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Tuexenia 41: 37–51. Göttingen 2021.
doi: 10.14471/2021.41.012, available online at www.tuexenia.de
Carpinus orientalis forests in Georgian Colchis:
First insights
Carpinus orientalis-Wälder in der georgischen Kolchis: Erste Einblicke
Pavel Novák1 * , Vladimir Stupar2 & Veronika Kalníková1, 3
1Department of Botany and Zoology, Faculty of Science, Masaryk University, Kotlářská 267/2,
61137 Brno, Czech Republic; 2Faculty of Forestry, University of Banja Luka, S. Stepanovića 75A,
78000 Banja Luka, Bosnia and Herzegovina; 3Beskydy Protected Landscape Area Administration,
Nádražní 36, 756 61 Rožnov pod Radhoštěm, Czech Republic
*Corresponding author, e-mail: pavenow@seznam.cz
Abstract
Colchis (Caucasus Ecoregion, Euxinian Province) is a region with unique Tertiary relict biota and
high species and vegetation diversity. However, its vegetation has been only little studied by Braun-
Blanquet methods so far. Based on original field data (20 phytosociological relevés), we describe
a novel vegetation type of calciphilous and thermophilous Carpinus orientalis forests in western Geor-
gia (Campanulo alliariifoliae-Carpinetum orientalis ass. nova). This species-rich community inhabits
sunny limestone slopes and is developed under a humid warm-temperate climate. We present the com-
munity in the context of relevés from the literature (n = 105 in total) of Carpinus orientalis dominated
or co-dominated forests across the whole Euxinian Province (southern Black Sea coast). Ordination and
unsupervised classification analyses revealed the main pattern in their species composition closely
linked to biogeography backed up by macroclimatic gradients and vegetation history. Eastwards, Bal-
kan and Mediterranean species decrease gradually, while Euxinian and Euxino-Caucasian species are
more frequent. Although the analysed forest communities were highly variable in species composition,
they all shared a subset of submediterranean and Euxinian species. Numerous Eastern Euxinian and
Euxino-Caucasian endemics (e.g. Campanula alliariifolia, Klasea quinquefolia) are characteristic of the
community recorded in Georgia. These are accompanied by evergreen species (e.g. Smilax excelsa,
Vinca major subsp. hirsuta) and common forest mesophytes (e.g. Campanula rapunculoides, Carex
digitata) both indicating a relatively mild and precipitation-rich climate. The association Erico-
Carpinetum described in NE Turkey was identified as the most similar unit to the new community. As
it is the type association of the alliance Castaneo sativae-Carpinion orientalis, we adopted this assign-
ment for the new association from Georgia.
Keywords: biogeography, Carpinus orientalis, Caucasus, Colchis, ecology, Euxinia, forest vegetation,
Georgia, phytosociology
Erweiterte deutsche Zusammenfassung am Ende des Artikels
1. Introduction
Colchis is an area encompassing the south-eastern shore of the Black Sea in Turkey,
Georgia and Russia. It covers the western slopes of the Greater and Lesser Caucasus, the
northern slopes of the Pontic range and the Colchic Lowland. It is well-known for its unique
Manuscript received 30 March 2021, accepted 10 August 2021
Published online 31 October 2021
Co-ordinating Editor: Péter Török
38
relict biota (ZAZANASHVILI et al. 2000, NAKHUTSRISHVILI et al. 2015). It is one of the most
important Tertiary refugia across the temperate zone of the northern hemisphere. The biolog-
ical legacy of those times has survived in only a few warm-temperate regions with an excep-
tionally stable environment (MILNE & ABBOTT 2002). Colchis is a part of the Caucasus
Ecoregion. The Caucasus is listed among the top 34 biologically richest and most endan-
gered biodiversity hotspots globally (MITTERMEIER et al. 2004, ZAZANASHVILI & MALLON
2009). In terms of phytogeography, Colchis represents an eastern sector of the Euxinian
Province encompassing the southern coast of the Black Sea between Bulgaria and Russia
(TAKHTAJAN 1986). Euxinia has diverse flora and vegetation reflecting its position between
the Balkans, the Mediterranean Basin, Anatolia and the Caucasus. Deciduous forests with
evergreen Tertiary relict shrubs (e.g. Ilex colchica, Prunus laurocerasus, Rhododendron
ponticum) are its unique feature (TAKHTAJAN 1986). They were identified as temperate rain-
forests by some authors (e.g. NAKHUTSRISHVILI et al. 2015). Recent phylogeographical and
phylogenetic research on forest vascular plants confirmed the unique biogeographical posi-
tion of the province within western Eurasia (e.g. GRIMM & DENK 2014, VOLKOVA et al.
2020). Euxinia is also a region of diverse climate regimes. Its Mediterranean character with
dry summer periods gradually disappears eastwards and the easternmost part of the province
is precipitation-rich throughout the year. The climate of Colchis is humid warm-temperate
(WALTER 1970). With annual precipitation of around 1800–2000 mm (but locally exceeding
4000 mm), it is one of the most humid regions of western Eurasia. The absence of regular
winter frosts at low elevations is another essential climatic feature of Colchis (DENK et al.
2001, NAKHUTSRISHVILI et al. 2015).
The forests of Euxinia have been a subject of phytosociological research mainly in Tur-
key (e.g. QUÉZEL et al. 1980, KORKMAZ et al. 2011, KAVGACI et al. 2012, ÇOBAN & WILL-
NER 2019) and Bulgaria (e.g. TZONEV et al. 2019). Studies from Georgia published to date
have dealt with mesophilous forest types (e.g. FILIBECK et al. 2004, NOVÁK et al. 2019),
while almost no attention has been paid to thermophilous types. The presented study is fo-
cused on Carpinus orientalis (hereafter “C. orientalis”) forests of the limestone massifs of
western Georgia. Carpinus orientalis is a submediterranean deciduous tree distributed in
south-eastern Europe, Anatolia, Syria, Crimea, Caucasus and northern Iran. It is a ther-
mophilous and xerophilous species preferring calcareous soils, well-known is its tolerance to
various traditional management practices (SIKKEMA & CAUDULLO 2016). Forests dominated
or co-dominated by C. orientalis occur across the entire Euxinia (QUÉZEL et al. 1980,
ÇOBAN & WILLNER 2019, AKHALKATSI 2019). Generally, they prefer sunny slopes on vari-
ous bedrock. However, they may represent a dominant forest type in coastal regions under
the influence of the Mediterranean climate. They form both low and open, as well as tall and
closed stands (QUÉZEL et al. 1980). Colchic C. orientalis forests are partly supposed to be
a regeneration stage after the degradation of oak or mixed forests (DOLUKHANOV 2010,
AKHALKATSI 2015). Locally, they served as coppices (AKHALKATSI 2015), similarly to the
Balkans (STUPAR et al. 2016) or Italy (BLASI et al. 2001).
The vegetation of thermophilous deciduous forests is still relatively understudied in Eux-
inia and the Caucasus, at least by Braun-Blanquet methods (cf. ÇOBAN & WILLNER 2019,
MUCINA et al. 2016). They have been investigated mainly in the western part of the area (see
citations above). Despite their broad distribution in the eastern part (GULISASHVILI et al.
1975, NAKHUTSRISHVILI 2013), they have been studied only marginally so far, although
some types are listed in national red lists of habitats (e.g. AKHALKATSI 2019). Within
39
the Caucasus Ecoregion, dry-mesic forests of C. orientalis and Zelkova carpinifolia have
recently been described phytosociologically in eastern Georgia (NOVÁK et al. 2020) and in
Hyrcania in northern Iran (GHOLIZADEH et al. 2020).
Due to the lack of knowledge on the phytosociology of thermophilous Colchic forests
and their position within the Euxinian forest vegetation, the goals of this study are (1) to
describe the ecology and species composition of the newly recorded C. orientalis forests in
western Georgia, and (2) to determine the ecological and floristic relationships between the
community studied here and the C. orientalis stands documented so far in Euxinia.
2. Study region
The study region stretches between the cities of Chiatura (Imereti Region) and Jvari
(Samegrelo Region) in western Georgia (42°17'–42°46' N, 42°02'–43°18' E). The sampling
sites were located at low elevations (90–570 m a.s.l.) of the limestone massifs of Kvira,
Senaki, Askhi, Tskaltubo-Kutaisi, Racha and Chiatura. Rendzic leptosol developed on Cre-
taceous limestones is the main soil type of the sampling sites (URUSHADZE & GHAMBA-
SHIDZE 2013, ASANIDZE et al. 2019). The region represents a transition zone between the
Colchic and Greater Caucasian climatic zones (BONDYREV et al. 2015). The sampling sites
have mean annual temperature of 11.8–14.8 °C and annual precipitation of 999–2074 mm
(KARGER et al. 2017). The temperature may drop slightly below freezing point in the winter.
Rainfall is distributed almost evenly over the year (Fig. 1) and is accompanied by frequent
horizontal precipitations brought by moist air masses from the Black Sea. However, the
water-permeable limestone bedrock presumably weakens the effect of humid climate on
vegetation (DENK et al. 2001, DOLUKHANOV 2010, BONDYREV et al. 2015).
The region is covered by extensive forests, generally deciduous (BOHN et al. 2000–2003,
NAKHUTSRISHVILI 2013). They harbour a significant diversity of understorey species, includ-
ing narrow endemics (KOLAKOVSKII 1961), both shrubs (e.g. Corylus colchica, Staphylea
colchica) and herbs (e.g. Cyclamen colchicum, Peucedanum adae). It is an area with one of
the highest concentrations of endemic plant species within Georgia (SLODOWICZ et al. 2018).
In the study region, forests dominated or co-dominated by Carpinus orientalis are reported
as a distinctive vegetation type of steep limestone slopes at low elevations (DOLUKHANOV
2010, NAKHUTSRISHVILI 2013). They generally occur up to 800 m, i.e. within the zone of
mixed Colchic forests (e.g. Carpinus betulus, Castanea sativa, Fagus orientalis, Quercus
petraea subsp. iberica) with evergreen shrubs.
The forests in the study region have been disturbed by human activities in many ways,
especially since the middle of the 20th century. This includes overlogging, setting forest
fires, building activities and overgrazing by domestic animals, all followed by soil erosion
and landslides, tree pathogen outbreaks and general degradation of forest ecosystems
(AKHALKATSI 2015). Only a negligible part of these forests lies within protected areas.
