1999 Courtesy of Editors Courtesy of Editors Courtesy of Editors Courtesy of Editors Courtesy of ...

80 Pages·2012·49.74 MB·English

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issn 0393 5434
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15
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y THE VEGETATION OF GEORGIA
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o Giorgi Nakhutsrishvili
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rt CAMERINO
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o 1999
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rt PUBLICATION DU DEPARTEMENT DE BOTANIQUE ET ECOLOGIE DE L'UNIVERSITE DE CAMERINO ET DE LA
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o STATION INTERNATIONALE DE PHYTOSOCIOLOGIE DE BAILLEUL SOUS L'EGIDE DE L'ASSOCIATION AMICALE
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DE PHYTOSOCIOLOGIE ET DE L'ASSOCIATION INTERNATIONALE POUR L'ETUDE DE LA VEGETATION

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dit EDITEURS:
E BRAUN-BLANQUETIA

of Jean-Marie Gehu
y Universite R. Descartes Paris et Un heritage est enrichissant et ouvre de nouvelles possibilites creatrices.
s
e Station Internationale de Phytosocio Mais il en decoule en contre partie I'obligation de ne pas gaspiller le
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ur logie, Haendries patrimoine reyu. Ceux qui, aujourd'hui etudient la vegetation grace a la
o phytosociologie peuvent utiliser des methodologies bien au point et
C F -59270 Bailleul
tirer profit d'un ensemble coherent de connaisances.

s
r C'est le resultat du travail methodique de nombreux chercheurs de
o Franco Pedrotti
t qualite pendant plusieurs decennies. Aujourd'hui, nous nous trouvons
di Dipartimento di Botanica ed Ecologia
E face a des problemes qui ne sont sans doute pas tout a fait nouveaux
dell'Universita, Via Pontoni, 5
f mais qui paraissent infiniment plus graves que dans le passe: primaute
o I -62032 Camerino (MC)
y de la technique, specialisation, penurie de matieres premieres. d'energie
s et d'espace, crise de I'environnement ...
e
rt Sandro Pignatti 11 se developpe ainsi des problemes specifiques divers pour lesaue s il
u Dipartimento di Biologia Vegetale
o est necessaire de trouver des reponses nouvelles. Les chercheurs sont
C Universita "La Sapienza"
places devant un veritable defi et il depend de leur savoir et de leur

rs Piazzale Aldo Moro imagination de montrer si la Science de la vegetation est capable
to I -00185 Roma d'apporter une contribution appreciable a la solution de ces problemes.
di
La tradition phytosociologique dans ce contexte constitue une base
E
f Salvador Rivas-Martinez essentielle. La conception typologique de la vegetation et la clarte du
o
Departamento de Botanica systeme qui en decoule, I'habitude des chercheurs de vivre en contact
y
s Facultad de Farmacia etroit avec la vegetation, les recherches basees sur I'observation
e
t Universidad Complutense condition antithetique de I'experimentation, sont les traits caracteristi
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o E -28040 Madrid ques de la phytosociologie.
C Les lignes directrices qui nous ont ete transmises par les maltres de la

rs Erich Hubl Science de la vegetation, Josias Braun-Elanquet et Reinhold Tuxen
o
t Botanisches Institut avant tout, constituent actuellement une part importante de notre
di patrimoine d'idees. Notre but est de valoriser cet heritage et d'honorer
E Universitat fOr Bodenkunde
f GymnasiumstraBe, 79 la memoire du premier de ces ma'i'tres et fondateur de la phytosociolo
o
gie moderne par une nouvelle serie de publications.
y A - 1190 Wien
s Pourront y trouver place des monographies etudiant concretement la
e
t vegetation selon les enseignements de J. Braun-Blanquet et R. Tuxen
r COMITE DE LECTURE:
u qui, a travers la creativite des auteurs, produiront de nouveaux fruits.
o
C Disciples nous-memes de J. Braun-Blanquet et ayant collabore a son
P.V. Arrigoni (Firenze)
s activite, nous pensons qu'a travers cette serie de publications son
r O. De Bolos (Barcelona)
o heritage restera vivant dans I'esprit originel et avec de nouvelles idees.
dit N. BoscaiL: (Cluj-Napoca)
E P. Bridgewater (Canberra)

