The annotated checklist of plant species that occur in the wetland habitats of Georgia (the Caucasus)
expand article infoGiorgi Tedoradze, Ilia Akobia, Zurab Janiashvili§, Tamar Jugeli, Nikoloz Lachashvili|, Nutsa Megvinetukhutsesi, Vasil Metreveli, Giorgi Mikeladze, Zaal Kikvidze
‡ Ilia State University, Tbilisi, Georgia
§ The Ministry of Environmental Protection and Agriculture, Tbilisi, Georgia
| National Botanical Garden of Georgia, Tbilisi, Georgia
Open Access


The checklist includes 270 species that belong to 80 families and 183 genera. Each species has been annotated with the following information: life form, wetland indicator status, and location. In this checklist, Angiosperms are represented by 252 species (93.3%), Bryophytes – 10 species (3.7%), Pteridophytes – 8 (3%), Gymnosperms – 1 (0.4%). The largest families by the number of species are Cyperaceae – 39 (14.4%), Poaceae – 29 (10.7%), Rosaceae – 19 (7.1%), Asteraceae – 17 (6.3%), Fabaceae – 11 (4.1%) and Juncaceae – 11 (4.1%). The checklist is dominated by 55 Palaearctic species (20.4%), followed by 46 Holarctic (17.1%), 31 Euro-Mediterranean (11.5%), 31 Cosmopolitan (11.5%), and 27 Euro-Siberian (10.03%) species. The endemism rate is 4.8%, and the proportion of invasive and naturalized plants is 8.5%. Obligate wetland plants, mainly belonging to the families Cyperaceae and Juncaceae, make up 34.2% of the floristic composition. This is the first comprehensive published checklist of the flora of Georgian wetlands, annotated with wetland indicator values.

Key words

Chorotypes, Floristic composition, Life Forms, Wetlands, Wetland Indicator Status


Wetland ecosystems are considered a natural resource of global importance. They provide many valuable ecosystem services, among them flood protection, water quality improvement, and carbon dioxide sequestration. Wetlands are home to thousands of plant and animal species (Verhoeven et al. 2006; Ramsar Convention Secretariat 2013), including endemic, relict, and rare species (Suppl. material 1, figs 1–3). Furthermore, the aesthetic and recreational benefits of wetlands are also important (Suppl. material 2, figs 1–3). Hence, the preservation of this ecosystem is of the utmost importance (Hazeu et al. 2010; Tedoradze et al. 2020). Many wetlands are threatened by eutrophication (Suppl. material 2: fig. 4) driven by both global and local factors, as well as habitat fragmentation and degradation (Verhoeven et al. 2006). Wetlands harbor many characteristic species, such as hydrophytes (Suppl. material 3, figs 1–2), which are often used as specific indicators for assessing the current condition of wetlands (Reed and Peters 1988; Lyon and Lyon 2011; Eggers and Reed 2015). Therefore, having a comprehensive checklist of wetland flora is the first necessary step for planning and executing the monitoring and conservation of wetlands in Georgia.

Georgia is a mountainous country adjacent to the sea, and such settings create steep climatic gradients, in which geographical location plays an important role (Kikvidze 2020). Therefore, we also performed a gradient analysis of the species composition of wetland communities across Georgia using geographical coordinates and altitude as environmental variables.

Here we present the recorded and assessed 270 species as a list of plants found in different wetlands in most regions of Georgia. It should be noted that number of studies have been published on wetlands in Georgia, relating to different periods (Kimeridze 1966, 1985; Melia 1985; Kikvidze and Oshawa 2001; Denk et al. 2001; BSSGW 2008; Matchutadze 2008; Krebs et al. 2009; Matchutadze et al. 2015). However, there are practically no studies covering all the major wetlands, nor do we have any studies on the indicator status of typical wetland species recorded. Therefore, our study addresses these two needs.

Materials and methods

Georgia is a climatically diverse country owing to the adjacent Black Sea and high mountain chains of various orientations. Through our sampling regions, the mean annual temperature varies from 14 °C in Colchis to 3 °C in Kazbegi; annual precipitation varies roughly between 600 and 900mm and its distribution changes from almost ideally uniform in Imereti (760mm ± 15SD) to strongly seasonal in Sioni (812mm ± 45SD). Therefore, the study regions represent quite a diverse climatic character (Fick and Hijmans 2017).

Study regions are located at different hypsometric heights and in different climatic zones. In total, we sampled 218 plots in seven regions across Georgia (Fig. 1): Kolkheti Lowland (121 plots), Javakheti Plateau (53 plots), mountainous Adjara – the highlands of Adjara (8 plots), Kazbegi region (7 plots), Iori River Gorge – the northwestern part of Kakheti province (9 plots), Upper Imereti (6 plots), Shida Kartli Lowland (6 plots), and Alazani Valley (8 plots).

Figure 1. 

Map showing study sites/plots in Georgia.

Fieldwork campaigns were conducted during the vegetation seasons of 2015-2017. The sampling locations were selected prior to our fieldwork using topographic maps of the Soviet period (1950-60s), which depicted mires, swamps, and other wetlands in a rather detailed manner. Before sampling, we determined the location and morphology of wetland bodies with 5m accuracy using RapidEye satellite multispectral imagery.

Sampling points on the pre-selected sites were located using navigation instruments − GPS and satellite images. First, we described the hydrological conditions of a given wetland by assessing its water surface: water not visible (0), well visible (1), and partly visible (2); we also assessed observable human impacts through three categories (low, intermediate, and high impacts).