3. Methods
3.1 Field sampling
During the field survey (2016–2019), we recorded 20 square-shaped phytosociological relevés
(100 m2). The aim was to sample forests dominated or co-dominated by Carpinus orientalis (except for
one relevé dominated by the ecologically similar Zelkova carpinifolia; DENK et al. 2001). In each
relevé, we assessed the percentage covers of tree (E3), shrub (E2), herb (E1) and moss (E0) layers and
40
covers of species of vascular plants in each layer using the nine-degree Braun-Blanquet scale
(DENGLER et al. 2008). Furthermore, we estimated mean heights of the tree, shrub and herb layers. For
each relevé, we determined slope inclination and aspect and cover of rocks. We collected soil samples
from a depth of 5–10 cm for soil pH measuring done in a suspension of a dried soil sample (particles
< 2 mm) with distilled water (2:5) by a portable Greisinger instrument. The geographical position
(WGS 84) and elevation of the relevés were acquired by a portable GPS device. The relevés were
stored in the Turboveg 2.1 database (HENNEKENS & SCHAMINÉE 2001) and processed in Juice 7.1
software (TICHÝ 2002). The vascular plant nomenclature was standardized following the Euro+Med
PlantBase (http://ww2.bgbm.org/EuroPlusMed/; accessed 2020–10–01) except for one ad hoc aggre-
gate (Carex muricata aggr. – Carex divulsa, C. muricata, C. spicata) and all Rubus taxa recorded dur-
ing the field survey were merged as Rubus subgen. Rubus (SOCHOR & TRÁVNÍČEK 2016). The relevés
were also stored in the AS-00–005 Transcaucasian Vegetation Database included in the European
Vegetation Archive (CHYTRÝ et al. 2016).
3.2 Dataset and data analyses
We compiled a dataset of original relevés from Georgia and relevés from the literature to compare
biogeographical and environmental aspects of the newly described community with previously de-
scribed C. orientalis forests described to date from the rest of Euxinia. We included relevés of associa-
tions dominated or co-dominated by C. orientalis. These were extracted from the following sources:
QUÉZEL et al. (1980) (Carpinetum betulo-orientalis, n = 11 relevés; Crataego curvisepalae-Quercetum
cerridis, n = 7; Erico arboreae-Carpinetum orientalis, n = 16; Rusco aculeatae-Carpinetum orientalis,
n = 7; missing association assignment of the community, n = 3), KUTBAY & KILINÇ (1995) (Carpino
orientalis-Quercetum cerridis, n = 15; Carpino orientalis-Phillyrietum latifoliae, n = 6), YARCI (2002)
(Querco cerridis-Carpinetum orientalis, n = 10) and KORKMAZ et al. (2011) (Corno mari-Quercetum
cerridis, n = 10). The dataset contained 105 relevés in total. Relevés from the literature were georefer-
enced based on descriptions of sampling sites. Climatic data were obtained from the CHELSA Bioclim
dataset (KARGER et al. 2017). For each relevé, average values of essential climatic variables (mean
annual temperature, annual precipitation, temperature seasonality, precipitation seasonality) were re-
trieved as a mean value from a circular buffer zone of 25 km2. We applied classification (flexible beta
clustering with parameter β = -0.2 and Bray-Curtis distance) and ordination (NMDS) analyses to show
ecological and floristic patterns in the dataset. Taxa determined to only the genus level were omitted,
records of same-name species were merged across the layers (FISCHER 2015) and percentage cover
values of species were square-root transformed prior to the analyses (TICHÝ et al. 2020). The ordination
analysis was computed using the package vegan 2.5-7 (OKSANEN et al. 2020) in the R 4.0.2 environ-
ment (R CORE TEAM 2020). Species-to-cluster fidelity was expressed as the phi coefficient (CHYTRÝ et
al. 2002). Diagnostic (Φ ≥ 0.25) and highly diagnostic (Φ ≥ 0.55) species are provided for each cluster.
Before its calculation, the number of plots per cluster was virtually equalized (TICHÝ & CHYTRÝ 2006).
Fisher’s exact test (p ≥ 0.05) was applied to omit species with non-significance occurrence from the list
of diagnostic species.
The new association was formally described following the 4th edition of the Code of Phytosociolog-
ical Nomenclature (THEURILLAT et al. 2021). Due to the many unresolved issues in the syntaxonomy of
the Euxinian forest vegetation, the first mention of a syntaxon is accompanied by an author citation,
except for classes whose nomenclature follows MUCINA et al. (2016).
4. Results and discussion
4.1 Carpinus orientalis community in Georgian Colchis
In the study region, Carpinus orientalis forests inhabit various topographic positions,
generally with soil water shortage. They favour steep slopes (mean inclination 30°) with
rugged rock outcrops (Fig. 1). They also occur on hilltops and plateau edges. Stony topsoil
41
Fig. 1. a) Carpinus orientalis forests below the Motsameta Monastery near Kutaisi, site of the type
relevé of the association Campanulo alliariifoliae-Carpinetum orientalis ass. nova. b) Carpinus orien-
talis forests near Tsutskhvati. c) Limestone landscape around Katski Pilar Monastery where Carpinus
orientalis forests dominate (Photos: P. Novák, a) July 2019 b) July 2017 c) May 2018). d) Walter-type
climadiagram for Motsameta (based on CHELSA Bioclim dataset; KARGER et al. 2017). Additional
photos are provided in Supplement E4.
Abb. 1. a) Carpinus orientalis-Wälder unterhalb des Motsameta-Klosters bei Kutaisi, Lokalität der
Typusaufnahme des Campanulo alliariifoliae-Carpinetum orientalis ass. nova. b) Carpinus orientalis-
Wälder bei Tsutskhvati. c) Kalkgesteinslandschaft im Bereich des Klosters Katski Pilar, wo Carpinus
orientalis-Wälder dominieren (Fotos: P. Novák, a) Juli 2019 b) Juli 2017 c) Mai 2019). d) Klimadia-
gram nach Walter für Motsameta (basierend auf dem CHELSA Bioclim Datensatz; KARGER et al.
2017). Zusätzliche Fotos sind in Anhang E4 zu finden.
of neutral to slightly alkaline reaction (pH 6.6–7.8) reflects the limestone bedrock. It is
a relatively species-rich community (30–45 species per 100 m2).
The tree canopy is rather closed (mean cover 83%), generally reaching around 10 m
in height. The dominant tree C. orientalis is often accompanied by other deciduous trees
(e.g. Acer campestre, Carpinus betulus, Fraxinus excelsior), and infrequently also endemic
and subendemic ones (e.g. Acer cappadocicum, Quercus hartwissiana and Zelkova carpini-
folia). Frequent multi-trunk C. orientalis trees indicate former coppice management. Smilax
excelsa, a widespread evergreen liana in the community, is a characteristic species of re-
gions with mild winters in Euxinia, the Caucasus and northern Iran (NAKHUTSRISHVILI 2013,
GHOLIZADEH et al. 2020, NOVÁK et al. 2020). In contrast to mesophilous Colchic for-
ests (NAKHUTSRISHVILI 2013, NOVÁK et al. 2019), endemic Hedera colchica is usually
a)
b)
c)
d)
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substituted by H. helix which has lower air humidity requirements (DOLUKHANOV 1980).
The shrub layer is mostly developed (mean cover 14%). It consists of species of the canopy
accompanied by deciduous shrubs (e.g. Cornus mas, Corylus avellana, Staphylea colchica).
Evergreen Colchic shrubs are generally absent except for Ilex colchica which is remarkably
tolerant to shallow calcareous soils (DOLUKHANOV 1980). Buxus sempervirens was previous-
ly common (cf. DOLUKHANOV 2010, Supplement E4). Its ongoing massive dieback induced
by a co-invasion of alien fungus and insect pathogens has seriously damaged a large portion
of Colchic Buxus population since 2009 when damages were firstly detected (MATSIAKH et
al. 2018). That will presumably influence affected forest ecosystems by increasing slope
erosion, modifying forest succession and loss of species obligate on Buxus (MITCHELL et al.
2018). The herb layer (mean cover 37%) usually lacks a clear dominant species. Evergreen
species including Ruscus aculeatus and Vinca major subsp. hirsuta are frequent and co-
dominate in places together with lianas. The community is rich in Caucasian endemic and
subendemic forest mesophytes (e.g. Campanula alliariifolia, Symphytum grandiflorum,
Veronica peduncularis). However, species diagnostic for the Colchic mesophilous forests
(e.g. Polystichum woronowii, Pteris cretica, Ruscus colchicus; NOVÁK et al. 2019) are al-
most absent. A group of xerophilous species also shows rather high endemism (e.g. Digitalis
schischkinii, Peucedanum adae, Klasea quinquefolia), alongside broadly distributed species
(e.g. Asplenium adiantum-nigrum, Teucrium chamaedrys). A mixture of mesophilous and
xerophilous species in the undergrowth of C. orientalis forests has also been reported in
other regions of their common occurrence, e.g. the Balkans (STUPAR et al. 2015), Italy
(BLASI et al. 2001) and Crimea (DIDUKH 1996). Submediterranean forest generalists are
represented by Dioscorea communis, Potentilla micrantha and Viola alba for instance.
Shade-tolerant species of rock crevices (e.g. Asplenium ruta-muraria, A. trichomanes) in-
habit rock outcrops. The humid climate supports ferns (Adiantum capillus-veneris, Aspleni-
um scolopendrium) which generally avoid C. orientalis forests in south-eastern Europe ex-
cept for the bottoms of limestone canyons (STUPAR et al. 2020). Local populations of pasture
weeds (e.g. Leontodon hispidus, Plantago lanceolata, Pteridium aquilinum) indicate occa-
sional wood-pasture management. Alien species (sensu KIKODZE 2010) are mostly rare in
the community. There are two exceptions – the stoloniferous grass Oplismenus hirtellus
subsp. undulatifolius which is common across Colchic forests (DOLUKHANOV 2010) and
Robinia pseudoacacia which used to be planted in Georgia and is now spreading spontane-
ously (AKHALKATSI 2015). The moss layer is mostly well-developed (mean cover 16%),
preferentially colonizing rock outcrops. Moreover, epiphytic bryophytes often cover tree
trunks, as a characteristic feature of Colchic forests, due to high rainfall and frequent fogs
(KÜRSCHNER et al. 2012).
