The environmental history of Georgia during the Late Miocene based of foraminifera and pollen

Full text

საქართველოს ეროვნული
მუზეუმი
G E O R G I A N
N A
T I O N A L
M U S E U M
RESEARCH INSTITUTE OF PALEOANTROPOLOGY AND PALEOBIOLOGY
TSU Aleksandre Janelidze
Institute of Geology
Georgian Technical
University
The environmental history of Georgia during the Late Miocene
based of foraminifera and pollen
Shatilova I.I., Maissuradze L.S., Koiava K.P.,
Kokolashvili I.M., Bukhsianidze M.G., Bruch A.A.
TBILISI-2020

UDC 502.7(479.22)
S - 53
Addresses of authors:
Shatilova
I.I.
-
Georgian National Museum Institute of
Paleobiology. Rustaveli av. 3, Tbilisi, Georgia
Maissuradze
L.S.
-
Georgian National Museum
Institute of Paleobiology. Rustaveli av. 3,Tbilisi,
Georgia
Koiava
K.P.
- Ivane Javakhishvili Tbilisi State
University, Aleksandre Janelidze Institute of Geology,
A. Politkovskaia str. 31, Tbilisi 0186, Georgia
Kokolashvili I.M.
- GeorgiaTechnicalUniversty,
Gamsakhurdia av.77., Tbilisi, Georgia
Bukhsianidze M.G.
-
Georgian National Museum
Institute of Paleobiology. Rustaveli av. 3, Tbilisi,
Georgia
Bruch
A.A.
-
ROCEEH Research Centre, Senckenberg
Research Institute, Frankfurt/M, Germany
Editor: David Lordkipanidze
Reader: Maya Grigolava
ISBN 978-9941-26-684-3
Georgian National Museum, 2020

3
Preface
In Georgia, observed changes in the evolution of vegetation and marine microfauna
during the Sarmatian to Pontian time can be correlated to the same stratigraphic levels. This
suggests significant environmental changes which effected both, marine and terrestrial
realms. Especially two stratigraphic events played a major role as turning-points for the
environmental history of Georgia in late Miocene.
One is the transition between the middle and upper Sarmatian, when important paleo-
geographical transformations took place in the Caucasus region. The Transcaucasus
intermontane depression became dry land with the Kura Bay in the East, which was
connected to the Caspian Sea, and the Rioni Bay in the West, which was the part of Black
Sea. In the East, continental deposits started to form, while in the West, along the Rioni Bay,
the uninterrupted accumulation of the marine deposits continued.
The second common turning-point in the environmental evolution of marine and
terrestrial biota is the boundary between Meotian and Pontian. This is marked by the
Eupatorian horizon with a pronounced impoverishment in the composition of marine fauna
and of vegetation, probably connected to a worsening of climatic conditions.
To study these bioevents in more detail, the objectives of this investigation are to
review and summarize information on the assemblages of foraminifers and pollen from
Georgian deposits of Sarmatian, Meotian and Pontian and compare their implications on
environmental changes. Furthermore, a comparison of the vegetation history in Eastern and
Western Georgia will reveal insights into the beginning of the diversification of terrestrial
habitats in the Caucasus area.
The following Sarmatian sections were studied: Aragvi, Nadarbazevi, Uplistsikhe,
Udabno, Gombori, Chachuna (Eastern Georgia), Chanistskali, Vake (Western Georgia).
Marine deposits of Meotian and Pontian age were studied in sections of Western
Georgia: Vake, Djapareuli, Gedjiri, Otapi Galidzga, Gudau, and Urta-Zana (Fig.1).
For all sections studied by pollen data two types of diagrams are prepared. The first
one shows changes of pollen abundances of the main woody plants for each section. The
diagrams are divided into pollen zones, distinguished on the basis of pollen assemblages
found within them. In this case the pollen zones are considered as stratigraphical unit.

4
The second type of diagram reflects the abundance of ecological groups of plants
indicating the changes in the major forest formations on territory of Georgia during the
Sarmatian, Meotian and lower Pontian.
The stratigraphical chart of late Neogene, a map of the studied sections, lists of
foraminifers and plants determined by pollen data, and two types of pollen diagrams are
enclosed to this monograph.

5
Fig. 1. Simplified Geological map of Georgia with locations of study sections
(Geological map adapted after G. Gudjabidze, I. Gamkrelidze, 2003)

6
1. Introduction
The Sarmatian regional stage of Eastern Paratethys is divided into three regiosubsta-
ges: Volhynian, Bessarabian and Khersonian (Table I). This division is based on lithological
and biofacial characteristics that are especially clearly expressed in the outcrops of Azov-
Black Sea region. The Volhynian substage is characterized by euryhaline foraminiferal
associations, followed by marine faunas of the Bessarabian substage indicating a large-scale
marine transgression. During the Khersonian reversed environmental changes took place and
the Eastern Paratethys turned into a closed basin (Nevesskaya et al. 2003).
Table I. Correlation of standard chronostratigraphy units and Eastern Paratethys regional
stages for Sarmatian-Pontian. Standard chronostratigraphy based on the International
Stratigraphic chart 2017/02 (ICS); Regional stages, substages, formations and beds modified
after Buleishvili, 1960, Goncharova, 1989; Nevesskaya et al., 2004, Vernyhorova, 2018;
Regional stages Ma Data: Vasiliev et al., 2011, van Baak et al., 2016, Palcu et al., 2017, 2019

7
The marine deposits of lower and middle Sarmatian are known in both regions,
Eastern and Western Georgia. While in the West the upper Sarmatian has a limited
distribution and is represented mainly by conglomerates (Buleishvili 1986). In the East it is
represented by shallow marine and continental deposits. Younger marine deposits of Meotian
regional stage are known only from Western Georgia.
Meotian deposits are represented in Abkhazia, Samegrelo and Guria and are divided
into two regional substages: Bagerovian and Akmanaian. The most complete outcrops are
found in Abkhazia. In Guria deposits mainly of the lower Meotian occur, while in Samegrelo
upper Meotian sections are present (Badzoshvili 1979).
The Meotian is overlain by Pontian. Continuous deposits from upper Meotian to
lower Pontian are found only in sections Otapy and Urta-Zana, where the Novorossian
substage begins by Eupatorian and is overlain by the Odessian horizon. The deposits of
Eupatorian are of minor thickness (2-4m). They are widely distributed in the Black Sea-
Caspian region, but not present in all sections of Pontian deposits. After Taktakishvili
(1984) the main reason of this phenomenon is the wide early Pontian transgression, which
eroded the thin Eupatorian layers.
The Eupatorian deposits of Western Georgia are characterized by impoverished
assemblages of brackish mollusks similar to those in same layers of Western Crimea,
indicating the strong freshening of basin. In both regions the main fossils are Prosodacna
littoralis (Eichwald) and Congeria novorossica (Sinzov). The other part of benthos is
composed by freshwater gastropods (Taktakishvili 1978, 1984; Taktakishvili et al. 2018).
The present research is based on rich fossil material from numerous sections in
Eastern and Western Georgia (Table II). Pollen and foraminifera come from sections that
cover the interval between lower Sarmatian and the boundary between Meotian and Pontian.
This allows us to reconstruct the history of marine and terrestrial biomes of Georgia during 7
million years spanning from ca. 12.65-5.6 Ma (van Baak et al., 2016; Palcu et al., 2017).

8
Table II. List of sections of Sarmatian to Pontian age in Georgia considered
in this study (F: foraminifera, P: pollen)
Name Stage(s) Latitude Longitude Reference
Eastern Georgia
Aragvi Sarmatian 41
o
51‘13“ 44
o
42‘35“ F: Koiava, 2006
a
P: Shatilova et al., 2012
Nadarbazevi Sarmatian 41
o
58‘03“ 44
o
19‘16“
F & P: Maissuradze et al. 2008
P: Shatilova & Kokolashvili, 2013
F: Ediberidze et al., 2019
Uplistsikhe Sarmatian 41
o
58’26” 44
o
14’27”
F & P: Shatilova et al., 2008
P: Shatilova & Kokolashvili, 2013
F: Ediberidze, 2019
Udabno Sarmatian 41
o
26’50” 45
o
22’33” P: Shatilova & Kokolashvili, 2013
Gombori Sarmatian 41
o
48’32” 45
o
08’27” F: Koiava 2006
P: Shatilova & Kokolashvili, 2013
Chachuna Sarmatian 41
o
12’13” 45
o
58’10” P: Kokolashvili et al. 2018
Western Georgia
Chanistskali Sarmatian 42
o
39‘24“ 42
o
07‘14“ P: Shatilova et al. 1999
Vake Sarmatian
/Meotian 41
o
59‘33“ 41
o
51‘27“ P: Shatilova et al. 1999
Galidzga Meotian
/Pontian 42
o
50‘10“ 41
o
36‘01“ F: Popkhadze, 1977
F
& P: Shatilova et al. 2008
Urta-Zana Meotian
/Pontian 42
o
21‘52“ 42
o
00‘48“ F: Popkhadze, 1977
F& P: Maissuradze et al. 2013
Gedjiri Meotian 42
o
53‘03“ 41
o
39‘14“ F& P: Shatilova et al.2018
Gudau Meotian 43
o
07‘01“ 40
o
38‘36“ F: Popkhadze, 1977
F
& P: Shatilova 1992
Otapi Meotian
/Pontian 42
o
53‘10“ 41
o
32‘29“ F & P: Maissuradze et al. 2013

9
2. A micropaleontological review
The foraminifers of Miocene deposits of Black Sea-Caspian region were studied by
many paleontologists, most importantly Bogdanowicz (1965), Didkovsky (1966),
Dzhanelidze (1970), Dzhanelidze et al. (1985). The foraminifers of Sarmatian and Meotian
deposits of Western Georgia were the subject of investigations of Maissuradze (1966,
1971, 1980, and 1988). The foraminifers of Sarmatian deposits of Eastern Georgia were
studied by Koiava (2004, 2006). According these authors the ancestors of Sarmatian
foraminifers were middle Miocene euryhaline taxa, which survived the decrease in salinity
in the basin at the end of the Konkian. During the Sarmatian three main stages are distingu-
ished in the history of foraminifers. The first stage reflects the process of formation of the
Sarmatian fauna, the middle is its maximal development, and the late stage is time of
extinction (Maissuradze et al. 2008).
The lower Sarmatian conformably overlies the Konkian deposits and is represented by
argillo-arenaceous facies. The thickness of the lower Sarmatian varies between 50-90 m in
the coastal region and between 250-350 m in the central part of the basin. The sections are
dated by foraminifers: Elphidium macellum (Fichtel & Moll), E. hauerinum (d’ Orbigny), E.
crispum (Linnaeus), E. optusum (d’ Orbigny), E. angulatum (Egger), Elphidiella artifex
(Serova). In lower numbers of specimens occurs Ammonia ex gr. beccarii (Linnaeus),
Nonion tumidulus Pishvanova, N. bogdanowiczi Voloshinova, Sinuloculina consobrina
(d’Orbigny), Porosononion martkobi (Bogdanowicz), P. subgranosus (Egger). The lower
Sarmatian is subdivided into layers with Varidentella reussi and with Elphidium aculeatum.
Below the list of foraminifes from Sarmatian deposits of Eastern Georgia is given
(Table III).

10
Table III. The distribution of Sarmatian foraminifera of Georgia
(based on Maissuradze, 1971 and Koiava, 2006)

11
The lower Sarmatian is overlain by the middle Sarmatian, with thicknesses from 100
to 1000m (Buleishvili 1960; Koiava 2006, 2006
a
). Deposits are dated by specific association
of foraminifers and are divided into three parts (Koiava et al., 2008): layers with Affinetrina
voloshinovae, layers with Porosononion aragviensis and layers with Porosononion hyalinus.
The lower part of the middle Sarmatian is represented by comparatively deep-water
argillaceous deposits. The foraminiferal complex consists of the following taxa: Affinetrina
guriana (Dzhanelidze), A. voloshinovae (Bogdanowicz), Sinuloculina angustioris Bogda-
nowicz, Varidentella floriformis Bogdanowicz and others.
In general the middle part of middle Sarmatian is the flourishing age of foraminifera.
They increase significantly in size and endemic forms achieve their maximum diversity.
Interestingly, in synchronous deposits of Eastern and Western Georgia the abundances of
miliolids and elphidiids differ: the miliolids are dominant in the West, while elphidiids
(Nonionidae and Elphidiidae) prevail in the East.
The foraminiferal complexes sharply impoverish from the final middle Sarmatian
onward: all middle Sarmatian endemics and species present since the early Sarmatian
disappear. This impoverished complex is composed of the following taxa: Porosononion
hyalinus Bogdanowicz, P. subgranosus subgranosus (Egger), P. subgranosus umboelata
Gerke, P. aragviensis (Dzhanelidze), Elphidium fichtelianum (d’ Orbigny), E. macellum
(Fichtel & Moll), E. crispum (Linnaeus). Thick-shelled Affinetrina guriana (Dzhanelidze)
and Ostracodae are rare. This assemblage is known as Porosononion hyalinus complex.
Orogenic movements started at the end of the middle Sarmatian and achieved a
maximum in the late Sarmatian. As a result a continental setting was established on most of
the Georgian territory. The Kartli depression and large parts of Kakheti transformed into a
region of accumulation of continental deposits. Only isolated basins remained in Eastern
Georgia (Koiava et al. 2012).
In Kartli thick series of continental deposits of the so-called Natskhori Formation, with
thicknesses from 1600 to 2500m, belong to the upper Sarmatian. This formation is widely
distributed on the slopes of depressions and forms parts of most syncline and anticline folds.
In Kakheti the upper Sarmatian mostly is represented by continental deposits of the
Eldari Formation with freshwater gastropods. However, towards the South-East the change
to continental depositional environments occurred later. Here the lower part of the upper
Sarmatian is still represented by marine sediments with Mactra (Maissuradze, Koiava 2006).

