).
tion to its limited distribution in Lebanon and Syria. It generally forms
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pure stands in southwestern Turkey (western Taurus Mountains) and
found on other mountains in the Mediterranean area, for instance,
is co-dominant with Abies cilicica in the central and eastern Taurus
the alliance Fraxino orni-Ostryion carpinifoliae in the Balkans (Čarni
Mountains. This geographical differentiation also resulted in the
et al., 2009; Stupar et al., 2016). We decided to suggest a new con-
syntaxonomic classification as Lonicero nummulariifoliae-Cedrion
cept for these Mediterranean Ostrya carpinifolia, Quercus trojana
libani in the western Taurus Mountains and Abieti cilicicae-Cedrion
and Quercus vulcanica forests and propose a new alliance, Querco
libani in the central and eastern Taurus Mountains (Akman et al.,
vulcanicae-Ostryion carpinifoliae (Appendix S2-9).
1978; Kavgacı & Čarni, 2012). Our classification supports this dis-
Quercus cerris, a species having a large distribution through-
tinction between Cedrus libani-dominated forests in Mediterranean
out the Balkans and central Mediterranean (Di Pietro et al., 2020;
Turkey (Appendix S2-17–18).
Terzi et al., 2020), forms forests that are geographically separated
from forests of the alliance Querco vulcanicae- Ostryion carpinifoliae. These forests have a large distribution range in Mediterranean
4.2.6 | Deciduous forests
Turkey, especially in the supra-mediterranean region, mainly between 800 and 1,300 m a. s. l. They occur on deep soils. We de-
Quercus ithaburensis subsp. macrolepis and Quercus infectoria forests
scribe these forests as a new alliance, Falcario vulgaris-Quercion
in Turkey have also been the subject of phytosociological studies
cerridis (Appendix S2-10).
(Duman, 1995; Kargıoğlu & Tatlı, 2005). These deciduous oak for-
Populus tremula is the second most widely distributed tree in the
ests of the eastern Mediterranean mainly appear above the meso-
world, after Pinus sylvestris (Caudullo & de Rigo, 2016). In Turkey,
mediterranean belt. They are grouped within the alliance Quercion
it generally occurs in the Euro-Siberian area, and to a lesser extent
macrolepidis of the order Quercetalia pubescenti-petraeae and the
in the Mediterranean region. Pinus tremula-dominated forests are
class Quercetea pubescentis (Bergmeier et al., 2018; Appendix S2-8).
grouped within the alliance Fragario vescae-Populion tremulae, rep-
Ostrya carpinifolia and Quercus cerris are diagnostic species of the
resenting relict extrazonal temperate deciduous birch and poplar
alliance Ostryo-Quercion pseudocerridis (Akman et al., 1978; Quézel
forests on mineral soil in Europe (Mucina et al., 2016; Willner et al.,
et al., 1978). Quercus pseudocerris is a synonym of Quercus cerris
2016). In agreement with Bergmeier et al. (2018), we classified these
(Euro+Med, 2006). In our case, Ostrya carpinifolia and Quercus cerris
forests in Turkey within this alliance (Appendix S2-14).
are distinguished as dominants of different groups. In the description
Temperate tree species (Carpinus betulus, Castanea sativa, Corylus
of the alliance Ostryo-Quercion pseudocerridis, the authors (Quézel
avellana, Fagus orientalis) show an extrazonal distribution pattern
et al., 1978) pointed out the similarity with the vegetation of the
in Mediterranean Turkey. These forests are the result of possible
Balkan Peninsula and recognized two subtypes within this alliance.
migration of Euro-Siberian floristic elements during past geological
The first subtype consists of Ostrya carpinifolia and Carpinus orientalis
ages (Davis, 1971). These forests have been classified under many
forests, while the other subtype is Quercus cerris-dominated forests.
different syntaxa (Akman et al., 1979a; Bekat & Oflas, 1990; Varol
The former subtype includes primary deciduous forests, while the
& Tatlı, 2001; Tatlı et al., 2005; Kavgacı et al., 2013). However, our
latter includes mainly secondary forests occurring on sites of conif-
classification showed that they are a relatively homogeneous group,
erous forests (Quézel et al., 1978). This distinction is also clearly seen
and they can be grouped under the same alliance. In our analysis,
in our data.