of M. Costa (Valencia) R. Pott (Hannover) Secretariat general de la publication:

y A. Damman (Storrs, Conn.) P. Quezel (Marseille)
s
e K. Dierssen (Kiel) F. A. Roig (Mendoza) Prof. Roberto Venanzoni
t
ur N. Donita (Bucuresti) R. Schumacker (Liege) Dipartimento di Botanica ed Ecologia
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C U. Eskuche (Corrientes) MAJ. Werger (Utrecht) Via Pontoni 5, 62032 Camerino (Italia)
s J. B. Falinski (Bialowieza) R. Wittig (Frankfurt a.M.) Tel. [phone] Fax [phone]
or M. Grandtner (Quebec) V. Westhoff (Nijmegen) e-mail: [email protected]
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di S. Grigore (Timisoara) O. Wilmanns (Freiburg i.Br.)
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L. IIijanic (Zagreb)
Secretariat d'edition: Laura Carimini
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o J. Izco (Santiago)
email: [email protected]
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s F. KI6tzli (Zurich)
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t A. Lacoste (Paris-Orsay) This volume has been written, edited and composed on a desktop publishing system using
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u D. Lausi (Trieste) Apple Macintosh'· PageMaker6.0®b y Laura Carimini. Proofs and final pages were created
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C A. Miyawaki (Yokohama) on the PostScript® Apple LaserWriter®.

s J. Moravec (Pruhonice)
or A. Noirfalise (Gembloux) © 1999 Dipartimento di Botanica ed Ecologia dell'Universita -Camerino et Station de
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di E. Oberdorfer (Freiburg i. Br.) Phytosociologie - Bailleul
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T. Ohba (Yokohama)
of Printed in Italy by Centro Audiovisivi e Stampa, Camerino, 1999.
A. Pirola (Pavia)
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BRAUN-BLANQUETIA
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y RECUEIL DE TRAVAUX DE GEOBOTANIQUE
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t REVIEW OF GEOBOTANICAL MONOGRAPHS
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THE VEGETATION OF GEORGIA

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Giorgi Nakhutsrishvili
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dit Drawnform aJ. pBhRoAtoUgNra-BphL AbNy QFUraEnTJ;,o 1is9e5 M4 . Dansereau
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dit BRAUN-BLANQuETIA, vat. 15, 1999 5
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y INTRODUCTION the need for irrigation systems, having ture, vegetable growing, etc.); 2) cattle
s proved so essential in the economy of breeding; 3) domestic craft (weaving,
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rt Georgia - an ancient Christian the Ancient East (Egypt, Babylon). Be wood work, blacksmith work, textile
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o country - occupies the central and sou sides, these rivers served as trade-rou production, pottery, ceramics, etc.).
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th-western parts of Caucasia - an isth tes. Due to their rapid current, the rivers Georgians developed plough-land culti

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r mus between the Black and the Caspian were used for transportation of timber vation, various forms of irrigation (es
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t Sea. In the west, it is contiguous to the (especially in Colchis). pecially in the mountain areas) aiming
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E Black Sea, in the east to the intermoun Georgia is rich in various mineral to increase crop capacity of natural gras
f tain depression of the River Mtkvari (= resources. In the mountains of Georgia slands (2 and 3 harvests per season). Of
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Kura) in the Caspian Lowlands (the Tran copper was extracted, essential for non 16 species of wheat 11 are represented in
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s scaucasian Depression), the northern ferrous metallurgy, which by that time Georgia.
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rt border follows the high mountain tops reached a high level of development. Ancient Georgian manuscripts and
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o of the Greater Caucasus, the southern Rich iron deposits prompted the pro special medical books ("Karabadini")
C one crosses the Minor Caucasus (in the gress of ferrous metallurgy. Magnetic contain information on various herbs.

