Christ's thorn formation (Paliureta spina-christi) in the Tbilisi environs (East Georgia, South Caucasus)

Christ's thorn formation (Paliureta spina-christi) in the Tbilisi environs (East Georgia, South Caucasus)

Annals of Agrarian Science 16 (2018) 189–195 Contents lists available at ScienceDirect Annals of Agrarian Science journal homepage: www.elsevier.com...

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Annals of Agrarian Science 16 (2018) 189–195

Contents lists available at ScienceDirect

Annals of Agrarian Science journal homepage: www.elsevier.com/locate/aasci

Christ's thorn formation (Paliureta spina-christi) in the Tbilisi environs (East Georgia, South Caucasus)

T

N.J. Lachashvilia,∗, M.N. Khachidzeb, N.V. Eradzec, L.D. Khetsurianib a

Ilia State University, Institute of Botany, Georgian National Botanical Garden, 1, Botanikuri Str., Tbilisi, 0114, Georgia Ilia State University, Institute of Botany, 1, Botanikuri Str., Tbilisi, 0114, Georgia c Georgian National Botanical Garden, 1, Botanikuri Str., Tbilisi, 0114, Georgia b

A R T I C LE I N FO

A B S T R A C T

Keywords: Tbilisi environs Paliureta spina-Christi Typological composition Plant community Distribution area Structural characteristics Floristic composition

Christ's thorn formation (Paliureta spina-christi) of Tbilisi environs is studied. This formation is one of the typical representatives of hemixerophilous shrubberies of shibliak type in the Caucasus region and one of the characteristic for vegetation cover of Tbilisi surroundings. Plant communities of Christ's thorn formation with different plots area are fragmentary spread almost all over the territory of Tbilisi environs from 400 to 800 (900) m above s.l.. Plant communities are developed on slopes and plane place with various exposure and inclination, mainly on the grey-cinnamonic and cinnamonic soils. The most xerophilous variants are developed on the erosive bare mother rocks as well. In Tbilisi environs the Christ-thorn's plant communities are either primary or secondary origin. Formation is characterized by rich typological and floristic composition. We identified 6 plant communities: (1) Paliureto-mixtofruticetum gramino-mixtoherbosum, (2) Paliureto-Rhmanetum gramino-mixtoherbosum, (3) Paliureto gramino-mixtoherbosum, (4) Paliureto bothriochlooso gramino-mixtoherbosum, (5) Paliuretum festuceto-bothriochloosum, (6) Paliureto-Astragaleto-Rhamnetum. For each separated plant communities the basic structural characteristics (general projective coverage, projective coverage, distribution and height of layers, sodding degree, dominant-edificator plants, characteristic species, number of species, moss cover, litter, species richness, spectrum of life forms), distribution area in the Tbilisi environs and main physicalgeographical conditions (topography, altitude, exposure, inclination, soil type) are given. 190 species of vascular plants, which belong to 40 families and 132 genera, were recorded. In the floristic spectrum leading families are: 1. Poaceae – 27 species (14,2%), 2. Asteraceae – 25 species (13,2%), 3. Fabaceae – 17 species (8,9%), 4-5. Lamiaceae and Rosaceae – 15-15 species (7,9-7,9%), 6-8. Apiaceae, Caryophyllaceae and Rubiaceae – 8-8 species (4,2-4,2%), 9. Brassicaceae – 7 species (3,7%), 10-11. Caprifoliaceae and Rhamnaceae – 4-4 species (2,1-2,1%). The life form spectrum is as follows: hemicryptophytes (including biennials) – 91 (47,9%), therophytes – 58 species (30,5%), phanerophytes – 21 (11,1%), chamaephytes – 7 (3,7%), geophytes – 13 (6,8%). The article is attached full list of recorded plants.

Introduction The part of basin of the Mtkvari River, which is spread from the village Dzegvi to the section between Ponichala and Rustavi, is considered in the environs of Tbilisi. Endings of the thick geographic units of various origins and geographic age are gathered in the vicinities of Tbilisi that make its topography considerably complicated. Privately, they are involved in the environs of Tbilisi: (1) extreme southern branching of Caucasian range – Saguramo-Ialno ridge. (2) Eastern part of Kvernaki low range (Skhaltba low range), (3) Eastern endings of Trialeti ridge (the ranges of Satskepela and Armazi, Mskhaldidi and

Lisi, Mtatsminda, Narikhala, Tabori, Telet-sakharaulo), (4) Western and extreme north-western part of Iori upland (Samgori, Vaziani, Tbilisi Sea and its adjacent territory, etc.), (5) Extreme north-western ending of Mtkvari-Arax lowland (plains of Ponichala and Kumisi-Tsalaskuri). Hypsometrical amplitude of the Tbilisi vicinity is from 350 m to 1875 m above s.l.. Types of low and middle-height mountain-gorge and stepped plain relief are highlighted in the topography [1 – 5]. In the environs of Tbilisi, two types of climate with the appropriate two zones of the climate are apportioned [2,3]: (1) The climate with insufficient humidity, dry and hot summer and mild but well expressed winter, (2) moderately humid climate with moderately warm summer

Peer review under responsibility of Journal Annals of Agrarian Science. ∗ Corresponding author. E-mail address: [email protected] (N.J. Lachashvili). https://doi.org/10.1016/j.aasci.2018.03.008 Received 9 January 2018; Accepted 23 March 2018 Available online 21 April 2018 1512-1887/ © 2018 Agricultural University of Georgia. Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/BY-NC-ND/4.0/).