12
The late Sarmatian regression was followed by a transgression during the Meotian.
Black Sea and Mediterranean Sea became connected and marine fauna penetrated into the
Eastern Paratethys. At the beginning of the Meotian, relatively stenohaline species of
foraminifers and many other organisms colonized the basin. A new stage of development of
Miocene fauna began. Most of the Meotian foraminifers reveal some similarities with recent
Mediterranean species and also with middle Miocene taxa of the Ponto-Caspian region
(Maissuradze, Koiava 2011).
The predominate fossils of microfauna in Meotian are foraminifers. Ostracods are
represented only by single tests. The picture changed in Eupatorian, in deposits of which the
abundance and variety of ostracods species increased. The genera Pontaniella and Bakunella
appeared here for the first time. The genera Leptocythere and Loxoconcha are represented by
numerous species. The number of foraminifers and their specific diversity is low. The fora-
miniferal assemblage is composed of euryhaline species, which morphologically differ
from taxa widely distributed in the late Meotian. The assemblage has a mixed Meotian-
Pontian composition.
In Odessian, species of Pontaniella and Bakunella, Lineocypris, Caspiocypris, and
some Leptocythere penetrated from several basins of the Paratethys into Rioni bay.
Meotian relicts fully disappeared but in general the diversity of ostracods became richer
(Maissuradze et al. 2013).
Below the list of framinifers from Meotian deposits of Western Georgia is given
(Table IV).

13
Table IV. The distribution of Meotian foraminifera of Georgia
(based on Maissuradze, 1988 and Maissuradze & Koiava, 2011
a
)
So, judging from the history of development of Meotian and early Pontian
foraminifers it is possible to conclude that a gradual deterioration of bionomical conditions
promoted the disappearance of foraminifers from the Pontian basins of Eastern Paratethys.
Their extinction began in the late Meotian and finished in the Eupatorian. The decrease of
salinity was the main factor that influenced this process. The composition of brackish and
freshwater mollusks distributed during these epochs and also the abundance of ostracods
characteristic for freshwater basins confirms this fact. The salinity of water in basins of the
late Meotian and Pontian is determined as 5
0
/
00
that was pernicious for foraminifers but
favorable for ostracods, which after Meotian became the dominant microfaunistical group
(Davitashvili 1933; Eberzin 1959; Dzhanelidze et al. 1985; Maissuradze 1980; Maissuradze,
Koiava 2008, 2011; Maissuradze et al. 2013; Taktakishvili 1984; Taktakishvili et al. 2018).

14
3. A review of the main components of the Sarmatian to Pontian floras of
Georgia
Historically, the knowledge about the Georgian flora of Sarmatian to Pontian was
based on macrobotanical remains (Uznadze 1965; Uznadze, Tsagareli 1979; Kolakovsky et
al. 1970; Kolakovsky, Shakryl 1976; Chelidze 1972, 1979, 1987; Purceladze, Tsagareli
1974). Since the last decades, also rich palynological material was studied from numerous
outcrops of Eastern and Western Georgia (Shatilova 1992; Shatilova et al. 2008, 2009, 2010;
Shatilova, Kokolashvili 2013).
One of the main characteristics of upper Miocene floras of Georgia is the exceptional
high diversity of ferns belonging to 18 families.
The family Schizaeaceae was represented by taxa determined as Microfoveolatosporis
neogranuloides Krutzsch and Reticulosporis polonicus Krutzsch. In Georgia, M.
neogranuloides is known only from the Sarmatian of Eastern Georgia. The spores of M.
neogranuloides are similar to recent Schizaea pennula Swardz, a taxon which is now
distributed in tropical America (Bolkhovitina, 1961). R. polonicus was found in the flora of
both Eastern and Western Georgia. This species persisted until Kimmerian in Western
Georgia. Its spores are comparable with the recent Schizaea pussila Pursh, which as well is a
tropical species, growing in the South of North America (Stuchlik et al. 2001).
The family Anemiaceae shows a high taxonomic diversity in the fossil record,
including the genera Cicatricosisporites Potonié & Gelletich, Radialisporis Krutzsch,
Anemia Swartz, Mohria Swartz, and Pelletieria Seward. Anemia, a plant today occurring in
tropical America, was represented by numerous species found mainly in Eastern Georgia.
From Sarmatian and Meotian deposits of Western Georgia the species Cicatricosisporites
chattensis Krutzsch and Radialisporis radiatus (Krutzsch) Jansonius & Hills are determined,
which are fossil taxa morphologically closely related to Anemia. The genera Mohria and
Pelletieria were components of the Sarmatian flora of both regions of Georgia. They
survived throughout the entire Meotian in Western Georgia.
The family Lygodiaceae is represented mainly by the genus Lygodium with six
species, which were widely distributed on the territory of Georgia during the Sarmatian and
Meotian. Later, only one species, L. reticulatum Schkuhr, prevailed in Western Georgia until

15
Kimmerian. Today, representatives of the genus Lygodium are plants of tropical regions
(Bolkhovitina 1961).
From the families Pteridaceae and Adiantaceae the following genera were determined:
Segmentizonosporites Kedves, Pteridacidites Sah, Anogramma Link, Cryptogramma
R.Braun and Pityrogramma Link. In Sarmatian floras the specific diversity of these taxa was
low. During the Meotian the number of species of genus Pteridacidites increased. But in
general, the flourishing of families Pteridaceae and Adiantaceae on the territory of Georgia
began only after the Meotian (Shatilova, Stuchlik 1996; Shatilova et al. 2016).
The family Gleicheniaceae is characteristic for the Sarmatian flora comprising
Clavifera triplex (Bolchovitina) Bolchovitina, C. tuberosa Bolchovitina, Gleicheniidites
carinatus (Bolchovitina) Bolchovitina and G. senonicus Ross, preserved on the territory of
Western Georgia until Pontian. Today the ferns of this family are distributed in tropical and
subtropical zones of both hemispheres (Bolkhovitina 1968).
In the family Polypodiaceae the genera Polypodium L. and Pyrrosia Mirbel are
determined. The abundances of Polypodium spores in the Sarmatian and Meotian floras were
low. More characteristic was the genus Pyrrosia, especially during the Sarmatian. Today the
genus Pyrrosia is a component of moist tropical mesophilous forests (Bobrov 1978).
The fossil species Verrucatosporites histiopteroides Krutzsch was the component of
the Neogene flora of Georgia with the widest distribution. It is compared to the modern
Histiopteris incisa (Thunberg) J. Sm., which is a common plant in Australia and New
Zealand (Krutzsch 1962; Stuchlik et al. 2001).
By botanical affinity the species Verrucatosporites favus (Potonié) Thomson & Pflug
belongs to family Dennstaedtiaceae and is compared with the recent genus Paesia A.
Saint-Hilaire, distributed in tropical America and Malaysia (Stuchlik et al. 2001).
Dicksoniaceae are respesented by a rich specific composition of the genus Dicksonia
L’Heritier. Some of the species determined from Sarmatian and Meotian assemblages
persisted in Western Georgia until the Gurian.
Spores of the genus Hemitelia R. Braun were described first from Sarmatian deposits
of Eastern Georgia (Shatilova et al. 2016). It is a woody fern, a component of the understory
in sclerophyllous forest of Southern Africa (Walter 1974).
In summary, most of the ferns in the Sarmatian to Pontian floral assemblages of
Georgia were plants, which recent relatives occur in the tropical and subtropical zones of

16
both hemispheres. Also, quite a large number of fern taxa are fully extinct and have been
determined by morphological nomenclature.
In the family Podocarpaceae the genera Dacrydium Solander, Podocarpus L’Heritier
ex Persoon, Pinipollenites Potonié and Podocarpidites Cookson ex Cooper are determined.
The genus Dacrydium comprised three species: D. aff. pierrei Hickel, Dacrydium sp.
1, and sp. 2. On the territory of Western Georgia the genus prevailed until the Gurian,
represented by D. aff. cupressinum Soland. Now this taxon is distributed in New Zealand and
on Pacific islands.
The genus Podocarpidites was represented by two species P. eocaenicus Krutzsch and
P. nageiaformis (Zaklinskaya) Krutzsch, known only from Sarmatian of Eastern Georgia. In
the Cenozoic the species P. nageiaformis was widely distributed in Europe (Nagy 1985;
Stuchlik et al. 2002). Today the genus Podocarpus comprises about 100 species, distributed
mainly in the Southern Hemisphere. The areal of Dacrydium is restricted to Malasya, New
Zealand and Tasmania (Murav’eva, Borkhvardt 1978).
The genus Phyllocladus was represented by two species P. aff. protractus (Ward.)
Pilg. and P. aff. trichomanoides D. Don. The first is known only from Sarmatian of Eastern
Georgia. The second species was distributed on the territory of Western Georgia from
Sarmatian until Gurian. Most of the 6 species of the genus Phyllocladus are inhabitants of
Tasmania and New Zealand. Those are plants of mountain vegetation, growing at an altitude
of 900-4000m (Murav’eva, Borkhvardt 1978).
From the family Pinaceae the most characteristic components of Sarmatian-Meotian
flora of Georgia are the genera Cathaya Chun et Kuang and Cathayapollis Ziembinska-
Tworzydlo. The genus Cathaya was represented by species C. aff. argyrophylla Chun et
Kuang, which from Middle Miocene until Meotian was a component of the Western Georgia
flora. The fossil genus Cathayapollis was represented by species C. potonié (Sivak)
Ziembinska-Tworzydlo and C. millaya (Sivak) Ziembinska-Tworzydlo, known only from
Sarmatian deposits. Today there are two species of the genus Cathaya distributed in south-
eastern China (Chavchavadze, Yatsenko-Khmelevsky 1978).
Besides the conifers noted above, it is necessary to mention the genus Pinus, which
was the main component of flora in Eastern Georgia during the whole Sarmatian.
So, in Sarmatian in both regions of Georgia the floristic composition of conifers was
nearly similar but the abundances of separate taxa differed. In Western Georgia the warm-

17
temperate genera were predominant, while in Eastern Georgia representatives of genus
Pinus prevailed.
Among the angiosperms the main components of the flora were the genera of the
families Myricaceae, Juglandaceae, Betulaceae, Fagaceae, Lauraceae, Magnoliaceae,
Hamamelidaceae, Icacinaceae, Araliaceae and Arecaceae.
The family Myricaceae comprised Comptonia Banks, Myrica Linnaeus and
Myricipites Wodehouse. The genus Myrica was represented by M. conspiqua Gladkova, M.
notabilis Gladkova, M. pseudogranulata Gladkova and M. intermedia Gladkova. Only the
last species prevailed until Pontian in Western Georgia. The genus Myricipites was
represented by M. bituitus (Potonié) Nagy, which was a component of the flora of Western
Georgia until Kuyalnician.
Of great interest is the presence of pollen grains of family Icacinaceae in Sarmatian
deposits of Eastern Georgia (Shatilova et al. 2018). By macro-remains two species
(Icacinaxylon citronelloides Shilkina, I. goderdzicum Shilkina) of this family are described
from deposits of Goderzi suite, which was dated to the upper Miocene (Sarmatian, Meotian)
based on the petrographic analysis of fossil ash (Skhirtladze 1958; Shilkina 1958). Today the
family Icacinaceae includes 58 genera, widely distributed in tropical and subtropical
mountain forests up to 2000-2500m.
In the family Fagaceae the genera Castanea Miller, Castanopsis (D. Don) Spach,
Lithocarpus Blume, Fagus Linnaeus, Quercus Linnaeus, Cupuliferoipollenites Potonié,
Faguspollenites Raatz and Quercopollenites Nagy were determined.
The species Cupuliferoipollenites pusillus (Potonié) Potonié is known from Eocene
until Sarmatian (Shatilova et al. 2018). Morphologically the pollen of this species is similar
to pollen of recent Castanopsis and Lithocarpus, which are evergreen plants distributed in
subtropical and tropical zones.
The species Faguspollenites verus Raatz, F. minor Nagy, F. subtilis Nagy and F. vivus
Nagy prevailed on territory of Western Georgia until Meotian. At the same time, the recent
species Fagus orientalis Lipsky already existed. Pollen data and fossil plant macro-remains
give evidence for its occurrence on the territory of Georgia already in the Sarmatian
(Shatilova et al. 2011).
One characteristic sign of Sarmatian and Meotian floras of Western Georgia is the rich
composition of the family Hamamelidaceae, represented by 32 species (including macrofossils)

