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Appl Veg Sci. 2024;26:e12753.
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https://doi.org/10.1111/avsc.12753
Applied Vegetation Science
wileyonlinelibrary.com/journal/avsc
Received:7March2023
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Revised:6August2023
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Accepted:13October2023
DOI:10.1111/avsc.12753
VEGETATION SURVEY
Classification of forest and shrubland vegetation in central and
eastern Euxine Turkey and SW Georgia
AliKavgacı1 | Mustafa Karaköse2 |EmineSedaKeleş3 |NeslihanBalpınar4 |
Münevver Arslan5 |ErkanYalçın6 | Pavel Novák7 |AndražČarni8,9
This is an op en access arti cle under the ter ms of the CreativeCommonsAttribution-NonCommercial License, which permits use, distribution and reproduction
in any medium, provided the original work is properl y cited an d is not use d for comm ercial purposes.
©2023TheAuthors.Applied Vegetation SciencepublishedbyJohnWiley&SonsLtdonbehalfofInternationalAssociationforVegetationScience.
1Food,AgricultureandLivestock
VocationalSchool,BurdurMehmetAkif
ErsoyUniversit y,Burdur,Turkey
2EspiyeVocationalSchool,Giresun
University,Giresun,Turkey
3ForestGeneralDirectoryofTurkey,
Ankara,Turkey
4FacultyofArtsandSciences,Burdur
MehmetAkifErsoyUniversity,Burdur,
Tur ke y
5ResearchInstituteforForestSoiland
Ecology,Eskişehir,Turkey
6FacultyofScience,BiologyDepartment,
OndokuzmayısUniversity,Samsun,Turkey
7DepartmentofBotanyandZoology,
Faculty of Science, Masar yk University,
Brno,CzechRepublic
8Research Center of the Slove nian
AcademyofSciencesandArts,Instituteof
Biology,Ljubljana,Slovenia
9SchoolforViticultureandEnology,
UniversityofNovaGorica,NovaGorica,
Slovenia
Correspondence
AliKavgacı,Food,Agricultureand
LivestockVocationalSchool,Burdur
MehmetAkifErsoyUniversity,Burdur,
Tur ke y.
Email: akavgaci@mehmetakif.edu.tr
Funding information
JagnaAgencijazaRaziskovalnoDejavnost,
RS;CzechScienceFoundation
Co- ordinating Editor:BorjaJiménez-
Alfaro
Abstract
Questions: What are the main vegetation types of forest and shrubland vegetation in
centralandeasternEuxineTurkeyandSWGeorgia?Whatarethemainenvironmental
factorsaffectingtheir diversity?Whatistheirsyntaxonomicposition?Canweinte-
gratethemintotheEuropeanvegetationclassificationsystem?
Location: CentralandeasternEuxineTurkeyandSWGeorgia.
Methods: We collected 3104 vegetation plots of forest and shrubland vegetation in
the study region andperformedTwo-WayIndicatorSpecies Analysis (TWINSPAN)
classification. We described vegetation types based on the classification results, ex-
pert knowledge and information from literature sources. We defined diagnostic spe-
cies and prepared distribution maps for each vegetation type. To determine the most
significant environmental variables on floristic differentiation, we used canonical cor-
respondence analysis. Detrended correspondence analysis with passive projection of
most significant environmental variables was run to interpret the environmental vari-
ation of vegetation types.
Results: The studied vegetation was divided into 29 vegetation types related to seven
main vegetation groups: relict Mediterranean forests and shrubland (mainly along
thecoastline,besidesome inlandlocalities),lowlandtosubmontaneforests,central
Euxine mountain forests,easternEuxine (Colchic) mountain forests,subeuxinefor-
ests,azonalriparianforestsandsubalpineandalpineshrubland.Elevationisthemost
importantfactorcausingthedifferentiationinvegetation.Itisfollowedbylongitude
andlatitude.Amongclimaticvariables,temperatureseasonality,annualprecipitation
and precipitation of the wettest quarter are the most significant factors for vegetation
differentiation. These factors correlate with the reduction of maritime climate and
geomorphological features.
Conclusions: Vegetation types mostly correspond to the syntaxa accepted in the
EuroVegChecklist. However, some of them do not appearin the EuroVegChecklist
since they appear only beyond Europe. We described three syntaxa as new: Abietion
equi-trojani, Querco cerridis-Carpinion orientalis and Piceo orientalis- Fagenion orientalis.
The study revealed high vegetation diversity of the region that should be taken into
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1 | INTRODUCTION
Biomes and phytogeographicalregions have beenmainly basedon
theclimaticvariationonEarth(Woodwardetal.,2004).Duetothe
locationattheintersectionofthree different climatic zones,three
differ ent phytogeog raphic region s appear in Turkey (Davis, 1971).
ThesearetheEuro-Siberianphytogeographicregioninthetemper-
ateclimate zoneinthe north, theMediterraneanphytogeographic
region in the areas dominated by a Mediterranean climate in the
south an d west, and the I rano-Turanian phy togeograph ical region
formed as a result of the continental climate in the central, eastern
and southeastern part s. This and the dif ferentiation in local topo-
graphical, geological and other environmental factors are the main
reasons for the rich flora and extraordinarily high vegetation diver-
sityinTurkey(Kaya&Raynal,20 01).
Vegetation classification atthe country scale hasnot yet been
doneindetail(Kavgacıetal.,2 014).However,revealingthevegeta-
tion diversity in detail is required for better ecosystem management,
especially in the context of determination of ecosystem ser vices and
products, biodiversityandnature conservation(Mace et al.,2012).
Vegetationdatabasesareusefulandvaluabletoolsforunderstand-
ing vegetat ion diversit y on a large scale (S chaminée et al ., 2009).
Efforts that begin at the regional scale and extend from there to
createnational-,continental-andworld-scalevegetationdatabases
have been making very valuable contributions to the understand-
ing of ecosystem richness and diversit y, on both the European
and global scales (Dengler et al., 2 011; Chytr ý et al., 2016, 2020;
Bruelheideetal.,2019).
As a result of these efforts, amongothers, the EVA(European
Vegetation Archive, Chytrý et al., 2016), habitat classification of
Europe(Chytrýetal.,2020),EuropeanVegetationChecklist(Mucina
et al., 2016)and maps of European vegetationallianceshavebeen
prepared(Preislerováetal.,2022).Oneofthevegetationdatabases
intheEVAistheForestVegetationDatabaseofTurkey(FVDT).The
database includes plots from vegetation studies carried out in the
forests of Turkey. Comprehensive vegetation research in Turkey
startedinthe1940s(Ekim&Akman,1990).However,it intensified
mainly in t he 1970s and later (Ak man et al., 1978 , 1979a, 1979b;
Aksoy,1978).Based on this research, studies revealingthegeneral
vegetation structure of the countr y's forests and syntaxonomic lists
have been pr epared (Quézel et al ., 1992; Ketenoğ lu et al., 2010;
Bergmeieretal.,2018).
TheFVDTprovidesanoppor tunitytorevealthebiologicaldiver-
sity and richness of forest s in Turkey. With the help of this database,
the vegetation structure of Mediterranean forest s and shrubland in
Turkeyhasbeenstudiedindetail(Kavgacıetal.,2021).Someother
vegetation classification studies of specific vegetation types have
also been carried out, and valuable information has been produced
to understand the biological richness and diversity of Turkish forests
(Kavgacı et al.,2012;Kavgacı &Čarni,2012).Similarly,someother
vegetation databases have also been created and significant find-
ings have been achieved for the veget ation classification of Turkey
(Uğurluetal.,2012; Çoban & Willner, 2019).
Recently, a new dat abase has been constituted to cover
Transcaucasica (GIVD Transcaucasian Vegetation Database AS-
00-005;Nováketal.,2023).Inadditiontostudiescarriedoutinthe
past, t he database has b een enriched w ith recent studi es (Novák
et al., 2019, 2021).Beyondrevealingtheuniquespeciesrichnessand
high vegetation diversity of the region, the database may contribute
to vegetation studies to be done on a larger scale.
The Euro-Siberian phytogeographical (Euxine) region is an im-
portantpartofTurkey'sandGeorgia'secosystemrichness(Yaltırık&
Efe, 198 9; Akhalkatsi&Tarkhnishvili,2012; Nakhutsrishvili, 2013).
A comprehensive numerical classification of forest vegetation of
western Euxine Turkey was made by Çoban and Willner (2019).
Conducting suchastudyfor the other parts of theregion(central
andeasternEuxine[Colchis])isrequiredtounderstandthediversity
and richness within the entire region. Determining the biodiversity
of the region will contribute not only to the knowledge of Turkish
forests but also to understanding the Caucasus biodiversity hotspot
onalargerscale(Myersetal.,2000).
Thisstudyaimedto:(a)determinethevegetationtypesthatbuild
the forest and shrubland vegetation in central and eastern Euxine
TurkeyandSWGeorgia;(b)definetheessentialenvironmentalfac-
tors affecting the diversity offorest and shrubland vegetation; (c)
prepare a syntaxonomic list of the vegetation types indicating their
biologicaldiversity;and(d)integratethesesyntaxonomicunitsinto
the European vegetation classification system.
2 | MATERIALSANDMETHODS
2.1 | Studyarea
The nor thern part of Turkey and t he adjoining par t of southwestern
GeorgiaaremainlycoveredbytheEuro-Siberianphytogeographi-
cal region, i.e., the Euxine province, covering the area of almost
allEuro-SiberianvegetationalongtheBlackSeacoast(Figure 1).
consideration in ecosystem management and used as a reference in restoration and
mitigation of the effects of global changes.
KEYWORDS
BlackSea,Colchis,coniferousforest,deciduousforest,Euro-Siberian,Euxine,Georgia,
numericalanalysis,plantcommunity,shrubland,Turkey,vegetationtype,vegetation-plot
database
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The province can be divided into three distinct parts: the eastern
par tcomprisingnort heasternTurkeyandwesternGeorgia,includ-
ing the adjacent slopes of the Caucasus, which is also known as
Colchis;andthewesternandcentralBlackSeacoastalregionand
adjacentareasthroughinnerThraceandAnatoliathataregener-
allydrier(Davis,1965–1985).Inadditiontothislongitudinal divi-
sion alongthe Black Sea coast, Mayerand Aksoy (198 6)divided
the Euxine province into two par ts, from the coast toward inner
Anatoliawithadecreasingmaritimeclimateeffectduetothepar-
allel course of the mountain range along the sea coast: Euxine and
Subeuxine.