Vegetation was sampled as follows: once a wetland was recognized and inspected, the team chose visually the most representative area(s) for sampling. A central point of circular plots was placed at random in a selected area, and radii for plots were delineated: a 9 meter-radius circular plot around a sampling point was used to sample trees; a 6 meter-radius plot was used for sampling shrubs, and a 3 meter-radius circle was used for sampling herbaceous plants (USACE 1987). Within these plots, we identified and recorded all detectable species and assessed their abundance by the modified scale of Braun-Blanquet (r, +, 1, 2m, 2a, 2b, 3, 4, 5) (Braun-Blanquet 1964; van der Maarel 2007). Plant species were identified using the Key to Plants of Georgia (Ketskhoveli et al. 1964, 1969) and the Excursion Flora of Germany (Exkursionsflora von Deutschland) (Jäger et al. 2013). The plant species that were not easily identifiable in the field were collected and later accurately identified by comparing them to the specimens in the National Herbarium (TBI) of the Institute of Botany, Ilia State University, Georgia.

In all, we collected 216 herbarium specimens of 132 plant species (herbarium specimens will be available at the TBI). Latin names of taxa (families, genera, and species) are based on nomenclatural checklists of ‘Flora of Georgia’ (Gagnidze 2005; Davlianidze et al. 2018) and are harmonized with international plant databases (Euro+Med 2006; GBIF 2022; IPNI 2022; POWO 2022; WFO 2022). Species that are alien to Georgia are listed according to Kikodze et al. (2010).

Each species was ranked according to its wetland indicator status on the following scale (USACE 1987; Reed and Peters 1988; Lyon and Lyon 2011; Lichvar et al. 2013; Eggers and Reed 2015):

  • Obligate wetland plants (OBL) – almost always occur in wetlands (estimated probability > 99%) under natural conditions.
  • Facultative wetland plants (FACW) – usually occur in wetlands (estimated probability 67% – 99%), but is occasionally found in non-wetlands.
  • Facultative plants (FAC) – equally likely to occur in wetlands (estimated probability 34% – 66%) or non-wetlands.
  • Facultative upland plants (FACU) – usually occur in non-wetlands (estimated probability 67% – 99%), but are occasionally found in wetlands (estimated probability 1% – 33%).
  • Obligate upland plants (UPL) – almost always occur (estimated probability > 99%) in non-wetlands under natural conditions.

Each species was assessed for its phytogeographical distribution (Komarov 1934-1960; Takhtajan 1954-2011, 2003-2012; Tutin et al. 1964-1993; Davis 1965-2001; Ketskhoveli et al. 1971-2011) and online international plant databases (GBIF 2022; Euro+Med 2006; POWO 2022). In addition, we used different regional works in this field to determine the chorotypes of the species (Gagnidze and Kemularia-Nathadze 1985; Portenier 1993, 2000; Gagnidze 2004; Shetekauri and Chelidze 2016; Lachashvili et al. 2020; Lachashvili et al. 2021; Tedoradze 2022). Plant life forms were determined using the LEDA Traitbase (Klotz et al. 2002). Endemics of the Caucasus are given after Solomon et al. (2013).

To analyze the variation of species composition on geographical gradients, we performed Canonical Correspondence Analysis (CCA). Statistical significance was computed with 999 permutation tests. We analyzed species with a frequency of occurrence in the community matrix equal to or greater than 5. For this purpose, we used PAST (v. 4.12) software (Hammer et al. 2001).

Results and discussion

We sampled wetlands across the diverse landscapes of Georgia: the Black Sea coastal zone with a maritime climate, the inland lowlands with a dry and continental climate, high dry plateaus, and subalpine environments. Our results permitted us to construct an annotated checklist of the wetland flora of Georgia, which includes 270 species sampled from 218 pre-selected sites. Each species on the resulting checklist (Table 1) is annotated with the following data: life form, wetland indicator status, chorotype, and location (study regions). The coordinates of each plot can be found in the supplementary information, where the species presence-absence matrix is also included as Suppl. material 4.

Table 1.

Annotated checklist of plant species that occur in the wetland habitats of Georgia. LF (Life Forms): F – Ferns; H – Herbaceous; H(A) – Herbaceous (Aquatic); M – Moss; W – Woody; W(L) – Woody (Liana). WIS (Wetland Indicator Status): FAC – Facultative; FACU – Facultative upland; FACW – Facultative Wetland; OBL – Obligate. Chorotypes: Aus. – Native Australia; CA – Caucaso-Anatolian; Cau. – Caucasian; Cau.(EC) – Caucasian (Endemic of Caucasia); Cau.(EG) – Caucasian (Endemic of Georgia); CE – Caucaso – Euxinian; Cosm. – Cosmopolitan; Cosm.(ExPR) – Cosmopolitan (except polar regions.); Cosm.(MO) – Cosmopolitan/Mediterranean origin; Cosm.(SAmI) – Cosmopolitan/South America origin/invasive; Cosm.(TAf) – Cosmopolitan/Probably native to tropical Africa; CSWA – Caucaso-SW Asian; E – European; EA – East Asian; EA(I) – East Asian/invasive; EA(N) – East Asian/naturalized; EMed. – Euro-Mediterranean; EMed.(Cond.) – Euro-Mediterranean conditionally; ES – Euro-Siberian; ESWA – Euro-SW Asian; Eux. – Euxinian; HE – Hyrcano-Euxinian; Hol. – Holarctic; IT – Irano-Turanian; Med. – Mediterranean; Med. (Cond.) – Mediterranean conditionally; MSWA – Mediterranean-SW Asian; MSWAES – Med.-S.W. As.-Euras. Steppe; NA – North American; NA(Adv) – North American/adventive; NA(I) – North American/invasive; NA(N) – North American/naturalized; Pal. – Palearctic; Pal.(W) – West Palearctic; Pal.(WW) – Palearctic/worldwide introduced; SA(N) – S American/naturalized; SEA – SE Asian; Study Regions in Georgia: 1 – Kolkheti lowland; 2 – Javakheti; 3 – Mtiani Adjara; 4 – Mtiuleti; 5 – Iori River gorge; 6 – Upper Imereti; 7 – Shida Kartli lowland and Alazani Valley.