18
belonging to 16 genera and 3 subfamilies (Shatilova et al. 2016
a
). Somewhat poorer was the
composition of this family in Eastern Georgia, where by pollen data the genera Corylopsis
Siebold & Zuccarini, Hamamelis Linnaeus, Fothergilla Murray, Parrotia C.A. Meyer,
Disanthus Maximowicz and Liquidambar Linnaeus were determined. Hamamelidaceae is
regarded as a more ancient family of angiosperms, and is considered as a link between
Trochodendrales and amentiferous orders Casuarinales, Urticales and Fagales. The family
includes about 31 genera and more than 100 species. Among them 15 are monotypic, from
which Disanthus is considered the most ancient (Takhtajan 2009). Today representatives of
Hamamelidaceae are evergreen or deciduous trees and shrubs, which are growing in
subtropical and warm-temperate forests. The largest and richest center of their distribution
(about 50% of species) is East and South-East Asia. The second important distribution area is
the Central America and the third is Eastern and tropical Australia. Most of the species are
endemics and only three genera (Hamamelis, Fothergilla and Liquidambar) are present both in
Asia and America The greatest number of Hamamelidaceae lives in Norhern hemisphere,
excepting Ostrearea, Neostrearia, Trichocladus and Dicoryphe, which occur in Southern
hemisphere (Bogle 1970; Skvorzova 1975; Takhtajan 2009).
The family Alangiaceae was represented by Alangium Lamarck and Alangiopollis
Krutzsch comprising the species A. eocaenicus Krutzsch, A. simplex Nagy and Alangiopollis
sp. The species A. eocaenicus is known in Georgia from Oligocene until Sarmatian
(Shatilova et al. 2018). The pollen grains of this species are compared with Alangium
grisolleoides Capuron (Krutzsch 1969). The second species Alangiopollis sp., which is
known only from Sarmatian can be compared with a taxon described from the Miocene
deposits of Hungary (Nagy 1969) and with a taxon from the lower Miocene of Slovenia
(Pacltova 1966). In the Meotian the family Alangiaceae was represented by Alangiopollis
simplex Nagy and Alanguim aff. kurzii Craib, which prevailed on the territory of Western
Georgia until Gurian. Today the genus Alangium is represented by about 20 species
distributed in western Africa and on Pacific islands (Grushvitsky 1980).
The fossil genus Fupingopollenites was a characteristic component of Sarmatian and
Meotian floras of Georgia, with the two species F. wackersdorfensis (Thiele-Pfeiffer) Liu
Geng-wu and F. minutus Liu Geng-wu. The genus occurred in Eastern Georgia until the end
of Sarmatian and in Western Georgia until the end of Meotian (Shatilova et al.2009).

19
The botanical affinity of Fupingopollenites remains unknown. Lu Geng-wu (1985)
compares it with Scabiosapollis Sung et Zheng and Convolvulus Linnaeus and assumes that
the plant, which produced Fupingopollenites pollen grains, lived in subtropical climate and
was a component of evergreen formations.
Table V. List of plants determined by pollen data in the Sarmatian, Meotian and lower
Pontian deposits of Georgia
Familia Species
Eastern
Western
Sarmatian
Sarmatian
Meotian
Eupatorian
Odessian
1 2 3 4 5 6 7
Sphagnaceae
Distverrusporis pliocenicus
(Krutzsch )
Grabowka X X X X
Stereisporites stereoides
Thomson et Pflug
X
Anthocerotaceae
Rudolphisporites rudolphii
(Krutzsch)
Krutzsch & Pacltova X
Lycopodiaceae
Retitriletes lusaticus
Krutzsch X X
Lycopodium serratum
Thunb.
X X
Selaginellaceae
Echinatisporis miocenicus
Krutzsch X
E. longechinus
Krutzsch X X X
Selaginella fusca
N
.
Mtchedlishvili X X X
X
S. eggersii
Sodiro X X X
S. speciosa
Krasnova X
S. selaginoides
(L.) Link X X X
X
Ophhioglossaceae
Bothrychium
simplex
Hitchcock
X
Osmundaceae
Osmunda cinnamomea
Linnaeus X X X
X
O. regalis
Linnnaeus
X
X
Todea
sp. X
Schizaeaceae
Reticulosporis polonicus
Krutzsch X X X
Microfaveolatosporis neogranuloides
Krutzsch X
Microfaveolatosporis
sp. Kohlman-Adamska X
Schizaea
sp. X X X
Anemiaceae
Cicadicosisporites chattensis
Krutzsch X X X
Radialisporis radiatus
(Krutzsch) Jansonius &
Hills X X X
Anemia
aff.
auriculata
Bayrich X

20
1 2 3 4 5 6 7
Anemiaceae
A.
aff.
imbricata
Sturm
X
A
. aff.
imperfecta
(Mal.) Bolchovitina X
A.
aff.
hirta
( L.) Swarz X
A
. aff.
schimperiana
Presl X
A.
aff.
mexicana
Klotzsch X
A.
aff.
filiformis
(Sw.) Swartz X
Mohria
sp. X X X
Pelletieria
aff.
pacifica
Bolchovitina X X X
Lygodiaceae
Lygodium digitatum
Presl X X X
L. conforme
Christ.
X X
L. multivallatum
(W. Krutzsch) Ramishvili X X
L. japonicum
(Thbg.) Swartz X X X
L. oligostachyum
(Willd.) Desv. X X X
L. reticulatum
Schkuhr
X
Pteridaceae
Cryptogramma acrostichoides
R. Braun
X
C. crispa
(L.) R. Braun
X
Segmentizonosporites paucirugosus
(Nagy)
Stuchlik X
S. latigracilis
(Krutzsch) Stuchlik X X
Polypodiaceoisporites corrutoratus
Nagy
X X
Pteridacidites helveticus
(Nagy) Stuchlik &
Shatilova X X
P.boerzsoenyensis
(Nagy) Stuchlik & Shatilova X X
P. dentatiformis
Shatilova & Stuchlik
X X
P.georgensis
Shatilova & Stuchlik
X
X
P. grandifoliiformis
Stuchlik & Shatilova X X X
X
P. guriensis
Shatilova & Stuchlik
X
P
.
longifoliiformis
Shatilova & Stuchlik
X X
X
P
.
pseudocreticus
Shatilova & Stuchlik
X
P.rarotuberculatus
Shatilova & Stuchlik
X
P.remotifolioides
Shatilova & Stuchlik
X
P. variabilis
Stuchlik & Shatilova
X
P. venustaeformis
Stuchlik & Shatilova X
X
P. verus
(N. Mtchedlishvili) Shatilova &
Stuchlik X X X
P. vittatoides
Stuchlik & Shatilova
X
Adiantaceae
Adiantum
sp. X X X
Anogramma
sp. X X X
Pityrogramma
sp. X X X
Onychium sp.
X

21
1 2 3 4 5 6 7
Gleicheniaceae
Gleichniidites senonicus
Ross
X X X X
G. carinatus
(Bolch.) Bolchovitina X
Clavifera triplex
(Bolch.) Bolchovitina X
C. tuberosa
Bolchovitina X
Polypodiaceae
Polypodium
aureum
Linnaeus
X
X
P. pliocenicum
Ramishvili X X X
X
P. verrucatum
Ramishvili X X X
X
Polypodium
sp.
X
Pyrrosia
sp. X X X
Thyrsopteridaceae
Cibotium guriensis
Purceladze
X
C.
aff.
glaucum
(Sw.) Hr. & Arn.
X X
Dicksoniaceae
Dicksonia luculenta
Purceladze
X X
D. reticulata
Purceladze X X X X
D. spanditocincta
Purceladze X X X
D. unitotuberata
Purceladze X X X
X
D. antarctica
R. Braun
X X
Cyatheaceae
Alsophilla
sp. X X X
Hemitelia
sp. X
Cyathea
sp. X X X
X
Dennstaedtiaceae
Verrucatosporites histiopteroides
W.Krutzsch X X X
X
V. megafavus
Krutzsch
X X
V. favus
(R. Potonié) Tomson & Pflug X X
Davalliaceae
Microlepia
sp.
X
Verrucatosporites alienus
(Potonié)Thomson
& Pflug X
Gingkoaceae
Gingko biloba
Linnaeus X X X
Podocarpaceae
Dacrydium
aff.
pierrei
Hickel X
Dacrydium
sp. 1
X X
Dacrydium
sp. 2 X
Pinipollenites libellus
Potonié X X X
Podocarpidites nageiaformis
(Zaklinskaya)
Krutzsch X X
P
. aff.
neriifolius
D. Don
X X
P. eocaenicus
Krutzsch
X X
Pyillocladaceae
Phyllocladus
aff.
protractus
(Ward.) Pilg. X
Ph.
aff.
trichomanoides
D. Don
X X
Araucariaceae
Araucaria
sp. X X X
X
Pinaceae
Abies alba
Miller X X X
X
A. ciliticaeformis
N. Mtchedlishvili X X X
X
A. nordmanniana
(Steven) Spach X X X X X

22
1 2 3 4 5 6 7
Pinaceae
Cathayapollis millaya
(Sivak) Ziembinska-
Tworzydlo
X
C. potoniei
(Sivak) Ziembinska-Tworzydlo X X
Cathaya
aff.
argyrophylla
Chun & Kuang X X X
X
Cedrus deodara
Loud X X X X X
C. sauerae
N. Mtchedlishvili X X X
X
C. libani
Laws X X X
Cedripites lusaticus
Krutzsch X X
X
C. dacrydioides
Krutzsch
X
Keteleeria caucasica
Ramishvili X X X X X
Picea complanataeformis
N. Mtchedlishvili X X X
P. minor
N.
Mtchedlishvili X X X
X
P. orientalis
(L.) Link
X X X
Pinus
sp. X X X X X
Pseudolarix
sp. X X X
X
Tsuga canadensis
(L.) Carriére
X
T. pattoniana
Engelmann X X X
X
T. diversifolia
(Maxim.) Mast.
X X X
T. patens
Downie
X
T. shatilovae
Mchedlishvili
X
X
Sciadopityaceae
Sciadopitys
aff
.verticillata
Siebold et
Zuccarini X X X
Taxodiaceae
Cryptomeria japonica
D. Don X X X
X
Glyptostrobus
aff.
pensilis
(Stauton) Koch X X X
Metasequoia
aff.
glyptostroboides
Hu & Cheng X X X
X
Sequoia
aff.
sempervirens
(Lamb.) Endlicher X X X X X
Sequoiadendron
aff.
giganteum
(Lindley)
Buchholz X X X
Taxodium
aff.
distichum
(L.) Richard
X X X
X
Cupressaceae
Cupressus sp.
X
Juniperus
sp. X
Ephedraceae
Ephedra
aff.
distachya
Linnaeus X X X
Ephedra
sp.
X X
X
Distachyapites berheidensis
(Krutzsch)
Grabowska & Wazynska X X X
D. eocenipites
(Wodehouse) Grabowska &
Wazynska X
Welwitschia
sp. X