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and moderately cold snowy winter. The first zone of the climate contains the whole left side of the river Mtkvari, plains of Ponichala and Kumisi-Tsalaskuri and the river floodplain as well. The secondary climate zone contains the mountainous part of the right side and the main part of Saguramo-Ialno range. Mainly grey-cinnamonic soil is developed on the western ending of the Iori upland and on the slopes of middle-height low range, which is skeletal in most cases. There are the same types, but slightly salinity soils are on the plains of Ponichala and Kumisi-Tsalaskuri, where the solonets and solonchak are speckled. Mainly different modifications of cinnamonic soil (cinnamonic leached, cinnamonic calcareous, cinnamonic light) are spread on the slopes of eastern endings of Trialeti range, on the slopes of southern exposure of Saguramo-Ialno ridge and partly on the Skhaltba low range. And, brown forest soil of various modifications is met in the upper parts of these ridges. Alluvial soils remained on the terraces of the rivers. Thin primitive skeletal soil and scree-stony are widely spread in the grey cinnamonic soil areal. Rocky bareness of groundless soil covering is also met. The section of clay and clay-sand badlands is met on the southern slope of Skhaltba low range [2,3,5 – 8]. Tbilisi environs are characterized by large biodiversity. Both ecosystem and floristic biodiversity are expressed [9 – 15]. From the point of landscapes creation and taken space, shrubberies of shibliak type are one of the important and are represented by diverse modifications. From them Christ's thorn formation (Paliureta spina-christi) is one of the characteristic for vegetation cover of Tbilisi surroundings [9]. Also, this formation is one of the typical representatives of hemixerophilous shrubberies of shibliak type on the foothills and in the lower mountain belt of the Caucasus, especially in the semiarid regions of South Caucasus. Though, the literature data about this formation are scanty In addition, in this scientific works is not presented typological composition and structural characteristics of plant communities of Christ's thorn formation [16 – 18].

fragmentary spread almost all over the territory of Tbilisi environs, on the both sides of the river Mtkvari. Their altitudinal range is from 400 to 800 (900) m above s.l.. Plant communities are developed on slopes and plane places with various exposure and inclination, mainly on the grey-cinnamonic and cinnamonic soils. Often soils are skeletal. Mainly, soils are middle or thin depth. The most xerophilous variants are developed on the erosive bare mother rocks as well. In the Tbilisi environs the Christ-thorn's plant communities are either primary or secondary origin. Nowadays, to draw the line between the primary and the secondary coenoses is impossible in most cases. Spreading of primary plant communities is linked with first climate zone (the climate with insufficient humidity, dry and hot summer and mild but well expressed winter) and various modifications of greycinnamonic soils which are developed in this zone. Secondary plant communities were formed by digressive succession processes of postforest vegetation. Their area is outside of first climate zone. They are spread on second climate type zone (the climate with insufficient humidity, dry and hot summer and mild but well expressed winter). Accordingly, secondary plant communities of Christ's thorn formation are included in the area of forests and xeromezophilous shrubberies of lower mountain belt [9,26].

Objectives and methods

Area in the Tbilisi environs: Sxaltba range (between Mtskheta and Shio-Mghvime); Altitude (m): 600-700; Topography: slope; Exposure: S-E; Inclination: from 15° to 32°; Soil: cinnamonic, middle depth; General projective coverage: 95 – 97%; I layer (shrubs)

Typological composition and geo-botanical characteristic We identified 6 Plant communities: (1) Paliureto-mixtofruticetum gramino-mixtoherbosum, (2) Paliureto-Rhmanetum gramino-mixtoherbosum, (3) Paliureto gramino-mixtoherbosum, (4) Paliureto bothriochlooso gramino-mixtoherbosum, (5) Paliuretum festuceto-bothriochloosum, (6) Paliureto-Astragaleto-Rhamnetum. Short geobotanical characteristic and area in the Tbilisi environs of identified plant communities is given bellow. (1) Paliureto-mixtofruticetum gramino-mixtoherbosum

The object of research is Christ's thorn formation (Paliureta spinachristi) of Tbilisi environs. The main aim of the research was to establish area, typological, floristic composition and spectrum of life of Christ's thorn formation in the Tbilisi environs; determination of area and main physical-geographical conditions of identified syntaxa (plant communities); study their geo-botanical structure and evaluation of ecological situation. Geo-botanical data were obtained by the route method. Geo-botanical surveys were carrying out on 25 m2 plots. During the geo-botanical descriptions, studying the structure of phytocoenoses and identification of syntaxa, we were guided the traditional geo-botanical methods [19 – 25]. On the each plots were registration general projective cover (in %) of plant communities, projective cover (in %), distribution, height and floristic composition of each layers, sodding degree (in %), general floristic composition, coenotic role of each species (projective cover in %), as well physical-geographical characteristics (topography, exposure, inclination, soil type, altitude). In the process of cameral work for each plant community were established frequency of occurrence of each species and were determined constant (typical) species, were calculated species richness on 25 m2 and spectra of life forms.