Family Species LF* WIS** Chorotypes Study Regions
in Georgia
Brachytheciaceae Brachythecium mildeanum W. P. Schimper M OBL Pal. 1
Climaciaceae Climacium dendroides Weber & D. Mohr M OBL Hol. 3
Mniaceae Plagiomnium ellipticum T.Koponen M OBL Hol. 3
Polytrichaceae Polytrichastrum longisetum G.L. Smith M OBL Hol. 3
Polytrichum strictum Menzies ex Bridel M OBL Hol. 3
Sphagnaceae Sphagnum capillifolium Hedwig M OBL Cosm. 1,3
Sphagnum imbricatum Hornschuch ex Russow M OBL Pal. 1,3
Sphagnum palustre L. M OBL Cosm. 1,3
Sphagnum papillosum Lindberg M OBL Hol. 1
Sphagnum platyphyllum Warnstorf M OBL Hol. 1
Aspleniaceae Asplenium trichomanes L. F FACU Cosm. 1
Athyriaceae Athyrium filix -femina (L.) Roth F FAC Hol. 1
Dennstaedtiaceae Pteridium aquilinum (L.) Kuhn (P. tauricum V. I. Krecz.) F FAC Pal. 1
Dryopteridaceae Dryopteris filix -mas (L.) Schott F FAC Hol. 7
Equisetaceae Equisetum palustre L. H FACW Hol. 1,2,3,4,5,6,7
Lycopodiaceae Lycopodiella inundata (L.) Holub (Lycopodium inundatum L.) F OBL Hol. 4
Osmundaceae Osmunda regalis L. F OBL EMed. 1
Salviniaceae Salvinia natans (L.) All. F OBL Pal.(WW) 1,7
Pinaceae Pinus pinaster Aiton (P. maritima Du Roi) W FAC Med. 1
Apiaceae Anthriscus sylvestris (L.) Hoffm. H FACU Pal. 4
Carum carvi L. H FACU Pal. 3
Daucus carota L. H FACU Pal.(W) 5
Eryngium caeruleum M.Bieb. H FACU IT 5
Eryngium campestre L. H FACU EMed. 1
Apocynaceae Periploca graeca L. W(L) FACW EMed. 1,7
Vinca major subsp. hirsuta (Boiss.) Stearn (V. pubescens D’Urv.) H FAC CA 1
Aquifoliaceae Ilex colchica Pojark. W(L) FAC Eux. 1
Araliaceae Hydrocotyle ramiflora Maxim. H OBL EA 1,3
Hydrocotyle vulgaris L. H OBL EMed. 1
Hedera colchica (K.Koch) K.Koch W(L) FACW CA 1
Hedera helix L. W(L) FAC ES 1,7
Asteraceae Achillea millefolium L. H FACU Hol. 2
Ambrosia artemisiifolia L. H FACU NA(I) 1
Arctium lappa L. H FAC Pal. 2,5
Artemisia absinthium L. H FAC Pal. 2
Bidens tripartita L. H OBL Hol. 1,3,5
Centaurea oxylepis (Wimm. & Grab.) Hayek H FACW EMed. 1
Cichorium intybus L. H FACU Cosm. 2,6
Cirsium arvense (L.) Scop. H FACU Cosm. 5
Cirsium obvallatum (M.Bieb.) M.Bieb. (C. obvallatum (M.Bieb.) Fisch.) H FACU Cau. 4
Cirsium vulgare (Savi) Ten. H FACU Cosm. 2
Eupatorium cannabinum L. H FACW EMed. 1,5
Inula helenium L. H FAC Pal.(W) 5
Pentanema britannicum (L.) D.Gut.Larr., Santos-Vicente, Anderb., E.Rico & M.M.Mart.Ort. (Inula britannica L.) H FACW ES 2,5
Petasites albus (L.) Gaertn. H FACW E 2
Solidago canadensis L. H FACU NA(N) 1,7
Taraxacum campylodes G.E.Haglund (Leontodon taraxacum L.; Taraxacum officinale Weber ex Wiggins) H FACU Pal. 2
Xanthium strumarium L. H FAC Cosm. 1
Betulaceae Alnus glutinosa subsp. barbata (C.A.Mey.) Yalt. (A. barbata C.A.Mey.) W FACW Cau.(EC) 1,5,7
Carpinus betulus L. (C. caucasica Grossh.) H FACU EMed. 1
Corylus avellana L. H FACU EMed. 1,7
Brassicaceae Capsella bursa -pastoris (L.) Medik. H FACU Cosm. 1,2
Nasturtium officinale R.Br. H OBL Cosm. 4
Caprifoliaceae Cephalaria gigantea (Ledeb.) Bobrov H FACU Cau. 2
Dipsacus laciniatus L. H FACU ESWA 5
Valeriana alliariifolia var. tiliifolia (Troitsky) V.E.Avet. (V. tiliifolia Troitsky) H FAC Cau.(EG) 5
Celastraceae Euonymus europaeus L. H FAC EMed. 1,2,5
Ceratophyllaceae Ceratophyllum demersum L. H(A) OBL Cosm. 2
Convolvulaceae Convolvulus arvensis L. H FAC Pal. 1,5
Cornaceae Cornus sanguinea subsp. australis (C.A.Mey.) Jáv. (Swida australis (C. A. Mey.) Pojark. ex Grossh.) W FAC Eux. 1,5,7
Droseraceae Drosera intermedia Hayne H OBL Hol. 3
Drosera rotundifolia L. H OBL Hol. 1
Ebenaceae Diospyros lotus L. W FAC SEA 1
Elaeagnaceae Elaeagnus angustifolia L. W FAC Pal. 7
Hippophae rhamnoides L. W FAC Pal. 4
Ericaceae Rhododendron luteum Sweet W FAC Eux. 1
Rhododendron ponticum L. W FAC Eux. 1
Vaccinium arctostaphylos L. W FACW CA 1
Fabaceae Amorpha fruticosa L. W FACW NA(I) 1,5
Colutea orientalis Mill. W FACU CSWA 5
Gleditsia triacanthos L. W FAC NA(N) 1
Glycyrrhiza glabra L. H FACU Pal.(W) 5,7
Ononis arvensis L. W FAC Pal.(W) 5
Robinia pseudoacacia L. W FACU NA(I) 1
Trifolium canescens Willd. H FACU CSWA 2
Trifolium pratense L. H FAC Pal. 2,4
Trifolium repens L. H FAC Pal. 1
Trifolium spadiceum L. H FACW ES 4
Vicia tenuifolia subsp. variabilis (Freyn & Sint.) Dinsm. (V. variabilis Freyn & Sint.) H FACU CSWA 1,5
Fagaceae Quercus hartwissiana Steven W FAC Eux. 1
Quercus petraea subsp. iberica (Steven ex M. Bieb.) Krassiln. (Q. iberica Steven) W FACU CSWA 5
Gentianaceae Gentiana septemfida Pall. H FAC CA 4
Swertia iberica Fisch. ex C.A. Mey. H OBL Cau.(EG) 4