2.2 | Datasetandanalysis
After checking the FVDT (h t t p : // w w w . g i v d . i n f o / I D / 0 0 - T R -
001), stored in the TURBOVEG database management program
(Hennekens & Schaminée, 2001), we digitalize d the missing pub-
lished and unpublished vegetation plots from forests and shrublands
of central and eastern Euxine Turkey. We also enriched the data set
withvegetationplotsfromSWGeorgianforests(Nováketal.,2 019,
2021) and supplemented them with unpublished plots from the
Transcaucasian Vegetation Database (h t t p s : / / w w w . g i v d . i n f o / I D /
A S - 0 0 - 0 0 5 ,Nováketal.,2023).Vegetationplotsfromtheintersec-
tions with other phytogeographical areas through the inland were
also included in the data set.
Allvegetationplotsinthedatasetweresampledaccordingtothe
Braun-Blanquetsamplingmethod(Braun-Blanquet,196 4).Thespe-
cies taxonomy and nomenclature in the database was unified to fol-
lowtheFloraofTurkey(Davis,1965–1985; Davis et al., 1988;Güner
et al., 2000)(seeAppendixS1forthevegetationdatasources).
For the analyses, we selected only veget ation plots containing
a tree or shrub layer with a cover of at least 25% and also deleted
vegetation plots with three or fewer species due to floristic inade-
quacies in numerical analysis. The selection resulted in 3104 vegeta-
tionplots.Allrecordsofspeciesrepresentedbyvarioussubspecies
and varieties in the data set in addition to records at species level
weremergedtothespecieslevel. However,taxarepresented only
at subspecies or variety level in the data set without records at spe-
cies level like Abies nordmanniana subsp. equi-trojani and A. nord-
manniana subsp. Nordmanniana, were ret ained. We removed species
determined to the genus or family level, as well as bryophytes and
lichens,becauseofinconsistentsampling. Additionally,we merged
species records from different layers into a single layer because this
information was not consistently recorded among studies. The data
set finally included 1804 species. These data were used for the sub-
sequent classification andordinationanalyses.Weusedthe JUICE
7.1program(Tichý,2002)fordatasetediting.
Classification was carried outusingTwo-WayIndicatorSpecies
Analysis(TWINSPAN).
(Hill,1979).Pseudospeciescutlevelsofspeciescoverswere0%,
2%, 5%, 25%, the maximum number of divisions was nine and the
minimumgroupsizewasthreeplots.Weevaluateddivisions,andthe
optimal number of vegetation units was determined by exper t opin-
ion, taking into account differences in site conditions and biogeogra-
phy.Vegetationtypesweremainlybasedontheseclusteringresults.
The clusters representing the vegetation types were determined ac-
cording to the dominant species of clusters conforming to the main
vegetationtypesat the alliance level. In somecases,wealso inte-
gratedexpertknowledgeandinformationfromtheliterature.Inthis
context, the clusters dominated by the same tree species, appearing
in similar ecological conditions, distributed in the same biogeograph-
ical pattern and therefore representing the same vegetation t ypes,
weremerged(Kavgacıetal.,2021).
Diagnostic species of each t ype were defined by calculating the
fidelityofeachspeciestoeachvegetationtype(Chytrýetal.,2002)
FIGURE 1 MapoftheEuxineprovinceinnorthernTurkeyandSWGeorgiaandborderofthestudyarea.Thephytogeograpicalmapwas
basedonDinersteinetal.(2017)
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using the φ-coefficientasthefidelitymeasure.Specieswithaφ-value
higher than 0.20 were considered to be diagnostic. The φ-coefficient
wascalculatedforanequalizedsizeofclusters(Tichý&Chytrý,2006).
Species whose occurrence concentration in the plots of a particu-
lar cluster was not significant at p < 0.05 (Fish er's exact te st) were
excluded fromthesetofdiagnosticspecies(Tichý & Chytrý,2006).
Species with a cover of more than 25% in at least 10% of the vegeta-
tion plots in each t ype were considered dominant species.
We used elevat ion, latitude , longitude and c limatic data (fro m
WorldClim 2; Fick & Hijma ns, 2017) as ex planatory vari ables. To
test their effec ts on the floristic differentiation of forest and shru-
bland veget ation in the whole study area, we used c anonical corre-
spondenceanalysis(CCA)withforward selection ofvariables and
aMonte Carlo test with 999 unrestricted permutations. Variables
were selected one by one to prevent autocorrelation between them.
TheoriginalBraun-Blanquetcovervaluesweretransformedtoper-
centage and square-rooted. CCA assumes aunimodal response of
species to environmental gradients, and we selected it because of
the heterogeneity of the species matrix (7.53SDunits).The most
significantenvironmentalvariablesobtainedbyCCAwerepassively
projected onto the ordination space of detrended correspondence
analysis (DCA) to interpret the environmental variation between
vegetation types. We performed the analysis using CANOCO 5
(Šmilauer&Lepš,2014).
We used the fourth edition of the International Code of
PhytosociologicalNomenclature(ICPN;Theurillat et al.,2021) for
nomenclature and t ypification of syntaxa.
3 | RESULTS
3.1 | Hierarchicalclassification
Classification showed a clear floristic differentiation of forest and
shrubland vegetation in central and eastern Euxine Turkey and SW
Georgia,mainlygroupedasMediterraneanrelictvegetation,subeux-
ine vegetation, lowland to submontane vegetation, Euxine mountain
vegetation,Colchic mountain vegetation,azonalriparianvegetation
andsubalpineandalpineshrubland(Figure 2).Asaresultofclassifi-
cation analysis, we obtained 36 clusters showing clear differences in
floristiccompositionandvegetationstructure.However,someclus-
ters dominated by the same tree species, sharing the same ecological
conditions and appearing in the same area were merged, since they
represent the same vegetation type. We, therefore, obtained 29 veg-
etation types, which mainly represent the phytosociological alliances
of forests and shrublands in central and eastern Euxine Turkey and
SWGeorgia(Table 1,AppendixS2 and Figure 3a,b).
Vegetation t ype 1 includ es Mediterr anean Tamarix smyrnensis-
dominate ds hrubl andsonriverbed sa nddunes(Figure 2 and Table 1).
There is a constant appearance of Vitex agnus-castus in this vegeta-
tiontype.Itischaracterizedbyahighnumberofdiagnosticspecies,
many of which are ruderals, such as Plantago lanceolata, Polygonum
aviculare and Xanthium spinosum.
Vegetation t ype 2 is represented by Mediter ranean Juniperus
oxycedrus subsp. macrocarpa-dominated shrublands (Figure 2 and
Table 1).Jasminum fruticans also appears with high cover. This com-
munity c olonizes sand d unes. The spe cies forming t he vegetation
are characteristic of this kind of ecosystem, such as Juncus capitatus,
Jurinea kilaea and Medicago marina. This community is also charac-
terizedbyahighnumberofdiagnosticspecies.
Vegetationtype 3represents Mediterranean evergreen sclero-
phyllous high shrublands and forest s dominated by Arbutus unedo,
Olea europaea, Erica arborea and Quercus ilex in the Euxine province
(Figure 2 and Table 1).Itison eofthetypi calMedi ter r ane anrelic tve g-
etation t ypes in northern Turkey. Phillyrea latifolia, Buxus sempervirens
and Laurus nobilis are also common species in this vegetation type.
OtherwoodyspeciesformingthisvegetationtypeareCistus creticus,
Spartium junceum, Ligustrum vulgare and Colutea cilicica. The appear-
ance of Buxus sempervirens and Ligustrum vulgare in this community
indicates the pseudomacchia character of the vegetation type.
Vegetation type 4is another shrubland vegetation type inthe
Euxine region showing a Mediterranean charac ter and is domi-
nated by the deciduous species Elaeagnus angustifolia(Figure 2 and
Table 1). Thiscommunity isalso represented by a high number of
diagnosticspecies.Agoodexampleofthisvegetationtypeisfound
throughtheKelkitRiverValleyinthesubeuxinearea.
Vegetation type 5 represents Mediterranean Pinus brutia for-
ests in Eu xine Turkey,r anging from the co ast to 700 m (Figure 2
and Table 1). The increased numberof Mediterranean evergreen
woody species is apparent in this community, including Cistus cre-
ticus, Juniperus oxycedrus subsp. oxycedrus, Phillyrea latifolia, Pistacia
terebinthus subsp. palaestina, Quercus infectoria subsp. infectoria,
Ruscus aculeatus, Laurus nobilis, Phillyrea latifolia, Arbutus andrachne
and Erica arborea. Some deciduous shrubs and trees, such as Cornus
sanguinea and Carpinus orientalis, can also be seen in the floristic
composition of this vegetation t ype.
Vegetation type 6 is characterized by the presence of
Mediterranean Pinus pinea-dominated forests, which is another
Mediterranean vegetation type in Euxine Turkey (Figure 2 and
Table 1).There aremanydiagnostic species in theseforests, many
of which are ruderal plants, such as Alyssum murale, Fumana arabica,
Onobrychis armena and Trifolium stellatum. Cistusspecies(C. creticus
and C. salviifolius)accompanyP. pinea forests in the scrub layer.
Vegetationt ype7 represents Mediterranean Juniperus excelsa
forest(woodland)andshrubland(Figure 2 and Table 1).Juniperus ex-
celsaisoftenco-dominatedbyFontanesia phillyreoides, Vitex agnus-
castus, Paliurus spina-christi and Pistacia terebinthus subsp. palaestina.
Berberis crataegina is also a characteristic element of this vegetation
in the scrub layer.
Vegetation type8isanotherMediterranean relictvegetation
in Euxine Turkey, dominated by Arbutus andrachne and Quercus
coccifera(Figure 2 and Table 1).Otherwoodyspeciesaccompa-
nying the floristic composition of this vegetation are Pistacia tere-
binthus subsp. palaestina, Phillyrea latifolia, Cistus creticus, Cotinus
coggygria, Juniperus oxycedrus subsp. oxycedrus and Jasminum
fruticans.
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FIGURE 2 Hierarchicaldendrogramofforestsandshrublandincentralandeastern(Euxine)TurkeyandSWGeorgia.Thenumbers(1–29)
indicate the vegetation types.
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TAB LE 1 Shortenedsynoptictableofthepercentagefrequenciesofthedominantanddiagnosticspeciesofthevegetationtypesclassifiedatthealliancelevel;speciesaresortedby
decreasing values of the φ-coefficient;onlythesixmostdiagnosticspeciesforthevegetationtypesareshown;afullsynoptictableisavailableinAppendixS2; the threshold value of the φ-
coefficient is 0.20; the names of the vegetation types are writ ten in rows along the t able.