Projective coverage: 35 – 40%; Distribution: uneven or more or less evenly; Average height (cm): 200-300; II layer (herbs, semi-shrubs & dwarf semi-shrubs) Projective coverage: 75 – 80%; Distribution: more or less evenly; III layer (moss cover) Projective coverage: 30 – 40%; Distribution: more or less evenly; Sodding degree: – ; Dominant-edificator: Paliurus spina-christi (projective coverage 30 – 32%); Subdominant-edificator: – ; Characteric species:

Results and analysis Area and distribution regularities This formation is one of the typical representatives of hemixerophilous shrubberies of shibliak type in the Caucasus region and one of the characteristic for vegetation cover of Tbilisi surroundings. Plant communities of this formation with different plots area are

• Shrubs: Juniperus oxycedrus, Rhamnus pallasii, Jasminum fruticans, Lonicera iberica, Spiraea hypericifolia, Prunus incana, Ephedra procera;

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• Perennial

• •

Litter: – ; Species richness on 25 m2: 40 species; Spectrum of life forms: Phanerophytes – 9 species (11,7%), Chamaephytes – 2 (2,6,0%), Hemicryptophytes (with biannual plants) – 47 (61,0%), Geophytes – 6 (7,8%), Therophytes – 13 (16,9%).

plants: Bothriochloa ischaemum, Dactylis glomerata, Dictamnus albus, Galium verum, Elytrigia intrermedia, Phleum phleoides, Koeleria cristata, Melica transsilvanica, Poa angustifolia, Stipa capillata, Cleistogenes serotina, Psephellus carthalinicus, Potentilla recta, Stachys atherocalyx, Polygala transcaucasica, Thalictrum collinum, Turritis glabra, Asparagus verticillatus, Onobrychis radiata; Annual plants: Bromus japonicus, Trigonella spicata, Linum corymbulosum, L. nodiflorum, Aegilops tauschii, Scabiosa micrantha, Helianthemum salicifolium, Xeranthemum cylindraceum, Trifolium campestre, Melampyrum arvense, Galium tenuissimum; Semi-shrubs & dwarf semi-shrubs: Teucrium nuchense, T. polium, Thymus coriifolius.

(3) Paliureto bothriochlooso gramino-mixtoherbosum Area in the Tbilisi environs: foothills of Saguramo-Ialno ranges; Altitude (m): 580-850; Topography: Slope; Exposure: S-E, S-W, E, S; Inclination:: 5°-10°; Soil: cinnamonic, grey-cinnamonic, skeleton, middle or thin depth; General projective coverage: 80 – 95%; I layer (shrubs)

Number of species: 105; Litter: 50 – 60%; Species richness on 25 m2: 48,5; Spectrum of life forms: Phanerophytes – 9 species (15,5%), Chamaephytes – 3 (5,2%), Hemicryptophytes (with biannual plants) – 30 (51,7%), Geophytes – 3 (5,2%), Therophytes – 13 (22,4%).

Projective coverage: from 35% to 60 – 75%; Distribution: more or less evenly; Average height (cm): 300-400;

(2) Paliureto-Rhamnetum gramino-mixtoherbosum II layer (herbs, semi-shrubs & dwarf semi-shrubs) Area in the Tbilisi environs: foothills of Armazi (surroundings of Mukhadgverdi cemetery) and Saguramo ranges; Altitude (m): 600-670; Topography: plane place, slopes with small inclination; Exposure: various; Inclination: 1°-4°; Soil: cinnamonic; skeleton, middle or thin depth; General projective coverage: 92 – 95%; I layer (shrubs)

Projective coverage: from 65% to 80 – 90%; Distribution: evenly; III layer (moss cover) Projective coverage: – ; Distribution: – ; Sodding degree: 10 – 20%; Dominant-edificator: Paliurus spina-christi (projective coverage from 35% to 60 – 70%); Subdominant-edificator: Bothriochloa ischaemum (projective coverage 20 – 40%); Characteric species:

Projective coverage: 55 – 70%; Distribution: more or less evenly; Average height (cm): 250-300; II layer (herbs, semi-shrubs & dwarf semi-shrubs)

• Shrubs: – ; • Perennial plants: Dactylis glomerata, Phleum phleoides, Elymus repens,

Projective coverage: 85 – 95%; Distribution: evenly;

Origanum vulgare, Inula aspera, Eryngium campestre, Salvia nemorosa, Potentilla recta, Rumex tuberosus, Euphorbia seguieriana, Euphorbia boissieriana, Euphorbia boissieriana, Linum austriacum, Stachys atherocalyx, Securigera varia, Falcaria vulgaris, Medicago caerulea, Dictamnus albus, Thalictrum collinum, Galium verum, Fragaria vesca, Cephalaria media, Hypericum perforatum, Cynosurus echinatus, Carex humilis and Carex divulsa; 1. Annual plants: Scabiosa micrantha, Petrorhagia prolifera, Helianthemum ledifolium, Helianthemum salicifolium, Lolium rigidum, Bromus japonicus, Erodium cicutarium, Ziziphora capitata, Daucus carota, Xeranthemum squarrosum, Trifolium campestre, Althaea hirsuta, Brachypodium distachyon; Semi-shrubs & dwarf semi-shrubs: Teucrium polium; Number of species: 95;