Hypericaceae Hypericum mutilum L. H FACW NA(N) 1,2
Hypericum perforatum L. H FACU Pal. 1
Juglandaceae Pterocarya fraxinifolia (Poir.) Spach (Juglans pterocarpa Michx.) W FACW HE 1,7
Lamiaceae Lycopus europaeus L. H OBL ES 1,2
Mentha aquatica L. H OBL Cosm. 1,7
Prunella vulgaris L. H FACU Hol. 1
Stachys palustris L. H OBL ES 1,6
Thymus collinus M.Bieb. H FACU Cau.(EC) 2,5
Lentibulariaceae Utricularia minor L. H OBL Hol. 1
Utricularia vulgaris L. H OBL Pal. 2
Lythraceae Lythrum hyssopifolia L. H OBL Cosm. 5
Lythrum salicaria L. H OBL Pal. 1,2,4,5,6,7
Trapa natans L. (T. colchica Albov) H(A) OBL Pal. 1
Malvaceae Hibiscus coccineus (Medik.) Walter H FAC NA(I) 1
Hibiscus ponticus Rupr. H FACW Cau.(EG) 1
Kosteletzkya pentacarpos (L.) Ledeb. H FACW EMed. (Cond.) 1
Menyanthaceae Menyanthes trifoliata L. H OBL Hol. 1,2,3
Moraceae Ficus carica L. W FACU IT 1
Morus alba L. W FAC EA 7
Morus nigra L. W FAC ESWA 1,7
Myrtaceae Eucalyptus cinerea F.Muell. ex Benth. W FACW Aus. 1
Nymphaeaceae Nuphar lutea (L.) Sibth. & Sm. H(A) OBL ES 1
Nymphaea alba L. H(A) OBL EMed. 1,2
Oleaceae Fraxinus excelsior L. W FACU ES 1,7
Onagraceae Epilobium montanum L. H FAC ES 4
Epilobium palustre L. H OBL Hol. 2,4
Orobanchaceae Euphrasia caucasica Juz. H FACU Cau.(EC) 4
Rhamphicarpa medwedewii Albov H FACW Cau.(EC) 1
Rhinanthus minor L. H FACU ES 4
Parnassiaceae Parnassia palustris L. H OBL Hol. 4
Phytolaccaceae Phytolacca americana L. H FACU NA 1,7
Plantaginaceae Plantago major L. H FAC Hol. 7
Plantago media L. H FAC Pal. 2
Polygonaceae Persicaria hydropiper (L.) Delarbre (Polygonum hydropiper L.) H OBL Cosm. 3
Persicaria maculosa Gray (Polygonum persicaria L.) H FACW Pal. 1,2,6
Persicaria perfoliata (L.) H.Gross (Polygonum perfoliatum L.) H FAC EA(I) 1
Persicaria thunbergii (Siebold & Zucc.) H.Gross (Polygonum thunbergii Siebold & Zucc.) H FACW EA(I) 1,7
Polygonum carneum K.Koch H FAC CA 2
Primulaceae Lysimachia vulgaris L. H FACW Pal. 1,2,5
Primula auriculata Lam. H FACW CSWA 3
Ranunculaceae Clematis vitalba L. H FAC EMed. 1
Ranunculus caucasicus M.Bieb. H FACU Cau.(EC) 2
Ranunculus ophioglossifolius Vill. H OBL EMed. 2,5
Ranunculus trichophyllus Chaix (Batrachium trichophyllum (Chaix) Bosch) H OBL Hol. 2
Rhamnaceae Frangula alnus Mill. H FAC ES 1,7
Paliurus spina -christi Mill. H FACU MSWA 1
Rosaceae Agrimonia eupatoria L. H FACU Pal.(W) 1
Alchemilla sericata Rchb. H FACU Cau.(EC) 4
Comarum palustre L. H OBL Hol. 1,2,3
Crataegus microphylla K.Koch H FAC ESWA 1
Crataegus monogyna Jacq. (C. kyrtostyla Fingerh. ex Schltdl.) H FACU ES 1
Filipendula vulgaris Moench H FACU Pal. 2
Fragaria vesca L. H FACU Hol. 7
Geum urbanum L. H FACU Pal.(W) 1,7
Malus sylvestris subsp. orientalis (Uglitzk.) Browicz (M. orientalis Uglitzk.) W FAC CSWA 1
Mespilus germanica L. W FAC EMed. 1
Potentilla erecta (L.) Raeusch. H FACU ES 1,3
Pyrus communis subsp. caucasica (Fed.) Browicz (P. caucasica Fed.) W FACU Cau. 1
Rosa canina L. W FACU Pal.(W) 1
Rubus caesius L. W FAC Pal.(W) 1
Rubus caucasicus Focke W FACU Cau.(EC) 1
Rubus caucasigenus (Sudre) Juz. W FAC Cau.(EC) 1
Rubus hirtus Waldst. & Kit. W FACU EMed. 1
Rubus silesiacus Weihe (R. candicans Weihe) W FAC E 1
Spiraea japonica L.f. H FACU EA 1
Rubiaceae Galium verum L. H FACU Pal. 2
Rutaceae Citrus trifoliata L. H FAC EA 1
Salicaceae Populus × canescens (Aiton) Sm. W FACW ES 7
Salix alba L. W FACW ES 5,7
Salix caprea L. W FAC ES 5
Sapindaceae Acer campestre L. W FACU EMed. 1,4,7
Acer negundo L. W FACW NA(Ad) 7
Simaroubaceae Ailanthus altissima (Mill.) Swingle W FACU EA(I) 5
Solanaceae Solanum dulcamara L. (S. persicum Willd.) H FACW Pal. 1
Ulmaceae Ulmus minor Mill. H FAC EMed. 1,7
Urticaceae Urtica dioica L. H FAC Pal. 5,7
Viburnaceae Sambucus ebulus L. W FAC EMed. 7
Sambucus nigra L. W FAC EMed. 1
Viburnum lantana L. W FACU EMed. 1
Viburnum opulus L. W FACW ES 1,7
Acoraceae Acorus calamus L. H OBL Hol. 1
Alismataceae Alisma plantago -aquatica L. H OBL Pal. 1,2,3,4,5,6
Sagittaria sagittifolia L. H OBL ES 1,2
Amaryllidaceae Allium sp. H FAC - 5
Pancratium maritimum L. H FACU Med. 1
Araceae Lemna minor L. H(A) OBL Cosm. 2,3,6
Lemna trisulca L. H(A) OBL Cosm. 1,2
Asparagaceae Asparagus officinalis L. H FACU ES 1
Ruscus aculeatus L. W FAC EMed. 1
Ruscus colchicus Yeo W FAC Cau.(EC) 1
Butomaceae Butomus umbellatus L. H OBL Pal. 2
Commelinaceae Commelina communis L. H FAC EA(N) 1
Cyperaceae Blysmus compressus (L.) Panz. ex Link H OBL Pal.(W) 2,3,4
Bolboschoenus maritimus (L.) Palla H OBL Cosm.(ExPR) 1,5,7
Carex acuta L. H OBL ES 2