23
1 2 3 4 5 6 7
Myricaceae
Myrica conspiqua
Gladkova
X X
M. notabilis
Gladkova X X
M. pseudogranulata
Gladkova X X X
M. intermedia
Gladkova
X
Myricipites bituitus
(Potonié) Nagy X X X
M. peregriniformis
(Gladkova) Grabowska-
Wazynska X X X
Comptonia imperfecta
Gladkova X X X
X
C. grandis
Gladkova X X X
Juglandaceae
Carya aquatica
(Michx.) Nutt. X X X X X
C.
cordiformis
(Wangenh.) K. Koch X X X
X
C. ovata
(Mill.) K. Koch X X X
C
. aff.
tomentosa
(Lam.) Nutt.
X
C.
aff.
texana
Buckley X X X
C. spackmania
Traverse
X X X
Caryapollenites simplex
(Potonié) Raatz ex
Potonié X X X
Engelhardia
aff.
wallichiana
Lindl.
X X
X
Momipites punctatus
(Potonié) Nagy X X X
M. quietus
(Potonié) Nichols
X X
M. gracilis
(Potonie) Kohlman-Adamska
X
Juglans nigra
Linnaeus
X
J. cinerea
Linnaeus X X X
X
J. regia
Linnaeus X X X X X
J. sigillata
Dode X X X
Platycaryapollenites miocaenicus
Nagy X X X
X
Pterocarya pterocarpa
(Michx.) Kunth X X X X X
P. stenoptera
DC. X X X
P. rhoifolia
Siebold et Zuccarini X X X
Oreomunnea
sp.
X X
Salicaceae
Salixipollenites capreaformis
Planderova X
Betulaceae
Alnus
aff.
incana
(L.) Moench
X X
Alnipollenites metaplasmosus
(Potonié)
Potonié X
Betula
aff.
pumila
Linnaeus
X
Betulaepollenites costataeformis
Grabowska &
Wazynska X
Carpinus orientalis
Miller X X X
X
C. betulus
Linnaeus X X X X
C. caucasica
Grossheim
X X
X
Corylus
aff.
ferox
Wall.
X
C. maxima
Miller
X
X
Tricolpopollenites megagranifer
(Potonié)
Thomson & Pflug X

24
1 2 3 4 5 6 7
Fagaceae
Castanea sativa
Miller
X X X X X
C.
aff.
crenata
Siebold et Zuccarini X
Cupuliferoipollenites pusillus
(Potonié)
Potonié X
Castanopsis
sp. X X X
X
Faguspollenites minor
Nagy X
F. verus
Raatz X X X
F. vivus
Nagy X X X
F. subtilis
Nagy
X
Fagus orientalis
Lipsky X X X X X
Quercuspollenites granulatus
Nagy X X X X X
Q. rubroides
Kohlman -Adamska
X X
Ulmaceae
Celtipollenites intrastructurus
(Krutzsch &
Vanhoorne) Thiele-Pfeiffer X
Ulmipollenites undulosus
Wolf X X X
X
U. stillatus
Nagy
X X X
Zelkova carpinifolia
(Pall.) Koch X X X X X
Z. serrata
(Thumb.) Makino X X X
Eucommiaceae
Eucommia ulmoides
Oliver X X X
Caryophyllaceae Caryophyllaceae gen indet. X X X
X
Chenopodiaceae Chenopodiaceae gen. indet. X X X X X
Magnoliaceae
Magnolia megafigurata
(Krutzsch) Ramishvili X X
M. grandiflora
Linnaeus X X
Magnoliaepollenites neogenicus
(Krutzsch)
Mohr X X
Liriodendron tulipifera
Linnaeus X X X
Liriodendroipollis semiverrucatus
Krutzsch X X
Anonaceae
Anona
sp. X X X
Nymphaeaceae
Nupharipollis echinatus
Krutzsch
X
Nymphaea
sp.
X
Hamamelidaceae
Corylopsis
aff.
cordata
Merril X X X
C
. aff.
pauciflora
Siebold et Zuccarini
X X
Chunia
aff.
bucklandioides
H.T. Chang
X
Hamamelis
aff.
japonica
Siebold et Zuccarini X X X
Disanthus cercidifolius
Maximowicz X X X
D. cercidifolius
Maxim. var.
minor
Shatilova et
Mchedlishvili X
Distyliopsis
aff.
dunii
(Hamsley) P.K. Endress
X X

25
1 2 3 4 5 6 7
Hamamelidaceae
Distylium
aff.
rasemosum
Siebold & Zuccarini
X X
Fortunearia
aff.
sinensis
Rehd. & Wils.
X X
Fothergilla
aff.
major
(Sims
.
)
Loddiges
X
F
. aff.
gardenii
Murrey X X X
Eustigma
aff.
oblongifolium
Gardn. & Champ
X X
Molinadendron
sp.
X X
Parrotia persica
(DC) C. A. Meyer X X X
Parrotiopsis jacquemontiana
(Decne.) Rehd.
X
Sycopsis colchica
Ramishvili
X X
S
. aff.
sinensis
Oliv.
X X
Altingia
aff.
excelsa
Nor.
X
Liquidambar styraciflua
Linnaeus X X X
X
L. formosana
Hance
X X
L. orientalis
Mill.
X X
L. turgaica
Kuprianova
X X
Hamamelidaceae gen indet.
X X
Fabaceae
Lathyrus
sp. X
Acacia
sp. X X X
Rutaceae
Phellodendron
aff.
amurense
Ruprecht
X X
Anacardiaceae
Cotinus
aff.
coggygria
Scop. X X X
Hippocastanaceae
Aesculus
aff.
hippocastanum
Linnaeus X X X
Aquifoliaceae
Ilex colchica
Pojarkova
X X
X
Ilexpollenites propinquus
(Potonié) Potonié X
Staphyleaceae
Staphylea
colchica
Stev. X X X
X
Vitaceae
Parthenocissus quinquefolia
(L.) Planch.
X
Icacinaceae Icacinaceae gen. indet. X
Tiliaceae
Intratriporopollenites instructus
(Potonié)
Thomson & Pflug X X
In. schkopauensi
s Krutzsch
X
In. profenensis
Krutzsch
X
Tilia
aff.
cordata
Miller X X X
X
T. platyphyllos
Scop. X X X
Sterculiaceae
Sterculia
sp.1 X
Sterculia
sp.2
X
Sterculiaceae gen. indet. X X
Sapindaceae
Sapindus
sp.
X
Myrtaceae
Myrtaceidites myrtiformis
Simoncsics
X
Elaegnaceae
Slovakipollis elaegnoides
Krutzsch X
Boenhlensipollis hohli
Krutzsch
X
Onagraceae
Epilobium
sp. X
Corsinipollenites parvus
(Doktorowicz-
Hrebnicka) Slodkowska X X X

26
1 2 3 4 5 6 7
Alangiaceae
Alangium
aff.
kurzii
Craib
X
Alangiopollis eocaenicus
Krutzsch
X
A. simplex
Nagy
X
Alangiopollis
sp. X
Nyssaceae
Nyssa
aff.
sylvatica
Marshall X X X
Nyssapollenites pseudocruciatus
(Potonié)
Thiergart X X
Araliaceae
Aralia
aff.
hispida
Michx.
X
Brassaiopsis
sp. X X
Hedera colchica
C. Koch X X X
X
Araliaceae gen indet. X
Cornaceae
Cornaceaepollis microscabratus
Sladkowska &
Ziembinska-Tworzydlo X X X
Edmundipollis edmindi
(Pot.) Konz.,
Sladkowska & Ziembinska-Tworzydlo X
Sapotaceae
Sapotaceoidaepollenites folliformi
s (Pflug)
Nagy X
S. manifestus
(Potonié) Potonié, Thomson &
Thiergart X
S. hungaricus
(Kedves) Nagy
X X
Apocynaceae Apocynaceae gen. indet. X
Symplocaceae
Symplocos
aff.
paniculata
Wall. X X
Ericaceae
Ericipites callidus
(Potonié) Krutzsch
X
E. politus
(Mamczar) Grabowska
X
Caprifoliaceae
Lonicerapollis gallwitzii
Krutzsch X X X
Poaceae Poaceae gen. indet. X
Arecaceae Arecaceae sp. 1, 2. X
Angiosperm plant
of unknown
affinity
Fupingopollenites wackersdorfensis
(Thiele-
Pfeiffer) Liu Geng - wu X X X
F. minutus
Liu Geng - wu X

27
4. The Sarmatian to Pontian deposits of Georgia and their palynological
characteristics
4.1. Eastern Georgia
In Eastern Georgia Sarmatian deposits are known from Kartli and Kakheti regions
(Table II). In Kartli the Aragvi, Nadarbazevi and Uplistsikhe sections were studied. By fauna
the Aragvi and Nadarbazevi sections are the most complete outcrops. In Uplistsikhe section
the succession is represented by lower and middle Sarmatian.
In Kakheti, the Udabno, Gombori and Chachuna sections were studied, which also do
not comprise the full succession. In Udabno section the deposits of middle Sarmatian are
represented. More full is the Gombori section, where by data of foraminifers the deposits of
middle and upper Sarmatian are distinguished (Maissuradze, Koiava 2006). In Chachuna
section the uninterrupted upper Sarmatian is represented. No pollen data for later than
Sarmatian is available in eastern Georgia.
On the left bank of river Aragvi the section begins by argillo-arenaceous layers of lower
Sarmatian, dated by mollusks (Gruzinskaya et al. 1986). The middle Sarmatian is fully
represented in the section and contains rich assemblages of foraminifers described by Koiava
(2006). The big sizes of shells of Porosononion hyalinus (Bogdanowicz) point to the presence
of the upper layers of middle Sarmatian (Koiava et al. 2017). The upper Sarmatian is divided
into two parts. The lower part composed by argillo-arenaceous deposits with prevalence of
clays, while the upper part is represented by sandy-argillaceous layers (Buleishvili 1960).
In Nadarbazevi section the lower Sarmatian is represented by blue-grey sandy clays,
containing intercalations of sandstones and oolitic limestone with shells of mollusks. The
thickness of lower Sarmatian deposits here is nearly 100m (Buleishvili, 1960). The middle
Sarmatian is represented mainly by clays with interbeds of sandstones, limestone and
conglomerates in the upper part of horizon. The thickness of middle Sarmatian is about
300m. The lower part of upper Sarmatian or Natskhori Formation is built by parti-colored
clays with a thickness of about 200m. The upper part is represented by blue-gray clays and
coarse-grained sandstones. Natskhori Formation is overlain by red clays and thick
conglomerates, which by the composition of pebbles are similar to those of Shiraki
Formation, dated as Meotian-Pontian (Buleishvili 1960).

28
Because the Aragvi and Nadarbazevi sections are the most complete pollen record for
the Sarmatian of Eastern Georgia, their pollen zonation is taken as a reference for the other
sites studied. In all other Sarmatian sections, the above described pollen zones are presented
only partially. Only pollen zone III can be recognized in all sections. At whole the Sarmatian
deposits of Eastern Georgia are divided into 6 pollen zones.
Pollen zone I corresponds to the lower layers of lower Sarmatian. Characteristic is the
high percentage of woody plants. Among conifers the main components are Pinus and
genera of the family Podocarpaceae. Broad-leaved plants are represented by Myrica,
Comptonia, Carya, Engelhardia, Juglans, Castanopsis, Quercus, Nyssa, Araliaceae and
Arecaceae. The abundance of herbaceous plants is low.
Pollen zone II corresponds to the upper part of lower Sarmatian. From zone I it differs
by lower contents of warm-temperate conifers and broad-leaved plants, both warm-temperate
and subtropical. The number of herbaceous plants increases.
Pollen zone III covers the lower part of the middle Sarmatian. It is characterized by the
richest composition of pollen assemblages, main components of which are woody plants.
Pollen zone IV corresponds to the middle part of middle Sarmatian. It sharply differs
from the preceding pollen zone by lower contents of pollen grains of broad-leaved plants and
warm-temperate conifers. The main components of pollen assemblages are pine and herbs.
Pollen zone V comprises the upper layers of middle and lower layers of upper
Sarmatian. In the composition of pollen assemblages the number of broad-leaved plants and
warm-temperate conifers increase. The percentage of Pinus and herbaceous plants is low and
increase in upper part of zone.
Pollen zone VI corresponds to the upper Sarmatian. The main component of pollen
assemblages are herbs, Pinus and warm-temperate deciduous plants (Figs 2-4).
The Udabno, Gombori and Chachuna sections are situated further east in the region of
Kakheti (Fig.1).
The main part of Udabno section is built by deposits of middle Sarmatian, which
thickness is about 300m. The deposits of Udabno section are divided into pollen zones III, IV
and V. In general, the pollen assemblages are similar to those of Sarmatian deposits in Kartli.
But the assemblages of zone IV are somewhat richer by pollen grains of woody plants (Fig. 5).
By pollen data the Gombori section is divided into pollen zones III, V, VI. Most of the
section corresponds to pollen zone III (Fig. 6). The upper part of middle Sarmatian and