III layer (moss cover) Projective coverage: – ; Distribution: – ; Sodding degree: 2– 3%; Dominant-edificator: Paliurus spina-christi (projective coverage 40 – 45%); Subdominant-edificator: Rhamnus pallasii (projective coverage from 10-15% to 30 – 35%); Characteric species: Shrubs: – ;

• •

• Perennial

• •

plants: Bothriochloa ischaemum, Securigera varia, Hypericum perforatum, Inula oculus-christi, I. germanica, Salvia nemorosa, Scabiosa columbaria, Koeleria cristata, Filipendula vulgaris, Thalictrum collinum, Dactylis glomerata, Elymus repens, Euphorbia boissieriana, Stachys atherocalyx, Stipa capillata, Melica transsilvanica, Potentilla recta; Annual plants: Helianthemum ledifolium, Bromus squarrosus, Xeranthemum squarrosum; Semi-shrubs & dwarf semi-shrubs: Teucrium nuchense, Teucrium polium;

Litter: – ; Species richness on 25 m2: 32,3 Spectrum of life forms: Phanerophytes – 9 species (9,5%), Chamaephytes – 1 (1,0%), Hemicryptophytes (with biannual plants) – 55 (57,9%), Geophytes – 2 (2,1%), Therophytes −28 (29,5%). (4) Paliureto gramino-mixtoherbosum Area in the Tbilisi environs: foothills of Saguramo, Lisi, Sxaltba and etc. ranges; Altitude (m): 550-850;

Number of species: 77; 191

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Projective coverage: 70 – 95% (100%); Distribution: evenly;

Topography: slope; Exposure: N, N-E, S, S-E, E; Inclination: from 3° to 25°; Soil: cinnamonic, grey-cinnamonic, skeleton, middle or thin depth; General projective coverage: I layer (shrubs)

III layer (moss cover) Projective coverage: – ; Distribution: – ;

Projective coverage: from 20-30% – 60%; Distribution: uneven or more or less evenly; Average height (cm): 200-300;

Sodding degree: 20 – 25%; Dominant-edificator: Paliurus spina-christi (projective coverage 25 – 30%); Subdominant-edificators: Festuca valesiaca (projective coverage from 5-7% to 25 – 35%), Bothriochloa ischaemum (projective coverage from 7-10% to 20 – 25%); Characteric species:

II layer (herbs, semi-shrubs & dwarf semi-shrubs) Projective coverage: from 30-40% to 80 – 100%; Distribution: evenly;

• Shrubs: Rhamnus pallasii; • Perennial plants – Eryngium campestre, Echium rubrum, Medicago

III layer (moss cover) Projective coverage: 15 – 17%; Distribution: Sodding degree: –, rare +; Dominant-edificator: Paliurus spina-christi (projective coverage from 20-30% to 40 – 45%); Subdominant-edificator: – ; Characteric species:







Shrubs: Rhamnus pallasii; 2. Perennial plants: Achillea neilreichii, Bothriochloa ischaemum, Cleistogenes serotina, Convolvulus cantabrica, Securigera varia, Dactylis glomerata, Euphorbia boissieriana, Medicago caerulea, Melica transsilvanica, Onobrychis cyri, Potentilla adenophylla, Potentilla recta, Salvia nemorosa, Stachys atherocalyx, Stipa capillata, Thalictrum collinum, Turritis glabra; Annual plants: Bromus squarrosus, Crucianella angustifolia, Helianthemum ledifolium, Helianthemum salicifolium, Petrorhagia prolifera, Linum corymbulosum, Scabiosa micrantha, Brachypodium distachyon, Trifolium angustifolium, Xeranthemum squarrosum; Semi-shrubs & dwarf semi-shrubs: Teucrium nuchense, Teucrium polium;

caerulea, Potentilla recta, Euphorbia seguieriana, Achillea neilreichii, Salvia nemorosa, Petrorhagia saxifraga, Galium verum, Falcaria vulgaris, Dactylis glomerata, Filipendula vulgaris, Verbascum formosum; Annual plants – Trifolium campestre, Medicago minima, Alyssum hirsutum, Bromus japonicus, Sideritis montana, Xeranthemum squarrosum, Scabiosa micrantha, Lolium rigidum, Echinaria capitata, Valerianella rimosa, Filago eriocephala, Filago arvensis, Helianthemum salicifolium, Helianthemum ledifolium; Semi-shrubs & dwarf semi-shrubs: Teucrium polium, Thymus coriifolius;

Number of species: 50; Litter: – ; Species richness on 25 m2: 36,5 species; Spectrum of life forms: Phanerophytes – 2 species (4,0%), Chamaephytes – 2 (4,0%), Hemicryptophytes (with biannual plants) – 23 (46,0%), Geophytes – 3 (6,0%), Therophytes – 42 (40,0%).