Carex aterrima subsp. medwedewii (Leskov) T.V.Egorova (C. medwedewii Leskov) H FAC CSWA 3
Carex canescens L. H OBL Hol. 1,2
Carex cespitosa L. H OBL ES 2
Carex diluta M.Bieb. H FACW IT 4
Carex disticha Huds. H OBL EMed. 2,3
Carex divulsa Stokes H FAC EMed. 17
Carex echinata Murray H OBL EMed. 2,3
Carex elata All. H OBL EMed. 2,4
Carex hirta L. H OBL EMed. 2,4,5
Carex lachenalii Schkuhr H OBL Hol. 2
Carex lasiocarpa Ehrh. H OBL Hol. 1,2
Carex leporina L. H OBL ES 4
Carex limosa L. H OBL Hol. 2
Carex muricata L. H FAC ES 1
Carex nigra subsp. juncea (Fr.) Soó (C. juncella (Fries) Th. Fries) H OBL ES 3
Carex nigra subsp. transcaucasica (T.V.Egorova) Jim.Mejías, G.E.Rodr.-Pal., Amini Rad & Martín-Bravo (C. transcaucasica T.V.Egorova) H OBL CSWA 2
Carex pallescens L. H FACW Hol. 2
Carex panicea L. H OBL ES 1
Carex pendula Huds. H FACW EMed. 2
Carex pseudocyperus L. H OBL Hol. 2
Carex riparia Curtis H OBL EMed. 1,2,4
Carex rostrata Stokes H OBL Hol. 1,2,3
Carex tristis M.Bieb. H FAC CA 4
Carex vesicaria L. H OBL Hol. 1,2,3,4,7
Cladium mariscus (L.) Pohl H OBL Cosm. 1,2,7
Cyperus flavescens L. (Pycreus flavescens (L.) P.Beauv. ex Rchb.) H FACW Cosm. 1
Cyperus longus subsp. badius (Desf.) Bonnier & Layens (C. badius Desf.) H FACW Med. (Cond.) 3
Cyperus pannonicus Jacq. (Juncellus pannonicus (Jacq.) C.B. Clarke) H FACW Pal. 1
Cyperus serotinus Rottb. H FACW Pal. 1
Eleocharis palustris (L.) Roem. & Schult. H OBL Hol. 2,3
Eleocharis uniglumis (Link) Schult. H OBL Hol. 2
Rhynchospora alba (L.) Vahl H OBL Hol. 1
Rhynchospora caucasica Palla H OBL Cau.(EG) 1
Schoenoplectiella mucronata (L.) J.Jung & H.K.Choi (Scirpus mucronatus L.) H FACW Cosm. 1,2
Schoenoplectus tabernaemontani (C.C.Gmel.) Palla (Scirpus tabernaemontani C. C. Gmel.) H OBL Cosm. 2
Scirpus sylvaticus L. H FACW Pal.(W) 1,3,5
Hydrocharitaceae Elodea densa (Planch.) Casp. H(A) OBL SA(N) 1
Hydrocharis morsus -ranae L. H(A) OBL ES 1,7
Iridaceae Iris pseudacorus L. H OBL Pal.(W) 1,6,7
Juncaceae Juncus acutus L. H FACW EMed. 1
Juncus articulatus L. H OBL Hol. 1,2,4
Juncus atratus Krock. H OBL ES 2
Juncus bufonius L. H FACW Cosm. 1,3
Juncus conglomeratus L. H OBL Pal.(W) 2,3
Juncus effusus L. H OBL Cosm. 1,2,3,6,7
Juncus filiformis L. H FACW Hol. 2
Juncus inflexus L. H FACW Pal.(W) 4
Juncus maritimus Lam. H FACW EMed. 1
Luzula multiflora (Ehrh.) Lej. H FACU Hol. 2
Luzula stenophylla Steud. (L. pseudosudetica (V.I.Krecz.) V.I.Krecz.) H FAC CSWA 2
Juncaginaceae Triglochin palustris L. H OBL Hol. 4
Orchidaceae Spiranthes sinensis (Pers.) Ames (S. amoena (M. Bieb.) Spreng.) H OBL SEA 1
Poaceae Agrostis capillaris L. (A. tenuis Sibth.) H FAC Pal. 2
Agrostis vinealis Schreb. H FAC CA 2,5
Alopecurus aequalis Sobol. H OBL Hol. 2
Alopecurus arundinaceus Poir. H FACW Pal. 2
Anthoxanthum odoratum L. H FAC EMed. 2
Beckmannia eruciformis (L.) Host. H FACW Pal. 2
Bromus variegatus M.Bieb. (Bromopsis variegata (M.Bieb.) Holub) H FACU CSWA 4
Calamagrostis arundinacea (L.) Roth H FAC Pal. 2,5,7
Calamagrostis epigejos (L.) Roth H FAC ES 1,2,5
Calamagrostis pseudophragmites (Haller f.) Koeler H FAC Pal. 2
Cynodon dactylon (L.) Pers. H FAC Cosm.(TAf) 1,6,7
Dactylis glomerata L. H FAC Pal. 2
Echinochloa crus-galli (L.) P. Beauv. H FACW Cosm. 1
Festuca drymeja Mert. & W.D.J. Koch H FAC MSWA 7
Koeleria luerssenii (Domin) Domin H FACU Cau. 4
Lolium rigidum Gaudin H FACU MSWA 2
Molinia caerulea (L.) Moench H FACU Pal.(W) 1,2,6,7
Nardus stricta L. H FACU Hol. 2,3,4
Oplismenus undulatifolius (Ard.) P.Beauv. H FACW Cosm. 1,7
Paspalum dilatatum Poir. H FACW Cosm.(SAmI) 1,6,7
Paspalum distichum L. H OBL SA(N) 1
Phleum phleoides (L.) H. Karst. H FACU Pal. 2
Phleum pratense L. H FACU Pal. 3,4
Phragmites australis (Cav.) Trin. ex Steud. H FACW Cosm. 1,2,4,5,6,7
Poa annua L. H FAC Pal.(WW) 4
Scolochloa festucacea (Wild.) Link H OBL Hol. 2
Sorghum halepense (L.) Pers. H FAC Cosm.(MO) 1
Sporobolus fertilis (Steud.) Clayton H FACW EA(I) 1
Stipa pulcherrima K.Koch H FACU MSWAES 2
Potamogetonaceae Potamogeton crispus L. H(A) OBL Cosm. 1,2
Potamogeton natans L. H(A) OBL Hol. 1,2
Zannichellia palustris L. H OBL Cosm. 2
Smilacaceae Smilax excelsa L. W(L) FACW CE 1,7
Typhaceae Sparganium emersum Rehmann H OBL Hol. 1,2,3,6,7
Sparganium erectum subsp. neglectum (Beeby) K.Richt. (S. neglectum Beeby) H OBL Pal.(W) 1
Sparganium erectum L. H OBL Pal.(W) 5
Sparganium natans L. (S. minimum Wallr.) H OBL Hol. 2
Typha angustifolia L. H OBL Hol. 1,2,4,5,6
Typha latifolia L. H OBL Cosm. 1,2,6,7
Typha laxmannii Lepech. H OBL Pal. 1