s eastern part) and separates the Minor sands along the Black Sea were of parti The fIrst serious scientific information
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t Caucasus from the adjacent mountain cular importance. Rivers, containing concerning the vegetation of Georgia
di ranges of north-eastern Asia Minor. Po gold dust, were known during the Greek was given by Vakhushti Bagrationi (l8th
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litically, Georgia borders on the Russian Epoch. century). According to several histori
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Federation (to the north), Azerbaijan (to Vakhushti Bagrationi, the promi cal sources and notes of foreign travel
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s the east), Turkey (to the south-west) and nent Georgian historian and geographer lers, the plants from different parts of
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t Armenia (to the south-east). The bor of the 18th century, distinguished two Georgia and from other countries were
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u ders have changed several times during botanical and agricultural zones within cultivated in the park of Georgian kings
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C the history of Georgia, especially in the Georgia: the mountainous and the low in Legvta Khevi (now Botanical Garden

s earlier stages of the formation of the land one. Since the Neolithic Revolu of Tbilisi). In the 18th century, foreign
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o Georgian nation. tion, these two zones represented one scientists became interested in the extre
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di Kavkasioni, i. e. the Greater Cau closely integrated system. The lowlands me diversity of Georgian vegetation and
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casus range, rises up to 4605 m (Mt. were characterized by rich harvest of flora. In 1852, on the basis of rich collec
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o Ushba, highest mountain within Geor grain crops, vineyards and orchards. In tions of Caucasian plants, the Caucasian

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s gia) and 5633 m s. m. (Mt. Elbrus, in the the mountainous areas the harvest of Museum was founded. In the beginning
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Russian Federation) i. e. over 1000 m grain crops was much poorer, vineyards of the 20th century, all the branches of
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u above the eternal snow-line. Mountain and orchards were absent. botanical science (anatomy, physiolo
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C passes at an altitude of 2200-3000 m s. Georgia covers an area of 69.500 gy, taxonomy, floristics, phytosociolo
s m. were used as trade-routes from the km2• The population is 5.5 million (2/3 gy, etc.) started to develop in Georgia.
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o earliest times. In the mountains of Ka of them being Georgians). The Geor The Institute of Botany, which belongs
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di vkasioni permanent settlements are gians call themselves "kartveli" (sing.) to the Academy of Sciences, was ope
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found up to an altitude of 2500 m s. m. and "kartvelebi" (pl.), hence the name of ned in 1933. Long-standing expeditions

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o Though the Minor (or Lesser) Caucasus the country in Georgian language is to different parts of the Caucasus and of

y mountains reach an altitude of3304 m s. "Sakartvelo". The Georgian language Georgia, as well as monographic studies
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e m., there are no glaciers and no eternal belongs to the group of Kartvelian lan of separate taxa served as a basis for the
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u snow. guages, included in the family of Ibe 8-volume "Flora of Georgia". In 1971,
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C The natural conditions of the so rian-Caucasian languages. It is sugge the fIrst volume of the totally revised
called intermontane lowlands of Geor sted that the Georgian language is rela second edition of "Flora of Georgia"
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or gia, consisting of the Colchic Lowlands ted to Anterior-Asian languages, the appeared (11 volumes have already been
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di (adjacent to the Black Sea, in West Ge Basque language (the Basque nation is published so far). Later, it was decided
E orgia) and the foothills between Kavka living in Spain, in the northern part of to publish "The Vegetation of Georgia"

of sioni (in the north) and the Minor Cauca the Iberain peninsula, close to the Pyre under the editorship of the present au

y sus (in the south-west), including the nees), connections to the Indo-Europe thor(NAKHUTsRIsHVIU, 1990, 1991). Only
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e Mtkvari Valley (= Kura valley, in East an languages, however, are uncertain. two volumes of this work have been
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ur Georgia), were favourable for settle Written Georgian dates back to the 3rd published till now (DoLUKHANov: "Fo
o ment and mixed farming. century B.C. (to the reign of king Parna rest Vegetation of Georgia", Part I, 1990,
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In the east, Colchic Lowlands are vaz). and SHATlLOVA and RANlsHvru "Mate
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r surrounded by the Imeretian Elevation Christianity, adopted as the official rials on the History of Flora and Vegeta
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dit which connects the Greater Caucasus religion in 337 A. D., promoted the tion of Georgia", 1991); several volu
E with the Minor Caucasus. The highest unification of Georgia and the develop mes are in print. Until its complete pu

of part of it, the Surami range, divides the ment of written Georgian language. blication, the author of the present book
y country into two major natural geo In ancient times, Georgian tribes decided to characterize briefly the main
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e graphical regions: Eastern and Western were known as skillful farmers, cattle types of vegetation of Georgia and to
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r Georgia, the first having been called breeders and metallurgists. Various ar publish it in English. This present publi
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o Iberia and the latter Colchis by ancient chaeological discoveries, as well as an cation bears a special purpose: During
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Greek and Roman writers. cient oriental and Georgian manuscripts the 75-year period of isolation of the