(6) Paliureto-Astragaleto-Rhamnetum



Area in the Tbilisi environs: Teleti range, surroundings of vil. Teleti (northern from village) Altitude (m): 850; Topography: Slope; Exposure (macro): S-W; Exposure: S; Inclination: 25°-28; Soil: bare mother rocks, screes and stones of sandstones; General projective coverage: 25 – 30%; I layer (shrubs)

Number of species: 112; Litter: Species richness on 25 m2: 35,7 species; Spectrum of life forms: Phanerophytes – 10 species (0,9%), Chamaephytes – 4 (3,6%), Hemicryptophytes (with biannual plants) – 56 (50%), Geophytes – 8 (7,1%), Therophytes – 34 (30,4%). (5) Paliuretum festuceto-bothriochloosum

Projective coverage: 20 – 25%; Distribution: more or less evenly; Average height (cm): from 80 to 100 to 150;

Area in the Tbilisi environs: eastern part of Teleti range (surroundings of vil. Teleti - northern from village); Altitude (m): 700-800; Topography: slope; Exposure (macro): S; Exposure (micro): E, S-E, E; Inclination: from 5° to 7° to 20-°-25°; Soil: dry, grey cinnamonic, skeleton, middle or thin depth; General projective coverage: 95 – 100%; I layer (shrubs)

II layer (herbs, semi-shrubs & dwarf semi-shrubs) Projective coverage: 5 – 10%; Distribution: uneven; III layer (moss cover) Projective coverage: – ; Distribution: – ;

Projective coverage: 25 – 30%; Distribution: uneven, rare, more or less evenly; Average height (cm): (150) 200-300;

Sodding degree: Dominant-edificator: Paliurus spina-christi (projective coverage 15 –

II layer (herbs, semi-shrubs & dwarf semi-shrubs) 192

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Seseli grandivittatum Schischk.

18%); Subdominant-edificator: Astracantha microcephala, Rhamnus pallasii (projective coverage 5% for each); Characteric species:

Asteraceae Achillea biebersteinii Afan. Achillea neilreichii A.Kern. (Achillea nobilis subsp. neilreichii (A.Kern.) Velen) Artemisia incana (L.) Druce Carduus hamulosus Ehrh. Carduus seminudus M.Bieb. ex M.Bieb. Carthamus lanatus L. Centaurea ovina Pall. ex Willd. Centaurea solstitialis L. Crepis foetida subsp. rhoeadifolia (M.Bieb.) Čelak. Crepis sancta (L.) Babc. Crupina vulgaris Pres. ex Carss. Filago arvensis L. Filago eriocephala Guss. Filago pyramidata L. Inula aspera Poir. Inula oculus-christi L. Inula germanica L. Galatella villosa (L.) Rchb.f. [Linosyris villosa (L.) DC.] Psephellus carthalinicus Sosn. Pyrethrum corymbosum (L.) Scop. Scorzonera biebersteinii Lipsch. Tragopogon graminifolium DC. Tragopogon tuberosus K.Koch Xeranthemum cylindraceum Sm. Xeranthemum squarrosum Boiss.

• Shrubs: Spiraea hypericifolia; • Perennial plants: Agropyron • • • •

cristatum subsp. pectinatum, Alyssum tortuosum, Dactylis glomerata, Dianthus crinitus, Euphorbia boissieriana, Poterium polygamum, Stachys atherocalyx, Stipa arabica; Biannual plants: Campanula sibirica subsp. hohenackeri, Carduus hamulosus, Lappula barbata; Annual plants: Asperula arvensis, Bromus squarrosus, Callipeltis cucullaris, Erodium cicutarium, Gypsophila elegans, Papaver arenarium; Semi-shrubs & dwarf semi-shrubs: Artemisia incana, Fumana procumbens, Teucrium nuchense, Teucrium. polium, Thymus coriifolius; Number of species: 26;

Litter: – ; Species richness on 25 m2: 17 species; Spectrum of life forms: Phanerophytes – 4 species (15,4%), Chamaephytes – 5 (19,2%), Hemicryptophytes (with biannual plants) – 11 (42,3%), Therophytes – 6 (23,1%). Floristic composition 190 species of vascular plants, which belong to 40 families and 132 genera, were recorded. In the floristic spectrum leading families are: 1. Poaceae – 27 species (14,2%), 2. Asteraceae – 25 species (13,2%), 3. Fabaceae – 17 species (8,9%), 4-5. Lamiaceae and Rosaceae – 1515 species (7,9-7,9%), 6-8. Apiaceae, Caryophyllaceae and Rubiaceae – 8-8 species (4,2-4,2%), 9. Brassicaceae – 7 species (3,7%), 10-11. Caprifoliaceae and Rhamnaceae – 4-4 species (2,12,1%). By number of species leading genera are Trifolium (5 species), Alyssum and Galium (4-4). 12 genera (Carex, Cotoneaster, Euphorbia, Filago, Inula, Linum, Potentilla, Rhamnus, Salvia, Stipa, Teucrium, Verbascum) are represented by 3 species each. The life form spectrum is as follows: hemicryptophytes (including biennials) – 91 (47,9%), therophytes – 58 species (30,5%), phanerophytes – 21 (11,1%), chamaephytes – 7 (3,7%), geophytes – 13 (6,8%).