these forests showed a close similarity to Pinus nigra-dominated
Quézel et al. (1978) and Akman et al. (1978) did not mention any
forests. Additionally, Pinus nigra commonly co-occurs in the floristic
association belonging to the alliance Ostryo-Quercion pseudocer-
composition. Kavgacı et al. (2012) carried out a large-scale assess-
ridis, which is not in accordance with Art. 8 of the ICPN (Theurillat
ment of Fagus orientalis forests in Turkey, including their extrazonal
et al., 2021). Moreover, the concept of this alliance is not entirely
distribution in the Mediterranean part of the country. In this work,
clear (Uğurlu et al., 2012). The same authors failed in the typifica-
these forests were classified within the alliance Cisto laurifolii-Pinion
tion of the alliance in 1993 (Quézel et al., 1993), since there is no
pallasianae, due to the high number of diagnostic species of this alli-
indication of the bibliographical source of Quézel et al. (1978) and
ance. Therefore, it also seems suitable for the forests dominated by
Akman (1973) for the type of the alliance (ICPN, Art. 2b; Theurillat
temperate tree species in Mediterranean Turkey to be classified, at
& Moravec, 1996). Despite these deficiencies, this alliance was used
least provisionally, under this alliance. However, these forests need
in recent studies (e.g. Bergmeier et al., 2018). The ecological and
further syntaxonomical studies (Appendix S2-13).
geographical distinction between Ostrya carpinifolia and Quercus
cerris-dominated forests, as well as the invalidity of the alliance
Ostryo-Quercion pseudocerridis, led us to abandon this concept and
4.3 | Syntaxonomical scheme
describe two new syntaxa for these two vegetation types.
The group of thermophilous deciduous forests dominated by
Based on the above syntaxonomic discussion on Mediterranean
Ostrya carpinifolia, Quercus trojana and Quercus vulcanica occurs
forests and shrublands in Turkey, we propose the following syn-
at higher elevations in the supra- and oro-mediterranean vegeta-
taxonomic scheme (EVC: Alliance present in the EuroVegChecklist
tion belts, where precipitation is more abundant, and temperature
(Mucina et al., 2016); notEVC: Alliance not present in the
seasonality is less pronounced. Analogous vegetation can also be
EuroVegChecklist; New: Newly described alliance).
Applied Vegetation Science |
KAVGACI et Al.
Cisto-Lavanduletea stoechadis Br.-Bl. in Br.-Bl. et al. 1940
Lavandulo stoechadis-Hypericetalia olympici Mucina in Mucina
et al. 2016
Cisto salviifolii-Lavandulion stoechadis Kavgacı, Balpınar, Öner,
Arslan, Bonari, Chytrý et Čarni 2021 all. nov. (Cluster 1 in Figure 1,
25 of 29
Adenocarpo complicati-Pinion pallasianae Quézel, Barbero et
Akman 1993 (Cluster 12 in Figure 1, Table 1-12, Figure 2-12) (notEVC)
Cisto laurifolii-Pinion pallasianae Akman, Barbero et Quézel ex
Quézel, Barbero et Akman 1993 (Cluster 13 in Figure 1, Table 1-13,
Figure 2-13) (notEVC)
Table 1-1, Figure 2-1) (New)
Helichryso barrelieri-Phagnalion graeci (Barbero & Quézel 1989)
R. Jahn in Mucina et al. 2009 (Cluster 3 in Figure 1, Table 1-3,
Figure 2-3) (EVC)
Galio floribundi- Cistion laurifolii Kavgacı, Balpınar, Öner, Arslan,
Bonari, Chytrý et Čarni 2021 all. nov. (Cluster 11 in Figure 1, Table 1-
Junipero-Pinetea sylvestris Rivas-Mart. 1965 nom. invers.
Berberido creticae-Juniperetalia excelsae Mucina in Mucina et al.