The community resembles the formerly described forest type “Carpinuleto-Querceta
ruscosa (Ruscus colchicus)”, and partly also “Carpinuleto-Querceta seslerietosa” inhabiting
more rocky sites, both reported from the study region (DOLUKHANOV 2010, NAKHUTSRISH-
VILI 2013). In terms of nature protection, stands with Buxus sempervirens or Zelkova
carpinifolia are particularly important as both are listed among the priority habitat types of
Georgia (AKHALKATSI 2019).
It should be emphasized that the C. orientalis forests of Georgian Colchis require further
phytosociological research, as their occurrence is also reported on well-drained soils of
riverine terraces in the Colchic Lowland as well as on limestones and sands in the Black Sea
coastal zone in the north-western part of the country (KOLAKOVSKII 1961, DOLUKHANOV
2010, NAKHUTSRISHVILI 2013).
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4.2 Colchic Carpinus orientalis forests in the context of Euxinian forest vegetation
The comparison of the newly recorded community (Supplements E1–E2) with previous-
ly reported Euxinian communities dominated or co-dominated by C. orientalis (Fig. 2–3,
Table 1) resulted in three biogeographically distinct main clusters recognized in the unsu-
pervised classification. We interpreted them at the alliance level.
The first main cluster involved C. orientalis forests of the eastern part of Euxinia. East-
ern Euxinian and Euxino-Caucasian endemics (e.g. Campanula alliariifolia, Klasea quin-
quefolia, Vinca major subsp. hirsuta) were among its highly diagnostic species. They were
accompanied by Mediterranean evergreen shrubs with a scattered distribution across the
whole of Euxinia (e.g. Arbutus andrachne, Buxus sempervirens, Cistus salviifolius, Erica
arborea; cf. DONNER 1990, DENK et al. 2001, NAKHUTSRISHVILI 2013). Its syntaxonomic
interpretation is discussed below. The second main cluster contained mainly C. orientalis
forests of the central part of Euxinia. Mesophytes, mostly Euxinian endemics and subendem-
ics (e.g. Asperula cimulosa, Cirsium hypoleucum), were diagnostic. Mesophilous tree spe-
cies (e.g. Carpinus betulus, Fagus orientalis) often co-dominated with C. orientalis. Numer-
ous mesophytes indicated its transitional position between thermophilous forests and meso-
philous Euxinian oak-hornbeam forests of the alliance Trachystemono orientalis-Carpinion
betuli Çoban & Willner 2019 (class Carpino-Fagetea). The third main cluster unified forests
of C. orientalis and Quercus cerris recorded predominantly in western Euxinia. Mediterra-
nean (e.g. Phillyrea latifolia, Styrax officinalis) and Balkan (e.g. Quercus cerris) species
with a limited distribution in Euxinia (cf. DONNER 1990) and nitrophytes (e.g. Alliaria petio-
lata, Viola odorata) were diagnostic. Following the classification presented by ÇOBAN &
WILLNER (2019), we assigned it under the alliance Quercion confertae Horvat 1958 (class
Quercetea pubescentis).
The classification and ordination analyses revealed biogeography as the key factor de-
termining the variability of the Euxinian C. orientalis forests. Species characteristic of the
Balkan dry forests and partly also Mediterranean species gradually retreat eastwards while
Eastern Euxinian and Euxino-Caucasian endemics become more frequent (see the examples
above). Analogous patterns have been reported, for instance, for Euxinian oak-hornbeam
forests (NOVÁK et al. 2019) and to some extent also for oriental beech forests (KAVGACI et
al. 2012). This shift in species composition seems to be an essential pattern of the Euxinian
flora (cf. DONNER 1990). Moreover, in Euxinia, the precipitation seasonality characteristic of
Fig. 2. Distribution of relevé sites of the dataset. Different symbols refer to their cluster assignment.
Abb. 2. Verteilung der Lokalitäten der Vegetationsaufnahmen des Datensatzes. Unterschiedliche Sym-
bole beziehen sich auf ihre Cluster-Zuordnung.
44
Fig. 3. NMDS analysis of the dataset with relevé-to-cluster assignment and centroids of the clusters
based on the classification analysis (see dendrogram in the right upper part). Vectors of geographical
position (in black) and climatic variables (in blue) were passively plotted. Stress value = 0.214.
Abb. 3. NMDS-Analyse des Datensatzes mit Zuordnung der Vegetationsaufnahmen zu Clustern und
Zentroiden der Cluster basierend auf der Klassificationsanalyse (s. Dendrogramm rechts oben). Vekto-
ren der geographischen Lage (in Schwarz) und der klimatischen Variablen (in Blau) wurden passiv
geplottet. Stresswert = 0,214.
the Mediterranean and the southern Balkans is decreasing towards the humid Colchis
(QUÉZEL et al. 1980, DENK et al. 2001). Therefore, we assume that the observed biogeo-
graphical pattern is driven by both vegetation history and macroclimatic gradients.
The finer division of the dataset into nine clusters (Fig. 2–3, Table 1) reproduced most of
the analysed associations and communities relatively well, indicating their distinctive floris-
tic composition (see Supplement E3). The Georgian community formed its own cluster
within the first main cluster. Across the dataset, it possessed a unique combination of East-
ern Euxinian and Caucasian species.
In the finer classification, the association Erico-Carpinetum Quézel et al. 1992 was iden-
tified as the association most similar to the Georgian community. Erico-Carpinetum is the
type association of the alliance Castaneo sativae-Carpinion orientalis Quézel et al. 1992. In
the EuroVegChecklist (MUCINA et al. 2016), the alliance was assigned to the order
45
Carpinetalia betuli P. Fukarek 1968, class Carpino-Fagetea. However, Çoban & Willner
(2019) emphasized that this classification was in contradiction with its typification per-
formed by QUÉZEL et al. (1992). As Erico-Carpinetum was identified as its type association,
the alliance should unify thermophilous and xerophilous forests. Therefore, we classify the
Georgian community under the alliance Castaneo-Carpinion. However, the position of the
alliance itself deserves further study, as QUÉZEL et al. (1992) designated it as the type of the
order Rhododendro pontici-Fagetalia orientalis Quézel et al. 1992, encompassing Euxinian
deciduous forests, whose syntaxonomic concept is disputed (cf. MUCINA et al. 2016).
In the context of the Caucasian vegetation, there are many striking differences between
the Colchic community and the C. orientalis stands of central and eastern Georgia (cf.
DOLUKHANOV 2010, NAKHUTSRISHVILI 2013). The Colchic community harbours numerous
Euxinian species, though it lacks characteristic flora of more arid regions of Transcaucasia
(e.g. Astragalus spp., Juniperus spp., Rhamnus pallasii, Spiraea hypericifolia). Compared to
C. orientalis forests reported in Hyrcania (GHOLIZADEH et al. 2020), there is a notable ab-
sence of the Hyrcanian floral element (e.g. Centaurea hyrcanica, Digitalis nervosa) and the
presence of Euxinian and Euxino-Caucasian species. However, some species are shared (e.g.
Acer cappadocicum, Quercus petraea subsp. iberica, Sanicula europaea).
4.3 Syntaxonomic outline
Based on the presented numerical comparison of Euxinian C. orientalis forests, we
describe the Georgian community as a new association and classify it within the alliance
Castaneo sativae-Carpinion orientalis.
Campanulo alliariifoliae-Carpinetum orientalis ass. nova hoc loco
Holotypus (hoc loco) of the association: Georgia, Motsameta (Imereti Region): a forest on
a limestone slope ca 0.1 km N of the Motsameta Monastery, 42.28257° N, 42.75938° E,
10 × 10 m2, 24 July 2019, elevation: 210 m, aspect: 40°, inclination: 30°, soil pH (H2O):
7.33, cover of rocks: 10%, author: P. Novák. Relevé 19 in Supplements E1 and E2.
E3 (85%, mean height = 9 m): Carpinus orientalis 5, Fraxinus excelsior 2b; E2 (1%, mean
height = 0.8 m): Hedera helix +, Smilax excelsa +, Staphylea colchica +; E1 (30%, mean
height = 0.2 m): Hedera helix 2a, Brachypodium sylvaticum 1, Primula acaulis 1, Ruscus
aculeatus 1, Sedum stoloniferum 1, Vinca major subsp. hirsuta 1, Viola alba 1, Acer cam-
pestre +, Asplenium adiantum-nigrum +, A. trichomanes +, Campanula alliariifolia +, Carex
muricata aggr. +, C. sylvatica +, Clinopodium umbrosum +, Klasea quinquefolia +, Lamium
galeobdolon +, Lapsana communis +, Lathyrus laxiflorus +, Orobanche laxissima +, Poly-
podium cambricum +, Quercus petraea subsp. iberica +, Sanicula europaea +, Silene bal-
ansae +, Smilax excelsa +, Teucrium chamaedrys +, Veronica peduncularis +, Gleditsia
triacanthos r, Prunus avium r, Schedonorus giganteus r, Trachycarpus fortunei r; E0 (15%):
indet.
Diagnostic species of the new association: Asplenium adiantum-nigrum, Buxus sempervi-
rens, Campanula alliariifolia, Carpinus orientalis, Hedera helix, Klasea quinquefolia,
Lathyrus laxiflorus, Potentilla micrantha, Ruscus aculeatus, Smilax excelsa, Teucrium
chamaedrys, Vinca major subsp. hirsuta.
46
Table 1. Shortened synoptic table summarizing the classification results, species percentage frequen-
cies are provided. Highly diagnostic species (Φ ≥ 0.55; grey shaded) for each cluster are shown. Highly
diagnostic species for three main clusters are provided separately or marked by an asterisk if concur-
rently highly diagnostic for some of the clusters. Full version of the table is stored in Supplement E3.
Tabelle 1. Gekürzte Übersichtstabelle, die die Klassifikationsergebnisse zusammenfasst, prozentuale
Stetigkeiten sind dargestellt. Hochdiagnostische Arten für die drei Hauptcluster werden separat darge-
stellt oder mit einem Sternchen gekennzeichnet, wenn sie gleichzeitig für einige der Cluster hochdiag-
nostisch sind. Die vollständige Version der Tabelle befindet sich in Anhang E3.
Cluster
1
2
3
4
5
6
7
8
9
Number of relevés
10
4
20
12
3
7
22
15
12
Main cluster 1
Campanula alliariifolia
60
50
80
.
.
.
.
.
.
Oplismenus hirtellus subsp. undulatifolius
.
50
60
.
.
.
.
.
.
Cluster 1
Arbutus andrachne
70
.
.
.