Vegetation type 12345678910 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Number of plots 10 10 102 10 135 22 67 55 93 10 10 318 177 10 19 15 275 157 221 88 10 0 462 10 464 74 94 40 947
Trees
Pinus brutia 16 99 11 1
Pinus pinea 100
Juniperus excelsa 25 254 4 3 3
Carpinus orientalis 40 12 24 27 18 11 84 29 620 11 5332 11 250 520
Quercus cerris 821 3932 32 17 5425 24 122
Quercus
pubescens
6 6 55 15 2 1 2
Pinus nigra subsp.
pallasiana
122 71 423 11
Fagus orientalis 5 6 93 36 16 26 234 81 41 23
Abies
nordmanniana
subsp.
equi-trojani
12 788 9 1 1
Pinus sylvestris 13 19 724 94 50 512 1
Quercus petraea
subsp. iberica
9 2 5 27 11 713 810 89 10 419 17 11
Alnus glutinosa
subsp. barbata
1199 11 22 326 33
Picea orientalis 2 4118 27 10 94 10 47
Abies
nordmanniana
subsp.
nordmanniana
35222 10 8
Castanea sativa 1 1 8 5 9 10 44 51 40
Carpinus betulus 10 6 1 8 14 27 38 10 140 916 36 26 70
Fraxinus
angustifolia
subsp.
oxycarpa
210 58 22
Platanus orientalis 1 1 3 1 1 67
Scrubs
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Vegetation type 12345678910 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Number of plots 10 10 102 10 135 22 67 55 93 10 10 318 177 10 19 15 275 157 221 88 10 0 462 10 464 74 94 40 947
Tamarix
smyrnensis
100 44
Juniperus
oxycedrus
subsp.
macrocarpa
100
Jasminum
fruticans
100 24 16 66 31 11 3 2 1
Arbutus unedo 61 23 7 3 1 7
Olea europaea 27 4 4
Erica arborea 34 15 4 2 3 1 34
Phillyrea latifolia 60 86 73 22 78 58 1 1 3
Elaeagnus
angustifolia
100 1
Fontanesia
phillyreoides
133
Quercus coccifera 344
Arbutus andrachne 14 21 91 58 28 1 7
Sarcopoterium
spinosum
3100
Hippophae
rhamnoides
subsp.
caucasica
40 2100
Epigaea
gaultherioides
100 1 1
Juniperus
communis
subsp. nana
110 100 112 30 110 123
Corylus avellana 2 2 1 4 2 100 925312 20 15 545
Betula litwinowii 2 1 100 1
Rhododendron
caucasicum
1 1 40 1100
Vaccinium
myrtillus
1100 1311 10 20 679
TAB LE 1 (Continued)
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Vegetation type 12345678910 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Number of plots 10 10 102 10 135 22 67 55 93 10 10 318 177 10 19 15 275 157 221 88 10 0 462 10 464 74 94 40 947
1) Tamarix smyrnensis shrubland
Digitaria
sanguinalis
90 7
Sophora
alopecuroides
60
Mentha pulegium 80 1 4
Centaurium
pulchellum
60 1 2 1
Rumex pulcher 40 1
Glycyrrhiza
echinata
30
2) Juniperus oxycedrus subsp. macrocarpa shrubland
Jurinea kilaea 100
Lagurus ovatus 100
Artemisia vulgaris 90
Draba muralis 100 1
Juncus capitatus 60
Linaria kurdica 80 2 1
3) Olea europaea–Arbutus unedo shrubland and sclerophyllous forest
Galium
verticillatum
27 2
Crucianella
latifolia
25 3 1
Parietaria
lusitanica
27 15 2
Micromeria
nervosa
18
Buxus
sempervirens
35 6 1 3 1 3 1 4 6
Spiraea crenata 14
4) Elaeagnus angustifolia shrubland
Artedia squamata 260
Crepis pulchra 40
Lysimachia dubia 30
TAB LE 1 (Continued)
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Vegetation type 12345678910 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Number of plots 10 10 102 10 135 22 67 55 93 10 10 318 177 10 19 15 275 157 221 88 10 0 462 10 464 74 94 40 947
Bryonia alba 30
Asparagus
acutifolius
250 1 2
Clematis viticella 20
5) Pinus brutia forest
Quercus infectoria
subsp.
infectoria
39 118 2 2 1
Rubia peregrina 12 25 1
Astragalus
sanguinolentus
10
Astragalus
viciifolius
10 4
Trigonella
monspeliaca
215 10 2 2 1
Ononis pusilla 217 2 9
6) Pinus pinea forest
Arceuthobium
oxycedri
73
Crucianella
gilanica
82 9 1 2
Trifolium stellatum 273 9 1
Silene armeria 27
Satureja spicigera 36 2 1
Punica granatum 36 4
7) Juniperus excelsa forest
Berberis
crataegina
140 1 4
Bromus diandrus 18
Geranium molle 20 628 1
Velezia rigida 119
Senecio vernalis 30 3 1 1 7
Melica ciliata 418 1
(Continues)
TAB LE 1 (Continued)
1654109x, 2023, 4, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/avsc.12753 by Cochrane Greece, Wiley Online Library on [01/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
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Vegetation type 12345678910 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Number of plots 10 10 102 10 135 22 67 55 93 10 10 318 177 10 19 15 275 157 221 88 10 0 462 10 464 74 94 40 947
8) Arbutus andrachne–Quercus coccifera shrubland and sclerophyllous forest
Linum hirsutum 47 1
Sideritis dichotoma 429
Astragalus
supruneri
22
Helianthemum
salicifolium
22
Fumana thymifolia 429 1
Muscari aucheri 733
9) Quercus cerris–Carpinus orientalis forest
Crataegus
microphylla
222 42 5217 2 43
Celtis australis 1 7210 3 1
10) Sarcopoterium spinosum phrygana
Phleum subulatum 470
Linum bienne 460
Carlina lanata 30
Echium italicum 30
Holcus lanatus 8 1 90 1 1 1 5 1 1
Trifolium
angustifolium
770 2
11) Hippophae rhamnoides subsp. caucasica shrubland
Elymus farctus 50
Conyza canadensis 30 100 4 1 2 3
Cionura erecta 40
Stachys maritima 40
Xanthium
strumarium
40
Aristolochia
clematitis
30
12) Quercus cerris–Quercus pubescens forest
Vicia cracca 110 46 33 611 15 15 4 3 1 1
Lathyrus digitatus 116 1 1
TAB LE 1 (Continued)
1654109x, 2023, 4, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/avsc.12753 by Cochrane Greece, Wiley Online Library on [01/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
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Vegetation type 12345678910 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Number of plots 10 10 102 10 135 22 67 55 93 10 10 318 177 10 19 15 275 157 221 88 10 0 462 10 464 74 94 40 947
Helianthemum
nummularium
19 45 431 235 27 1 3 5 1 30 1
Coronilla varia 2 1 4 4 10 35 19 11 3510 14 153
Dactylis glomerata 18 34 64 25 29 27 60 55 46 53 612 27 28 250 1445
Linaria corifolia 113 3
13) Pinus nigra subsp. pallasiana forest
Lathyrus
tukhtensis
534 3410
Asyneuma
limonifolium
8 2 16
Anthyllis
vulneraria
215 2 1
Polygala anatolica 1 2 7 22 3251
Dorycnium
graecum
16 873 33 314 41 5611 35 3 1 2 3
Cistus laurifolius 210 1
14) Epigaea gaultherioides shrubland
Carex oligantha 90 1
Gypsophila
silenoides
370 1 2 1
Carduus adpressus 30 10 1
Poa angustifolia 71100 1 3 1 7 3 1 2
Epilobium
ponticum
20 1
Rubus saxatilis 40 2
15) Juniperus communis subsp. nana shrubland
Stipa
ehrenbergiana
79
Stachys spectabilis 47
Minuartia recurva 42
Alyssum
dasycarpum
42
Nonea atra 42
Nonea ventricosa 37
(Continues)
TAB LE 1 (Continued)
1654109x, 2023, 4, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/avsc.12753 by Cochrane Greece, Wiley Online Library on [01/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
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Vegetation type 12345678910 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Number of plots 10 10 102 10 135 22 67 55 93 10 10 318 177 10 19 15 275 157 221 88 10 0 462 10 464 74 94 40 947
16) Corylus avellana shrubland and forest
Lamium macrodon 47
Rosa gallica 73 4 2
Cerastium
anomalum
347 1
Campanula
peregrina
40 2
Paeonia mascula 1 1 40 2 1
Plantago maritina 33 2
17) Fagus orientalis forest
Lathyrus aureus 2 1 7 37 30 10 2 8 1 1 14 1
Cardamine
bulbifera
135 24 111 833
Melica uniflora 110 13 4 1 1 1 5
Neottia nidus- avis 114 10 1 8 2 2
Acer hyrcanum
subsp.