Boraginaceae Aegonychon purpurea-coeruleum Holub. Echium rubrum Forssk. Lappula barbata (M.Bieb.) Gürke Brassicaceae (Cruciferae) Alyssum alyssoides (L.) L. Alyssum hirsutum M.Bieb. Alyssum murale Waldst. & Kit. Alyssum tortuosum Willd. Rapistrum rugosum (L.) All. Thlaspi orbiculatum steven Turritis glabra L. Campanulaceae Campanula sibirica subsp. hohenackeri (Fisch. & C.A.Mey.) Damboldt

Full list of recorded plants is given bellow.

Caprifoliacea (Dipsacaceae) Lonicera iberica M.Bieb. Cephalaria media Litv. Scabiosa columbaria L. Scabiosa micrantha Desf.

GYMNOSPERMAE Cupressaceae Juniperus foetidissima Willd. Juniperus oxycedrus L. (Juniperus rufescens Link) Ephedraceae Ephedra procera C.A.Mey. [Ephedra major subsp. procera (C.A.Mey.) Bornm.]

Caryophyllaceae Arenaria serpyllifolia L. Dianthus crinitus Sm. Dianthus subulosus Freyn & Conrath Gypsophila elegans M.Bieb. Petrorhagia prolifera (L.) P.W.Ball. & Heywood [Kohlrauschia prolifera (L.) Kunth] Petrorhagia saxifraga (L.) Link Silene latifoia Poiar. [Melandrium latifolium (L.) Maire; M. boissieri Schishk.] Silene italica (L.) Pers.

ANGIOSPERMAE DYCOTYLEDONEAE Anacardiaceae Rhus coriaria L. Apiaceae Bupleurum marschallianum C.A.Mey. Bupleurum rotundifolium L. Daucus carota L. Eryngium campestre L. Eryngium caeruleum M.Bieb. (E. caucasicum Trautv.) Falcaria vulgaris Bernh. Peucedanum ruthenicum M.Bieb.

Cistaceae Fumana procumbens (Dunal) Gren. & Godr. Helianthemum ledifolium (L.) Mill. [H. lasiocarpum Jacques & Herinca] 193

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Jasminum fruticans L. Ligustrum vulgare L.

Helianthemum salicifolium (L.) Mill. Convolvulaceae Convolvulus cantabrica L.

Orobanchaceae Melampyrum arvense L.

Betulaceae (Corylaceae) Carpinus orientalis Mill.

Papaveraceae Glaucium corniculatum (L.) Curtis Papaver arenarium M.Bieb.

Crassulaceae Sedum caucasicum (Gross.) Boriss.

Plantaginaceae Plantago lanceolata L. Veronica multifida L. Veronica orchidea Crantz. [Veronica spicata subsp. orchidea (Crantz.) Hayek]

Euphorbiaceae Euphorbia boissieriana (Woronow) Prokh. Euphorbia helioscopia L. Euphorbia seguieriana Neck. Fabaceae (Leguminosae) Astracantha microcephala (Willd.) Podlech (Astragalus microcephalus Willd.) Dorycnium herbaceum Vill. [D. pentaphyllum subsp. herbaceum (Vill.) Rouy] Lathyrus aphaca L. Lathyrus cicera L. Lotus corniculatus L. Medicago caerulea Less. ex Ledeb. Medicago minima (L.) L. Onobrychis cyri Grossh. Onobrychis radiata (Desf.) M.Bieb. Securigera varia (L.) Lassen (Coronilla varia L.) Trifolium angustifolium L. Trifolium arvense L. Trifolium campestre Schreb. Trifolium echinatum M.Bieb. Trifolium tumens M.Bieb. Trigonella spicata Sm. Vicia angustifolia

Polygalaceae Polygala transcaucasica Tamamsch. Polygonaceae Rumex tuberosus L. Ranunculaceae Ranunculus illyricus L. Ranunculus oxyspermus Willd. Thalictrum collinum Wallr. Rhamnaceae Paliurus spina-christi Mill. Rhamnus catarthica L. Rhamnus pallasii Fisch. & C.A. Mey. Rhamnus spathulifolia Fisch. & C.A. Mey. Rosaceae Agrimonia eupatoria L. Cotoneaster meyeri Pojark. Cotoneaster morulus Pojark. Cotonoaster saxatilis Pojark. Crataegus kyrtostyla Fingerh. Crataegus meyeri Pojark. Filipendula vulgaris Moench Fragaria vesca L. Potentilla adenophylla Boiss. Potentilla inclinata Vill. Potentilla recta L. Poterium polygamum WAldst. & Kit. Prunus incana (Pall.) Batsch [Cerasus incana (Pall.) Spach] Rosa canina L. Spyraea hypericifolia L.