2016
Juniperion excelso-foetidissimae Em ex Matevski et al. 2010
(Cluster 16 in Figure 1, Table 1-16, Figure 2-16) (EVC)
11, Figure 2-11) (New)
Quercetea pubescentis Doing-Kraft ex Scamoni et Passarge 1959
Ononido-Rosmarinetea Br.-Bl. in A. Bolòs y Vayreda 1950
Hyperico empetrifolii- Genistetalia acanthocladae Mucina in
Mucina et al. 2016
Origano syriaci-Hypericion thymifolii Mucina et Theurillat in Mucina
et al. 2016 (Cluster 2 in Figure 1, Table 1-2a, Figure 2-2a) (EVC)
Querco- Cedretalia libani Barbero et al. 1974
Abieti cilicicae- Cedrion libani Quézel, Barbero et Akman 1993
(Cluster 17 in Figure 1, Table 1-17 Figure 2-17) (notEVC)
Lonicero nummulariifoliae-Cedrion libani Quézel, Barbero et Akman
1993 (Cluster 18 in Figure 1, Table 1-18, Figure 2-18) (notEVC)
Querco vulcanicae- Ostryion carpinifoliae Kavgacı, Balpınar,
Loiseleurio procumbentis-Vaccinietea Eggler ex Schubert 1960
Öner, Arslan, Bonari, Chytrý et Čarni all. nov. (Cluster 9 in Figure 1,
Vaccinio microphylli-Juniperetalia nanae Rias-Mart. et M. Costa
Table 1-9, Figure 2-9) (New)
1998
Daphno oleoidis-Juniperion alpinae Stanisci 1997 (Cluster 15 in
Figure 1, Table 1-15, Figure 2-15) (EVC)
Falcario vulgaris- Quercion cerridis Kavgacı, Balpınar, Öner, Arslan,
Bonari, Chytrý et Čarni all. nov. (Cluster 10 in Figure 1, Table 1-10,
Figure 2-10) (New)
Quercetalia pubescenti-petraeae Klika 1933
Quercetea ilicis Br.-Bl. ex A. Bolòs et O. de Bolòs in A. Bolòs y
Vayreda 1950
Quercetalia ilicis Br.-Bl. ex Molinier 1934
Quercion macrolepidis (Zohary 1973) Kavgacı, Balpınar, Öner,
Arslan, Bonari, Chytrý et Čarni nom. nov. (Cluster 8 in Figure 1,
Table 1-8, Figure 2-8) (EVC)
Arbuto andrachnes- Quercion cocciferae Barbero et Quézel 1979
(Cluster 4 in Figure 1, Table 1-4a, Figure 2-4a) (EVC)
Cyclamini cretici- Quercion ilicis Barbero et Quézel ex Quézel,
Barbero et Akman 1993 (Cluster 4 in Figure 1, Table 1, 4b, Figure 24b) (EVC)
Pistacio lentisci-Rhamnetalia alaterni Rivas-Mart. 1975
Brachypodio pinnati-Betuletea pendulae Ermakov et al. 1991
Fragario vescae-Populetalia tremulae Willner et Mucina in Willner
et al. 2016 nom. inval.
Fragario vescae-Populion tremulae Willner et Mucina 2016 ined.
(Cluster 14 in Figure 1, Table 1-14, Figure 2-14) (EVC)
Ceratonio siliquae-Pistacion lentisci Zohary et Orshan 1959
(Cluster 5 in Figure 1, Table 1-5, Figure 2-5) (EVC)
Quercetalia cocciferae Zohary 1955 nom. corr.
4.4 | Descriptions of the new syntaxa
Quercion cocciferae Zohary 1955 nom. corr. (Cluster 6 in Figure 1,
Table 1-6, Figure 2-6) (EVC)
Cisto salviifolii-Lavandulion stoechadis Kavgacı, Balpınar, Öner,
Arslan, Bonari, Chytrý et Čarni all. nov.
Pinetea halepensis Bonari et Chytrý in Bonari et al. 2021
Pinetalia halepensis Biondi, Blasi, Galdenzi, Pesaresi et Vagge
2014
Pinion pineae Feinbrun 1959 (Cluster 2 in Figure 1, Table 1-2b,
Figure 2-2b) (EVC)
Styraco officinalis-Pinion brutiae Bonari, Chytrý, Çoban, Kavgacı
et Sağlam in Bonari et al. 2021 (Cluster 7 in Figure 1, Table 1-7a,
Figure 2-7a) (notEVC)
Pinion brutiae Feinbrun 1959 (Cluster 7 in Figure 1, Table 1-7b,
Figure 2-7b) (EVC)
Corresponding name: Cisto salviifolii-Lavandulenion stoechadis
Barbero et al. 1989 nom. inval. (ICPN, Art. 4 and Art. 5).