hyrcanum
3 4 1 12 2 1 5 1
Lathyrus hirsutus 10 4 1 3 17 13 2 1 1 22 2
18) Abies nordmanniana subsp. equi- trojani forest
Myosotis sylvatica 3 5 2 3 25 1 1 1
Ranunculus brutius 3 7 24 4 1 10
Veronica
chamaedrys
7 8 16 39 17 7 2 1 1
Cirsium
hypoleucum
815 4 3 1 19 42 710 25 322 2
Valeriana
alliariifolia
132 5 9 12 80 3 4
Orthilia secunda 33 119 3
19) Pinus sylvestris forest
Cirsium trachylepis 410
20) Pinus sylvestris–Quercus petraea subsp. iberica forest
Bromus squarrosus 2 2 22
Trifolium aureum 11 119
TAB LE 1 (Continued)
1654109x, 2023, 4, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/avsc.12753 by Cochrane Greece, Wiley Online Library on [01/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
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(Continues)
Vegetation type 12345678910 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Number of plots 10 10 102 10 135 22 67 55 93 10 10 318 177 10 19 15 275 157 221 88 10 0 462 10 464 74 94 40 947
Argyrolobium
biebersteinii
2 4 4 11 11 2211 41 2
Scutellaria
velenovskyi
2 1 13 1
Lathyrus roseus 3 5 2 1 22 2
Origanum
rotundifolium
4 1 2 18 3
21) Colchic Alnus glutinosa subsp. barbata forest
Cardamine
raphanifolia
20 1
Angelica sylvestris 16 1
Petasites hybridus 131 6 4 11
Mentha longifolia 2 1 15 1
Impatiens
noli- tangere
29 8 3 5
Prenanthes
cacaliifolia
1128 5 8
22) Abies nordmanniana subsp. nordmanniana- –Picea orientalis forest
Oxalis acetosella 112 514 60 12 43
Galium
rotundifolium
7 6 33 30 2563 911
Cardamine
impatiens
925 4310 54 15 17
Veronica officinalis 1 2 8487323 311
Polypodium
vulgare
10 1 4 1315723 7414 3
Lonicera caucasica 6 1 5 11 53 11 22 31835 40 13
23) Betula litwinowii shrubland
Aconitum nasutum 170
Anemone
narcissiflora
2100 11
Veratrum album 1 3 90 2 6
Lilium carniolicum 10 2 1 60 4
Pedicularis
condensata
250 4
TAB LE 1 (Continued)
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Vegetation type 12345678910 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Number of plots 10 10 102 10 135 22 67 55 93 10 10 318 177 10 19 15 275 157 221 88 10 0 462 10 464 74 94 40 947
Cephalaria
gigantea
20
24) Picea orientalis–Castanea sativa–Fagus orientalis forest
Ulmus minor
subsp. Minor
5 1 19 8511 11 34 51 50 20 18
Laurocerasus
officinalis
8 3 18 528 27 33
Rhododendron
luteum
7 2 1 1 1 35 10 21 34 11 27 60 51 65 51
Ilex colchica 2 2 1 3 22 11 31821 35 58 33
Blechnum spicant 44514 5
Ruscus colchicus 1 1 11 17
25) Carpinus orientalis- Castanea sativa forest
Ruscus
hypoglossum
2 1 2 3 1 32 11
Epimedium
pubigerum
9 3 5 40 934621 542 15
Oenanthe
pimpinelloides
12 4 8 50 41 21 132 925
Vitis vinifera 3 3 1 1 15 2 5
Eragrostis collina 415 9
Quercus petraea
subsp. petraea
2 3 11 1
26) Castanea sativa- Carpinus betulus forest
Oplismenus
undulatifolius
218 6448 5
Pteris cretica 8 7 38
Vinca major 120
Hypericum
xylosteifolium
1 1 4 4 27
Carex muricata 1 1 16
Diospyros lotus 9 1 21 5
27) Alnus glutinosa subsp. Barbata- Fraxinus angustifolia floodplain forest
Leucojum aestivum 648
Polygonum
persicaria
235
TAB LE 1 (Continued)
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Vegetation type 12345678910 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Number of plots 10 10 102 10 135 22 67 55 93 10 10 318 177 10 19 15 275 157 221 88 10 0 462 10 464 74 94 40 947
Galium palustre 7 1 2 1 53
Fraxinus excelsior 1 2 1 1 1 33
Iris pseudacorus 20
Carex riparia 20
28) Platanus orientalis forest
Salix alba 1 1 1 67
Populus nigra
subsp. nigra
1 1 44
Tripleurospermum
tenuifolium
1 1 1 33
Veronica persica 111
Raphanus
raphanistrum
111
Matricaria
chamomilla
111
29) Vaccinium myrtillus- Rhododendron caucasicum shrubland
Vaccinium
uliginosum
166
Carex atrata 34
Geranium
platypetalum
130
Deschampsia
flexuosa
1 3 1 38
Carex umbrosa 21
Festuca woronowi 21
Other species with a freq uency higher t han 10% in the whole d ata set
Lapsana communis 9724 338 38 10 26 22 27 25 33 26 20 13 16 16 25 4
Viola sieheana 4 8 12 11 21 33 24 45 31 18 26 39 16 31 18 10
Daphne pontica 613 9 4 18 100 10 0 40 41 47 48 18 13 821 34 24
Pteridium
aquilinum
18 27 14 115 29 24 29 17 14 13 30 43 46 3
Brachypodium
sylvaticum
10 610 215 14 411 18 24 17 20 39 23 16 14 33 13
(Continues)
TAB LE 1 (Continued)
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Vegetation type 9 is formed by Carpinus orientalis-dominated
forests(Figure 2 and Table 1).ItisalsoaMediterraneanvegetation.
Carpinus orientalis is the only diagnostic species of this vegetation
type; in some communities Quercus cerris can be a dominant spe-
cies. It isdistributed in lower mountain belts withthe optimum of
its distribution between200 and 500 m. The floristic composition
of this vegetation type is enriched by Mediterranean elements, such
as Arbutus andrachne, A. unedo, Cistus creticus, C. salviifolius, Erica ar-
borea, Laurus nobilis, Myrtus communis and Phillyrea latifolia.
Vegetation type 10 represents Mediterranean Sarcopoterium
spinosum phrygana, which is a low, thorny and chamaephytic com-
munity ( Figure 2 and Table 1). It appe ars below 200 m a .s.l. along
the coast al line. Since this is the only phrygana cluster in the data
set, there is a high number of diagnostic species, many of which are
ruderals, such as Carlina lanata, Cichorium intybus, Echium italicum,
Eryngium campestre, E. creticum, Holcus lanatus, Linum bienne, L. trigy-
num, Rumex conglomeratus, and Trifolium angustifolium.
Vegetation type 11 includes Hippophae rhamnoides subsp.
Caucasica-dominatedshrubland(Figure 2 and Table 1).Itisdistrib-
utedalmostonlyatsealevelandischaracterizedbythehighappe ar-
ance of ruderal species as diagnostic s, such as Conyza canadensis,
Crepis foetida, Daucus carota, Hordeum murinum, Sinapis arvensis,
Xanthium strumarium, etc.
Vegetation type 12 corresponds to oak-dominated forests
in subeuxine Turkey formed by Quercus pubescens and Q. cerris
(Figure 2 and Table 1).Carpinus orientalis and Pinus nigra subsp. pal-
lasiana locally occur in the floristic composition of these forests as
dominant s. This vegetation t ype is distributed between elevations of
800to1700 m.Pinus sylvestris can also be seen locally in the floristic
composition of these communities. Some relict Cedrus libani forests
in the subeuxine region are also grouped within this cluster.
Vegetationtype13representsthesubeuxineconiferousforests
dominated by Pinus nigra subsp. pallasiana and P. sylvestris(Figure 2
and Table 1).Thisvegetationtypethrivesathigheraltitudesthanoak
forests in subeuxine areas. While P. nigra mostly forms pure forests,
P. sylvestris is locally accompanied by Juniperus excelsa and Carpinus
betulus.
Vegetation type14includesEpigaea gaultherioides shrubland in
thesubalpinevegetationbelt,above1800 m(Figure 2 and Table 1).
Daphne pontica, Rhododendron ungernii, Vaccinium myrtillus are other
shrub species that are partly also among the diagnostic species oc-
curring in the floristic composition of this vegetation type.
Vegetationtype15includesanothersubalpineshrublandvege-
tation t ype dominated by Juniperus communis subsp. nana(Figure 2
and Table 1).This vegetation type ranges from1700 mto 2000 m.
Daphne pontica thrives in this community as a constant species.
Other constant wo ody species, suc h as Sorbus torminalis, Corylus
colurna, Prunus divaricata and Rubus hirtus, are also diagnostic.
Vegetationtype16iscomposedofCorylus avellana shrublands
andforests in Euxinemountainareas (Figure 2 and Table 1).This
vegetation type is rich in terms of the number of woody species,
some of which are also diagnostic for it, such as Euonymus verru-
cosus, Sorbus umbellata, Berberis vulgaris, Cotoneaster nummularia,
Vegetation type 12345678910 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Number of plots 10 10 102 10 135 22 67 55 93 10 10 318 177 10 19 15 275 157 221 88 10 0 462 10 464 74 94 40 947
Clinopodium
vulgare
7 4 18 28 13 30 22 36 19 14 24 13 16 17 58110
Cyclamen coum 60 53416 16 15 33 32 25 14 11 221 516 10
Primula vulgaris 4 7 22 711 11 53 13 38 22 27 914 623 26 35
Lathyrus laxiflorus 1 1 2 5 19 20 47 31 29 30 43 210 2 1
Rosa canina 30 940 445 12 925 25 36 79 19 816 27 35 11113
Trachystemon
orientalis
413 8224 17 119 719 47 47 25
Teu criu m
chamaedrys
80 16 27 82 46 53 20 40 22 24 32 33 3610 2 1 2 3
Campanula
rapunculoides
42515 22 14 17 820 19 614 522 4
Tamus communis 80 5 3 15 1 2 16 11 3225 13 13 26 35 8
Greyshadingistoindicatethediagnosticspeciesofeachgroup.
TAB LE 1 (Continued)
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FIGURE 3 Distributionmapsofvegetationtypesbasedontheanalysisofadatasetof3104vegetationplots.Numberscorrespondto
vegetation types in the tex t and in Table 1.
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FIGURE 3 (Continued)
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Taxus baccata and Prunus spinosa subsp. dasyphylla, whereas
Viburnum lantana, Rosa gallica and Lonicera caucasica are constant
species.
The deciduous forests in central and western Euxine mountain
areas are mainly dominated by Fagus orientalis and its mixed forests
with Abies nordmanniana subsp. equi- trojani and Carpinus betulus.
The classification analysis showed that pure deciduous forests of
F. orientalis are grouped under separate clusters from pure A. nord-
manniana subsp. equi- trojani and its mixed forests. We therefore
treated pure deciduous forests of F. orientalis as a different group
from A. nordmanniana subsp. equi- trojani- dominated pure and
mixed forests.
Vegetation type 17represents pureFagus orientalis-dominated
forestsinthelowlandandsubmontaneareas(Figure 2 and Table 1).
Howeveritcanbefoundalsoatthemountainbelts.Carpinus betulus
appears in the floristic composition as a dominant tree species lo-
cally. This group appears more of ten in western and central Euxine
regions but can also be found in the lowlands of the eastern Euxine
region. Daphne pontica, Rhododendron luteum and Rubus hirtus ap-
pear with high constancy in the floristic composition.
Vegetation type 18 includes Abies nordmanniana subsp. equi-
trojani-dominatedforests(Figure 2 and Table 1).Pinus sylvestris and
Fagus orientalis represent its local companion. This vegetation t ype
can be found in the central and western Euxine mountain areas and
were accepted as one type comprising pure and mixed Abies nord-
manniana subsp. equi- trojani forests. These forest s are distributed
at higher elevational belts than the pure Fagus orientalis-dominated
forests of central and western Euxine regions.
Vegetation type 19 contains Euxine mountain pure Pinus
sylvestris-dominated forests (Figure 2 and Table 1). The optimum
distribution ofthis vegetation type is between 1500 and2000 m,
but it can also appear outside this range.
Vegetation t ype 20 consist s of Quercus petraea subsp. iberica-
dominated forests (Figure 2 and Table 1). Pinus sylvestris can also
be found as a dominant species in these forests. They appear in the
Euxine mountains and have their optimum at elevations between
900and1300 m.