The checklist includes 270 species belonging to 80 families and 183 genera. Angiosperms are the core of the floristic composition – 251 species (93%). Bryophytes are represented by 10 species (3.7%), Pteridophytes – 8 (3%), Gymnosperms – 1 (0.4%). Leading families by the number of species are Cyperaceae – 39 (14.4%), Poaceae – 29 (10.7%), Rosaceae – 19 (7.1%), Asteraceae – 17 (6.3%), Fabaceae – 11 (4.1%) and Juncaceae – 11 (4.1%). The floristic composition of Georgian wetlands is dominated by Palaearctic species 55 (20.4%), followed by Holarctic 46 (17.1%), Euro-Mediterranean 31 (11.5%), Cosmopolitan 31 (11.5%) and Euro-Siberian 27 (10.03%) species.

Out of 270 species, half are represented by only one or two sites. The most frequent species are Alnus glutinosa subsp. barbata – (69 sites), Molinia caerulea – (56 sites), Juncus effusus – (42 sites), Carex vesicaria – (40 sites), Phragmites australis – (37 sites), Persicaria thunbergii – (36 sites), Sphagnum palustre – (35 sites), Lythrum salicaria – (32 sites), Cladium mariscus – (29 sites), Frangula alnus – (29 sites). Among the species listed above, five are OBL in wetlands, three – FACW, and of the remaining two, one – FACU and the other – FAC. The presence of the latter, among the ten most frequent species, indicates the current state of the wetland, which is often expressed by fragmentation and the spread of species uncharacteristic for wetlands.