Holotypus: Salvio aramiensis-Hypericetum triquetrifolii Barbero et
Quézel ex Kavgacı, Balpınar, Arslan, Öner, Bonari, Chytrý et Čarni
2020 ass. nov. (holotypus of the association: Barbero & Quézel, 1989,
p. 52, table 6, relevé 12).
Diagnostic species: Cistus monspeliensis, Cistus salviifolius,
Lavandula stoechas and Sideritis pisidica.
Ecology: This alliance encompasses the phrygana vegetations
thriving over non-carbonate bedrock in Mediterranean Turkey and
adjacent parts of the Mediterranean region.
Erico-Pinetea Horvat 1959
Erico-Pinetalia Horvat 1959 nom. conserv. propos.
Galio floribundi- Cistion laurifolii Kavgacı, Balpınar, Öner, Arslan,
Bonari, Chytrý et Čarni all. nov.
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KAVGACI et Al.
Holotypus: Hyperico heterophylli- Cistetum laurifolii Sağlam ex
Kavgacı, Balpınar, Arslan, Öner, Bonari, Chytrý et Čarni ass. nov. (holotypus of the association: Sağlam, 2013, table 5, relevé 23).
Synonym: Quercion macrolepidis anatolicum Zohary 1973 (Zohary,
1973: p. 516) nom. illeg. (ICPN, Art. 34a).
Holotypus: Quercetum macrolepidis anatolicum Zohary 1973.
Diagnostic species: Acinos rotundifolius, Allium flavum, Alyssum
Zohary (1973, p. 516, 517) described the association Quercetum mac-
desertorum var. desertorum, A. strigosum, Arabis nova, Bromus squarro-
rolepis anatolicum and indicated four relevés published by Schwarz
sus, Bromus tectorum, Cistus laurifolius, Galium floribundum, Herniaria
(1936b, p. 392) as stands of this association. The name of the asso-
glabra, Holosteum umbellatum, Hypericum olympicum, Hypericum orig-
ciation is illegitimate (ICPN, Art. 34a) but can be used as a type of
anifolium, Juniperus oxycedrus, Linaria simplex, Logfia arvensis,
the alliance (ICPN, Art. 17). As Zohary (1973, p. 517) mentioned the
Minuartia mesogitana, Phleum exaratum, Salvia tomentosa, Sedum
species composition of relevé 67 in Schwarz (1936b, p. 392), it can
pallidum, Silene supina subsp. pruinosa, Trifolium arvense, Verbascum
be understood as the holotypus of the association. Nevertheless, we
insulare, Veronica grisebachii, Vulpia ciliata and Ziziphora taurica.
propose a new name Quercetum macrolepidis (Zohary 1973) Kavgacı,
Ecology: This alliance represents the supra-mediterranean gar-
Balpınar, Öner, Arslan, Bonari, Chytrý et Čarni nom. nov., based
rigue communities dominated by Cistus laurifolius resulted from the
on the basionym Quercetum macrolepidis anatolicum Zohary 1973
degradation of Pinus nigra subsp. pallasiana forests.
(Zohary, 1973: 516) nom. illeg. (ICPN, Art. 34a).
Diagnostic species (Zohary, 1973): Quercus ithaburensis subsp.
Querco vulcanicae- Ostryion carpinifoliae Kavgacı, Balpınar, Öner,
Arslan, Bonari, Chytrý et Čarni all. nov.
Holotypus:
Asyneumo
michauxioidis- Quercetum
macrolepis.
Ecology: Sparse meso- and supra-mediterranean oak forests
trojanae
Ocakverdi et Çetik ex Kavgacı, Balpınar, Öner, Arslan, Bonari, Chytrý
dominated by Quercus ithaburensis subsp. macrolepis and Quercus infectoria on different bedrocks of the Eastern Mediterranean.
et Čarni 2020 ass. nov. (holotypus of the association: Ocakverdi &
Çetik, 1987, table 17, relevé 53).