29
layers, dated as upper Sarmatian belong to pollen zones V and VI.
The Chachuna section is essential because it enables observing changes of pollen
assemblages through the uninterrupted and well constrained entire upper Sarmatian. Two
lithofacial unites, shallow marine and continental (Eldari Formation), were established in this
section. They conformably are overlain by deposits of Shiraki Formation, dated as Meotian -
Pontian (Bukhsianidze et al. 2018; Kokolashvili et al. 2018; Bukhsianidze, Koiava 2018). By
pollen data the upper Sarmatian in Chachuna section is divided into zones V and VI (Fig. 7).
Pollen zone V corresponds to shallow marine deposits. The main components of pollen
assemblages are warm-temperate conifers, subtropical and warm-temperate broad-leaved
plants. The proportion of herbaceous plants is low. Pollen zone VI corresponds to the
deposits of Eldari Formation. The pollen assemblages are dominated by Pinus, broad-leaved
warm-temperate plants and herbs, which percentages significantly increase. Subtropical
plants and ferns are fully absent.
Based on these pollen data, the Sarmatian deposits of Eastern Georgia can be divided
into 6 pollen zones (Fig. 8). The pollen record starts with pollen zone I, which is
characterized by a prevalence of pollen grains of woody plants. In pollen zone II the
abundances of herbs increase and pollen grains of woody plants, especially broad-leaved
subtropical plants are represented by lower percentages. Pollen zone III occurs in deposits of
the lower part of middle Sarmatian. It is the most widely distributed zone, which is
represented in almost all sections of Kartli and Kakheti: Aragvi, Nadarbazevi, Uplistsikhe,
Udabno and Gombori. Pollen zone III is characterized by the richest composition of pollen
assemblages, dominated by warm-temperate and subtropical plants. The number of pollen
grains of herbs is low. Quite different are the pollen assemblages of pollen zone IV (middle
part of middle Sarmatian), main components of which are herbaceous plants. The deposits of
this pollen zone are represented in sections Aragvi, Nadarbazevi, Udabno. Pollen zone V
corresponds to the upper part of middle and the lower part of upper Sarmatian. The
percentage of herbs decrease but as a whole the composition of pollen assemblages became
poorer. In Kartli pollen zone V is described in Aragvi and Naderbazevi sections whereas in
Kakheti it occurs in sections Udabno, Gombori and Chachuna. The deposits of pollen zone V
are overlain by pollen zone VI. Both pollen zones are represented best in sections of Kakheti
and especially in Chachuna section.

30
Fig. 2. Diagram of pollen abundance of main woody plants in Sarmatian deposits of Aragvi section (pollen zones I - VI)

31
Fig. 3. Diagram of pollen abundance of main woody plants in Sarmatian deposits of Nadarbazevi section (pollen zones I – VI)

32
Fig. 4. Diagram of pollen abundances of main woody plants in Sarmatian deposits of Uplistsikhe section (pollen zones I, III)

33
Fig. 5. Diagram of pollen abundance of main woody plants in Sarmatian deposits of Udabno section (pollen zones III, IV, V)

34
Fig. 6. Diagram of pollen abundance of main woody plants in Sarmatian deposits of Gombori section (pollen zones III, V, VI)

35
Fig. 7. Diagram of pollen abundance of main woody plants in upper Sarmatian deposits of Chachuna section (pollen zones V, VI)

36
Fig. 8. Summary diagram of pollen abundance of main woody plants and herbs in Sarmatian deposits of Eastern Georgia (pollen zones I-VI)

37
4.2. Western Georgia
In Western Georgia the deposits of Sarmatian are widely distributed. Most of sections
are studied by macro- and microfauna, including the Chanistskali section which is considered
as stratotypical (Maissuradze 1966, 1971; Muskhelishvili 1980).
In section Chanistskali the layers of Volhynian regiosubstage are characterized by
similar pollen assemblages. Therefore, pollen data for the whole lower Sarmatian is assigned
to one single pollen zone I-II, which by stratigraphical position is synchronous to pollen
zones I and II in Eastern Georgia.
The assemblages of pollen zone I-II are mainly composed of warm-temperate
conifers, i.e. Gingko, Dacrydium, Podocarpus, Cedrus, Keteleeria and representatives of
the family Taxodiaceae. The number of pollen grains of temperate conifers, especially of
genera Abies and Tsuga is low (Fig. 9). The percentage of herbaceous plants is low during
the whole Sarmatian.
Pollen zone III embraces most part of the middle Sarmatian deposits. The percentages
of subtropical broad-leaved plants and ferns increase. The pollen zone III is synchronous to
zone III in Eastern Georgia. In both regions the pollen assemblages are characterized by a
high diversity of subtropical and warm-temperate woody plants.
Pollen zone IV corresponds to the end of the middle Sarmatian. The number of pollen
grains of warm-temperate broad-leaved plants and temperate conifers increase here. Pollen
zone IV is synchronous to the same zone in Eastern Georgia, but the character of changes in
the composition of pollen assemblages is quite different. In Western Georgia the main
components of pollen assemblages remain to represent woody plants, only their diversity
becomes poorer. In Eastern Georgia the same pollen zone reflects an increase of pollen
grains of herbaceous plants.

38
Fig. 9. Diagram of pollen abundance of main woody plants in Sarmatian deposits of Chanistskali section (pollen zones I-II, III, IV)

39
In Western Georgia the upper Sarmatian has a very limited distribution and is
represented mainly by conglomerates (Buleishvili 1986). Thus, between middle Sarmatian
and Meotian a lack of pollen data embraces the whole Khersonian regiosubstage (pollen
zones V and VI in Eastern Georgia).
The late Sarmatian regression was followed by a transgression in the Meotian.
Deposits of this stage are widely distributed in Western Georgia, and are divided by pollen
data into pollen zones VII and VIII.
Pollen zone VII corresponds to lower Meotian. Main components of pollen
assemblages are warm-temperate conifers: Dacrydium, Podocarpus, Araucaria, Cedrus,
Keteleeria, and genera of the family Taxodiaceae. Broad-leaved plants, both warm-temperate
(Carya, Alnus, Fagus) and subtropical (Comptonia, Myrica, Engelhardia, Platycarya), are
represented nearly by equal number of pollen grains (Fig. 10).
Pollen zone VIII corresponds to upper Meotian. Compared to zone VII, it differs by
smaller percentages of pollen of warm-temperate conifers and subtropical broad-leaved
plants. The number of pollen grains of deciduous trees increases. But at the same time also
the percentage of pollen grains of family Hamamelidaceae increases (Fig. 11).
Pollen zone IX reflects the transition from the rich pollen flora of upper Meotian to
poor assemblages of the Eupatorian horizon, with a predominance of pine. Pollen zone X
corresponds to deposits of the Odessian horizon. The percentages of pollen grains of Pinus
decrease but in general the composition of flora became poorer (Fig. 12).

40
Fig. 10. Diagram of pollen abundance of main woody plants in lower Meotian deposits of Djapareuli section (pollen zone VII)

41
Fig. 11. Diagram of pollen abundance of main woody plants in lower and upper Meotian deposits of Gedjiri, Galidzga, Otapi and Gudau
sections (pollen zones VII and VIII)

42
Fig. 12. Diagram of pollen abundance of main woody plants in upper Meotian - lower Pontian deposits of Urta- Zana section
(pollen zones VIII, IX, X)

43
5. Main stages of vegetation development in Georgia during the Sarmatian to
Pontian
5.1. Eastern Georgia
In the development of vegetation on the territory of Eastern Georgia six stages are
distinguished during the Sarmatian. The basis of such division is the reduction of subtropical
forests and their full extinction, the decrease of forest formations in general, and the spread
of herbaceous plants and open landscapes (Figs 13 -16).
Stage I corresponds to the time of accumulation of the lower part of early Sarmatian.
During this time the subtropical and warm-temperate plants had an equal share in the
composition of vegetation. The area covered by temperate conifers was small and remained
nearly unchanged during the whole Sarmatian. The components of forests, which occupied
most part of dry land, were Gingko, Podocarpus, Dacrydium, Keteleeria, Cedrus, Pinus,
Myrica, Comptonia, Carya, Engelhardia, Platycarya, Ulmus, Zelkova, Quercus, Nyssa,
Symplocos, Araliaceae and Arecaceae. The lower layer of forests was built mainly by ferns
(Anemia, Lygodium, Gleichenia, Dicksonia, and representatives of the fossil genus
Verrucatosporites). The area covered by herbaceous plants was small.
Stage II is the time of accumulation of deposits of the upper part of lower Sarmatian.
In the composition of forests the part of subtropical and warm-temperate broad-leaved plants
decreased. The area covered herbaceous plants enlarged.
Stage III corresponds to the lower part of middle Sarmatian, and is the most
widespread on the territory of Eastern Georgia. In stage III the area covered by subtropical
communities received its maximum. The role of pine and herbaceous plants decreased. It was
the time of the climatic optimum, when temperature and humidity reached their peaks.
Stage IV corresponds to the middle part of middle Sarmatian. It abruptly differed from
the previous stage. The diversity of warm-temperate conifers and broad-leaved trees,
deciduous and subtropical, decreased and the area covered by ferns also retracted. The role of
pine and herbaceous plants increased. In southern Kakheti (section Udabno) simultaneously
with the spread of herbaceous plants shrubs became an important part of the plant
communities, specifically represented by the genera Juniperus and Myrica.

44
Fig. 13. Diagram of abundances of ecological groups of plants indicating the changes in the
major forest formations during the stages I-VI (Aragvi section)

45
Fig. 14. Diagram of abundances of ecological groups of plants indicating the changes in the
major forest formations during the stages I - VI (Nadarbazevi section)
During stage V the area covered by forests increased and their main components
became deciduous plants.
Stage VI is characterized by a sharp decrease in abundances of broad-leaved plants
and conifers, excluding pine. Probably, some woody plants whose pollen grains occur in
assemblages of upper Sarmatian deposits were represented by shrubs. Disanthus cercidifolius
Maximowicz var. minor Shatilova & Mchedlishvili can be cited as an example. The pollen
grains of this taxon are of much smaller size than those of the species Disanthus cercidifolius
Maximowicz. Probably, the plant developed the growth-type of a shrub under unfavorable
climatic conditions with low humidity. Similar small pollen grains of Disanthus are
described as D. kuprianova Ananova var. minor Ananova from Sarmatian deposits of Kerch
peninsula, and connected to the process of xerophytisation which expanded over most of the
Russian plain (Ananova, 1982).

46
Fig. 15 Diagram of abundances of ecological groups of plants indicating the changes in the
major forest formations during the stages III, IV, V (Udabno section)
So, on the territory of Eastern Georgia during the Sarmatian six stages are distin-
guished in development of flora and vegetation. The basis of such division is: the reduction
of subtropical plant areas and their fully extinction; decrease of forest formation and
distribution of herbaceous plants. One of the main factors influenced the evolution of flora
and vegetation of Eastern Georgia was the oscillation of humidity, began at the end of early
Sarmatian (stage II). The following stage III was the climatic optimum, when the
temperature and humidity achieved the highest indices, after which the humidity sharply
reduced and the area of warm-temperate conifers and broad-leaved plants decreased (stage
IV). In the following stage V, which corresponds partly to middle and upper Sarmatian, the

47
prevalence of woody plants was restored but the composition of flora became poorer. The
process of xerophytisation received the sharpest character in stage VI (upper part of upper
Sarmatian), after which the prevalence of open landscapes began in most of Eastern Georgia.
The woody plants were mainly the components of riparian forest and open woodlands. Most
of them probably were represented by shrubs.
Fig. 16. Diagram of abundances of ecological groups of plants indicating the changes in major
forest formations during the stages V, VI (Chachuna section)

48
5.2. Western Georgia
In Western Georgia the Sarmatian regiostage is represented mainly by deposits of
Volhynian and Bessarabian regiosubstages. Following the dynamics of development of plant
communities, in early and middle Sarmatia the following stages can be distinguished: I-II,
III, IV (Fig. 17).
Stage I-II corresponds to early Sarmatian (pollen zone I-II). The predominant plant
community was warm-temperate conifer forest. The area covered by temperate conifers was
small. Warm-temperate and subtropical broad-leaved plants had nearly the same share in the
composition of vegetation. Geochronologically and by its common character, which is a
vegetation poorer than those of the following stretch of middle Sarmatian, stage I-II of
Western Georgia can be compared with stages I and II in Eastern Georgia.
Stage III embraced most part of middle Sarmatian, when the communities of
subtropical broad-leaved plants and warm-temperate conifers occupied the greatest area. It
was a time of climatic optimum, which was synchronous to same phenomenon described
from Eastern Georgia.
Stage IV corresponds to end of the middle Sarmatian. Both, the areas covered by
warm-temperate broad-leaved plants and temperate conifers, increased. In Western Georgia
stage IV was the time of mass extinction of thermopilous plants. Still, the preserved
subtropical plants continued to occupy large areas and the general type of vegetation was not
changed. In comparison, the dynamics of vegetation development on the territory of Eastern
Georgia was quite different. The pollen assemblages of stage IV here reflect the dominance
of unstable climatic conditions, causing towards xerophytisation, as a result of which the area
covered by herbaceous plants sharply increased.