The percent share of endemic species in the checklist is 4.8 %; the rate of endemism is low compared to the flora of Georgia, where the endemism rate is approximately 31% (Solomon et al. 2013). There are 13 endemic species, four of which are endemic to Georgia and nine to the Caucasus. At the same time, we recorded 23 non-native species, which gives a rate (8.5%) higher than that of endemics. The low endemism rate in wetland vegetation is not surprising since the typical wetland species usually have a very wide distribution; wetland vegetation is often considered to be azonal (sensu Sieben 2019), with very little variable composition across large gradients within subcontinents such as Mexico (Ramos-Tapia et al. 2022) or South Africa (Mucina et al. 2006) or even between so distant places such as the western and eastern extremities of Eurasia (Kikvidze and Ohsawa 2001; see also Denk et al. 2001).

In the checklist, species that almost always occur in wetlands (OBL) and species that are commonly (FACW) found in wetlands together made up 52.1% of the total number of species (OBL – 92 and FACW – 48 species). Species that are equally likely to occur in wetlands or non-wetlands (FAC) made up 68 species (25.3%). The plants that usually occur in non-wetlands but are occasionally found in wetlands (FACU) consisted of 62 species (23.1%).

Out of the 218 plots, 65 were taken in seasonally dry areas (water not visible), 37 were taken in fully water-covered habitats (water well visible), 113 – partially water-covered habitats (water hardly visible) and for three plots no data. It should be noted that we canceled a certain number of plots since we found different types of agricultural fields in places where, according to the old Soviet maps, there should have been wetlands.

According to the degree of human impact on the habitat, 46 plots were assessed as low (insignificant or not observed at all), 82 plots were assessed as intermediate (low impact rate), 89 plots were assessed as high (high anthropogenic impact), and for one plot there was no data.

During the fieldwork, we recorded Drosera rotundifolia for the first time in the Samegrelo region, where, together with the Rhynchospora alba, they make a unique plant community in the middle of the urbanized area (village Namikolaevo). This location is not mentioned for this species in the ‘Flora of Georgia’ (Ketskhoveli et al. 1971-2011), nor in the other reviewed literature. Regretfully, this unique habitat is in a very poor state owing to grazing by domestic animals and fragmentation, and it is possible that without proper protective measures it will disappear soon (see Suppl. material 2, fig. 5).

The presence of FAC and FACU species among the ten most frequently occurring species, the relatively low level of endemism against a high percentage of non-native species, and the assessment of 89 out of 218 sites as being under heavy human influence are indicative of the current challenging status of wetlands. The remaining undamaged wetlands with their highly diverse vegetation, distinctive relics, unique flora, and picturesque landscapes are impressive and can attract many visitors, provided such recreational and touristic activities are duly organized.

CCA axis 1 explained 63% of the variation in species distribution along geographical gradients, this axis was statistically highly significant (p = 0.005). The second axis explained 37% of this variation, yet it was insignificant (p = 0.34). Axis 3 could be dismissed as its explanatory power was very low (less than 1%, p = 0.53). The triplot (Fig. 2) shows that axis 1 represents variation in altitude and longitude. Variation in latitude apparently does not play an important role as it is associated with the insignificant axis 2. Altitude in Georgia is tightly correlated with the mean annual temperature and can serve as its precise proxy, whils longitude can be a proxy of precipitation (Kikvidze 2020). Therefore, perhaps the main climatic driver of species composition in the wetlands of Georgia is temperature, while precipitation might play a secondary role since the main feature of the wetlands is waterlogging, which can only weakly depend on precipitation.

Figure 2. 

Wetland vegetation species composition along geographical gradients as analyzed with the CCA ordination; all data are well within the 95% confidence interval (encircled). “Eastern lowlands” include the wetlands in Shida Kartli Lowland and Alazani Valley; Iori Valley stands for the entire Iori River gorge.