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r Diversity of climate and relief of testify to an early economical, social USSR and of Georgia in particular, it
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t Georgia has been highly important for and cultural development of the Geor was not so easy for foreign scientists to
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E its economic development since the ear gian people. According to these data, the get adequately acquainted with the inte
f ly stages of human activities. Numerous leading branches of economy of the co un resting plant life of Georgia and to com
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rivers (more than 25.000), which pro try must have been: 1) agriculture (field pare the Caucasian flora and plant com
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s moted intensive agriculture, eliminated crop cultivation, vine making, horticul- munities with those of other countries.
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o6 The vegetation of Georgia (Caucasus)
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Historico-Geographical Regions of Georgia
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Russia

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to 1. Abkhazeti 11. Tush-Pshav -Khevsureti
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Co 5. Imereti 15. Gardabani

rs 6. Guria 16. Trialeti
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of 8. Shiga Kartli 18. Javakheti

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Today, Georgian botanists are glad to be Sea) to some of the highest peaks of the 1) the soil province of Western Georgia;
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s able to collaborate with scientists from Greater Caucasus, reaching 4695 m (Mt. 2) the soil province of Eastern Georgia;
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rt many other countries. Ushba), 5068 m (in the Shkhara massif), 3) the soil province of Southern Geor
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o and 5047 m s. m. (Mt. Kazbegi, Kaz gia.
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bek). From the geological point of view, The main types of soil are allocated

rs l. ENVIRONMENT ALCONDITIONS this area consists, mainly, of Meso- to in accordance with the altitudinal zones.
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t Cenozoic deposits. AncientPrecambrian In the Western Georgia soil province
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Georgia occupies an interesting and Paleozoic formations are poorly re (between the Black Sea and Likhi Mts.)
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f geobotanical position as a part of Cauca presented and of secondary importance. it is possible to distinguish the bog and
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sia - the region which links Europe with podzolic soil zone in the lowlands, the
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s Asia. The country is characterized by 1.2 Climate krasnozem and zheltozem zone in the
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t rather contrasting natural conditions hilly piedmonts, the zone of mountain
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o which account for the extremely high The climate is one of extremes. It is forest and the mountain-meadow soils.
C degree of di vergence of plant communi possible to distinguish several climatic The soil province of Eastern Geor

s ties within this comparatively small area. zones from the humid, almost subtropi gia comprises the plains, piedmonts and
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The landscape of the country includes cal climate to the climate of permanent mountain massifs, situated eastwards
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di different types of desert and semi-desert snows and glaciers. Such a considerable from Likhi mountain range. Chestnut
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vegetation mainly in the eastern parts of range of climatic conditions is caused soils, chemozems, brown humic-sulpha
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Georgia, luxuriant Colchic forests of both by the orographic structure and the tes, saline soils of steppes and semi
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s moist, almost subtropical climate in the presence of the Black Sea and the Ca deserts, as well as intermediate forest
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t west, and high-mountain plant commu spian Sea. The Black Sea and high moun steppe and mountain-meadow soils oc
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u nities in the north and in the south. tain ranges of the Greater Caucasus are cur in this province.
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C The complex orographic structure the most important orographic factors The Southern Georgia soil provin
s of Georgia and its geographical position determining the climate of Georgia by ce includes Javakheti, Tsalka-Dmanisi
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o account for the geographical and ecolo preventing the invasion of cold air mas and Erusheti uplands, the hollow of
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di gical isolation of certain plant commu ses from the north. The climate of the Akhaltsikhe, etc. A considerable part of
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nities, which has resulted in a high ratio country has been formed by the air mas this area is covered both with the moun
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o of local endemism (particularly ende ses blowing from sea, as well as by the tain chernozems (which are formed at