.
.
.
.
.
Cistus salviifolius
60
.
.
.
.
.
.
.
.
Laurus nobilis
60
.
5
.
.
14
.
.
.
Rhododendron ponticum
60
25
.
.
.
.
5
.
.
Cluster 2
Buxus sempervirens
.
75
25
.
.
.
.
.
.
Vinca minor
.
50
.
.
.
.
.
.
.
Vincetoxicum nigrum
.
50
.
.
.
.
.
.
.
Datisca cannabina
10
50
.
.
.
.
.
.
.
Rhamnus imeretina
.
50
.
.
.
14
.
.
.
Hypericum xylosteifolium
10
50
.
.
.
.
5
.
.
Cluster 3
Vinca major subsp. hirsuta*
.
.
90
.
.
.
.
.
.
Viola alba*
.
.
85
.
.
.
.
.
.
Klasea quinquefolia*
.
.
75
.
.
.
.
.
.
Carex digitata
.
.
60
.
.
.
.
.
.
Clinopodium umbrosum
.
.
55
.
.
.
.
.
.
Veronica peduncularis
.
.
50
.
.
.
.
.
.
Diospyros lotus
.
.
40
.
.
.
.
.
.
Asplenium trichomanes
.
.
50
.
.
.
14
.
.
Viola reichenbachiana
.
.
35
.
.
.
.
.
.
Main cluster 2
Epimedium pubigerum
20
25
.
83
67
57
.
13
8
Salvia forsskaolei
20
50
.
83
33
71
.
13
8
Asperula cimulosa
30
25
.
83
33
57
.
.
33
Carpinus betulus
.
.
35
58
100
86
9
.
8
Cirsium hypoleucum
30
25
.
75
33
57
5
.
17
Fagus orientalis
.
.
15
67
100
14
.
7
.
Cluster 4
Euphorbia oblongifolia*
.
.
.
75
.
14
.
.
.
Cluster 5
Sesleria phleoides
.
.
.
.
100
.
.
.
.
Dictamnus albus
.
.
.
8
100
.
.
.
.
Cotinus coggygria
.
.
5
.
100
.
5
7
.
Tilia platyphyllos
.
.
.
8
100
14
.
.
.
Quercus pubescens
.
.
.
.
100
.
.
.
25
Frangula alnus
.
.
.
.
67
.
.
.
.
Pimpinella tripartita
.
.
5
.
67
.
.
.
.
Corylus avellana
20
.
20
17
100
14
27
.
.
Colutea cilicica
.
.
.
.
67
.
5
.
25
Campanula glomerata
.
.
.
17
67
.
.
.
17
Tanacetum poteriifolium
.
.
.
8
67
.
9
.
25
47
Cluster
1
2
3
4
5
6
7
8
9
Number of relevés
10
4
20
12
3
7
22
15
12
Fraxinus ornus
.
25
.
8
67
14
.
.
.
Cota tinctoria
.
.
.
.
33
.
.
.
.
Cluster 6
Hedera colchica
.
.
20
8
.
57
.
.
.
Main cluster 3
Crataegus monogyna
.
.
.
.
.
.
59
93
.
Quercus cerris
.
.
.
58
33
57
82
100
100
Cluster 7
Potentilla reptans
.
.
.
.
.
.
36
.
.
Cluster 8
Rubus ulmifolius
.
.
.
.
.
.
.
40
.
Cluster 9
Vicia cracca
.
.
.
17
.
.
5
.
83
Crataegus rhipidophylla
.
.
.
.
.
.
.
.
42
Lathyrus roseus
.
.
.
.
.
.
.
.
42
Nepeta nuda subsp. albiflora
.
.
.
.
.
.
.
.
42
Sorbus umbellata
.
.
.
.
.
.
.
.
42
Cephalanthera rubra
.
.
.
17
.
.
9
.
58
Aristolochia pallida
.
.
.
.
.
.
.
.
33
Silene latifolia
.
.
.
.
.
.
.
.
33
Erweiterte deutsche Zusammenfassung
Einleitung – Die Wälder der Kolchis (Ökoregion Kaukasus, euxinische Provinz) stellen ein einzig-
artiges Refugium tertiärer Reliktbiota dar (NAKHUTSRISHVILI et al. 2015). Sie wurden jedoch haupt-
sächlich in Bulgarien und der Türkei pflanzensoziologisch untersucht (z. B. QUÉZEL et al. 1980), wäh-
rend in Georgien nur eine begrenzte Anzahl von Studien existiert (z. B. NOVÁK et al. 2019). Eine pflan-
zensoziologische Untersuchung mit Schwerpunkt auf thermophilen Laubwäldern der georgischen Kol-
chis fehlte bisher. Daher waren die Ziele dieser Studie (1) die Ökologie und Artenzusammensetzung der
Carpinus orientalis-Wälder der georgischen Kolchis als ein Beispiel für kolchische thermophile Wälder
zu beschreiben; (2) die ökologischen und floristischen Beziehungen zwischen der hier untersuchten
Gesellschaft und analogen, bisher in der euxinischen Provinz erfassten Gesellschaften zu ermitteln.
Untersuchungsgebiet – Die Stichprobenerhebung konzentrierte sich auf mesozoische Kalksteinge-
biete in den Ausläufern des Großkaukasus im westlichen Georgien (BONDYREV et al. 2015). Das Un-
tersuchungsgebiet hat ein feuchtes warm-gemäßigtes Klima, die Probenahmestellen hatten eine mittlere
Jahrestemperatur von 11,8–14,8 °C und einen jährlichen Niederschlag von 999–2074 mm. Innerhalb
Georgiens gilt es als das Gebiet, das außerordentlich reich an endemischen Gefäßpflanzenarten ist
(SLODOWICZ et al. 2018).
Methoden – Wir erhoben 20 Vegetationsaufnahmen der Waldvegetation, die von Carpinus orienta-
lis dominiert oder mitdominiert wurde. Anschließend haben wir einen Datensatz sowohl dieser Vegeta-
tionsaufnahmen, als auch der Aufnahmen analoger Assoziationen und Gesellschaften aus dem türki-
schen Teil der euxinischen Provinz (n = 105 Aufnahmen insgesamt) zusammengestellt. Wir haben
flexibles Beta-Clustering (β = -0,2, Bray-Curtis-Distanz) und eine NMDS-Ordinationsanalyse ange-
wendet, um Hauptmuster in ihrer Artenzusammensetzung zu erkennen.
Ergebnisse und Diskussion – Die in Georgien erfassten Carpinus orientalis-Wälder besetzten
normalerweise steile und oft felsige Hänge mit steinigem Oberboden mit neutraler bis schwach alkali-
scher Reaktion (pH 6,6–7,8). In der Baumschicht wurde C. orientalis häufig von anderen Laubbaumar-
ten begleitet. Das Unterholz enthielt zahlreiche immergrüne Arten (z. B. Hedera helix, Ruscus aculea-
tus, Smilax excelsa, Vinca major subsp. hirsuta). Waldmesophyten (z. B. Carex digitata, Veronica
48
peduncularis) waren in der Krautschicht häufig, begleitet von Xerophyten (z. B. Klasea quinquefolia,
Teucrium chamaedrys). Chasmophyten waren ebenfalls vorhanden (z. B. Asplenium scolopendrium,
A. trichomanes).
Die Clusteranalyse des Datensatzes ergab drei Hauptcluster, die eng mit der Biogeographie ver-
knüpft sind. Bei dem ersten, dem östlichsten, handelte es sich um Typen mit signifikantem kaukasi-
schem und ostauxinischem floristischem Einfluss. Er enthielt die georgische Gesellschaft und die Asso-
ziation Erico-Carpinetum, Typusassoziation des Verbandes Castaneo sativae-Carpinion orientalis. Der
zweite Hauptcluster war auf den zentralen Teil der euxinischen Provinz beschränkt. Aufgrund seines
eher mesophilen Charakters, der oft von Carpinus betulus oder Fagus orientalis mitdominiert wird,
scheint er einen Übergang zwischen thermophilen Wäldern und Eichen-Hainbuchen-Wäldern des
Verbandes Trachystemono orientalis-Carpinion betuli darzustellen. Die dritte Hauptgruppe umfasste
insbesondere Mischwälder aus C. orientalis und Quercus cerris. Sie wurden fast ausschließlich im
westlichen Teil der Provinz nachgewiesen und enthielten mehr Balkan- oder Mittelmeerarten, die auf
den westlichen Teil der euxinischen Provinz beschränkt sind. Diese Einheit wurde nach der Klassifizie-
rung von ÇOBAN & WILLNER (2019) dem Verband Quercion confertae zugeordnet. In der feineren
Unterteilung des Datensatzes waren die meisten beteiligten Assoziationen und Gesellschaften gut
erkannt, was auf ihre charakteristische floristische Zusammensetzung hinweist. Diese zeigte auch viele
einzigartige floristische Merkmale der georgischen Gesellschaft, die daher als neue Assoziation Cam-
panulo alliariifolae-Carpinetum orientalis ass. nova hoc loco beschrieben wurde (Verband Castaneo-
Carpinion). Die Ordinationsanalyse betonte auch die Bedeutung der Biogeographie für die Diversität
der euxinischen Carpinus orientalis-Wälder, die vermutlich durch das Makroklima und die Vegetati-
onsgeschichte bedingt wird.
Acknowledgements
We thank Goffredo Filibeck and one anonymous reviewer for their helpful comments on a previous
version of this study, Kryštof Chytrý, Anna Hlaváčková, Jakub Salaš and Dominik Zukal for help with
the field research, Ali Kavgacı for providing the phytosociological literature from Turkey and Renata
Piwowarczyk for determination of herbarium specimens of the genus Orobanche. This study was con-
ducted in the Centre for European Vegetation Syntheses (CEVS) funded by the Czech Science Founda-
tion (project no. 19-28491X).
Author contributions
P.N. led the writing and performed analyses. P.N. and V.S. conceived the idea of the research. P.N
and V.K. participated in the field sampling. All the authors critically revised the manuscript.
ORCID iDs
Veronika Kalníková https://orcid.org/0000-0003-2361-0816
Pavel Novák https://orcid.org/0000-0002-3758-5757
Vladimir Stupar https://orcid.org/0000-0003-0835-2249
Supplements
Additional supporting information may be found in the online version of this article.
Zusätzliche unterstützende Information ist in der Online-Version dieses Artikels zu finden.