Geraniaceae Erodium cicutarium (L.) L’Her Geranium dissectum L. Hypericaceae Hypericum perforatum L. Labiatae (Labiatae) Ajuga genevensis L. Clinopodium vulgare L. Origanum vulgare L. Phlomis pungens Willd. Salvia nemorosa L. Salvia verticillata L. Salvia viridis L. Scutellaria orientalis L. Sideritis montana L. Stachys atherocalyx K.Koch Teucrium nuchense K.Koch [Teucrium chamaedris subsp. nuchense (K.koch) Rech.f.] Teucrium orientale L. Teucrium polium L. Thymus coriifolius Ronniger Ziziphora capitata L.

Rubiaceae Asperula arvensis L. Callipeltis cucullaris (L.) DC. Crucianella angustifolia L. Galium humifusum M.Bieb. [Asperula humifusa (M.Bieb.) Besser] Galium rivale (Sibth. & Sm.) Griseb. (Asperula rivalis Sibth. & Sm.) Galium tenuissimum M.Bieb Galium verum L. Rubia tinctorum L. Rutaceae Dictamnus albus L. [Dictamnus caucasicus (Fisch. & C.A.Mey.) Grossh.]

Linaceae Linum austriacum L. Linum corymbulosum Rchb. Linum nodiflorum L.

Scrophulariaceae Verbascum orientale (L.) All. (Celsia orientalis L.) Verbascum phoeniceum L. Verbascum formosum Fisch. ex Schrank

Malvaceae Althaea hirsuta L.

Valerianaceae Valerianella rimosa Bastard

Oleaceae 194

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Violaceae Viola alba Besser

gramino-mixtoherbosum, (2) Paliureto-Rhmanetum gramino-mixtoherbosum, (3) Paliureto gramino-mixtoherbosum, (4) Paliureto bothriochlooso gramino-mixtoherbosum, (5) Paliuretum festuceto-bothriochloosum, (6) Paliureto-Astragaleto-Rhamnetum. 189 species of vascular plants, which belong to 40 families and 132 genera, were recorded. In the floristic spectrum leading families are: 1. Poaceae – 27 species (14,2%), 2. Asteraceae – 25 species (13,2%), 3. Fabaceae – 17 species (8,9%), 4-5. Lamiaceae and Rosaceae – 15-15 species (7,9-7,9%), 6-8. Apiaceae, Caryophyllaceae and Rubiaceae – 8-8 species (4,2-4,2%), 9. Brassicaceae – 7 species (3,7%), 10-11. Caprifoliaceae and Rhamnaceae – 4-4 species (2,1-2,1%). The life form spectrum is as follows: hemicryptophytes (including biennials) – 91 (47,9%), therophytes – 58 species (30,5%), phanerophytes – 21 (11,1%), chamaephytes – 7 (3,7%), geophytes – 13 (6,8%).

MONOCOTYLEDONEAE Amaryllidaceae (Alliaceae) Allium pseudoflavum Vved. Allium rotundum L. Asparagaceae (Hyacinthaceae) Asparagus officinalis L. Asparagus verticillatus L. Ornithogalum pyrenaicum L. Cyperaceae Carex divulsa Stokes Carex humilis Leyss. Carex liparocarpos subsp. bordzilowskii (V.I.Krecz.) T.V.Egorova

References

Orchidaceae Anacamptis pyramidalis (L.) Rich.