49
Fig. 17. Diagram of abundances of ecological groups of plants indicating the changes in the
major forest formations during the stages I-II, III, IV (section Chanistskali)
As noted above, in Western Georgia data for pollen zones V and VI, which would
correspond to the Khersonian substage in Eastern Georgia, are lacking. Therefore, the history
of vegetation continues with the Meotian-Pontian stages VII, VIII, IX and X (Figs 18, 19). In
Eastern Georgia these correspond to the Shiraki Formation.
Stage VII corresponds to the early Meotian. During this entire time subtropical and
warm-temperate broad-leaved plants covered nearly equal areas. Big territories were
occupied by warm-temperate conifers. In late Meotian (stage VIII) the correlation between
the main coenoses of forest was changed and significant increased the part of pine. It was the
beginning of a phenomenon, which took place in early Pontian, Eupatorian time (stage IX),
when pine became the dominant tree in the vegetation. In the later early Pontian, in Odessian
time (stage X), the whole character of vegetation changed. The middle and lower mountain

50
belts were occupied by warm-temperate conifers and broad-leaved plants. The area covered
by subtropical plants significantly decreased. Probably, they were distributed mainly in the
lower mountain belt. The temperate conifers formed a separate formation in the upper
mountain belt.
Fig. 18. Diagram of abundances of ecological groups of plants indicating the changes in the
major forest formations during the stages VII, VIII
(Gedjiri, Galidzga, Otapi and Gudau sections)

51
Fig. 19. Diagram of abundances of ecological groups of plants indicating the changes in the
major forest formations during the stages VIII, IX, X (Urta-Zana section)

52
6. Comparison of the environmental history of Eastern and Western Georgia
The record of the Neogene floras of Georgia begins in early Miocene (Chokrakian,
Karaganian and Konkian), and is studied paleobotanically by pollen data and macro-remains
(e.g., Avakov, 1979, 2008, 2010; Ramishvili 1982). These data document the long-term
development of flora and vegetation in the region.
Among the conifers, pine dominated in the assemblages of Chokrakian, Then, the
predominance of representatives of family Podocarpaceae began (Fig. 20), and the genera
Dacrydium and especially Podocarpus became the main components of conifer forests until
nearly the Pontian. The role of Abies, Picea and Tsuga, the genera being typical for younger
floras, was insignificant.
The vegetation of the lower mountain belt was formed by the most thermophilous
plants. Two formations, sclerophyllous typical of areas with relatively dry climate and the
moist-subtropical one, existed there. More mesophilous formations of deciduous plants of
warm-temperate and temperate climate inhabited hypsometrical higher levels. Such stands of
broad-leaved plants were probably mixed with conifers.
At a whole, the Miocene floras of Georgia and contemporaneous floras of the
Mediterranean (Nagy 1985, 1992) developed evenly without major perturbation, especially
in early and middle Miocene. Their development was not as monotonous as during the
Paleogene, but more homogeneous than the floras of the periods to follow, when increasing
differentiation led to the formation of separate phytogeographical provinces.
As it was already noted, the end of middle Sarmatian was the turning-point in the
Neogene history of Caucasus. The severe paleogeographical changes, the forming of high
mountain relief; the division of the territory of Georgia into two geographical regions, and
the isolation of Colchis refuge, were confined to this time.
On basis of the given about data, it is possible to conclude that during the early and
middle Sarmatian forests were the main plant communities on the territory of Eastern Georgia,
in spite of temporary reductions of forested areas in some stretches of time (stages II and IV).

53
Fig. 20. Summary diagram of pollen abundances of main woody plants in the Early Miocene Chokrakian, Karaganian and Konkian
deposits of Georgia (by data of Ramishvili 1982)

54
The littoral and lower mountain belt was occupied mainly by thermospilous and
hygrophilous plants: Myrica, Engelhardia, Platycarya, Castanopsis, Magnolia, Alangium,
the representatives of families Araliaceae, Sapotaceae, Lauraceae, Icacinaceae, and
Arecaceae. The lower layer of forest was built by subtropical ferns: Lygodium, Anemia,
Gleichenia, Hemitelia, and Dicksonia.
Hypsometrically higher, deciduous plants (Juglans, Carya, Castanea, Quercus, Carpi-
nus, Ulmus, Zelkova, and Tilia) and warm-temperate conifers (Dacrydium, Podocarpus,
Keteleeria, and Cedrus) were distributed. Temperate conifers played a minor role in the
composition of vegetation.
In Sarmatian in both regions of Georgia the floristic composition of conifers was nearly
similar but the abundances of separate taxa differed. In Western Georgia the warm-temperate
genera were predominant, while in Eastern Georgia representatives of genus Pinus prevailed.
Rich macrofossil material from lower and middle Sarmatian deposits of Eastern
Georgia indicates that Magnolia, Potagonium, Myrtus, Apocynophyllum and representatives
of the family Lauraceae were the prevalent components of the vegetation.They represent a
subtropical flora, in which taxa of hard-leaved (sclerophyllous) and moist-subtropical
formations were represented nearly by equal parts. In the hard-leaved formation the role of
Mediterranean elements was high (Chelidze 1972, 1979, 1987).
Paleobotanical material from synchronous deposits of Western Georgia indicates the
prevalence of moist-subtropical forests. The coastal plains were covered by marsh forests
with Taxodium, Alnus, Carya, Nyssa, Liquidambar and other plants.
The vegetation of lower mountain zones was represented by evergreen forests with
predominance of Lauraceae and Myrsinaceae. Here Castanopsis, Symplocos, Fothergilla,
some Araliaceae and ferns Lygodium, Dicksonia, Cibotium were also growing. A large area
was covered by thermophilous deciduous plants, like Ocotea and Persea, which now occur
in Central and Southern America, Mediterranean regions and Southern Asia (Kolakovsky &
Shakryl 1976).
Warm-temperate and temperate plants occurred in varied assemblages, distributed on
elevated dry land and possibly in cooler valleys as well. Carpinus, Castanea, Juglans,
Cathaya, Cedrus, Keteleeria, Podocarpus, Dacrydium belonged to this ecological group.
Hemixerophytes played a minor role and included Arbutus elegans Kolakovsky, Celtis
magnifica Kolakovsky, Smilax aspera Linnaeus, Quercus pseudorobur Kováts, and Pinus

55
paraeuxina Kolakovsky. The upper mountain belt was occupied by Abies, Picea and Tsuga.
However, judging from the composition of pollen assemblages, the role of these plants in the
coenosis of early and middle Sarmatian was insignificant.
Based on pollen data the difference between flora of Eastern and Western Georgia
took place already from the beginning of Sarmatian, when signs of lower humidity started to
be reflected in the flora of Eastern Georgia. The differences shown in both, in the floristic
compositions and in the development of vegetation. As noted above, in Western Georgia
during whole early and middle Sarmatian forests were the main plant communities, which
developed more slowly and quite than in Eastern Georgia. The evolution was mainly directed
towards the extinction of subtropical elements. But this process went without sharp changes
in the general structure of vegetation.
In Eastern Georgia the reduction of areas covered by woody plants and the expansion of
herbaceous coenoses began in early Sarmatian (stage II). One of the main factors that
influenced the evolution of flora and vegetation of Eastern Georgia was the oscillation of
humidity. The following stage III was the climatic optimum, when temperature and humidity
achieved their peaks. After that, humidity sharply declined and the area covered by warm-
temperate conifers and broad-leaved trees decreased. The area of herbaceous plants increased
(stage IV). In stage V, which corresponds to the transition from middle to upper Sarmatian, the
prevalence of woody plants was restored but the composition of flora became poorer. The
process of xerophytisation reached its most pronounced character in stage VI, when the
prevalence of open landscapes began in most parts of Eastern Georgia (Fig.21). Woody plants
became mainly components of riparian forests and open woodlands. Most of them probably
were represented as shrubs. Similar changes took place in south-eastern Europe, Ukraina,
Turkey, Bulgaria, and the northern part of Black Sea (Taman peninsula). Palaeobotanical data
from Bessarabian and Khersonian deposits of these regions suggest the appearance of open
landscapes and the development of xerophytic plant communities. These initial open
landscapes were probably of the so-called parkland type, where a mosaic of open landscapes
and smaller or larger stands of trees exist (Ivanov et al. 2010; 2014; Razumkova, 2012).
The importance of humidity is also evident in the spatial differences in the pollen
signals. In all sections studied in Kartli, the pollen assemblages of lower and middle Sar-
matian deposits reflect similar vegetation, both by composition and structure, which
developed under conditions of decreasing humidity. In Kakheti however, conditions were

56
slightly different, especially in the region of Gombori (Fig. 22). Here, during most parts of
the Sarmatian the main components of vegetation were subtropical plants. Thus, it is possible
to conclude that in early and middle Sarmatian the climate of Eastern Georgia was generally
subtropical, but humidity was not equally distributed in the region.
Fig. 21. Summary diagram of abundances of ecological groups of plants
indicating the changes in the major forest formations during the
Sarmatian of Eastern Georgia

57
Fig. 22. Diagram of abundances of ecological groups of plants indicating the changes in the
major forest formations during the Sarmatian on the territory of Kakheti (section Gombori)
The vegetation changes, which took place after the Sarmatian, are possible to judge by
pollen data of section Vake (Western Georgia), where the middle Sarmatian is overlain by
lower Meotian (Badzoshvili 1979). Here, between middle Sarmatian and Meotian warm-
temperate conifer forests retracted, deciduous trees became a more important part of broad-
leaved forests, while temperate conifer forests became more stable (Figs 23, 24).

58
Fig. 23. Diagram of pollen abundance of main woody plants on Sarmatian – Meotian boundary (pollen zones IV, VII)
in Western Georgia (section Vake)

59
Fig. 24. Diagram of abundance of ecological groups of plants indicating the changes in the
major forest formations on Sarmatian – Meotian boundary
(stages IV, VII ) in Western Georgia (section Vake)
Judging by habitat conditions of recent plant-equivalents, the vegetation cover on the
territory of Western Georgia in the Meotian can be restored. In the lowlands, big areas were
occupied by communities analogous to recent swamp forests, distributed today in regions
with subtropical humid climate in South-Eastern North America. Those swamp forests were
composed of Taxodium, Carya, Nyssa, Liquidambar, Fothergilla and other plants. Macrobo-
tanical data provide further evidence of the occurrence of Glyptostrobus, Dalbergia, Diospy-
ros, Eugenia, which all are characteristic for coastal swamp forests of South-Eastern China
and Viet-Nam. The valleys of rivers were covered by Pterocarya, Ulmus and Alnus, which
also spread up to mountains.

60
The lower belt of mountains was occupied by subtropical forests composed of
evergreen and deciduous plants, including representatives of families Lauraceae, Fagaceae,
Araliaceae and Hamamelidaceae. In the middle belt the communities of subtropical conifers
(Araucaria, Cedrus, Cryptomeria, Cathaya, Keteleeria, Podocarpus, Dacrydium) and
deciduous plants (Quercus, Carpinus, Juglans, Fagus, Zelkova and Tilia) were distributed.
The ferns Dicksonia, Cyathea, Pteris, Polypodium and Cryptogramma were connected with
both formations. The genera Anemia, Lygodium, Cibotium and Gleicheniа, typical represen-
tative of Sarmatian floras became more rare components of forest communities.
The upper mountain belt was occupied by Abies, Picea and Tsuga. After the
Sarmatian the area covered by these plants increased and temperate conifer forests became a
distinct zonal type of vegetation.
On the basis of all data, the climate of Colchis during Meotian can be interpreted as
supposedly subtropical. During the whole time subtropical plants remained a major
component of the vegetation. Although the abundance of Podocarpus decreased, it continued
to be a main component of the warm-temperate forest. Not less significant is the presence of
Araucaria, which now is a component of subtropical rain forest in America, Australia and on
islands of Pacific (Walter 1968).
Pollen assemblages of the Eupatorian horizon show the transition from a rich and
diverse vegetation of Meotian to poor communities with a predominance of pine that
probably was connected with changes in humidity. After this phase of lower humidity
during the Eupatorian, in Odessian the general character of vegetation was restored but the
floristic diversity was considerably reduced. So, the vegetation changes during the
Eupatorian in Western Georgia can be considered as the transition between typical
Miocene and Pliocene floras.
In summary it is possible to conclude that after the middle Sarmatian, the vegetation of
Eastern and Western Georgia developed independently under the influence of different
climatic conditions. While in the East a certain degree of xerophytization influenced the
vegetation already from late Sarmatian onward, in the West stable humid conditions
prevailed. Only after Meotian a stepwise reduction of the floristic diversity began. This
asynchronous development of vegetation can be connected to orogenic movements, which
led to important paleogeographical changes, mainly the draining of the Transcaucasus strait,
the division of the territory of Georgia into two regions, and isolation of the Colchis.