Supplement E1. Relevé table of the association Campanulo alliariifoliae-Carpinetum orientalis.
Anhang E1. Tabelle der Vegetationsaufnahmen des Campanulo alliariifoliae-Carpinetum orientalis.
49
Supplement E2. Header data for the original relevés of Supplement E1.
Anhang E2. Kopfdaten der Vegetationsaufnahmen von Anhang E1.
Supplement E3. Full synoptic table and correspondence between associations and clusters.
Anhang E3. Vollständige Übersichtstabelle und Übereinstimmung zwischen den Assoziationen und
Clustern.
Supplement E4. Additional photos of the vegetation of the association Campanulo alliariifoliae-
Carpinetum orientalis.
Anhang E4. Zusätzliche Fotos der Vegetation des Campanulo alliariifoliae-Carpinetum orientalis.
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Novák et al.: Carpinus orientalis forests in Georgian Colchis: First insights
Relevé number 1 2 3 4 5678910 11 12 13 14 15 16 17 18 19 20
Tree layer cover (%) 95 90 90 90 85 85 70 70 85 70 80 80 90 70 85 80 90 80 85 85
Shrub layer cover (%) 8720 320 12 15 20 35 25 10 30 8 3 1 30 515 1 2
Herb layer cover (%) 30 35 60 15 30 35 55 65 40 25 30 25 30 60 45 35 30 40 30 30
Moss layer cover (%) 4 2 5 6 520 20 10 10 40 30 0512 6230 40 15 50
Tree layer
Carpinus orientalis 3354 3 4 3 5 3 4 4 4 5 4 5 . 5 5 5 5
Carpinus betulus 2b 3.42b 3 . . . . . . 2a .2a .....
Acer campestre 12a . . ..1...2a .2a ..1 . . . 2a
Quercus petraea subsp. iberi ca 2b .........2a . . 2b ......
Hedera helix 1.....+....1........
Fagus orientalis . . . . 2b .2a .2a ...........
Fraxinus excelsior ......1...1.......2b .
Acer cappadocicum . . . . 2a ...4...........
Robinia pseudoacacia .... . 2m .........2a . . . .
Ulmus glabra .... . . . 1 . . . 2a ........
Zelkova carpinifolia ............. . 2b 5 . . . .
Shrub layer
Smilax excelsa .+2a + 1 1 + 2a 1 1 . . 2a 1 . . 1 + + .
Carpinus orientalis . 1 2a .1++.2a 2a 1.+..2b .1..
Crataegus sp. . + . . 11+1..1.........
Buxus sempervirens 1 . .. 1 . . . . . . 2b . . + . . . . +
Cornus sanguinea .+.. . . . . . + . . . + . . + . . .
Quercus petraea subsp. iberi ca . . . . 1...++...1......
Hedera helix ......+... . . . + + . . . + .
Corylus avellana .... . . . 2a 2a 1..+.......
Ilex colchi ca 2m +...1..............
Crataegus pentagyna . . . 1 ............11..
Ligustrum vulgare ......+.. . + . . . . . . + . .
Castanea sativa +... . . . . 2b ...........
Rosa sp. . + . . ..r.............
Diospyros lotus . . + . ...1......... . . .
Staphylea colchica .....1. . . . . . . . . . . . + .
Ficus carica .....+..+...........
Pyrus communis .... . . 1 . . . . . . . . . . 1 . .
Philadelphus coronarius .........1+.........
Cornus mas ...... . . . . 1 . . . . . . 1 . .
Herb layer
Vinca major subsp. hirsuta 1 + 1 + +1+2b 2a 2a 2m +1+.++.1+
Brachypodium sylvaticum + + 1 + ++++++++.. 1 1 1 + 1 1
Hedera helix 1 1 2b 1 + . 2b 1 + 1 . 1 r . 1 2b + + 2a 1
Primula acaulis 112m 1 . 1 1 r . 1 1 . 1 + 1 1 1 1 1 +
Viola alba 1+++ + 1 . . . + + + + + 1 1 + 1 1 1
Smilax excelsa + + 2a +1111..+1 1 1 1 1 + + + .
Campanula alliariifoli a + + . + . + + 2a . 1 2m . + + + + 1 1 + +
Klasea quinquefolia +.++ + + + + . . + + + . . + + + + +
Ruscus aculeatus . . 2a +12a +r.+++..2b 12b 311
Lathyrus laxiflorus + + r . .+.+.+2a . . + + + + 1 + +
Sanicula europaea 2a 1++ + . 1 + . + + + + . . . . . + .
Dioscorea communis +.rr + + + + + + + + + . . . . . . .
Potentilla micrantha +..+ + + + + r 1 + . + r . . + . . .
Oplismenus hirtellus subsp. undulatifolius . + + 1 .2a +2a 1+ . + . . 2a + + . . .
Campanula rapunculoides .+.. + + . + 1 r . + + . + + + . . 1
Carex digitata ..++ + . 1 1 1 1 + . + . + . 1 1 . .
Clinopodium umbrosum + + + + .+....++..++.+ + .
Asplenium adiantum-nigrum . + + + +++. . . 1 . . . + . + . + 1
Veronica peduncularis .+++ . + . . . . + . . r . . + + + +
Drymochloa drymeja . . + + 2a .+2b 2b .++12a ......
Asplenium trichomanes . . + + ..r+.++...+. + + + .
Carex sylvatica 1 . . + .11 . . + . + . . + . . . + +
Carex muricata aggr. .++. . . . . . . . + . . 1 + + + + 1
Viola reichenbachiana +... . + + + r + . + . . . . . . . .
Leontodon hispidus . . 1 + ...r......+ 1 1 1 . .
Symphytum grandiflorum 2m 1+. + . . + . . . 2m ........
Helleborus orientalis + + . . ... . . . . + . . . 1 1 + . .
Lathyrus vernus + . . + ....r.. + + . . . + . . .
Lapsana communis . + ++ + . . . . . . . . . . . . . + +
Asplenium scolopendrium . + . r .++.r+..... . . . . .
Teucrium chamaedrys .r.. . . + . . . . . . . r . + + + .
Trachystemon orientalis ..+. + . + 1 1 + . . . . . . . . . .
Prunella vulgaris ..+. . + + . . . + r . . . . + . . .
Hieracium sabaudum . . . . +.+r..+..r . . . r . .
Rubus subgen. Rubus . . + . . + . . . 1 . . + . . . + . . .
Carpesium cernuum . . . + .+.. . . . . . . + + + . . .
Pteridium aquilinum .... + . 1 . + + . . . . . . + . . .
Digitalis schischkinii . . . . +..r1++.........
Carex michelii +.+. . . . . . . . . + . . + . . . .
Luzula forsteri + . . . ++. . . r . . . . . . . . . .
Ajuga reptans +... . + . . . . . + . . . + . . . .
Euphorbia macroceras . 1 . . +...... . + + . . . . . .
Calystegia silvatica . + +............++...
Plantago lanceolata . . + . .r . . . . . . . . . . r + . .
Carex flacca subsp. serrulata .... + . + . . . . . . + . . . + . .
Physospermum cornubiense . . . . +..... + . + + . . . . . .
Schedonorus giganteus .....++....... . + . . r .
Sedum stoloniferum . . . . . + . . . . + . . . . . . . 1 1
Silene balansae .... . . . . . . + . . . . + . . + +
Salvia glutinosa . + . . ...+ . + . . . . . . . . . .
Hedera colchica . + . . ......1. . 2a ......
Trifolium repens . . + . . + . . .... . . . . + . . .
Epimedium pinnatum subsp. colchicum . . . . 1 . . . . . . . 2b 2a .. . ...
Lonicera caprifolium . . . . + . . . . . + . . + . . . . . .
Peucedanum adae . . . . + . . . . . . . 1 1 . . . . . .
Ruscus colchicus .... . ++ . . . .. . + . . . . . .
Medicago lupulina . . . . . + . . . . . . . . .. + + . .
Poa angustifolia . . . . . . + . . . + . . . . + . . . .
Securigera varia ......r..+.... . . . r . .
Pimpinella saxifraga .......r . .... . . + . + . .
Origanum vulgare .........+...... + + . .
Peucedanum caucasicum ............+ . . + . + . .
Paeonia caucasica + . . . . . . . . . . + . . . . . . . .
Carex sp. . + .... . .. + . . ........
Potentilla indica . . + + . . . . . . . . . . ... . . .
Periploca graeca ..+..... . .. . . . . r . . . .
Fragaria vesca . . . . + . . . . . . + . . . . . . . .
Hypericum androsaemum ......+.+...... . . . ..
Solidago virgaurea .......++..... . . . . . .
Arabis nordmanniana ... . . . . + . r... . . . . . . .
Galium valantioides .......+.... . + . . . . . .
Euphorbia squamosa .......+.......+ . ...
Lamium galeobdolon . . . . . + . . + . . . . . . . . . + .
Clematis vitalba . . . . . . . . . r . . + . . .. . . .
Polypodium vulgare ..........+ . . . . . . r . .
Sesleria alba ............+2a .... . .
Brunnera macrophylla . . . . . . . . . . . . + + . . . . . .
Cephalanthera longifolia ............rr . .. . . .
Trifolium pratense . . . . . . . . .... . + . . . + . .
Prunella ×intermedia ..............+. + . . .
Cruciata glabra ............ . . . . + + . .
Polypodium cambricum ............... . . . + 1
Orobanche laxissima ....... . .. . . . . . . . . + r
Juveniles
Carpinus orientalis . + + + . . + . 1 + . + . + . . + 1 . .
Quercus petraea subsp. iberi ca . . . r . r + + + + + r + . . . . . + .
Crataegus sp. r . +.++ + + . r+ . . + . . . . . .
Diospyros lotus r . + + r + . + . . . . . . . . r . . .
Ficus carica . . . r . + . r r + . . . . . + . + . .
Acer campestre + . . . . . . . . + . + + ... + . + .
Cornus sanguinea . + + . + . . . . r .r . . . . + . . .
Carpinus betulus + + . . + + . . . . . .. . + . . . . .
Acer cappadocicum + + . . + . . . 1 . . + . . . . . . . .
Ligustrum vulgare + + + . . . + . . . . . . . . . . . . .
Tilia begoniifolia . + . . . . . . + . . . + . .... . .
Crataegus pentagyna ...+....... . . . . . + + . .
Fraxinus excelsior . . . r . . + . . . . . + . . . . . . .
Taxus baccata . . . . + . . . . r . . . r . . . . . .