[1] M. Maruashvili, Physical Geography of Georgia, Tsodna, Tbilisi, 1964 (in Georgian). [2] K.V. Kavrishvili, Physical-geographical Characteristics of Tbilisi Surroundings, Metsniereba, Tbilisi, 1964 (in Russian). [3] D. Ukleba, Physical-geographical Zoning of East Georgia, Metsniereba, Tbilisi, 1968 (in Georgian). [4] D. Ukleba, Landscapes and Physical-geographical Districts of Mountain Regions of East Georgia, Metsniereba, Tbilisi, 1974 (in Georgian). [5] Z. Tatashidze, Tbilisi, in: Z. Tatashidze (Ed.), Geography of Georgia, Part I, Metsniereba, Tbilisi, 2000, pp. 200–207 (in Georgian). [6] M. Sabashvili, Soils of Georgian SSR, Metsniereba, Tbilisi, 1965 (in Georgian). [7] T. Urushadze (Ed.), Soil Map of Georgia, Cartography, Tbilisi, 1999. [8] T.F. Urushadze, V. Blum, Soils of Georgia, Mtsignobari, Tbilisi, 2014 (in Russian). [9] N.J. Lachashvili, M.N. Khachidze, L.D. Khetsuriani, N.V. Eradze, Typology and distribution regularities of the vegetation of Tbilisi environs (East Georgia, South Caucasus), J. Ann. Agrar. Sci. 13 (3) (2013) 55–62. [10] N.J. Lachashvili, M.N. Khachidze, L.D. Khetsuriani, N.V. Eradze, Commuties of juniper forest (Junipereta; J. foetidissima, J. polycarpos) in Shida Kartli (East Georgia, South Caucasus), J. Ann. Agrar. Sci. 12 (2) (2014) 49–65. [11] N.J. Lachashvili, M.N. Khachidze, N.V. Eradze, L.D. Khetsuriani, Steppe of Tbilisi environs (East Georgia, South Caucasus), J. Ann. Agrar. Sci. 15 (3) (2017) 118–129. [12] N.J. Lachashvili, N.V. Eradze, L.D. Khetsuriani, Conspectus of trees and shrubs of Tbilisi environs (East Georgia, South Caucasus), J. Ann. Agrar. Sci. 15 (1) (2017) 118–129. [13] N.J. Lachashvili, N.V. Eradze, Trees and Shrubs of Tbilisis Environs (East Georgia, South Caucasus), Universal, Tbilisi, 2017 (in Georgia). [14] A. Makashvili, Flora of Tbilisi Environs, vol. I, Publishing House of the Stalin Tbilisi State Uni versity, Tbilisi, 1952 (in Georgian). [15] A. Makashvili, Flora of Tbilisi Environs, vol. II, Publishing House of the Stalin Tbilisi State Uni versity, Tbilisi, 1953 (in Georgian). [16] N. Ketskhoveli, Vegetation Cover of Georgia, Publishing House of the Academy of Sciences of the Georgian SSR, Tbilisi, 1960 (in Georgian). [17] L.I. Prilipko, Vegetation Cover of Azerbaijan, ELM, Baku, 1970 (in Russian). [18] N. Lachashvili, M. Khachidze, Typology and distribution pattern of vegetation of Kiziki (East Georgia, South Caucasus), Cauc. Geogr. Rev. 10 (2009) 90–96 (in Georgian). [19] A.A. Korchagin, Species (floristic) composition of plant communities and the methods of its investigation, in: E.M. Lavrenko, A.A. Korchagin (Eds.), Field Geobotany, III, Nauka, Moscow-Leningrad, 1964, pp. 39–62 (in Russian). [20] V.M. Ponyatovskaya, Estimation of abundance and distribution of species in natural plant communities, in: E.M. Lavrenko, A.A. Korchagin (Eds.), Field Geobotany, III, Nauka, Moscow-Leningrad, 1964, pp. 209–299 (in Russian). [21] A.A. Yunatov, The types and the scope of geobotanical investigations, the selection of sample areas and the construction of ecological profiles, in: E.M. Lavrenko, A.A. Korchagin (Eds.), Field Geobotany, III, Nauka, Moscow-Leningrad, 1964, pp. 9–36 (in Russian). [22] A.P. Shennikov, Introduction to Geobotany, Publishing House of the Leningrad University, Leningrad, 1964 (in Russian). [23] Braun-Blanquet J. Pflanzensoziologie, Grundzüge der Vegetationskunde. 3 Aufl, Springer, Wien, New-York, 1964. [24] T.A. Rabotnov, Phytocenology, Publishing House of the Moscow University, Moscow, 1983 (in Russian). [25] V.Ch Vasilevich, Concerning methods of vegetation classification, Botulinum J. 70 (12) (1985) 1596–1604 (in Russian). [26] N. Lachashvili, M. Khachidze, L. Khetsuriani, Successions of Tbilisi Environs PostForest Vegetation, Univrersal, Tbilisi, 2015 (in Georgian).

Poaceae (Gramineae) Aegilops ovata L. (Ae. neglecta Req. ex Bertol.) Aegilops tauschii Coss. Agropyron cristatum subsp. pectinatum (M.Bieb.) Tzvelev Avena sterilis subsp. ludoviciana (Durieu) Gillet & Magne Bothriochloa ischaemum (L.) Keng Brachypodium distachyon (L.) P.Beauv. [Trachynia distachya (L.) Link.] Brachypodium sylvaticum (Huds.) P.Beauv. Bromus japonicus Thunb. Bromus squarrosus L. Cleistogenes serotina (L.) Keng Cynosurus echinatus L. Dactylis glomerata L. Echinaria capitata (L.) Desf. Elymus repens (L.) Gould Elytrigia intermedia (Host.) Nevski [Elymus hispidus (Opiz) Melderis] Festuca valesiaca Schleich. ex Gaudin Hordeum bulbosum L. Koeleria cristata (L.) Pers. Lolium rigidum Gaudin Melica transsilvanica Schur Phleum paniculatum Huds. Phleum phleoides (L.) H.Karst. Poa angustifolia L. Poa bulbosa subsp. vivipara (Koeler) Arcang. Stipa arabica Trin. & Rupr. Stipa capillata L. Stipa pennata L. Conclusion Area of Christ's thorn formation (Paliureta spina-christi) in Tbilisi environs is in foothills and lower mountain belt, approximately 400 – 800 (900) m above s.l.. Plant communities of formation are developed on slopes and plane places with various exposure and inclination, mainly on the grey-cinnamonic and cinnamonic soils. Often soils are skeletal. Mainly, soils are middle or thin depth. The most xerophilous variants are developed on the erosive bare mother rocks as well. In Tbilisi environs the Christ-thorn's plant communities are either primary or secondary origin. Typological composition of Christ's thorn formation (Paliureta spina-christi) of Tbilisi environs is such: (1) Paliureto-mixtofruticetum

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