Vegetationtype21representseastern(Colchic)EuxineAlnus glu-
tinosa subsp. barbata-dominatedforests(Figure 2 and Table 1).They
appearmainlybetween900and1400 m a.s.l.butcanalsobefound
outsidethisrange.Aldermostlybuildspurestandsandischaracter-
izedbysoilhumidity-tolerantandnutrient-demandingspeciessuch
as Angelica sylvestris, Athyrium filix-femina, Geranium robertianum,
Petasites hybridus and Urtica dioica.
Vegetationtype22encompassesPicea orientalis and Abies nord-
manniana subsp. nordmanniana-dominatedpure coniferous forests
oftheColchic region (Figure 2 and Table 1).This vegetation type
rangesfrom1000to2000 mintheeastern(Colchic)Euxinemoun-
tains. Locally, Fagus orientalis can be found as a dominant species.
Rhododendron ponticum often occurs in the shrub layer.
Vegetation t ype 23 correspon ds to eastern (Colchic) Euxinia n
upland Betula litwinowii-dominatedshrubland(Figure 2 and Table 1).
These deciduous shrublands thrive in the subalpine belt and loc ally
form the t imberline in t he eastern (Col chic) Euxine reg ion. Sorbus
aucuparia and Acer trautvetterii can be found in this group as diag-
nostic species, while Rhododendron luteum is a constant one.
Vegetation type 24 is formed by Fagus orientalis forests dis-
tributedin the eastern (Colchic) Euxine region that are locally co-
dominated or dominated by Picea orientalis, Castanea sativa and
Alnus glutinosa subsp. barbata(Figure 2 and Table 1).Castanea sa-
tiva, Rhododendron ponticum and Rubus platyphyllos, all of which are
woody species, are diagnostic species of this group. These forests
canbefoundatallelevational beltsoftheeastern(Colchic)Euxine
region,withtheiroptimumelevationbetween700and1500 m.
Vegetation type 25 indicates Castanea sativa and Carpinus
orientalis-dominatedforeststhriving throughouttheEuxine region,
fromthe lowlandtothesubmontane vegetation belt(Figure 2 and
Table 1). In addition toRhododendron ponticum, Smilax excelsa and
Ilex colchica are also diagnostic species, appearing densely in the
scrub layer of the forest. This communit y occurs from sea level up to
700 m,withtheoptimumat300 m.
Vegetation type 26 is formed by lowland mesophilous Carpinus
betulus and Castanea sativa-dominated forests representing lowland
tosubmontanedeciduousforests(Figure 2 and Table 1).Somevegeta-
tion plots dominated by Alnus glutinosa subsp. barbata were also placed
within this cluster due to their relatively mesophilous character. This
vegetationtypeismainlydistributedupto500 m.Corylus avellana and
Rhododendron luteum often appear as dominant species of the shrub
layer.
Vegetation t ype 27 corresp onds to Fraxinus angustifolia subsp.
oxycarpa and Alnus glutinosa subsp. barbata-dominated floodplain
forestsincentralEuxineTurkey(Figure 2 and Table 1).Fraxinus ex-
celsior, Pterocarya fraxinifolia, Periploca graeca, Crateagus curvisepala,
Quercus hartwissiana and Alnus glutinosa subsp. glutinosa are diag-
nostic species of these forests.
Vegetation type 28 represents Platanus orientalis-dominated
riverine azonal forests (Figure 2 and Table 1). Salix alba, Populus
nigra subsp. nigra, Alnus glutinosa subsp. glutinosa, Juglans regia and
Tamarix smyrnensis, all of which are woody species, appear in these
communities as diagnostic species.
Vegetation type 29 is formed byalpine Rhododendron caucasi-
cum and Vaccinium myrtillus-dominated shrublands in the eastern
(Colchic)Euxineregion(Figure 2 and Table 1).Theyoccur ateleva-
tionshigherthan2000 mandformthevegetationabovethetimber
lineintheeastern(Colchic)Euxinianmountains.
3.2 | Environmentalgradients
Canonicalcorrespondenceanalysis(CCA)showedthatenvironmen-
tal variables have clear effects on floristic differentiation in central
andeasternEuxineTurkeyandSWGeorgia(AppendixS3).Elevation
is the most important ecological factor, and it is followed by longi-
tudeandlatitude(Figure 4).Additionally,someclimaticvariablesalso
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appeared as significant factors for the floristic differentiation in the
study area. These are temperature seasonality, annual precipitation
and precipitation of the wettest quarter.
The DCA ordination of the 29 forest and shrubland vegeta-
tion ty pes in central an d eastern Euxine Turkey and S W Georgia
(Figure 5) showed clear grad ients along both a xes. However, the
gradient at axis 1 results mainly from the differentiation between
Mediterranean relict vegetation types and the gradient along axis
2 is much clearer for interpreting the environmental variation be-
tween vegetation t ypes of the whole study area. The effects of ele-
vation, longitude and latitude on floristic differentiation are evident.
Vegetationgroups,whichweredeterminedby classificationanaly-
ses, are well differentiated on the ordination plane in accordance
with their ecological and environmental differences.
Mediterranean vegetation types located in the lower elevation
belts and mostly on the coast are positioned at the right and right
upper corner of the ordination plane. Subeuxine forests dominated
by Carpinus orientalis, Quercus cerris, Q. pubescens, Pinus nigra and P.
sylvestris can be found at the bottom right of the ordination plane.
Central Euxine mountain vegetation t ypes formed of Carpinus bet-
ulus, Fagus orientalis, Quercus petraea, Abies nordmanniana subsp.
equi- trojani and P. sylvestris occupy a central position on the ordi-
nation plane. Deciduous forests of Carpinus betulus, Castanea sativa
and Alnus glutinosa subsp. barbata forming lowland to submontane
forests in the whole area are placed at the left upper position of the
ordination. Colchic mountain forests dominated by Alnus glutinosa
subsp. barbata, Betula litwinowii, Fagus orientalis, Abies nordmanni-
ana subsp. nordmanniana and Picea orientalis can be found in the left
part ofthe ordination.Alpineshrublands ofEpigaea gaultherioides,
Rhododendron caucasicum and Vaccinium myrtillus are the most re-
mote left part. Riparian forests dominated by Fraxinus angustifolia
and Platanus orientalis from lowland areas are placed in the upper
part of the ordination plane.
4 | DISCUSSION
4.1 | Ecologicalinterpretation
CentralandeasternEuxineTurkeyandSWGeorgiahaveahighfor-
est and shrubland diversity. Elevation is the most important envi-
ronmental factor causing vegetation differentiation and is followed
by longitude, latitude, temperature seasonality, annual precipitation
and precipitation of the wettest quarter.
Elevation is known to be an important ecological factor affect-
ing climatic differences on a regional scale resulting in the variation
of vegetatio n (Blondel & Aro nson, 1999; Quézel & Médail , 2003;
Nakhutsrishvili,2013; Gauquelin et al., 2016; Médail etal., 2019).
These differences cause the development of different vegeta-
tion ty pes in central a nd eastern Eux ine Turkey and SW Georg ia,
which were p reviously def ined by studies i n the region (Mayer &
Aksoy,1986). Similarly, the effects of other geomorphological dif-
ferences and, in connection with them, macroclimatic variations are
determining environmental factors for vegetation differentiation of
easternEuxine Turkey.Our results supportthe phytogeographical
division proposed by Davis (1965–1985), who indicated that the
eastern part of the Euxine region comprising nor thwestern Turkey
isconnectedwithGeorgia,whichisalsoknownasColchisandpos-
sesses a distinct vegetation cover from the western, generally drier
part,consistingofthewesternandcentralBlackSeacoastandadja-
centareasininnerThraceandAnatolia.Thisgeographicalvariation
of vegetation was also defined by our study, in which Colchic vege-
tation showed some specific differences from the rest of the Euxine
region: forests dominated by Abies nordmanniana subsp. nordman-
niana, Alnus glutinosa subsp. barbata, Betula litwinowii, and Picea
orientalis and shrubland of Epigaea gaultherioides, Rhododendron
caucasicum and Vaccinium myrtillus mainly appear in Colchis. The
highermountains,reachingmorethan3000 mandwithmorehumid
conditions,withalmost2500 mmofprecipitationperyearinColchis
(above 40 00 mm in the vicinity of Batumi), are the m ain reasons
for the vegetation differences with the rest of the Euxine region,
aswellasstrongerbiogeographicallinkstotheCaucasus(Yaltırık&
Efe, 198 9;Nakhutsrishvili,2013).
MayerandAk soy(1986)alsodidafurt herdivisi onoftheEuxine
region based on theeffect of theBlackSea onmountain slopes.
Due to the parallel course of the mountain range along the sea-
coast, locally reaching elevations of over 200 0–3000 m dire ctly
by the sea, the effect of the maritime climate decreases toward
the interior of Anatolia, causing theoccurrence ofdifferences in
vegetation. They therefore called the areas in the hinterland of
the Euxine region, with a decline in the maritime ef fect, subeux-
ine. This differentiation was also defined in our work, and espe-
cially in the central Euxine region; the subeuxine vegetation with
oak forests of Quercus pubescens and Q. cerris, both of which are
characteristicspeciesofsemiaridandaridzonesinAnatolia(Mayer
& Aksoy, 1986), and pin e forests of Pinus nigra and P. sylvestris,
was clear ly different iated from the res t of the region. Howeve r,
no such clear appearance of subeuxine vegetation was detec ted
forthe Colchic region.Here, a differentiation emerged primarily
due to elevation. Due to the higher mountains in Colchis, there is a
rapidtransitionfromtheColchicvegetationtotheIrano-Turanian
phytogeographical region (Davis, 1965–1985), unlike in central
and western Euxine regions. The disappearance of Pinus nigra in
Colchis is also clear evidence of the abrupt transition from the
EuxinetotheIrano-Turanianregion,sinceitisacharacteristicspe-
ciesof thetransitionalzoneamong Euro-Siberian, Irano-Turanian
and Mediterranean phytogeographical regions in other parts of
Anatolia(Çoban&Willner,2019;Kavgacıetal.,2021).
Asaresultoftheelevational,biogeographicalandmacroclimaticvari-
ations, seven main different vegetation groups were determined in central
andeasternEuxineTurkeyandSWGeorgia:(1)relictMediterraneanfor-
ests and shrublands mainly appeared along the coastline, besides some
inlandlocalities;(2)lowlandandsubmontane forests;(3)centralEuxine
mountainforests;(4)eastern Euxine(Colchic)mountainforests;(5) sub-
euxine forests; (6) azonal riparian forests;and (7) subalpine and alpine
shrubland. We will discuss them according to this list.