Our study presents the most comprehensive checklist of Georgia's wetland flora compared to any previously published work. Importantly, we added an assessment of an indicator status, life forms, and chorotypes to each species, and the coordinates to each site in supplementary data in the form of a presence-absence matrix. The present list of plant species is based only on the results of field studies. The checklist includes 270 species belonging to 80 families and 183 genera. Among them are some non-native species that are not yet included in the ‘Flora of Georgia’ (e.g., Eucalyptus cinerea, Pinus pinaster, and Hibiscus coccineus). Angiosperms are the core of the floristic composition – 251 species (93%). Bryophytes are represented by 10 species (3.7%), Pteridophytes – 8 (3%), Gymnosperms – 1 (0.4%). Leading families by the number of species are Cyperaceae – 39 (14.4%), Poaceae – 29 (10.7%), Rosaceae – 19 (7.1%), Asteraceae – 17 (6.3%), Fabaceae – 11 (4.1%), and Juncaceae – 11 (4.1%). The floristic composition of Georgian wetlands is dominated by Palaearctic species 55 (20.4%), followed by Holarctic 46 (17.1%), Euro-Mediterranean 31 (11.5%), Cosmopolitan 31 (11.5%) and Euro-Siberian 27 (10.03%) species. Endemism is 4.8% and the proportion of invasive and naturalized plants is 8.5%. Obligate wetland plants, mainly belonging to the families Cyperaceae and Juncaceae, make up 34.2% of the floristic composition. Out of 270 species, half are represented by only one or two sites. The five most frequent species are Alnus glutinosa subsp. barbata, Molinia caerulea, Juncus effusus, Carex vesicaria and Phragmites australis. 52.1% of species belong to the OBL and FACW categories. Therefore, the main climatic driver of species composition in the wetlands of Georgia seems to be temperature, while precipitation plays a secondary role since the main feature of the wetlands is waterlogging, which can only weakly depend on precipitation.

Overall, our results indicate that the wetlands are under strong anthropogenic influence. We hope that this checklist will facilitate further work on completing the Georgian wetlands plant inventory.


ჩვენი კვლევა წარმოადგენს საქართველოს ჭარბტენიან ჰაბიტატებში გავრცელებული სახეობების ყველაზე სრულ ნუსხას ადრე გამოქვეყნებულ შრომებთან შედარებით. რაც მთავარია, ჩვენ თითოეულ სახეობას მივანიჭეთ ჭარბტენიანი ტერიტორიების ინდიკაციის სტატუსი, სასიცოცხლო ფორმები და ქოროტიპები. ასევე მოცემულია თითოეული სახეობისთვის გავრცელების კოორდინატები ყოფნა-არყოფნის მატრიქსის სახით. ამასთან, მცენარეთა სახეობების არსებული სია ეყრდნობა მხოლოდ ველზე ჩატარებული აღწერების შედეგებს. ნუსხა მოიცავს 270 სახეობას, რომელიც გაერთიანებულია 80 ოჯახსა და 183 გვარში. ენდემიზმის პროცენტული მაჩვენებელი შეადგენს 4.8 %-ს, ხოლო ინვაზიური და ნატურალიზებული სახეობები შეადგენს 8.5 %-ს. ჭარბწყლიანი გარემოსთვის დამახასიათებელი სახეობები შეადგენს საერთო რაოდენობის 34.2%-ს და ძირითადად მიეკუთვნება Cyperaceae და Juncaceae-ს ოჯახებს.


The authors thank Deutsche Gesellschaft für Internationale Zusammenarbeit (GIZ) and its regional program – Integrated Biodiversity Management, South Caucasus (IBiS) for their financial support. We wish to thank Prof. Alexander Gavashelishvili, Prof. David Tarkhnisvili and Prof. Zurab Javakhishvili of the Institute of Ecology of Ilia State University for their assistance in conducting this study. In addition, thanks to the staff of the National Herbarium of Georgia (TBI) and Department of Cryptogams and Fungi, among them, a special thanks to Dr. Ketevan Tigishvili, for her particular contribution to the identification of mosses. We would like to thank the Director of the Institute of Botany, Mr. Shalva Sikharulidze and Mr. Piran Chkheidze, without whose assistance it would have been very difficult to successfully accomplish the South Georgian expedition. We also thank to Mr. David Kikodze and stress the importance of the "Dzelkva" Report of the Botanical Survey of South Georgian Wetlands (2008), which provided a useful guideline for our project and contributed the data of six mires to our database.

Author contributions

GM planned the research; GT, GM, IA, NM, VM and ZJ conducted the field sampling; GT and TJ harmonized the nomenclature of species with international databases; GT and NL defined chorotypes; ZK performed the statistical analyses and led the writing; all authors critically revised the manuscript.


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Supplementary materials

Supplementary material 1 

Images of some rare wetland species in Georgia

Tedoradze G 2023

Data type: image

Explanation note: Images of some rare wetland species in Georgia are given, including Drosera rotundifolia, Rhynchospora alba, R. caucasica, Salvinia natans, and Trapa natans.

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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Supplementary material 2 

Some wetland panoramas from different regions in Georgia

Tedoradze G 2023

Data type: image

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (332.73 kb)
Supplementary material 3 

Some characteristic wetland species in Georgia

Tedoradze G 2023

Data type: image

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (122.07 kb)
Supplementary material 4 

Coordinates of sampling sites, wetland characteristics

Tedoradze G et al. 2023

Data type: species data

Explanation note: Coordinates of sampling sites, wetland characteristics (such as water cover and anthropogenic impact), and presence-absence data for all plant species given in the check-list are provided.

This dataset is made available under the Open Database License ( The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (187.12 kb)
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