61
7. Comparison of floristic stages with phases of foraminiferal development on the
territory of Georgia during the Sarmatian to Pontian
The correlation of floristic stages with phases of development of foraminifers on the
territory of Eastern Georgia was conducted on the basis of data from Aragvi and
Nadarbazevi sections, which were studied by both methods (Koiava 2006; Shatilova et al.
2008; 2009; Shatilova et al. 2012).
In the lower part of lower Sarmatian the assemblages of foraminifers are characterized
by genera Quinqueloculina, Sinuloculina, Varidentella, Affinetrina, Nonion, Elphidium,
Porosononion, and Ammonia. The genera Bolivina, Discorbis, Bulimina, Cibicides and
Fissurina are relatively rare. Their morphology does not differ noticeably from their middle
Miocene ancestors. The lower part of early Sarmatian is distinguished as the layers with
Varidentella reussi. They reflect the first phase of development of foraminifers.
Assemblages of foraminifers from the upper part of early Sarmatian are characterized
by reduced diversity because of the disappearance of Discorbis, Bulimina and Cibicides.
Nevertheless a large number of genera survived and adapted to the new environment. They
are characterized by strong intraspecific variability and a potential for speciation. The upper
part of early Sarmatian is distinguished as the layers with Elphidium aculeatum. They reflect
the second phase of development of early Sarmatian foraminifers.
By data on foraminifers, the middle Sarmatian is divided into three parts. The first is
characterized by assemblages very different from those observed in the early Sarmatian. The
new genera, such as Dogielina, Meandroloculina, Sarmatiella, contributed a considerable
number of species. Foraminifers of these assemblages are characterized by comparatively
large sizes.
The second part of the middle Sarmatian comprises the richest assemblages of
endemic foraminifers with the highest numbers of individuals and new species (Maisuradze
et al., 2009), and test sizes reaching their maximum.
The third part of middle Sarmatian is distinguished by a decrease in the abundances of
genera, species and individuals. Because of the worsening of bionomic conditions only very
few representatives of most euryhaline families survived: Elphidium, Porosononion,
Ammonia and very rarely Affinetrina and Varidentella. Among them, Porosononion is
characterized by large sizes and additional ornamentations on very coarse walls of tests.

62
Biostratigraphically the three parts of middle Sarmatian are defined by the occurrences
of Affinetrina voloshinovae in the lower layers, Porosononion aragviensis in the middle, and
Porosononion hyalinus in the upper layers.
Some species of Elphidium, Nonion, Porosononion and Ammonia of middle Sarmatian
also occur in upper Sarmatian. These taxa show deformations, irregular cameras or immature
shell development, and are presented only in some parts of the late Sarmatian basin. These
anomalies of shells point to the existence of not optimal conditions for foraminifers.
In total, in the history of development of foraminifers in Sarmatian basins of Eastern
Georgia five phases are distinguished, which cover the lower and middle Sarmatian. They
reflect different stages of abundances and diversity and thus environmental conditions. No
information on upper Sarmatian environments is available due to the lack of foraminifers in
the low salinity basin at that time. These five pahses can be compared to the stages of
development of flora and vegetation during the Sarmatian.
The vegetation stages I and II (pollen zones I, II) correspond to the deposits of lower
Sarmatian. On this basis they can be correlated with the first and second phases of development
of foraminifers. The conditions of this time were not optimal, neither for flora nor for
foraminifers (Fig. 25).
During the middle Sarmatian three stages (phases) are distinguished in the
development of flora and foraminifers, but stratigraphically they do not coincide completely.
The appearance of new taxa and increase of test sizes served as the common sign of the first
and second phases of development of foraminifers in middle Sarmatian. In the second phase
these phenomena were more distinct, indicating the existence of optimal conditions. We
correlate both microfaunistic phases with vegetation stage III, which reflects the conditions
of a climatic optimum.
The third phase of development of middle Sarmatian foraminifers (the layers with
Porosononion hyalinus) can be correlated with vegetation stage IV, when the impoverishment
of Sarmatian flora took place. It was the time of sharp changes in the composition of the marine
and terrestrial biocenosis and in the conditions of their existence.

63
Fig. 25. Comparison of floristic stages and phases of development of foraminifers during the
Sarmatian of Eastern Georgia

64
As mentioned above, stages V and VI cannot be compared with the foraminiferal re-
cord. Only after the late Sarmatian regression, which was the cause of this lack, a transgre-
ssion in Meotian restored the connection between Black Sea and Mediterranean regions,
which promoted the penetration of marine fauna into the Eastern Paratethys. At the begin-
ning of the Meotian, relatively stenohaline species of foraminifers colonized the basin, and
new stage in the development of Miocene fauna began.
Within the Black Sea-Caspian basin, the lower Meotian deposits of Western Georgia
are distinguished by the richest composition of foraminifers (Maissuradze 1988; Maissura-
dze, Koiava 2011; Shatilova et al. 2019). Quinqueloculina seminulum maeotica Gerke, Sinu-
loculina ex gr. consobrina (d’Orbigny), Cycloforina aff. gracilis (Karrer), Elphidium macel-
lum (Fichtel & Moll), E. feodorovi (Bogdanowicz), and Ammonia ex gr. beccarii (Linnaeus)
had wide distributions. Species with narrow distributions were Quinqueloculina akneriana
d’Orbigny, Q. maeotica Maissuradze, Q. vermicularis Karrer, Triloculina aff. intermedia
Karrer, and Miliolinella circularis (Bornemann).
In early Meotian basins of Western Georgia there probably were specific conditions
which promoted the origin of endemic species and subspecies, like Quinqueloculina iberiae
Bogdanowicz, Q. disparilis galidzgensis Bogdanowicz, Q. djanelidzeae Maissuradze,
Hauerina iljinae Bogdanowicz. Due to the interruption of oceanic connection the salinity of
the late Meotian Sea decreased and subsequently a large number of marine foraminifers
disappeared. They were replaced by euryhaline species, predominantly Ammonia beccarii
liliae Popchadze, and ostracods.
So, during the Meotian two phases can be distinguished in the development of
foraminifers in Western Georgia corresponding to the Bagerovian and Akmanaian regional
substages. In the early Meotian complex, stenohaline species of foraminifers dominated, and
in the late Meotian euryhaline species were abundant.
As noted above, in some sections of Western Georgia (Otapi, Urta-Zana) the transition
between upper Meotian and Pontian is gradual. The lower part of Pontian is represented by
the Eupatorian horizon, which is characterized by rare occurrences of Quinqueloculina
seminulum maeotica Gerke, Elphidium ex gr. ponticum Dolgopolskaya & Pauli,
Porosononion ex gr. subgranosum (Egger), Nonion aff. matagordanus Kornfeld, Ammonia

65
beccarii (Linnaeus). The history of development of late Cenozoic foraminifers ends with the
early Pontian, which is the Eupatorian (Maissuradze et al. 2013).
The Pontian deposits, which cover the Eupatorian horizon, accumulated in closed
basin with low salinity. In these deposits foraminifers are absent. This process of isolation
continued until the end of the Miocene and, therefore, foraminifers are absent in post-Pontian
basins of the Eastern Paratethys. The biotopes of these basins were inhabited by brackish
ostracods, mollusks and other euryhaline organisms. Endemic species of mollusks and
ostracods that immigrated from adjoining regions are dominant (Nevesskaya et al. 1986).
The complete absence of foraminifers in post-Miocene deposits can be explained at
first by a sharp decrease of salinity. Except salinity also the strong competition with ostrac-
ods must be taken into account, organisms of higher organization and more enduring of low-
salinity conditions than foraminifers. Both occupy the same biotope and use the same food.
Ostracods were numerous and widely distributed in the basins of the Pliocene.
Another factor promoting the extinction of foraminifers was unstable climatic conditi-
ons, especially the worsening of climate in Eupatorian time. Consequently, the combination
of decrease in salinity, absence of a stable and warm environment, and shortage of food
created unfavorable conditions for foraminifers.

66
8. Conclusion
The lower and middle Sarmatian deposits of Georgia accumulated in Trancaucasus
intermountain Area, which was entirely covered by sea. In the late Sarmatian this depression
was transformed into dry land with two basins, the Kura Bay, which was connected with the
Caspian Sea, and the Rioni Bay, which was part of Black Sea. Since the upper part of late
Sarmatian and during most of the following time, the territory of Georgia belonging to Kura
bay was dry land.
To the West, in the Trans-Black sea depression area of Rioni Bay, the accumulation of
marine deposits continued until the end of the Pleistocene. Today this is a stratotypical region
of the Eastern Paratethys, where the late Cenozoic deposits are fully represented and are well
characterized by marine fauna.
The following Sarmatian sections were studied: Aragvi, Nadarbazevi, Uplistsikhe,
Udabno, Gombori, Chachuna (Eastern Georgia), Chanistskali, Vake (Western Georgia).
Marine deposits of Meotian and Pontian age were studied in sections of Western
Georgia: Vake, Djapareuli, Gedjiri, Otapi Galidzga, Gudau, and Urta-Zana.
For all sections studied by pollen data two types of diagrams are prepared. The first
one shows changes of pollen abundances of the main woody plants for each section. The
diagrams are divided into pollen zones, distinguished on the basis of pollen assemblages
found within them. In this case the pollen zones are considered as stratigraphical unit.
The second type of diagram reflects the abundance of ecological groups of plants
indicating the changes in the major forest formations on territory of Georgia during the
Sarmatian, Meotian and lower Pontian.
The comparison of pollen data from Eastern and Western Georgia revealed differences
and similarities in the composition of flora and the dynamics of vegetation. In Eastern
Georgia during the Sarmatian six stages (I-VI) were distinguished in the development of
vegetation, among which stages I-IV correspond to stages (I-II, III, IV) in Western Georgia.
In both regions the time of climatic optimum (stage III) was synchronous, after which in
Eastern Georgia the area covered by forests sharply decreased while in the western part of
the country the development of forest vegetation continued (stage IV).
Consequently, after the middle Sarmatian, the vegetation of Eastern and Western
Georgia developed independently under the influence of different climatic conditions. This

67
phenomenon can be connected with orogenic movements, which led to important
paleogeographical changes, the draining of the Transcaucasus strait, the division of the
territory of Georgia into two regions and isolation of the Colchis. After the Sarmatian,
marine conditions were preserved only in Western Georgia.
In Western Georgia the Sarmatian is overlain by Meotian. There are distinct
differences between the vegetation cover during these two stages of Late Miocene. The
boundaries between coenosis of separate mountain belts became more distinct, temperate
conifer forests expanded and deciduous warm-temperate broad-leaved plants became
important parts of the polydominant lowland forests.
During the Meotian two main stages are distinguished. The early Meotian was the
time when subtropical plants still dominated the composition of vegetation, whereas in the
late Meotian warm-temperate communities spread and covered nearly the same amount of
space. These stages correspond to the two regional substages: Bagerovian and Akmanaian.
The Sarmatian deposits of Georgia are characterized by frequent changes of facies,
which was the main factor that influenced the composition of fauna of foraminifers. In
Eastern Georgia during the Sarmatian several phases in the development of foraminifers can
be distinguished.
The lower Sarmatian is characterized by different assemblages, which depended on
lithology and depth of the basin. The lower Sarmatian deposits are divided into two parts,
layers with Varidentella reussi and layers with Elphidium aculeatum.
The lower Sarmatian is overlain by the Middle Sarmatian, which is divided into three
parts: the layers with Affinetrina voloshinovae, the layers with Porosononion aragviensis and
the layers with Porosononion hyalinus.
The middle part of middle Sarmatian is a time of flourishing of foraminifers. The sizes
of foraminifers increase significantly and the number of endemic forms achieves a
maximum.
To the end of the middle Sarmatian, the faunal diversity of foraminifers sharply
impoverished. Orogenic movements, which began at the end of middle Sarmatian, achieved
a maximum in the late Sarmatian, and in most of Georgia a continental regime was
established. In the late Sarmatian foraminfers were practically absent.
The late Sarmatian regression was followed by a transgression in the Meotian. The
connection between the Black Sea and the Mediterranean regions was restored and relatively