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FIGURE 4 ForestandshrublandtypesincentralandeasternEuxineTurkeyandSWGeorgiaalongtheelevational,longitudinaland
latitudinal gradients. See Figure 2 and Table 1 for an explanation of the vegetation type numbers.
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4.2 | Syntaxonomyofforestsandshrublandsin
central and eastern Euxine Turkey and SW Georgia
4.2.1 | RelictMediterraneanforestsandshrublands
Western and southern Anatolia is mainly dominated by
Mediterranean vegetation, since these regions are under the effect
of a Mediterranean climate. The syntaxonomy and synecology of
forest and shrubland vegetation in these regions, based on numeri-
calclassificationandordination,waselaboratedindetailbyKavgacı
etal. (2021).However,relictMediterraneanvegetationisalsoseen
in northern Anatolia within the Euro-Siberian phytogeographical
region. This is a result of climate warming after the last glacial age
(Çoban&Willner,2019).
In addition to pine forests (Pinus brutia and P. pinea), sclero-
phyllous forests and shrublands dominated by Arbutus unedo, A.
andrachne, Laurus nobilis, Juniperus excelsa, Olea europea, Quercus
coccifera, Sarcopoterium spinosum and deciduous forests of Carpinus
orientalis are the typical representatives of Mediterranean vegeta-
tioninEuro-SiberianTurkey(Bonarietal.,2021;Kavgacıetal.,2021).
In some veget ation types, evergreen species such as Buxus sem-
pervirens and deciduous species such as Cornus sanguinea, C. mas,
Corylus avellana, Ligustrum vulgare etc., are represented in the floris-
tic composition as dominant or accompanying species. This kind of
vegetation is called pseudomacchia, representing mixed sclerophyl-
lous evergreen and deciduous shrub thickets of the periphery of the
rangeofMediterraneansclerophyllousshrubland(Čarnietal.,2018;
Chytr ý et al., 2020). Thepresenceof manyMediterranean ecosys-
tems in the Euxine region can be a guide in terms of the monitoring
of vegetation changes and preparation of mitigation measures in
management in the region within the scope of the ongoing climate
change.
Tamarix smyrnensis, Juniperus oxycedrus subsp. macrocarpa,
Eleagnus angustifolia and Hippophae rhamnoides-dominated shrub-
lands occurring on lowland sand dune or riverbed sites were also
grouped under relict Mediterranean vegetation. Tamarix smyrnensis-
dominated vegetation onriverbeds innorthernAnatolia was clas-
sified within Rubo sancti- Nerion oleandri of Tamericetalia africanae
and Tamaricetea(Vegetationtype1)(Mucinaetal.,2 016; Mumcu &
Korkmaz,2021).
Juniperus oxycedrus subsp. macrocarpa is a threatened taxon
growingincoastalareasoftheMediterraneanregion(Díez-Garretas
&Asensi,2014).ItscommunitiesincentralEuxineTurkeywereclas-
sified under Juniperion turbinatae(Yalçınetal.,2020).However,this
alliance was identified as an alliance of thermomediterranean tall ju-
niper scrub on coastal dune systems of the western Mediterranean
seaboardsbyMucinaetal.(2016).Díez-GarretasandAsensi(2014 )
divided the alliance Juniperion turbinatae into two suballiances, one
in the western part of the Mediterranean as Juniperenion turbina-
tae, and Asparago orientalis- Juniperion turbinaeappearinginGreece,
Turkeyand Cyrenaica. Mucina(inMucina etal., 2016)changedthe
position of these suballiances to alliances. The thermomediterra-
nean juniper scrub of the coastal dune systems of the central and
eastern Mediterranean seaboards was therefore classified within
the alliance Asparago orientalis-Juniperion macrocarpae of the order
Pistacio lentisci-Rhamnetalia alaterni and the class Quercetea ilicis
(Vegetationtype2)(Mucinaetal.,2016).
Sclerophyllous forests and scrubland of Mediterranean veg-
etation dominated by Olea europaea, Buxus sempervirens, Arbutus
unedo and Laurus nobilis often appear not only along the coast-
line of the Euxine region but also in low elevational belt s of inner
Anatolia (Quézel etal., 1980 ; Yurdakulol etal., 1992 , 2002;Kılınç
&Karaer,1995; Özen &Kılınç, 19 95; Korkmaz et al., 2008, 2011;
Karaeretal.,2010).Althoughthiskindofvegetationwaspreviously
classifiedunderdifferentalliances,Bergmeieretal.(2018)proposed
to classify Laurus nobilis and Arbutus unedo-dominated vegetation
within the alliance of Arbuto unenodis-Laurion nobilis (Vegetation
type3).ThisallianceisplacedwithintheorderQuercetalia ilicis and
class Quercetea ilicis(Mucinaetal.,2016).
Elaeagnus angustifolia-dominated vegetation occurring in
alluvial and colluvial sites in the Euxine region was classified
under Molinio-Arrhenatheretea and Arrhenatheretalia (Karaer
et al., 1999).However,wethinkthatthisclassificationisnotjusti-
fied, since Molinio-Arrhenatheretea is a class of anthropogenic pas-
tures,meadowsandtall-herbmeadowfringesonfertiledeepsoils
FIGURE 5 Detrendedcorrespondenceanalysis(DCA)
ordination of forest and shrubland vegetation t ypes in central and
easternEuxineTurkeyandSWGeorgia.Numbersindicatethe
centroid of vegetation types. Elevation, longitude, latitude and
BIO4 — temperatureseasonality,BIO12 — annualprecipitation,
BIO16 — precipitationofwettestquarterasBioClimdatawere
passively projected onto the ordination diagram, since they were
theonlysignificantvariablesintheCCAtest.Foranexplanationof
vegetation type numbers, see Figure 2 and Table 1.
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at low and mid-alt itudes (rarel y also high elevati ons) of Europe
(Mucinaetal.,2016).Althoughthereisnota clearsyntaxonomi-
cal classification of Elaeagnus angustifolia-dominated vegetation
in Europe (Mucina et al., 2016), we followed the classification
ofShibanova andOvcharova (2021), whoclassified this type of
vegetation under the alliance Galio humifusi-Tamaricion ramosis-
simae, representing xeromesophytic communities dominated by
Elaeagnus angustifolia. However, this alliance is a synonym for
Artemisio scopariae-Tamaricion ramosissimae(Vegetationtype4)
(Mucinaetal.,2016).ThealliancewasplacedwithinTamaricetalia
and Nerio-Tamaricetea.
Pinus brutia and P. pinea exhibit an extensive distributional range
inMediterranean Turkey(Bonarietal.,2020;Kavgacıetal.,2021).
AcomprehensivenumericalclassificationofMediterraneanlowland
to submontane pine forests including P. brutia and P. pinea was car-
ried out byBonari etal. (2021).In thiswork, P. brutia and P. pinea
forests in Euxine Turkey were placed within the alliances Styraco-
Pinion brutiae(Vegetationtype5)andPinion pineae(Vegetationtype
6), respec tively, which was also f ollowed by Kavgacı et a l. (2021)
Both allia nces are included u nder Pinetalia halepensis and Pinetea
halepensis.
Juniperus excelsa-dominatedvegetationshowingMediterranean
characteristics, with the dense appearance of Pistacia terebinthus,
Jasminum fruticans and Paliurus spina-christi, has been classified
under dif ferent syntaxonomical units by different authors, such
as Astragalo-BrometeabyKaraeretal.(1999), Querco calliprini-
Juniperion excelsaebyHamzaoğluetal.(2002) and Astragalo
karamasici-Gypsophilion eriocalycisbyKorkmazetal.(2011).
However,Mucina etal. (2016) classified this kindof vegetation as
the alliance Jasmino-Juniperion excelsae(Vegetation type7),which
consists of relict woods showing a strong Mediterranean influence
inthe Euro-Siberian region. Thisallianceis placedunderBerberido
creticae-Juniperion excelsae and Junipero- Pinetea sylvestris.
Mediterranean Arbutus andrachne and Quercus coccifera-
dominated macchia and forest s have been classified under the al-
liance Arbuto andrachnes-Quercion cocciferae(Kavgacıet al.,2021).
Bergmeie retal.(2018)indicatedthatthisalliancealsoappearsinthe
Euxine region. Arbutus andrachne and Quercus coccifera-dominated
vegetation in our study was therefore also placed within this alli-
ance, belonging to Quercetalia ilicis and Quercetea ilicis(Vegetation
type8).
Carpinus orientalisbuildscommunitiesinthreevegetationzones.
Itappearsin coastal vegetation with a strongMediterraneaninflu-
ence, in the subeuxine region under a continental impact and in the
eastern Euxine under precipitation-rich climatic conditions. Each
example of Carpinus orientalis-dominated vegetation shouldthere-
fore be assessed separately from a syntaxonomical point of view.
Mediterranean Carpinus orientalis forests with the constant appear-
ance of Mediterranean elements such as Cistus creticus, Pistacia
terebinthus subsp. palaestina and Phillyrea latifolia were classified
under the alliance Carpino-Acerion (Özen & Kılınç, 1995). This alli-
ancewasfirstpublishedbyAkmanetal.(1979a),butthenamewas
invalid, since there was no association classified within this alliance
(Art.8 oftheICPN — Theurillatetal.,2021).Thesame authorsalso
failedinthetypificationofthealliancein1992(Quézeletal.,199 2;
Theurillat & Moravec, 1996).Wetherefore describe a new alliance
for Carpinus orientalis forests with a strong Mediterranean influence
that also locally mix with oak species: Querco cerridis-Carpinion orien-
talis under Quercetalia pubescenti- petraeae and Quercetea pubescen-
tis(Vegetationtype9).
A well-established example of Sarcopoterium spinosum-
dominated phrygana is seen in the Sinop peninsula in the study area.
Ithasbeen suggestedthat these communities should be classified
under the class Quercetea ilicis or Cisto-Micromerietea due to the
higher proportion of Mediterranean elements in the floristic com-
positionofthecommunityintheEuxineregion(Karaeretal.,1999).
However,Kavgacıetal.(2021)classifiedthesekindsofcommunities
under the alliance Cisto salvifolii-Lavandulion stoechadis(Vegetation
type 10), belonging to Lavandulo stoechadis-Hypericetalia olympici
and Cisto-Lavandulatea stoechadis,intheirlarge-scalesyntaxonomi-
cal assessment of Mediterranean vegetation in Turkey.