Zelkova carpinifolia ..............11 + . . .
Castanea sativa . + . . . . + . . .... . . . . . . .
Crataegus germanica . + . . . . . . . . . . r . . . . . . .
Ulmus glabra . r . . . . + . . . . . . . . . . . . .
Robinia pseudoacacia .. + r . . . . . . . ....... . .
Prunus avium ... r . ........... . . r .
Sorbus torminalis . . . . + . . . . . r . . . . . . . . .
Staphylea colchica . . . . . + . . . . . . . + . . . . . .
Anhang E1.Tabelle der Vegetationsaufnahmen des Verbandes Campanulo alliariifolae-Carpinetum orientalis ass. nova hoc loco. Die Arten sind nach
Vegetationsschichten und innerhalb der Schichten nach abnehmender Häufigkeit sortiert. Die in einer Vegetationsaufnahme erfassten Arten sind unter der
Tabelle aufgeführt. Kopfdaten werden im Anhang E2 aufgeführt.
Supplement E1. Relevé table of the association Campanulo alliariifoliae-Carpinetum orientalis ass. nova hoc loco. Species are sorted according to
vegetation layer and within layers according to decreasing frequency. Species recorded in one relevé are provided below the table. Header data are stored
in Supplement E2.
Species recorded in one relevé: Tree layer:Pyrus communis 5: 1; Hedera colchica 6: 2a; Corylus avellana 8: 2a; Tilia begoniifolia 9: 3; Pinus
sylvestri s 12: 2m; Prunus avium 13: 2a; Smilax excelsa 13: 1; Quercus hartwissiana 20: 2a; Diospyros lotus 20: 1. Shrub layer:Carpinus betulus
2:
+; Prunus avium 2: +; Punica granatum 3: +; Rhododendron luteum 5: 2a; Sorbus torminalis 5: 1; Robinia pseudoacacia 6: +; Sambucus nigra 8: 1;
Tilia begoniifolia 9: 1; Acer cappadocicum 9: +; Euonymus latifolius 10: +; Fraxinus excelsior 13: +; Acer campestre 13: +; Hedera colchica 14: +;
Zelkova carpinifoli a 16: 2a; Laurus nobilis 16: 1; Rosa canina 17: +. Herb layer:Pulmonaria mollis 1: r; Arum sp. 2: +; Geranium robertianum
2:
+; Poa nemoralis 2: +; Sison amomum 2: +; Torilis japonica 2: +; Medicago sp. 2: r; Trifolium diffusum 2: r; Cirsium vulgare 3: +; Dactylis
glomerata 5: +; Euphorbia amygdaloides 5: 1; Viola sp. 5: r; Aristolochia pontica 6: +; Euphorbia stricta 6: +; Plantago major 6: +; Lotus
corniculatus 7: +; Ranunculus polyanthemos 7: +; Mercurialis perennis 8: +; Vicia sepium 8: +; Hypericum perforatum 9: +; Pachyphragma
macrophyllum 10: 1; Taraxacum sect. Taraxacum 10: +; Silene sp. 10: r; Campanula raddeana 11: 1; Cardamine parviflora 11: +; Galium album
11: +; Hieracium murorum 11: +; Laser trilobum 11: +; Polygonatum glaberrimum 11: +; Ranunculus bulbosus 11: +; Cyclamen coum 11: r;
Pimpinella tripartita 11: r; Primula veri s subsp. macrocalyx 12: 1; Epipactis sp. 12: r; Brachypodium pinnatum 14: 2a; Carex humi lis 14: 2m;
Melampyrum elatius 14: +; Asplenium ruta-muraria 14: r; Jacobaea vulgaris 15: +; Erigeron annuus 16: +; Adiantum capillus-veneris 17: +;
Lactuca muralis 17: +; Petrorhagia saxifraga 17: r; Scabiosa sosnowskyi 18: +; Leptopus chinensis 20: +; Tanacetum partheniifolium 20: r.
Juveniles:Clematis vitalba 2: +; Cotinus coggygria 2: +; Malus sylvestris 2: r; Morus alba 3: +; Fagus orientalis 7: r; Lonicera caucasica 9: +;
Euonymus latifolius 10: +; Corylus avellana 10: +; Rosa sp. 11: +; Euonymus latifolius 14: r; Pyru s sp. 14: r; Laurus nobilis 16: r; Cornus mas 18: r;
Ulmus minor 18: r; Gleditsia triacanthos 19: r; Trachycarpus fortunei 19: r.
Novák et al.: Carpinus orientalis forests in Georgian Colchis: First insights
Supplement E2. Header data (location, area, site conditions, vegetation height and date) for the original relevés of Supplement E1.
Anhang E2. Kopfdaten (Ort, Region, Standortsbedingungen, Vegetationshöhe und Datum) der Vegetationsaufnahmen von Anhang E1.
1Banoja Imereti 42.30944 42.67333 100 7.02 450 170 10 012 20.3 2016-07-22
2Chiatura Imereti 42.28556 43.30944 100 7.14 490 240 15 115 20.9 2017-07-03
3Khidi Imereti 42.42111 42.49528 100 7.57 250 140 35 311 30.75 2017-07-09
4Martvili Imereti 42.45361 42.37472 100 7.61 220 35 10 315 10.5 2017-07-09
5Nokalakevi Samegrelo 42.36528 42.19417 100 6.53 110 180 35 1 8 3 0.7 2017-07-10
6Senaki Samegrelo 42.29222 42.04139 100 6.83 90 250 25 713 30.5 2017-07-10
7Tsutskhvati Imereti 42.27333 42.85417 100 7.5 410 315 30 30 19 60.35 2017-07-04
8Khidi Imereti 42.42222 42.495 100 7.76 250 270 30 20 10 40.35 2017-07-09
9Nokalakevi Samegrelo 42.49 42.41806 100 7.67 130 10 30 5 8 2 0.4 2017-07-10
10 Nokalakevi Samegrelo 42.37194 42.18778 100 7.71 145 360 35 30 10 40.6 2017-07-10
11 Katskhi Imereti 42.28444 43.21556 100 7.2 570 280 35 25 8 2 0.5 2018-05-02
12 Banoja Imereti 42.30972 42.67417 100 6.59 461 180 5 0 10 20.2 2016-07-22
13 Jvari Samegrelo 42.75639 42.04361 100 6.75 510 135 30 3 8 1.5 0.25 2019-07-20
14 Jvari Samegrelo 42.75417 42.04333 100 6.87 490 200 30 381.2 0.35 2019-07-20
15 Martvili Imereti 42.45306 42.37639 100 7.09 220 225 35 112 1.5 0.35 2019-07-22
16 Matkhoji Imereti 42.38944 42.44722 100 7.71 220 180 40 022 2.5 0.2 2019-07-23
17 Matkhoji Imereti 42.39111 42.45083 100 7.16 270 180 35 8 7 1.2 0.2 2019-07-23
18 Matkhoji Imereti 42.39167 42.44889 100 7.33 270 180 40 271.5 0.3 2019-07-23
19 Motsameta Imereti 42.28257 42.75938 100 7.33 210 40 30 10 90.8 0.2 2019-07-24
20 Motsameta Imereti 42.28194 42.76 100 7.41 240 80 55 510 1.5 0.25 2019-07-24
Soil pH
#
Municipality
Region
N (°)
E (°)
area
(m2)
Date
Slope
(°)
Elevation
(m)
Aspect
(°)
Cover of
rocks
(%)
Mean
height E3
(m)
Mean
height E2
(m)
Mean
height E1
(m)
Novák et al.: Carpinus orientalis forests in Geo rgian Colc his: First insights
Cluster 1 2 3 4 5 6 7 8 9
Number of relevés 10 4 20 12 3 7 22 15 12
Cluster 1
Arbutus andrachne 70 . . . . . . . .
Cistus salviifolius 60 . . . . . . . .
Laurus nobilis 60 . 5 . . 14 . . .
Rhododendron ponticum 60 25 . . . . 5 . .
Bituminaria bituminosa 40 . . . . . 9 . .
Trifolium campestr e 30 . . . . . . . .
Castanea sativa 70 25 20 17 . 14 . . .
Paliurus spina-christi 20 ........
Ligustrum vulgare 50 . 30 17 . 14 . . .
Cornus sanguinea 50 50 35 . . 14 . 7 .
Rubia tinctorium 20 . . . . 14 . . .
Hyper icum calycinum 30 25 . . . 14 . . .
Vaccinium arctostaphylos 50 50 . 33 . 29 . 7 .
Tilia begoniifolia 30 25 15 . . 14 . . .
Dorycnium pentaphyllum 30 25 . 25 . . 5 .8
Brachypodium pinnatum 60 50 5 42 33 43 18 . 42
Cluster 2
Buxus sempervirens .75 25 . . . . . .
Vinca minor .50 . . . . . . .
Vincetoxicum nigrum .50 . . . . . . .
Datisca cannabina 10 50 . . . . . . .
Rhamnus imeretina .50 . . . 14 . . .
Hyper icum xylosteifolium 10 50 . . . . 5 . .
Briza media 10 50 . . . 14 . . .
Pyrus communis .50 15 .....17
Arbutus unedo . 25 . . . . . . .
Calluna vulgaris . 25 . . . . . . .
Drymochloa drymeja 50 100 50 50 33 43 . . .
Cluster 3
Vinca major subsp. hirs uta . . 90 . . . . . .
Viola alba . . 85 . . . . . .
Klasea quinquefolia . . 75 ......
Carex digitata . . 60 . . . . . .
Clinopodium umbrosum ..55 ......
Veronica peduncularis ..50 ......
Diospyros lotus . . 40 . . . . . .
Asplenium trichomanes . . 50 ...14 ..
Viola reichenbachiana ..35 ......
Carex muricata aggr. . . 45 . . . . . 17
Asplenium scolopendrium . . 30 . . . . . .
Hieracium sabaudum . . 30 . . . . . .
Lathyrus vernus ..30 ......
Symphytum grandiflorum . . 30 . . . . . .
Oplis menus hirtellus subsp. undulatifolius . 50 60 . . . . . .
Carpesium cernuum ..25 ......
Digitalis schischkinii . . 25 . . . . . .
Campanula rapunculoides ..60 ..14 .20 42
Brachypodium sylvaticum 30 25 90 42 . 43 27 20 17
Carex sylvatica . 25 45 . . 14 ...
Prunella vulgaris . . 30 . . . 5 7 .