68
stenohaline species of foraminifers colonized the basin. A new phase of development of
Miocene fauna began.
In Meotian two phases can be distinguished in the development of foraminifers in
Western Georgia. In the early Meotian, stenohaline species of foraminifers dominated the
assemblages and in the late Meotian euryhaline species were abundant.
During the Sarmatian and Meotian two common turning-points in the evolution of
marine and terrestrial biocoenosis can be distinguished. The relation of them to the same
stratigraphical boundaries permits to presume some significant changes in the environments,
which led to these bioevents.
On the territory of Eastern Georgia one such boundary is the transition between the
middle and upper Sarmatian, when significant changes are observed in the composition of
the marine biocoenosis, owing to a further freshening of the basin. The same level was a
turning-point in the development of terrestrial flora as well with its distinct opening of the
landscape and increasing xerophytisation.
The second boundary is documented in Western Georgia. It is the transition between
the Meotian and Pontian, i.e. the Eupatorian time, when the impoverishment in the
composition of marine organisms and of vegetation took place in parallel. The pollen
assemblages of Eupatorian show the transition from a rich and diverse flora of upper
Meotian to poor assemblages with a predominance of pine. These changes in the
composition of vegetation probably were connected with a decrease in humidity.
Subsequently, during Pontian the main character of vegetation was restored but the floristic
diversity became poorer.
As for the microfauna, in Meotian predominate fossils are foraminifers. Ostracods are
represented only by single tests. This picture changes for Eupatorian deposits, in which the
abundance and specific diversity of ostracods considerably increased, while the number of
foraminifers and their specific composition diminished. The assemblages were composed of
euryhaline species, which differ morphologically from earlier taxa widely distributed in the
late Meotian. The history of development of late Cenozoic foraminifers ends with the early
Pontian, the Eupatorian.
So in the Black Sea region the Eupatorian horizon (the lower part of Pontian) can be
considered as the level, after which the typical Miocene marine and terrestrial biocoenosis
were replaced by Pliocene one.

69
საქართველოს ტერიტორიის ეკოლოგიური გარემო
გვიანმიოცენურ დროში
ფორამინიფერებისა და პალინოლოგიური ანალიზის საფუძველზე
რეზიუმე
დღევანდლამდე ჩვენი ცოდნა საქართველოს სარმატული და მეოტური
ფლორისა და მცენარეულობის შესახებ ეფუძნებოდა მხოლოდ მაკრობოტანიკურ
კვლევებს. ვფიქრობთ, ჩვენს ხელთ არსებულმა მდიდარმა პალინოლოგიურმა მა-
სალამ აღმოსავლეთი და დასავლეთი საქართველოს სინქრონული ნალექებიდან
საშუალება მოგვცა მეტ-ნაკლებად ამოგვევსო ეს ხარვეზი.
საქართველოს ტერიტორიაზე, ისევე როგორც მთელს აღმოსავლეთ პარატე-
თისში, სარმატული რეგიოსართული იყოფა სამ რეგიოქვესართულად: ვოლი-
ნური, ბესარაბული და ხერსონული. სარმატულის ამგვარი დაყოფა უკავშირდება
ნალექების ლითო- და ბიოფაციესურ ცვალებადობას, რაც განსაკუთრებით კარგა-
დაა გამოხატული აზოვისა და შავიზღვისპირეთის რეგიონებში. აღნიშნულ ტერი-
ტორიაზე ქვედა- და შუასარმატული ნალექების დაგროვების არეალს ამიერკავკა-
სიის მთათაშუა არე წარმოადგენს.
ძირულის შვერილის უმეტესი ნაწილი და დასავლეთით მოსაზღვრე
ოკრიბის ამაღლება ადრესარმატულში ხმელეთს წარმოადგენდა, რომლის დასა-
ვლეთით ხდებოდა რიონის, ხოლო აღმოსავლეთით - მტკვრის მთათაშუა რო-
ფების ფორმირება. ტექტონიკური მოძრაობის ინტენსივობა მაქსიმუმს შუასარ-
მატულის ბოლოს აღწევს, ხოლო გვიანსარმატულის დასაწყისიდაან კი მტკვრის
მთათაშუა როფის უდიდეს ნაწილში კონტინენტური რეჟიმი ყალიბდება. გვიანსა-
რმატულიდან ადრეაღჩაგილურამდე აღმოსავლეთი საქართველოს დეპრესია
(მტკვრის მთათაშუა როფი) მთლიანად კონტინენტური ნალექების აკუმულაციის
არეალს წარმოადგენდა. რაც შეეხება დასავლეთ საქართველოს (რიონის მთათა-
შუა როფი), აქაც დეპრესიის უმეტეს ნაწილში კონტინენტური რეჟიმი იყო გაბა-

70
ტონებულია. გამონაკლისია მხოლოდ შავიზღვისპირეთი, სადაც ზღვიური ნალე-
ქების დაგროვება პლეისტოცენის ბოლომდე გრძელდება. გვიანკაინოზოური
ნალექები აქ სრულადაა წარმოდგენილი, კარგადაა დათარიღებული ფაუნით და
დღესდღეისობით აღმოსავლეთი პარატეთისისთვის სტრატოტიპულ რეგიონს
წარმოადგენს.
საქართველოს ტერიტორიაზე ზღვიური მეოტური ნალექები ცნობილია
მხოლოდ დასავლეთ საქართველოში, სადაც ისინი ფაუნის მონაცემებზე დაყრდ-
ნობით იყოფა ორ რეგიოქვესართულად - ბაგეროვული და აკმანაიური.
მეოტურს სტრატიგრაფიულად ზევით, ხშირად კუთხური უთანხმოებით,
აგრძელებენ პონტური ნალექები. გამონაკლისს წარმოადგენს - ოტაფისა და ურთა
- ზანას ჭრილები, სადაც გადასვლა მეოტურსა და პონტურს შორის თანდათანო-
ბითია. აღნიშნულ ჭრილებში პონტურის ქვედა დონე წარმოდგენილია ევპატო-
რიული ჰორიზონტით, რომელიც შეიცავს ღარიბ ევრიჰალურ ფაუნას: მოლუს-
კებს, ფორამინიფერებს, ოსტრაკოდებს (Taktakishvili et al. 2018; Maissuradze et al.
2013).
ამრიგად, კვლევის პროცესში ჩვენს მიერ შეგროვილი და შესწავლილია პა-
ლეონტოლოგიური მასალა ქვედა სარმატულიდან -ქვედა პონტურის ჩათვლით
(ევპატორიული ჰორიზონიტი), რაც ქრონოსტრატიგრაფიულად 12.7 - 6.5Ma
ინტერვალს მოიცავს.
მასალის დამუშავების შედეგად შემოთავაზებულია ორი ტიპის პალინო-
ლოგიური დიაგრამა: პირველი ასახავს ფლორის შემადგენლობას, ხოლო მეორე
მცენარეულობის განვითარების ეტაპებს. პალინოლოგიური დიაგრამების შედა-
რებამ გამოავლინა როგორც მსგავსება, ასევე განსხვავება აღმოსავლეთი და დასავ-
ლეთი საქართველოს ფლორის შემადგენლობასა და მცენარეულობის განვითა-
რებაში
დასავლეთ საქართველოს ტერიტორიაზე მთელი გვიანკაინოზოურის განმა-
ვლობაში გაბატონებული იყო ტყის ფორმაცია. იცვლებოდა მხოლოდ ცალკეული
ცენოზის შემადგენლობა. განსხვავებულია მცენარეულობის განვითარების

71
დინამიკა აღმოსავლეთი საქართველოს ტერიტორიაზე. ამ რეგიონის ადრე-
შუასარმატულის პალინოლოგიური კოპლექსები მიუთითებენ არასტაბილური
კლიმატის არსებობაზე. შუასარმატულის ბოლოს ადგილი ჰქონდა ტყის არეალის
შემცირებას და ბალახოვანი ასოციაციების გაფართოებას. აღმოსავლეთი საქარ-
თველოსგან განსხვავებით, დასავლეთ საქართველოში ტყის ფორმაციების განვი-
თარების პროცესი გაგრძელდა. ამავე დროს, მიუხედავად განსხვავებისა, ორივე
რეგიონში კლიმატის ოპტიმუმი შუასარმატულ პერიოდში დაფიქსირდა.
შუასარმატულის ბოლოს ორივე რეგიონში ხდება მკვეთრი პალეოგეოგრა-
ფიული ცვლილებები: მაღალმთიანი რელიეფის სწრაფი ფორმირება, საქართვე-
ლოს ტერიტორიის ორ იზოლირებულ გეოგრაფიულ ნაწილად გაყოფა და კოლ-
ხეთის რეფუგიუმის ჩამოყალიბება.
სარმატულისა და მეოტურის განმავლობაში, აღმოსავლეთი და დასავლეთი
საქართველოს ტერიტორიაზე ზღვის და ხმელეთის ბიოცენოზების განვითარება-
ში გარდატეხის ეტაპი დგება სხვადასხვა დროს. აღმოსავლეთ საქართველოში ეს
არის საზღვარი შუა- და გვიანსარმატულს შორის. ამ დროს, ზღვიურ ბიოცენოზე-
ბში დიდ ცვლილებებს ჰქონდა ადგილი, რაც დაკავშირებული იყო აუზის გამტკ-
ნარებასთან. დროის ამავე მონაკვეთში მოხდა გარდატეხა მცენარეულობის განვი-
თარებაშიც.
დასავლეთ საქართველოში გარდატეხის ეტაპი მოდის მეოტურისა და პონ-
ტურის საზღვარზე (ევპატორიული დრო), როცა მდიდარი და მრავალფეროვანი
მცენარეული საფრის ნაცვლად მთიან რეგიონებში თითქმის ყველა სარტყელში
განვითარდნენ ცენოზები, რომლებშიც ფიჭვია გაბატონებული. ოდესურ დროს
მცენარეულობის საერთო ხასიათი შენარჩუნდა, თუმცა ფლორის შემადგენლობა
გაღარიბდა.
რაც შეეხება მიკროფაუნას, მეოტურ აუზში დომინანტური მდგომარეობა
ეკავათ ფორამინიფერებს, ხოლო ოსტრაკოდები წარმოდგენილია მხოლოდ რამ-
დენიმე სახეობით. სურათი იცვლება ევპატორიულში, სადაც ოსტრაკოდების სის-
ტემატიკური შემადგენლობა და საერთო რაოდენობა იზრდება, ხოლო ფორამი-

72
ნიფერები წარმოდგენილია მცირე რაოდენობის ევრიჰალური სახეობებით, რომ-
ლებიც მორფოლოგიურად განსხვავდებიან ზედამეოტური ტაქსონებისგან. არსე-
ბული მონაცემების საფუძველზე შეიძლება ითქვას, რომ ფორამინიფერების გან-
ვითარების გვიანკაინაზოური ისტორია ევპატორიულში მთავრდება.
ამრიგად, შავი ზღვის რეგიონში ევპატორული (ქვედა პონტური) ჰორიზო-
ნტი შეიძლება ჩაითვალოს იმ საზღვრად, რომლის შემდგომაც იწყება ტიპური
მიოცენური ზღვიური და ხმელეთის ბიოცენოზების ჩანაცვლება პლიოცენურით.

73
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Contents
Preface ………………………………………………………………………… 3
1. Introduction …………………………………
.
………………………........ 6
2. A micropaleontological review ..………….…………………………........ 9
3. A review of the main components of the Sarmatian to Pontian floras of
Georgia ………………………………………………………………….. 14
4. The Sarmatian to Pontian deposits of Georgia and their palynological
characteristics …………………………………………………………… 27
4.1. Eastern Georgia ...…………………………………………………… 27
4.2. Western Georgia……………………………………………………... 37
5. Main stages of vegetation development in Georgia during the Sarmatian to
Pontian ………………………………………………………………….. 43
5.1. Eastern Georgia ……………………………………………………... 43
5.2. Western Georgia …………………………………………………….. 48
6. Comparison of the environmental history of Eastern and Western Georgia
…………………………………………………………………………… 52
7. Comparison of floristic stages with phases of foraminiferal development on
territory of Georgia during the Sarmatian to Pontian …………………... 61
8.
Conclusion ……………………………………………………………… 66
რეზიუმე ………………….…………………………………………………. 69
References ………….………………………………………………………… 73