Hippophae rhamnoides subsp. caucasica-dominated scrublands
on sand dunes in Euxine Turkey were classified within the alliance
Ammophilion australis of Ammophiletalia and Ammophiletea(Korkmaz
et al., 2012). However, Mucina et al. (2016) classified Hippophae
rhamnoides-d ominated shrubla nds on dunes under t he alliance of
Pyracantho coccineae-Hippophaeion fluviatilis of Salicetalia arenariae
and Salicetea arenariaeasinourcase( Vegetationtype11).Rareveg-
etation with dominance of Hippophae rhamnoides inhabiting a spe-
cifichabitatofrivergravelbarsinthemountainsinwesternGeorgia
was assigned to the association Salici eleagni- Hippophaëtum rham-
noidis (Kalní ková et al., 2020) This ass ociation belo ngs to the alli-
ance Salicion eleagno- daphnoidis, order Salicetalia purpureae and class
Salicetea purpureae(Kalníková et al.,2021).However,these stands
show significant differencies in species composition compared to
the dune scrublands, for example by the absence of coastal dunes
specialists(cf.Kalníkováetal.,2020).
4.2.2 | Lowlandandsubmontaneforests
The numerical analysis clearly distinguished lowland to submon-
tane Fagus orientalis-dominated forests of central and eastern
Euxine regions from mountainous pure and mixed beech forests
with conif erous specie s. Kavgacı et al . (2012) identifie d five alli-
ances of Fagus orientalis- dominated forests in Euxine Turkey and
Bulgaria:Staphyleo-Buxion, Fagion orientalis, Carpinio- Fagion, Violo-
Fagion and Veronico-Fagion. The last represents Colchic Fagus orien-
talis forests, while the remaining alliances are from the western and
centralEuxin eregions.Mucinaetal.(2016)lateracceptedCarpinio-
Fagion and Violo-Fagion as synonyms of Fagion orientalis. Willner
etal.(2017 )andÇobanandWillner(2019)alsofollowedthesame
approach and classified Fagus orientalis forest in the Euxine region
under a single alliance, Fagion orientalis. They suggested that only
basophilous but not acidophilous beech forests can be found in
Anatolia. They therefore included all thermophilous, mesic and
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fir–beech forests within a single alliance, Fagion orientalis under
Rhododendro pontici-Fagetalia orientalis and Carpino- Fagetea syl-
vaticae. We also follow this approach and classify lowland to
submontane Fagus orientalis-dominated forests of central and
western Euxine regions as Fagion orientalis, which corresponds to
the suballiance Fagenion orientalis(Vegetationtype17)(Çoban&
Willner, 2019).Thissuballiancecanlocallybeplacedatthemoun-
tain belt of central Euxine and also be found in the lowland to sub-
montanebeltsoftheeasternEuxine(Colchic)region.
EasternEuxine(Colchic)lowlandCastanea sativa- and Carpinus
orientalis-dominated forests have been classified under the alli-
ance Castaneo-Carpinion orientalisbyQuézeletal.(198 0, 1992 ).
Nováketal.(2021)alsoclassifiedthese forests underthisalliance.
Therefore, we followed this classification and assigned these forests
under Castaneo-Carpinion orientalis (Vegetation type 25). Quézel
et a l. (198 0)classifiedtheallianceunderRhododendro pontici-Fagetalia
orientalis with the type association Erico-Carpinetum. Later, in the
EuroVegChecklist (Mucin a et al., 2016), the alliance was gro uped
under Carpinetalia betuli. However, Çoban and Willner (2019) in-
dicated the contradiction with the typification of this alliance in
Quézeletal.(1980 ).Inrelationtothat, Novák etal.(2021)pointed
out the need of further study on the syntaxonomical position of the
alliance.Althoughthesyntaxonomicalpositionoftheallianceisnot
still clear, we provisionally group it under Quercetalia pubescenti-
petraeae and Quercetea pubescentis due to the dominance of ther-
mophilous species.
Carpinus betulus and Castanea sativa-dominatedforestsarevery
common elements of lowland to submontane forests all along the
Euxine coast (Vegetation type 26).This type of vegetation isclas-
sified under the alliance Trachystemono orientalis-Carpinion betuli
(Çoban&Willner,2019),whichisplacedunderCarpinetalia betuli and
Carpino-Fagatea sylvaticae.
4.2.3 | CentralEuxinemountainforests
InEuxineTurkey,Corylus avellana appears as the dominant species
insomelocalities,suchasTavşanmountaininthecentralEuxinere-
gion(Yıldırım et al.,2019).These kinds of vegetation are classified
under the alliance Astrantio- Corylion avellanae of Prunetalia spinosae
and Crataego-Prunetea (Veget ation type 16 — Mu cina et al., 2016),
althoughYıldırımetal.(2019)placedthiscommunityunderthealli-
ance Carpino betuli- Acerion hyrcani.
Inouranalyses,puredeciduousforestsofFagus orientalis are
grouped separately from Abies nordmanniana subsp. equi- trojani-
dominatedforests.ÇobanandWillner(2019)mergedallpureand
mixed forests of Fagus orientalis and Abies nordmanniana subsp.
equi- trojani under the alliance Fagion orientalis. We accept the
opinionofÇobanandWillner(20 19)andclassifypuredeciduous
Fagus orientalis-dominated forests mainly distributed in central
and western Euxine region within a single alliance, Fagion orienta-
lis.However,ÇobanandWillner(2019)pointedoutdifferencesof
Fagus-dominatedforests from montaneandsubalpineAbies and
Picea-domin ate dfor estsa ndd escr ibedaneword erf orth esek inds
of forests in the Euxine region as Abieti nordmannianae-Piceetalia
orientalis. However, they descri bed only one allia nce under this
order, Lonicero caucasicae-Piceion orientalis, encompassing conif-
erousforestsfromeasternEuxine(Colchis).
Our analysis showed that central Euxine Abies nordmanniana
subsp. equi- trojani forests are clearly differentiated from eastern
Euxine(Colchic) Abies nordmanniana subsp. nordmanniana and Picea
orientalis forests. Abies nordmanniana subsp. equi- trojani is distributed
in the central and western Euxine regions, while Abies nordmanniana
subsp. nordmannianaspreadsovertheeasternEuxine(Colchic)region
andtheCaucasus(Mayer&Aksoy,1986).Additionally,Picea orientalis
is distributed only in the eastern Euxine region and does not appear
in central and western Euxine regions. These are among the most
importantfloristic differencesoftheeasternEuxine(Colchic)region
that distinguish it from the rest of the Euxine region. The diagnostic
species of Abies nordmanniana subsp. equi- trojani forests also indicate
their difference from from the eastern Abies and Picea forests. These
species are Alchemilla heterophylla, Campanula olympica, Cirsium hy-
poleucum, Euphorbia amygdaloides, Galium radulifolium, Hieracium
oblongum, Lilium martagon, Mercurialis perennis, Myosotis sylvatica,
Orthilia secunda, Pyrola uniflora, Ranunculus brutius, Rubus hirtus,
Sanicula europaea, Saxifraga rotundifolia, Valeriana alliariifolia, Veronica
chamaedrys, and V. montana. Floristic differentiation and the most re-
cent syntaxonomic revision of Abies and Picea forests in the Euxine
region(Çoban&Willner,2019)ledustodescribeanewalliancefor
the central and western Euxine A. nordmanniana subsp. equi- trojani
forests, as Abietion equi-trojani under Abieti nordmannianae-Piceetalia
orientalis and Vaccinio- Piceetea(Vegetationtype18).
Pinus sylvestris has a very broad distribution along the north-
ern Anatolian Mountains (Mayer & Aksoy, 1986). Our analysis
showed that P. sylvestris forests on the Euxine mountains repre-
sent a dif ferent vegetatio n type from sub euxine fores ts. In the
precipitation-rich Euxine mountains, P. sylvestris forests appear
ondeep soils(Quézel et al., 1980 ).They wereclassifiedwithin a
differentalliancethan subeuxine pineforests spreadinginsemi-
arid conditions classified under Cisto-Pinion pallasianae (Akman
et al., 1978). Therefore, we classifiedEuxinemountain P. sylves-
tris forest s into the alliance Geranio iberici-Pinion sylvestris as was
alsosuggestedbyAkmanetal.,1978andBergmeieretal., 2018
(Vegetationtype19).
Quercus petraea is a widely distributed oak species in Turkey
(Yılmaz,2020).It forms pure ormixed forestsin Euxine Turkey.
Its forests have been classified under different alliances,
such as Carpino-Acerion (Yurdakulol et al., 2002; Ketenoğlu
et al., 2010)andQuercion frainetto(Kavgacıetal.,2010; Çoban &
Willner, 2 019). Ouranalysesindicated thatthese thermophilous
oak forests on the Euxine mountains fit well the concept of the
alliance Quercion frainetto, representing thermophilous decidu-
ousoakforestsonslightlyacidicdeepsoils(Vegetationtype20)
(Mucinaetal.,2016).
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4.2.4 | EasternEuxine(Colchic)mountainforests
Alnus glutinosa subsp. barbata is widespread in the floristic composi-
tionofforestvegetationin theColchicregion.Itcanalso be found
inthe floodplain forests ofthecentralEuxine region.InColchis,in
addition to hygrophilous sites and alluvial riverbeds, Alnus glutinosa
subsp. barbataalsoinhabits ravinesand mountainouszonalvegeta-
tion. This is completely related to the humid climate and site condi-
tions of Colchis, which allows Alnus glutinosa subsp. barbata to have
a larger ecological amplitude than in the rest of the Euxine region
(Kavgacıetal.,2016;Nováketal.,2019).Alnus glutinosa subsp. bar-
bata does not therefore form only pure stands but also builds mixed
stands with Carpinus betulus and Castanea sativa in ravines and with
Picea orientalis and Fagus orientalis in moist mountainous forests.
Alnus glutinosa subsp. barbata-dominated forests in Colchis were
classified under the alliance Alnion barbatae of the Rhododendro-
Fagetalia and class Carpino-Fagetea (Quézel et al., 1980, 1992;
Ermakov et al., 2021),andwefollowedthisclassification(Vegetation
type21).
Picea orientalis and Abies nordmanniana subsp. nordmanniana
appear onlyinColchicTurkeyandGeorgia.They are themost sig-
nificant suppor t for the floristic differentiation of Colchis to be
accepted as a separate phytogeographical subunit from the rest
ofEuxineregion(Yaltırık & Efe, 1989).Weclassifiedthese conifer-
ous forests within the alliance Lonicero caucasicae-Piceion orientalis
under Vaccinio- Piceetea, representing Colchic–Caucasian montane
humid spruce and fir forest s as suggested by Çoban & Willner, 2019
(Vegetationtype22).