Diagnostic species: Acer hyrcanum, Acer monspessulanum, Arabis
5 | CO N C LU S I O N S
caucasica, Asyneuma michauxioides, Campanula pterocaula, Carex divulsa, Cerastium fragillimum, Clinopodium vulgare, Cornus sanguinea,
We collected vegetation data on forest and shrubland in
Coronilla emerus subsp. emeroides, Elymus panormitanus, Epipactis
Mediterranean Turkey and present the first synthetic overview
condensata, Eragrostis cilianensis, Erysimum goniocaulon, Festuca het-
of their classification. We classified 21 vegetation types at the al-
erophylla, Fraxinus ornus, Fumaria parviflora, Galium lucidum, Geum
liance level. Among them, four alliances were described as new.
urbanum, Gladiolus atroviolaceus, Lapsana communis, Laser trilobum,
Twelve of them were already recognized in EuroVegChecklist, while
Lathyrus aureus, Myrrhoides nodosa, Opopanax hispidus, Ostrya
the other five alliances were formally described earlier but not in-
carpinifolia, Paeonia mascula, Papaver apokrinomenon, Physospermum
cluded in EuroVegChecklist. This confirms that the Mediterranean
cornubiense, Quercus trojana, Rubia rotundifolia, Scaligeria hermonis,
vegetation of Turkey has a close similarity to the vegetation of the
Serratula grandifolia and Silene alba subsp. eriocalycina.
European part of the Mediterranean. However, the vegetation of
Ecology: This alliance represents the supra-mediterranean Ostrya
Mediterranean Turkey also shows some differences because of its
carpinifolia, Quercus trojana and Quercus vulcanica-dominated forests
location in the eastern Mediterranean, the influence of the Irano-
occurring at precipitation-rich sites with low temperature seasonality
Turanian flora and almost 30% endemicity. A future integration of
and deep soils in the central and western Taurus Mountains.
Turkish syntaxa into EuroVegChecklist would be desirable for a better understanding of European and Mediterranean vegetation.
Falcario vulgaris- Quercion cerridis Kavgacı, Balpınar, Öner, Arslan,
Bonari, Chytrý et Čarni all. nov.
Holotypus: Trifolio physodis-Quercetum cerridis Tatlı et al. 2005
(holotypus in Tatlı et al., 2005, table 5, relevé 33).
Diagnostic species: Agrimonia eupatoria, Bunium ferulaceum,
AC K N OW L E D G E M E N T S
The data set (GIDV ID: 00-TR- 001) of this study was prepared at the
Southwest Anatolia Forest Research Institute and is managed by A.
Kavgacı. We thank Iztok Sajko for preparation of the figures.
Carex divulsa, Cephalorrhynchus tuberosus, Cerastium fragillimum,
Coronilla varia, Crataegus monogyna, Falcaria vulgaris, Festuca valesi-
AU T H O R C O N T R I B U T I O N S
aca, Galium verum, Lathyrus digitatus, Papaver apokrinomenon, Prunus
AK and AČ conceived of the idea; AK, NB, HHÖ and MA prepared
spinosa, Quercus cerris and Trifolium physodes.
Ecology: This alliance represents the supra-mediterranean
Quercus cerris-dominated forests at deep soils along whole
the data set; AK and AČ performed the statistical analyses; AK and
AČ wrote the manuscript with essential contributions of MC, GB,
NB and MA.
Mediterranean Turkey.
DATA AVA I L A B I L I T Y S TAT E M E N T
Quercion macrolepidis (Zohary 1973) Kavgacı, Balpınar, Öner,
Arslan, Bonari, Chytrý et Čarni nom. nov.
Vegetation-plot data used in this study are available in the Forest
Vegetation Database of Turkey, a part of the European Vegetation
Applied Vegetation Science |
KAVGACI et Al.
Archive (EVA). They can be obtained upon request from the first
author.
ORCID
Ali Kavgacı
https://orcid.org/0000-0002-4549-3668
Neslihan Balpınar
Hafize Handan Öner
https://orcid.org/0000-0002-4469-8629
https://orcid.org/0000-0003-2565-5030
Münevver Arslan
https://orcid.org/0000-0003-2645-1486
Gianmaria Bonari
https://orcid.org/0000-0002-5574-6067
Milan Chytrý
https://orcid.org/0000-0002-8122-3075
Andraž Čarni
https://orcid.org/0000-0002-8909-4298
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S U P P O R T I N G I N FO R M AT I O N
Additional supporting information may be found online in the
Supporting Information section.
Appendix S1. Vegetation data sources of each alliance
Appendix S2. Full synoptic table of the vegetation types at the alliance level in Mediterranean Turkey
How to cite this article: Kavgacı A, Balpınar N, Öner HH,
et al. Classification of forest and shrubland vegetation in
Mediterranean Turkey. Appl Veg Sci. 2021;24:e12589.
https://doi.org/10.1111/avsc.12589