Calystegia silv atica . . 20 . . . . . .
Carex flacca subsp. serrulata ..20 ......
Carex michelii . . 20 . . . . . .
Euphorbia macroceras ..20 ......
Plantago lanceolata ..20 ......
Robinia pseudoacacia . . 20 . . . . . .
Sedum stoloniferum . . 20 . . . . . .
Schedonorus giganteus . . 20 . . . . . .
Silene balansae ..20 ......
Ulmus glabra . . 20 . . . . . .
Primula acaulis 10 25 85 42 . 57 9 27 17
Leontodon hispidus ..35 ...5.17
Fraxinus excelsior 10 . 25 . . . . . .
Epimedium pinnatum subsp. colchicum ..15 ......
Lamium galeobdolon ..15 ......
Lonicera caprifolium . . 15 . .....
Medicago lupulina . . 15 . . . . . .
Peucedanum adae . . 15 . . . . . .
Peucedanum caucasicum ..15 ......
Pimpinella saxifraga . . 15 . . . . . .
Poa angustifolia ..15 ......
Ruscus colchicus ..15 ......
Staphylea colchica . . 15 . . . . . .
Taxus baccata . . 15 . . . . . .
Trifolium r epens . . 15 . . . . . .
Zelkova carpinifolia ..15 ......
Ajuga reptans . . 20 . . . . 7 .
Origanum vulgare . . 15 . . . 5..
Arabis nordmanniana ..10 ......
Brunnera macrophylla . . 10 . . . . . .
Cephalanthera longifolia . . 10 . . . . . .
Cruciata glabra . . 10 . . . . . .
Euonymus latifolius . . 10 ......
Euphorbia squamosa . . 10 . . . . . .
Galium valantioides . . 10 . . . . . .
Hypericum androsaemum ..10 ......
Orobanche laxissima . . 10 . . . . . .
Paeonia caucasica ..10 ......
Periploca graeca ..10 ......
Philadelphus coronarius . . 10 . . . . . .
Polypodium cambricum . . 10 . . . . . .
Potentilla indica . . 10 . . . . . .
Prunella ×intermedia ..10 ......
Sesleria alba . . 10 . . . . . .
Solidago virgaurea . . 10 . . ....
Prunus avium ..20 17 .....
Ficus carica 30 25 35 . . . 9 . .
Cluster 4
Euphorbia oblongifolia ...75 . 14 . . .
Crataegus pentagyna 20 25 15 92 .43 ..33
Rhododendron luteum 10 50 5 67 . 29 . . .
Aegonychon purpurocaeruleum . . . 58 33 . . 13 25
Acer cappadocicum 10 25 25 58 .14 ...
Lysimachia punctata . . . 25 . . . . 8
Galium rotundifolium ...17 .....
Phlomis samia ...17 .....
Viola sieheana . 50 . 92 33 71 41 33 33
Bromopsis benekenii . . . 17 . . . . 8
Luz ula for steri . . 20 33 . 14 . . 25
Silene compacta ...17 .14 ...
Asyneuma rigidum 40 . . 58 67 29 9 . 42
Daphne pontica 40 25 . 58 33 43 .53 .
Cluster 5
Sesleria phleoides . . . . 100 . . . .
Dictamnus albus . . . 8 100 . . . .
Cotinus coggygria . . 5 . 100 . 5 7 .
Tilia platyphyllos ...8100 14 ...
Quercus pubescens . . . . 100 . . . 25
Frangula alnus . . . . 67 . . . .
Pimpinella tripartita ..5.67 ....
Corylus avellana 20 . 20 17 100 14 27 . .
Colutea cilicica ....67 .5.25
Campanula glomerata ...17 67 ...17
Tanacetum poteriifolium . . . 8 67 . 9 .25
Fraxinus ornus . 25 . 8 67 14 . . .
Cota tinctoria . . . . 33 . . . .
Milium vernale ...33 67 29 ..8
Cluster 6
Heder a colchica . . 20 8.57 ...
Oenanthe pimpinelloides 30 25 .33 .86 .40 8
Trachystemon orientalis 20 50 30 8 . 86 5 47 .
Asplenium adiantum-nigrum 50 75 55 . . 86 5 27 .
Staphylea pinnata . 25 . . . 29 . . .
Galium paschale .25 .17 33 43 ...
Ruscus hypoglossum . 25 . . . 29 . 7 .
Cluster 7
Potentilla r eptans ......36 ..
Agrimonia eupatoria . . . . . . 27 . .
Festuca jeanpertii ......27 ..
Hypericum perforatum ..5...32 ..
Geranium purpureum . . . . . . 23 ..
Lonicera etrusca . . . . . . 23 . .
Styrax officinalis . . . . . . 23 . .
Arum maculatum ......18 ..
Chaerophyllum nodosum . . . . . . 18 . .
Rostr aria cristata ......18 ..
Teucrium polium ......18 ..
Doronicum orientale ......23 .8
Epipactis helleborine . . . . . . 14 . .
Alliaria petiolata ......23 .17
Scutellaria albida . . . . . . 23 . 17
Viola odorata . . . . . . 23 . 17
Phillyrea latifolia . . . . . . 23 20 .
Rosa canina . . 5 . . .23 .17
Carlina corymbosa ......9..
Cynosurus echinatus . . . . . . 9 . .
Daucus carota . . . . . . 9 . .
Erysimum cuspidatum . . . . . . 9 . .
Geranium dissectum . . . . . . 9 . .
Neottia nidus-avis ......9..
Piptather um coer ulescens . . . . . . 9 . .
Rubus sanctus ......9..
Tilia tomentos a ......9..
Torilis arvensis ......9..
Trifolium pannonicum ......9..
Vicia tenuifolia subsp. dalmatica . . . . . . 9 ..
Juniperus oxycedrus 10 .....27 .25
Stachys officinalis . . . 8 . . 18 7 .
Galium aparine . . . . . . 18 . 17
Campanula rapunculus . . . . . . 14 . 8
Digitalis lamarckii ......14 .8
Galium verum ......14 .8
Teucrium chamaedrys 10 50 30 . . . 41 . 17
Cluster 8
Rubus ulmifolius . . . . . . . 40 .
Cirsium pseudopersonata . . . . . . . 20 .
Arum italicum .......13 .
Veronica serpyllifolia . . . . . . . 13 .
Cluster 9
Vicia cracca . . . 17 . . 5 . 83
Crataegus rhipidophylla ........42
Lathyrus roseus . . . . . . . . 42
Nepeta nuda subsp. albiflora . . . . . . ..42
Sorbus umbellata ........42
Cephalanthera rubra . . . 17 . . 9 . 58
Aristolochia pallida . . . . . . . . 33
Silene latifolia . . . . . . . . 33
Asperula involucrata ......14 .42
Cephalanthera damasonium ........25
Vicia abbreviata . . . . . . . . 25
Silene italica . . . . . . 9.33
Trifolium m edium . . . 8 . . 5 . 33
Poa nemoralis . . 5 33 67 . . . 67
Astragalus ornithopodioides ........17
Astragalus ponticus . . . . . . . . 17
Hippocrepis emerus . . ......17
Moehringia trinervia . . . . . . . . 17
Onobrychis arenaria subsp. cana . . . . . . . . 17
Prunus mahaleb . . . . . . . . 17
Tanacetum parthenium 20 25 . . . .27 .50
Astragalus glycyphylloides .25 .8.14 ..42
Cornus mas 10 25 10 75 100 57 50 7 100
Dianthus calocephalus . . . . . . 5 . 17
Geranium molle . . . . . . 5 . 17
Hedysarum varium . . . . . . 5 . 17
Salv ia verticillata ......5.17
Clematis vitalba 10 25 15 8 . . 5 7 33
Securigera varia 20 50 15 8 33 .5.42
Specie s diagnostic for two or more clusters
Erica arborea 100 100 . . . 14 5 . .
Cistus creticus 60 75 ...14 97.
Campanula alliariifolia 60 50 80 . . . . . .
Vitis v inifera 50 50 .8.43 5..
Iris lazica 40 75 .......
Pteridium aquilinum 70 50 25 42 . 71 5 33 8
Rubus subgen. Rubus 40 25 35 8 . . 5 7 8
Ilex colchica 40 . 15 25 . 57 5..
Heder a helix 90 75 90 50 67 14 987 50
Genis ta tinctor ia 10 75 . 67 . . . . .
Verbascum lagurus subsp. ponticum . 50 . . . 43 . ..
Potentilla m icrantha 30 . 60 50 . 43 . . 8
Sanicula europaea 10 . 60 42 . 86 . . .
Acer cam pestre ..55 ..71 45 40 .
Dioscorea communis 20 50 60 58 . 57 . . 17
Festuca heterophylla . . . 58 ...53 .
Sorbus torminalis 10 . 10 83 . 29 32 . 83
Asperula cimulosa 30 25 . 83 33 57 . . 33
Epimedium pubigerum 20 25 . 83 67 57 . 13 8
Salvia forsskaolei 20 50 . 83 33 71 . 13 8
Prunus domestica subsp. insititia ...33 ....25
Digitalis lanata . . . 33 . . . . 25
Cirsium hypoleucum 30 25 . 75 33 57 5 . 17
Fagus orientalis . . 15 67 100 14 . 7 .
Lathyrus aureus 20 50 .58 .43 ...
Lathyrus laxiflorus 20 .70 92 67 100 32 .75
Stellaria holostea . . . 50 33 . 41 . 50
Cardamine bulbifera . . . 33 67 . 5 . .
Helleborus orientalis 10 50 30 75 33 86 5 40 50
Geranium asphodeloides . . . 33 . 43 . 13 25
Lathyrus tukhtensis ....67 ...42
Carpinus betulus . . 35 58 100 86 9 . 8
Clinopodium vulgare 10 ...67 .18 .50
Physospermum cornubiense . . 20 8 67 . . . 50
Cyclamen coum . . 5 42 100 71 14 33 50
Laser trilobum . . 5 17 67 43 . . 17
Lapsana communis 30 25 30 58 100 43 27 . 75
Euonymus europaeus .25 .17 67 71 ..25
Quercus hartwissiana . . 5 . . 57 27 . .
Crataegus monogyna . . . . . . 59 93 .
Euphorbia amygdaloides . . 5 . . . 36 60 8
Quercus petraea 80 25 65 75 .29 .78
Ruscus aculeatus 90 100 75 8.100 36 100 .