In Colchis, another vegetation type of subalpine mountain
vegetation is formed by Betula litwinowii (Terzioğlu et al., 2007;
Nakhutsrishvili, 2013). These deciduous shrublands are classified
within the alliance Rhododendro caucasici-Betulion litwinowii of
Rhododendro- Betuletalia litwinowii and Betulo carpaticae-Alnetea vir-
idis(Vegetationtype23)(Mucinaetal.,2016).
Colchic Fagus orientalis-dominatedforests,inwhichPicea orien-
talis may also occur as dominant species, have been classified under
the alliance Veronico-Fagion(Kavgacıetal., 2012).However,Çoban
an d W illner( 2019)showedthatthisalliancewasnotvalidlypublished
because the association Parido orientalis-Piceetum orientalis validat-
ingthealliancewasnotcorrectlytypified(Quézeletal.,199 2).Since
all F. orientalis-dominatedforestsareclassifiedunderthesinglealli-
ance Fagion orientalis, we decided to classify these Colchic forests
of Fagus at the suballiance level: Piceo orientalis-Fagenion orientalis
(Vegetation type 24). Betula medwediewii forests, which were pre-
liminarily classified within Geranio- Pinion(Eminağaoğluetal.,2006),
were also classified within this unit since they were grouped under
the same cluster.
4.2.5 | Subeuxineforests
Oak-dominated forests in subexine areas have been traditionally
classified to different alliances, such as Carpino-Acerion (Kutbay
et al., 1998–1999;Karaer etal., 1999; Cansaran et al., 2010 ;Kaya
et al., 2010; Korkmaz et al.,2011)and Quercion anatolicae(Kutbay
et al., 1998–1999; Hamzaoğlu e t al., 2002). The ori ginal study on
these forests (Akmanetal.,1979a) classified themwithinQuercion
anatolicaeandarecentnumericalstudybyÇobanandW illner(2019)
also approved this classification. In our study, we used the valid
name of Quercus anatolica: as Quercus pubescens subsp. crispata
(Euro+Med, 2006) and named the alliance as Quercion crispatae
(Vegetationtype12).
Pinus nigra subsp. pallasianaisgenerallycharacterizedasa
tree species of transitional areas from maritime to continental cli-
maticregions(Mayer&Aksoy,1986).Weacce ptedthepropos alof
Akmanetal .(1979 a)andclassifiedP. nigra subsp. pallasiana forests
of the subeuxine areas within the alliance Cisto laurifolii-Pinion pal-
lasianae under Erico-Pinetalia and Erico-Pinetea (Vegetationtype
13). We also grouped P. sylvestris- dominated forests from the
subeuxine province under this alliance due to their floristic sim-
ilarity to P. n igra fores ts and their coniferouscha racter (Akman
et al., 1979b).
4.2.6 | Azonalriparianforests
Typical floodplain forests in the study area, appearing in habitats
where the water table is usually at or near the surface and the sur-
face is periodically or at least occasionally flooded by shallow water,
can be found in the central Euxine region (Kavgacı et al., 2016).
These floodplain forests dominated by Fraxinus angustifolia and
Alnus glutinosa subsp. barbata are classified within the alliance
Periploco graecae-Fraxinion angustifoliae under Populetalia albae
and Alno glutinosae-Populetea albae (Vegetation ty pe 27) (Kavgacı
et al., 2016).
Platanus orientalis is a typical tree species of gallery for-
estsalongstreamsand rivers andmainlycolonizespoorly stabi-
lized alluvial sitesrich ingravel orstone depositsin theeastern
Mediterranean (Mandžkovski et al., 2021). These forests are
classified within the alliance Platanion orientalis under Populetalia
albae and Alno glutinosae-Populetea albae (Vegetation type 28)
(Doudaetal.,2016).
4.2.7 | Subalpineandalpineshrubland
The subalpine and alpine shrublands in the study area are built
by Epigaea gaultherioides, Juniperus communis subsp. nana,
Rhododendron caucasicum and Vaccinium myrtillus-dominated
plant communities. Epigaea gaultherioides-dominated shrub-
lands are classified within Vaccinion myrtilli(Vegetationtype14),
while Juniperus communis subsp. nana communities are under
Daphne oleoidis-Juniperion alpinae (Vegetati on type 15) (Kavga cı
et al., 2021) and Rhododendron caucasicum-dominated com-
munities are assigned to Rhododendrion caucasici (Vegetation
type 29). A ll of these three all iances are placed un der Vaccinio
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Applied Vegetation Science
KAV GAC I et a l.
microphylli-Juniperetalia nanae and Loiseleurio procumbentis-
Vaccinietea(Mucinaetal.,2016).
4.2.8 | Syntaxonomicalscheme
Accordingtotheabovesyntaxonomicdiscussiononcentralandeast-
ern(Colchic)EuxineforestsandshrublandsinTurkeyandSWGeorgia,
the follow ing syntaxo nomic schem e is proposed (E VC: alliance pr e-
sent in theEuroVegChecklist (Mucina et al., 2016); notEVC: alliance
notpresentintheEuroVegChecklist;New:Newlydescribedalliance).
Loiseleurio procumbentis- Vaccinietea microphylli Eggler ex
Schubert 1960
Rhododendro ferruginei- Vaccinetalia microphylli Br.-Bl. in
Br.-Bl.etJenny1926
Vaccinion myrtilli Krajina 1933 (Vegetation type 14 in
Figure 2 and Table 1)(EVC)
Rhododendrion caucasici Onipchenko 2002 (Vegetation
type 29 in Figure 2 and Table 1)(EVC)
Vaccinio microphylli- Juniperetalia nanae Rivas-Mart. et M.
Cost a 1998
Daphno oleoidis- Juniperion alpinae Stanisci 1997
(Vegetationtype15inFigure 2 and Table 1)(EVC)
Vaccinio- PiceeteaBr.-Bl.inBr.-Bletal.1939.
Abieti nordmannianae-Piceetalia orientalis Çoban et Willner
20 19.
Geranio iberici- Pinion sylvestris Quézel et al. ex Quézel
et al. 199 2 (Vegetation type 19 in Figure 2 and Table 1)
(notEVC).
Lonicero caucasicae- Piceion orientalis Çoban et Willner
2019(Vegetationtype22inFigure 2 and Table 1)(notEVC).
Abietion equi- trojani all. nova (Vegetation type 18 in
Figure 2 and Table 1)(New).
Carpino- Fagetea sylvaticaeJakucsexPassarge1968
Rhododendro pontici-Fagetalia orientalisPassarge1981.
Fagion orientalisSoó1964(EVC).
Fagenion orientalis(Vegetationtype17inFigure 2 and
Table 1).
Piceo orientalis- Fagenion orientalis suball. nova
(Vegetationtype24inFigure 2 and Table 1)
CarpinetaliabetuliP.Fukarek1968
Trachystemono orientalis- Carpinion betuli Çoban and
Willner 2019(Vegetationtype26inFigure 2 and Table 1)
(notEVC)
Quercetea pubescentisDoing-KraftexScamonietPassarge1959
Quercetalia pubescenti- petraeaeKlika1933
Quercion frainetto Horvat 1958 nom. corr. (Vegetation
type 20 in Figure 2 and Table 1)(EVC)
Castaneo sativae- Carpinion orientalis Quézel et al. ex
Quézel et a l. 1992 ( Vegetation typ e 25 in Figure 2 and
Table 1)(EVC).
Querco cerridis- Carpinion orientalis all. nova (Vegetation
type 9 in Figure 2 and Table 1)(New)
Quercion crispataeAkmanetal.exQué zeletal.19 92 nom.
corr.(Vegetationtype12inFigure 2 and Table 1)(notEVC)
Carpino-Prunetea Tx. 1962
Prunetalia spinosae Tx. 1952
Astrantio- Corylion avellanae Passarge 1978 (Vegetation
type 16 in Figure 2 and Table 1)(EVC)
Salicetea arenaria Weber 1999
Salicetalia arenariaePreisingetWeber1997
Pyracantho coccineae- Hippophaion fluviatilis de Foucault
etJulve2001(Vegetationtype11inFigure 2 and Table 1)
(EVC)
Junipero-Pinetea sylvestrisRivas-Mart.1965nominvers.
Berberido creticae-Juniperetalia excelsae Mucina in Mucina
et al. 2016
Jasmino- Juniperion excelsae Didukh, Vakarenko et
Shelya g-Sosonko e x Didukh 1996 (Vegetation t ype 7 in
Figure 2 and Table 1)(EVC)
Erico-PineteaHorvat1959
Erico-PinetaliaHorvat1959nom.conserv.prop.
Cisto laurifolii- Pinion pallasianae Ak man et al. ex Qué zel
et al. 199 2 (Vegetationtype 13 in Figure 2 and Table 1)
(notEVC)
Betulo carpaticae-Alnetea viridisRejmáneketal.exBoeufetal.in
Boefetal.2014
Rhododendro caucasici-Betuletalia litwinowii Mucina in
Mucina et al. 2016
Rhododendro caucasici- Betulion litwinowii
Onipchenko 2002 ( Vegetation type 23 in Figure 2 and
Table 1)(EVC)
Quercetea ilicisB r.-Bl. ex A . et O. Bolòs in A . Bolòs y Vayreda
1950.
Quercetalia ilicisBr.-Bl.exMolinier1934
Arbuto unedonis- Laurion nobilis Rivas-Mart. et al. 1999
(Vegetationtype3inFigure 2 and Table 1)(EVC)
Arbuto andrachnes- Quercion cocciferae Barbero et
Quézel1979(Vegetationtype8inFigure 2 and Table 1)
(EVC)
Pistacio lentisci-Rhamnetalia alaterniRivas-Mart.1975.
Asparago orientalis- Juniperion macrocarpae(DiezGarretas
etAsensi2014)MucinainMucinaetal.2016(Vegetation
type 2 in Figure 2 and Table 1)(EVC).
Pinetea halepensisBonarietChytrýinBonarietal.2021
Pinetalia halepensisBiondietal.inBiondietal.2014
Styraco officinalis- Pinion brutiae Bonari et al. 2021
(Vegetationtype5inFigure 2 and Table 1)(notEVC).
Pinion pineaeFeinb run1959( Vegetatio nty pe6inFigure 2
and Table 1)(EVC)
Cisto- Lavanduletea stoechadisBr.-Bl.inBr.-Bl.etal.1940
Lavandulo stoechadis- Hypericetalia olympici Mucina in Mucina
et al. 2016
1654109x, 2023, 4, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/avsc.12753 by Cochrane Greece, Wiley Online Library on [01/03/2024]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License