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s mics of the Greater Caucasus Range and latitudinal position between southern altitudes from 1200-2200 m) and mea
e of the Colchis) and of a variety of spe Russia and the Inner-Anatolian moun dow chernozem-like soils. In highlands
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u cies of different phytochorological tains system. they are replaced by mountain-meadow
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C groups, like Iberian and Anterior-Asian The highest mean annual tempera soils. Besides, the alluvial soils, redzi
s group, etc.). The flora of the country ture amounts to 15° C (Sukhumi in nas, brown as well as the meadow-brown
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o includes about 4.200 species of wild Abkhazia, on the Black Sea coast), the soils occur here, with the predominance
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di flowering-plant species, and over 8500 lowest in the Kavkasioni Ridge (altitude of brown forest type of soil in the moun
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species of spore-bearing ones (75 pteri 5000 m). The warmest winter (5-7° C, in tain forest belt.

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o dophytes, about 800 mosses, 600 li January) is in Colchis (West Georgia, on

y chens, 5000 fungi, 2000 algae). the Black Sea coast).
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e Annual precipitations in Georgia 2. HISTORY OF FLORA AND VEGE
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ur 1.1 Orography range from over 4500 to 400 mm or less. TATION
o
Increase of altitude e.g. in Svaneti (nor
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Geologically, the territory ofGeor th-western Georgia) and Javakheti (sou The history of flora and especially
s
or gia belongs to the Alpine System of th-western Georgia) is paralleled by re the vegetation of Georgia is very com
dit Eurasia. Its geological and geomorpho duction of precipitation, while in other plicated and, despite the intensive pale
E logical structure reveals a great genetic districts (Kazbegi region, Kakheti, etc.) obotanical researches of the recent past,

of diversity, as a result of tectonic, volca the amount of rainfall increases with still needs to be studied.

y nic, petrological, gravitational, erosio elevation. The highest amounts of preci The earliest records of fossil flora
s
e nal and other processes. pitation occur at the following altitudes: stem from the Paleozoic. Species of Le
t
r Structurally, the area can be divi from 300-500 m to 3500 m s. m. in pidodendron (large trees reaching a hei
u
o ded into the following major landforms: Western Georgia, and from 1200 to 3500 ght of nearly 40 m), Sigillaria (high
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1) The range of the Greater Caucasus m in the eastern part of the country. plants with erect stems and characteri
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r (Kavkasioni); The climatic conditions of several stic rhizophore system-stigmarias), Aste
o
dit 2) the Georgian Intermontane Area districts of Georgia are demonstrated rocalamites were determined from the
E (between the Greater and the Minor diagrammatically (fig. 1). Notwith Lower and Middle Carboniferous depo
f Caucasus); standing the short distance between the sition of the Khrarni crystalline massif
o
y 3) the Mountain System of the Minor se districts (40 km between Batumi and (South Georgia). From the same period
s
e Caucasus (Meskheti-Trialetian Ran Bakhmaro, 380 km between Batumi and are also known Cordaitales - a group
rt ge), including the South Georgian Shiraki, 155 km between Shiraki and with an atomic structures (tracheids, well
u
o Volcanic Upland. Kazbegi), their climatic conditions are developed pith) very similar to that of
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Georgia comprises the southern noticeably different. conifers (SHATTLOVA & RAMISHVILI,

s
r side of the middle part of the Kavkasioni 1990).
o
t as well as the north-western parts of the 1.3 Soils In the Early Jurassic, almost the
di
E Transcaucasian Depression, the Minor entire territory of Georgia was covered
f Caucasus, i. e. the Transcaucasian Uplan. There is a great diversity of soil by sea, except for Khrami, Loki and
o
Altitudes in Georgia vary conside types in Georgia. The following soil Dzirula ancient massifs.
y
s rably from the sea level (at the Black provinces can be distinguished: The climate of this period was
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o8 The vegetation of Georgia (Caucasus)
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rt Akhalkalaki 2000m 4.9 °C 621mm Bakhmaro 18S0m 4.0°C 1869mm
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o Batumi Om 14.0°C 2718mm Kazbegi 1850m 4.7°C 806mm
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s II III IV V VI VII VIII IX X XI XII
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o Tbilisi 450m 12.7°C 559mm Shiraki SOOm 10.3 °C 544mm

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y Fig. 1 - Climadiagrams of several regions of Georgia.
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