Smilax excelsa 80 75 95 25 . 100 5 47 .
Quercus cerris . . . 58 33 57 82 100 100
Other species
Carpinus orientalis 80 100 100 83 100 100 100 100 100
Crataegus germanica 40 50 10 25 33 43 5 27 8
Geum urbanum 20 50 . 33 . 29 32 . 25
Dactylis glomerata ..533 ..27 725
Fragaria vesca 20 . 10 25 33 . 5 7 25
Melica uniflora . . . 25 33 29 . . .
Pyracantha coccinea 20 . . 8 . 14 5 . .
Salvia glutinosa . 25 15 . . 14 . . .
Salvia tomentosa ....33 .18 ..
Trifolium pratense . . 10 . . . 5 . 17
Geranium lucidum . . . . . . 9 . 17
Geranium robertianum 10 . 5 8 . 14 . . .
Veronica chamaedrys . . . . . . 9 7 8
Athyrium filix-femina ......9.8
Campanula involucrata . . . . . . 9 . 8
Lactuca muralis ..5...9..
Petrorhagia saxifraga . . 5 . . . 9 . .
Polypodium vulgare . . 10 . . . . 7 .
Vicia hirsuta ... . . . 9 . 8
Achillea bis errata . 25 . . . . 5 . .
Anthemis kotschyana ......5.8
Anthemis tinctor ia . . . . . . 5 . 8
Astragalus leucothrix . . . . . . 5 . 8
Calystegia sepium . . . 8 . 14 . . .
Celtis australis 10 .....5..
Circaea lutetiana 10 25 .......
Epipactis condensata . . . . . . 5 . 8
Epipactis pontica . . . . . . 5 . 8
Hieracium murorum . . 5 . . . . . 8
Lathyrus nissolia . . . . . . 5 . 8
Ochlopoa annua . . . . . . 5 . 8
Ononis pusilla . . . . . . 5 . 8
Pimpinella tragium ...8. 14 . . .
Poa bulbosa .. . . . . 5 . 8
Securigera cretica . . . . 33 . . . 8
Sideritis montana . . . . . . 5 . 8
Ulmus minor . . 5 . . 14 . . .
Vicia sativa subsp. nigra .. . . . . 5 . 8
Acer trautvetteri 10 . . . . . . . .
Adiantum capillus-veneris . . 5 . . . . . .
Alnus glutinosa subsp. barbata 10 . . . . . . . .
Argyrolobium biebersteinii ..... . . 7 .
Aristolochia parvifolia ........8
Aristolochia pontica . . 5 . . . . . .
Asplenium ruta-muraria ..5......
Blackstonia perfoliata .... . . 5 . .
Campanula raddeana ..5......
Cardamine parviflora ..5......
Carex humilis ..5......
Cirsium vulgare ..5. . . . . .
Cleistogenes serotina 10 . . . . . . . .
Crepis reuteriana . . . . . . 5 . .
Cyanus depressus . . . . . . . . 8
Echinops spinosissimus . . . . . . 5 . .
Epilobium montanum . . . . . . . 7 .
Epilobium parviflorum . . . . . . . 7 .
Erigeron annuus . . 5 . . . . . .
Erysimum pulchellum . . . . . . 5 . .
Euphorbia orientalis . . . . . . 5 . .
Euphorbia stricta . . 5 . . . . . .
Fallopia convolvulus . . . . . . 5 . .
Filago pyramidata . . . . . . . 7 .
Galium album ..5 . . . . . .
Geranium sanguineum . . . . . . . 7 .
Gledits ia triacanthos . . 5 . . . . . .
Hordeum bulbosum . . . . . . . . 8
Jacobaea vulgaris ..5 . . . . . .
Jasminum fruticans . . . . . . 5 . .
Legousia speculum-veneris . . . . . . 5 . .
Leptopus chinensis . . 5 .. . .. .
Lonicera caucasica . . 5 . . . . . .
Lotus corniculatus . . 5 . . . . . .
Malus sylvestris ..5 . . . . . .
Medicago ×v aria ......5 . .
Melampyrum elatius . . 5 . .. . . .
Melilotus officinalis .. . . . . 5 . .
Mercurialis perennis . . 5 . . . . . .
Morus alba . . 5 .. . . . .
Myosotis sylvatica ..... . . . 8
Ornithogalum sigmoideum . . . . . . . 7 .
Paeonia mascula .. . . . . . 7 .
Pachyphragma macrophylla . . 5 . . .. . .
Pinus sylvestris . . 5 . .. . ..
Plantago major . . 5 . . . . . .
Polygonatum glaberrimum . . 5 . . . . . .
Primula veris subsp. macrocalyx ..5 . . . . . .
Pulmonaria dacica . . 5 .. . ...
Punica granatum . . 5 . . . . . .
Ranunculus bulbosus . . 5 . . . . . .
Ranunculus polyanthemos ..5 . . . . . .
Rhagadiolus stellatus .... . .. . 8
Rhamnus alaternus . . . . . 14 . . .
Sambucus nigra . . 5 . . . . . .
Scabiosa sosnowskyi . . 5 . . . . . .
Sison amomum . . 5 . . . . ..
Smilax aspera .... . 14 . ..
Sonchus asper . . . . . . 5 . .
Sorbaria tomentosa . . . 8 . . . . .
Sorbus aucuparia . . . 8 . . . . .
Tanacetum partheniifolium . . 5 . . . . . .
Taraxacum sect. Taraxacum . . 5 . . . . . .
Thalictrum minus . . . . . . 5 . .
Torilis japonica . . 5 . . . . . .
Trachycarpus fortunei . . 5 . . . . . .
Trachynia distachya . . . . .. 5 . .
Trifolium diffusum . . 5 . .. . . .
Trifolium ochroleucon .. . . . . 5 . .
Veronica officinalis . . . . . 14 . . .
Veronica orientalis ..... . 5 . .
Vicia sepium . . 5 . . . . . .
Number of relevés per associati on
Campanulo alliariifoliae-Carpinetum orientalis . . 20 . . . . . .
Carpino betulo-orientalis . . . 11 . . . . .
Carpino or ientalis-Phillyrietum latifoliae .... . . 6 . .
Carpino orientalis-Quercetum cerridis . . . . . . . 15 .
Corno maris-Quercetum cerridis . . . . . . 5 . 5
Crataego curvisepalae-Quercetum cerridis . . . . . . . . 7
Erico arboreae-Carpinetum orientalis 10 4 . . . 1 1 . .
missing associat ion assignment . . . . 3 . . . .
Querco cerridis-Carpinetum orientalis . . . . . . 10 . .
Rusco aculeate-Carpinetum orientalis .. . 1 . 6 . . .
Supplement E3. Full synop tic table. Species p ercentage frequencies in the clusters are provided. Diagnostic (Φ ≥ 0.2,
gre y shaded) and highly diagnostic (Φ ≥ 0.55, gr ey shaded, in bold) species for each cluster are in the upper part of the
table, sorted by decreasing fidelity . Other species are provided below, sorted by decreasing frequency in the dataset.
Correspondence between associations and clusters is at the bottom of the table.
AnhangtE3. Vollständige Übersichtstabelle und Übereinstimmung zwischen den Assoziationen und Clust ern.
Diagnostische (Φ ≥ 0.2, grau schattiert ) and hoch diagrnostische (Φ ≥ 0.55, grau schattiert, in Fettdruck) Art en für
jeden Cluster sind im oberen Teil der T abelle, sortiert nach abnehmender Treue. Andere Arten sind darunter
aufgeführt, nach abnehmender Frequenz im Dat ensat z sortiert. Die Übereinstimmung zwischen Assoziationen und
Clustern findet sich am Fuß der Tabelle.
Novák et al.: Carpinus orientalis forests in Georgian Colchis: First insights
Supplement E4. Additional photos of the vegetation of the association Campanulo alliariifoliae-Carpinetum orientalis ass. nova hoc loco and
its sites in western Georgia. a) Forested landscape of the southern macroslope of the Racha Massif. Carpinus orientalis forests occupy mainly
limestone rock outcrops here, while mixed forests of Carpinus betulus, Castanea sativa and Fagus orientalis dominate on zonal sites (July
2017). b) River Tskaltsitela limestone canyon above the Motsameta Monastery with extensive Carpinus orientalis forests (July 2019). c)
Forests of Carpinus orientalis colonizing the upper part of a sunny limestone rock near the city Tkibuli (July 2017). d) Mixed Carpinus betulus
and C. orientalis forests on a limestone ridge near the Sataplia Cave above the city Tskaltubo (July 2016). e) Carpinus orientalis forest below
the Motsameta Monastery near the city Kutaisi. Recent massive dieback of Buxus sempervirens shrubs is apparent (July 2019). f) Undergrowth
of an open Carpinus orientalis forest near the city Martvili. Festuca drymeja, Laser trilobum and Leptopus chinensis are visible in the
understorey (July 2017). All photos by P. Novák.
Anhang E4. Zusätzliche Fotos der Vegetation des Campanulo alliariifolae-Carpinetum orientalis ass. nova hoc loco und ihrer Standorte in
Westgeorgien. a) Waldlandschaft des südlichen Abhangs des Racha-Massivs. Carpinus orientalis-Wälder besetzen hier hauptsächlich
Kalksteinfel-sen, während Mischwälder aus Carpinus betulus, Castanea sativa und Fagus orientalis an zonalen Standorten dominieren (Juli
2017). b) Kalksteinschlucht des Flusses Tskaltsitela oberhalb des Klosters Motsameta mit ausgedehnten Carpinus orientalis-Wäldern (Juli
2019). c) Carpinus orientalis-Wälder, die den oberen Teil eines sonnigen Kalksteinfelsens in der Nähe der Stadt Tkibuli besiedeln (Juli 2017).
d) Mischwälder aus Carpinus betulus und C. orientalis auf einem Kalksteinrücken in der Nähe der Sataplia-Höhle oberhalb der Stadt Tskaltubo
(Juli 2016). e) Carpinus orientalis-Wald unterhalb des Klosters Motsameta in der Nähe der Stadt Kutaisi. Das kürzliche massive Absterben
von Buxus sempervirens-Sträuchern ist offensichtlich (Juli 2019). f) Unterholz eines offenen Carpinus orientalis-Waldes in der Nähe der Stadt
Martvili. Festuca drymeja, Laser trilobum und Leptopus chinensis sind im Unterwuchs sichtbar (Juli 2017). Alle Fotos von P. Novák.
